Primate
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Primates is an order of
mammal A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...
s, which is further divided into the strepsirrhines, which include lemurs, galagos, and lorisids; and the haplorhines, which include tarsiers and
simian The simians, anthropoids, or higher primates are an infraorder (Simiiformes ) of primates containing all animals traditionally called monkeys and apes. More precisely, they consist of the parvorders New World monkey, Platyrrhini (New World mon ...
s (
monkey Monkey is a common name that may refer to most mammals of the infraorder Simiiformes, also known as simians. Traditionally, all animals in the group now known as simians are counted as monkeys except the apes. Thus monkeys, in that sense, co ...
s and apes). Primates arose 74–63 million years ago first from small terrestrial mammals, which adapted for life in
tropical forest Tropical forests are forested ecoregions with tropical climates – that is, land areas approximately bounded by the Tropic of Cancer, tropics of Cancer and Tropic of Capricorn, Capricorn, but possibly affected by other factors such as prevailing ...
s: many primate characteristics represent adaptations to the challenging environment among tree tops, including large
brain size The size of the brain is a frequent topic of study within the fields of anatomy, biological anthropology, animal science and evolution. Measuring brain size and cranial capacity is relevant both to humans and other animals, and can be done by wei ...
s,
binocular vision Binocular vision is seeing with two eyes. The Field_of_view, field of view that can be surveyed with two eyes is greater than with one eye. To the extent that the visual fields of the two eyes overlap, #Depth, binocular depth can be perceived. Th ...
, color vision, vocalizations, shoulder girdles allowing a large degree of movement in the
upper limb The upper Limb (anatomy), limbs or upper extremities are the forelimbs of an upright posture, upright-postured tetrapod vertebrate, extending from the scapulae and clavicles down to and including the digit (anatomy), digits, including all the musc ...
s, and opposable thumbs (in most but not all) that enable better
grasping A grasp is an act of taking, holding or seizing firmly with (or as if with) the hand. An example of a grasp is the handshake, wherein two people grasp one of each other's like hands. In zoology Zoology ( , ) is the scientific study of an ...
and
dexterity Fine motor skill (or dexterity) is the coordination of small muscles in movement with the eyes, hands and fingers. The complex levels of manual dexterity that humans exhibit can be related to the nervous system. Fine motor skills aid in the growt ...
. Primates range in size from Madame Berthe's mouse lemur, which weighs , to the eastern gorilla, weighing over . There are 376–524
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
of living primates, depending on which classification is used. New primate species continue to be discovered: over 25 species were described in the 2000s, 36 in the 2010s, and six in the 2020s. Primates have large
brain The brain is an organ (biology), organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It consists of nervous tissue and is typically located in the head (cephalization), usually near organs for ...
s (relative to body size) compared to other mammals, as well as an increased reliance on
visual acuity Visual acuity (VA) commonly refers to the clarity of visual perception, vision, but technically rates an animal's ability to recognize small details with precision. Visual acuity depends on optical and neural factors. Optical factors of the eye ...
at the expense of the sense of smell, which is the dominant sensory system in most mammals. These features are more developed in monkeys and apes, and noticeably less so in lorises and lemurs. Some primates, including
gorilla Gorillas are primarily herbivorous, terrestrial great apes that inhabit the tropical forests of equatorial Africa. The genus ''Gorilla'' is divided into two species: the eastern gorilla and the western gorilla, and either four or five su ...
s,
human Humans (''Homo sapiens'') or modern humans are the most common and widespread species of primate, and the last surviving species of the genus ''Homo''. They are Hominidae, great apes characterized by their Prehistory of nakedness and clothing ...
s and
baboon Baboons are primates comprising the biology, genus ''Papio'', one of the 23 genera of Old World monkeys, in the family Cercopithecidae. There are six species of baboon: the hamadryas baboon, the Guinea baboon, the olive baboon, the yellow ba ...
s, are primarily ground-dwelling rather than arboreal, but all species have adaptations for climbing trees.
Arboreal locomotion Arboreal locomotion is the animal locomotion, locomotion of animals in trees. In habitats in which trees are present, animals have evolution, evolved to move in them. Some animals may scale trees only occasionally (scansorial), but others are e ...
techniques used include leaping from tree to tree and swinging between branches of trees ( brachiation); terrestrial locomotion techniques include walking on two hindlimbs (
bipedal Bipedalism is a form of terrestrial locomotion where an animal moves by means of its two rear (or lower) limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning 'two feet' (from Latin ''bis'' ...
ism) and modified walking on four limbs ( quadrupedalism) via knuckle-walking. Primates are among the most
social Social organisms, including human(s), live collectively in interacting populations. This interaction is considered social whether they are aware of it or not, and whether the exchange is voluntary or not. Etymology The word "social" derives fro ...
of all animals, forming pairs or family groups, uni-male harems, and multi-male/multi-female groups. Non-human primates have at least four types of
social systems In sociology, a social system is the patterned network of relationships constituting a coherent whole that exist between individuals, groups, and institutions. It is the formal Social structure, structure of role and status that can form in a smal ...
, many defined by the amount of movement by adolescent females between groups. Primates have slower rates of development than other similarly sized mammals, reach maturity later, and have longer lifespans. Primates are also the most cognitively advanced animals, with humans (genus ''
Homo ''Homo'' () is a genus of great ape (family Hominidae) that emerged from the genus ''Australopithecus'' and encompasses only a single extant species, ''Homo sapiens'' (modern humans), along with a number of extinct species (collectively called ...
'') capable of creating complex
language Language is a structured system of communication that consists of grammar and vocabulary. It is the primary means by which humans convey meaning, both in spoken and signed language, signed forms, and may also be conveyed through writing syste ...
s and sophisticated
civilization A civilization (also spelled civilisation in British English) is any complex society characterized by the development of state (polity), the state, social stratification, urban area, urbanization, and symbolic systems of communication beyon ...
s, while non-human primates have been recorded using tools. They may communicate using facial and hand gestures, smells and vocalizations. Close interactions between humans and non-human primates (NHPs) can create opportunities for the transmission of zoonotic diseases, especially virus diseases including herpes,
measles Measles (probably from Middle Dutch or Middle High German ''masel(e)'', meaning "blemish, blood blister") is a highly contagious, Vaccine-preventable diseases, vaccine-preventable infectious disease caused by Measles morbillivirus, measles v ...
,
ebola Ebola, also known as Ebola virus disease (EVD) and Ebola hemorrhagic fever (EHF), is a viral hemorrhagic fever in humans and other primates, caused by ebolaviruses. Symptoms typically start anywhere between two days and three weeks after in ...
,
rabies Rabies is a viral disease that causes encephalitis in humans and other mammals. It was historically referred to as hydrophobia ("fear of water") because its victims panic when offered liquids to drink. Early symptoms can include fever and abn ...
and
hepatitis Hepatitis is inflammation of the liver parenchyma, liver tissue. Some people or animals with hepatitis have no symptoms, whereas others develop yellow discoloration of the skin and whites of the eyes (jaundice), Anorexia (symptom), poor appetite ...
. Thousands of non-human primates are used in research around the world because of their psychological and physiological similarity to humans. About 60% of primate species are threatened with extinction. Common threats include
deforestation Deforestation or forest clearance is the removal and destruction of a forest or stand of trees from land that is then converted to non-forest use. Deforestation can involve conversion of forest land to farms, ranches, or urban use. Ab ...
, forest fragmentation, monkey drives, and primate hunting for use in medicines, as pets, and for food. Large-scale tropical forest clearing for agriculture most threatens primates.


Etymology

The English name ''
primates Primates is an order of mammals, which is further divided into the strepsirrhines, which include lemurs, galagos, and lorisids; and the haplorhines, which include tarsiers and simians ( monkeys and apes). Primates arose 74–63  ...
'' is derived from
Old French Old French (, , ; ) was the language spoken in most of the northern half of France approximately between the late 8th [2-4; we might wonder whether there's a point at which it's appropriate to talk of the beginnings of French, that is, when it wa ...
or French , from a noun use of Latin , from ('prime, first rank'). The name was given by
Carl Linnaeus Carl Linnaeus (23 May 1707 – 10 January 1778), also known after ennoblement in 1761 as Carl von Linné,#Blunt, Blunt (2004), p. 171. was a Swedish biologist and physician who formalised binomial nomenclature, the modern system of naming o ...
because he thought this the "highest" order of animals. The relationships among the different groups of primates were not clearly understood until relatively recently, so the commonly used terms are somewhat confused. For example, ''ape'' has been used either as an alternative for ''monkey'' or for any tailless, relatively human-like primate. Sir Wilfrid Le Gros Clark was one of the primatologists who developed the idea of trends in primate evolution and the methodology of arranging the living members of an order into an "ascending series" leading to humans. Commonly used names for groups of primates such as ''
prosimian Prosimians are a group of primates that includes all living and extinct Strepsirrhini, strepsirrhines (lemurs, Lorisoidea, lorisoids, and Adapiformes, adapiforms), as well as the Haplorhini, haplorhine tarsiers and their extinct relatives, the Om ...
s'', ''
monkey Monkey is a common name that may refer to most mammals of the infraorder Simiiformes, also known as simians. Traditionally, all animals in the group now known as simians are counted as monkeys except the apes. Thus monkeys, in that sense, co ...
s'', '' lesser apes'', and ''
great ape The Hominidae (), whose members are known as the great apes or hominids (), are a taxonomic family of primates that includes eight extant species in four genera: '' Pongo'' (the Bornean, Sumatran and Tapanuli orangutan); '' Gorilla'' (the ...
s'' reflect this methodology. According to our current understanding of the evolutionary history of the primates, several of these groups are
paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
, or rather they do not include all the descendants of a common ancestor. In contrast with Clark's methodology, modern classifications typically identify (or name) only those groupings that are
monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...
; that is, such a named group includes ''all'' the descendants of the group's common ancestor. All groups with scientific names are
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
s, or monophyletic groups, and the sequence of scientific classification reflects the evolutionary history of the related lineages. Groups that are traditionally named are shown on the right; they form an "ascending series" (per Clark, see above), and several groups are paraphyletic: * Prosimians contain two monophyletic groups (the suborder Strepsirrhini, or lemurs, lorises and allies, as well as the tarsiers of the suborder Haplorhini); it is a paraphyletic grouping because it excludes the Simiiformes, which also are descendants of the common ancestor Primates. * Monkeys comprise two monophyletic groups, New World monkeys and Old World monkeys, but is paraphyletic because it excludes hominoids, superfamily Hominoidea, also descendants of the common ancestor Simiiformes. * Apes as a whole, and the
great ape The Hominidae (), whose members are known as the great apes or hominids (), are a taxonomic family of primates that includes eight extant species in four genera: '' Pongo'' (the Bornean, Sumatran and Tapanuli orangutan); '' Gorilla'' (the ...
s, are paraphyletic if the terms are used such that they exclude humans. Thus, the members of the two sets of groups, and hence names, do not match, which causes problems in relating scientific names to common (usually traditional) 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 remedy is to create a new common name, in this case ''hominoids''. Another possibility is to expand the use of one of the traditional names. For example, in his 2005 book, the
vertebrate Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...
palaeontologist Paleontology, also spelled as palaeontology or palæontology, is the scientific study of the life of the past, mainly but not exclusively through the study of fossils. Paleontologists use fossils as a means to classify organisms, measure geolo ...
Benton wrote, "The apes, Hominoidea, today include the
gibbon Gibbons () are apes in the family Hylobatidae (). The family historically contained one genus, but now is split into four extant genera and 20 species. Gibbons live in subtropical and tropical forests from eastern Bangladesh and Northeast Indi ...
s and
orangutan Orangutans are great apes native to the rainforests of Indonesia and Malaysia. They are now found only in parts of Borneo and Sumatra, but during the Pleistocene they ranged throughout Southeast Asia and South China. Classified in the genus ...
... the
gorilla Gorillas are primarily herbivorous, terrestrial great apes that inhabit the tropical forests of equatorial Africa. The genus ''Gorilla'' is divided into two species: the eastern gorilla and the western gorilla, and either four or five su ...
and
chimpanzee The chimpanzee (; ''Pan troglodytes''), also simply known as the chimp, is a species of Hominidae, great ape native to the forests and savannahs of tropical Africa. It has four confirmed subspecies and a fifth proposed one. When its close rel ...
... and
human Humans (''Homo sapiens'') or modern humans are the most common and widespread species of primate, and the last surviving species of the genus ''Homo''. They are Hominidae, great apes characterized by their Prehistory of nakedness and clothing ...
s"; thereby Benton was using ''apes'' to mean hominoids. In that case, the group heretofore called ''apes'' must now be identified as the non-human apes. , there is no consensus as to whether to accept traditional (that is, common), but paraphyletic, names or to use monophyletic names only; or to use 'new' common names or adaptations of old ones. Both competing approaches can be found in biological sources, often in the same work, and sometimes by the same author. Thus, Benton defines ''apes'' to include humans, then he repeatedly uses ''ape-like'' to mean 'like an ape rather than a human'; and when discussing the reaction of others to a new fossil he writes of "claims that '' Orrorin'' ... was an ape rather than a human".


Classification of living primates

Order Primates was established by
Carl Linnaeus Carl Linnaeus (23 May 1707 – 10 January 1778), also known after ennoblement in 1761 as Carl von Linné,#Blunt, Blunt (2004), p. 171. was a Swedish biologist and physician who formalised binomial nomenclature, the modern system of naming o ...
in 1758, in the tenth edition of his book ''
Systema Naturae ' (originally in Latin written ' with the Orthographic ligature, ligature æ) is one of the major works of the Sweden, Swedish botanist, zoologist and physician Carl Linnaeus (1707–1778) and introduced the Linnaean taxonomy. Although the syste ...
'', for the genera ''
Homo ''Homo'' () is a genus of great ape (family Hominidae) that emerged from the genus ''Australopithecus'' and encompasses only a single extant species, ''Homo sapiens'' (modern humans), along with a number of extinct species (collectively called ...
'' (humans), '' Simia'' (other apes and monkeys), ''
Lemur Lemurs ( ; from Latin ) are Strepsirrhini, wet-nosed primates of the Superfamily (biology), superfamily Lemuroidea ( ), divided into 8 Family (biology), families and consisting of 15 genera and around 100 existing species. They are Endemism, ...
'' (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 Henri Marie Ducrotay de Blainville (; 12 September 1777 – 1 May 1850) was a French zoologist and anatomist. Life Blainville was born at Arques-la-Bataille, Arques, near Dieppe, Seine-Maritime, Dieppe. As a young man, he went to Paris to study a ...
, following Linnaeus and aping his nomenclature, established the orders Secundates (including the suborders Chiroptera,
Insectivora The Order (biology), order Insectivora (from Latin ''insectum'' "insect" and ''vorare'' "to eat") is a now-abandoned biological grouping within the class of mammals. Some species have now been moved out, leaving the remaining ones in the order ...
and
Carnivora Carnivora ( ) is an order of placental mammals specialized primarily in eating flesh, whose members are formally referred to as carnivorans. The order Carnivora is the sixth largest order of mammals, comprising at least 279 species. Carnivor ...
), Tertiates (or Glires) and Quaternates (including Gravigrada, Pachydermata and
Ruminantia Ruminants are herbivorous grazing or browsing artiodactyls belonging to the suborder Ruminantia that are able to acquire nutrients from plant-based food by Enteric fermentation, fermenting it in a specialized stomach prior to digestion, principa ...
), but these new taxa were not accepted. Before Anderson and Jones introduced the classification of Strepsirrhini and Haplorhini in 1984, (followed by McKenna and Bell's 1997 work ''Classification of Mammals: Above the species level''), Primates was divided into two superfamilies: Prosimii and
Anthropoidea The simians, anthropoids, or higher primates are an infraorder (Simiiformes ) of primates containing all animals traditionally called monkeys and apes. More precisely, they consist of the parvorders Platyrrhini (New World monkeys) and Catar ...
. Prosimii included all of the
prosimian Prosimians are a group of primates that includes all living and extinct Strepsirrhini, strepsirrhines (lemurs, Lorisoidea, lorisoids, and Adapiformes, adapiforms), as well as the Haplorhini, haplorhine tarsiers and their extinct relatives, the Om ...
s: Strepsirrhini plus the
tarsier Tarsiers ( ) are haplorhine primates of the family Tarsiidae, which is the lone extant family within the infraorder Tarsiiformes. Although the group was prehistorically more globally widespread, all of the existing species are restricted to M ...
s. Anthropoidea contained all of the
simian The simians, anthropoids, or higher primates are an infraorder (Simiiformes ) of primates containing all animals traditionally called monkeys and apes. More precisely, they consist of the parvorders New World monkey, Platyrrhini (New World mon ...
s. The
cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...
below shows one possible classification sequence of the living primates: groups that use common (traditional) names are shown on the right.


Phylogeny and genetics

Order Primates is part of the clade Euarchontoglires, which is nested within the clade
Eutheria Eutheria (from Greek , 'good, right' and , 'beast'; ), also called Pan-Placentalia, is the clade consisting of Placentalia, placental mammals and all therian mammals that are more closely related to placentals than to marsupials. Eutherians ...
of Class
Mammal A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...
ia. Recent molecular genetic research on primates, colugos, and treeshrews has shown that the two species of colugos are more closely related to primates than to treeshrews, even though treeshrews were at one time considered primates. These three orders make up the
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
Euarchonta. The combination of this clade with the clade Glires (composed of
Rodent Rodents (from Latin , 'to gnaw') are mammals of the Order (biology), order Rodentia ( ), which are characterized by a single pair of continuously growing incisors in each of the upper and Mandible, lower jaws. About 40% of all mammal specie ...
ia and Lagomorpha) forms the clade Euarchontoglires. Variously, both Euarchonta and Euarchontoglires are ranked as superorders. Some scientists consider Dermoptera to be a suborder of Primates and use the suborder Euprimates for the "true" primates.


Evolutionary history

The primate lineage is thought to go back at least near the Cretaceous–Paleogene boundary or around 74–63 ( mya). The earliest possible primate/proto-primate may be '' Purgatorius'', which dates back to
Early Paleocene The Danian is the oldest age (geology), age or lowest stage (stratigraphy), stage of the Paleocene Epoch or series (stratigraphy), Series, of the Paleogene Period or system (stratigraphy), System, and of the Cenozoic Era or Erathem. The beginnin ...
of North America ~66mya. The oldest known primates from the fossil record date to the Late Paleocene of Africa, c.57 mya ('' Altiatlasius'') or the Paleocene-Eocene transition in the northern continents, c. 55 mya ('' Cantius'', '' Donrussellia'', '' Altanius'', '' Plesiadapis'' and '' Teilhardina''). 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 Cladistics ( ; from Ancient Greek 'branch') is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is ...
reckoning, the order Primates is
monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...
. The suborder
Strepsirrhini Strepsirrhini or Strepsirhini (; ) is a Order (biology), suborder of primates that includes the Lemuriformes, lemuriform primates, which consist of the lemurs of Fauna of Madagascar, Madagascar, galagos ("bushbabies") and pottos from Fauna of A ...
, the " 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 strepsirrhine families are the five related
lemur Lemurs ( ; from Latin ) are Strepsirrhini, wet-nosed primates of the Superfamily (biology), superfamily Lemuroidea ( ), divided into 8 Family (biology), families and consisting of 15 genera and around 100 existing species. They are Endemism, ...
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 The Eocene ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...
, 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 demonstrated that '' Darwinius masillae'' fits into this grouping. The latter was closely related to tarsiers, monkeys, and apes. How these two groups relate to extant primates is unclear. Omomyids perished about 30 mya, while adapiforms survived until about 10 mya. According to genetic studies, the lemurs of Madagascar diverged from the lorisoids 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, a founding lemur population of a few individuals is thought to have reached Madagascar from Africa via a single rafting event between 50 and 80 mya. Other colonization options have been suggested, such as multiple colonizations from Africa and India, but none are supported by the genetic and molecular evidence. Until recently, the
aye-aye The aye-aye (''Daubentonia madagascariensis'') is a long-fingered lemur, a Strepsirrhini, strepsirrhine primate native to Madagascar with rodent-like teeth that perpetually grow and a special thin middle finger that they can use to catch grubs ...
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 Haplorhini (), the haplorhines (Greek language, Greek for "simple-nosed") or the "dry-nosed" primates is a suborder of primates containing the Tarsiiformes, tarsiers and the simians (Simiiformes or anthropoids), as sister of the Strepsirrhini ("m ...
, the simple-nosed or "dry-nosed" primates, is composed of two sister clades.
Prosimian Prosimians are a group of primates that includes all living and extinct Strepsirrhini, strepsirrhines (lemurs, Lorisoidea, lorisoids, and Adapiformes, adapiforms), as well as the Haplorhini, haplorhine tarsiers and their extinct relatives, the Om ...
tarsiers in the family Tarsiidae (monotypic in its own infraorder Tarsiiformes), represent the most basal division, originating about 58 mya. The earliest known haplorhine skeleton, that of 55 MA old tarsier-like '' Archicebus'', was found in central China, supporting an already suspected Asian origin for the group. The infraorder Simiiformes (simian primates, consisting of monkeys and apes) emerged about 40 mya, possibly also in Asia; if so, they dispersed across the Tethys Sea from Asia to Africa soon afterwards. There are two simian clades, both parvorders:
Catarrhini The parvorder Catarrhini (known commonly as catarrhine monkeys, Old World anthropoids, or Old World monkeys) consists of the Cercopithecoidea and apes (Hominoidea). In 1812, Geoffroy grouped those two groups together and established the name ...
, which developed in Africa, consisting of Old World monkeys, humans and the other apes, and Platyrrhini, which developed in South America, consisting of New World monkeys. A third clade, which included the eosimiids, developed in Asia, but became 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. The anthropoid primates possibly traversed the Atlantic Ocean from Africa to South America during the
Eocene The Eocene ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...
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 Continental drift is a highly supported scientific theory, originating in the early 20th century, that Earth's continents move or drift relative to each other over geologic time. The theory of continental drift has since been validated and inc ...
, the Atlantic Ocean was not nearly as wide at the time as it is today. Research suggests that a small 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 The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...
. Soon after, the lorises and tarsiers made the same journey. The first hominin 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 discovered recently, 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 ''Mammal Species of the World: A Taxonomic and Geographic Reference'' is a standard reference work in mammalogy giving descriptions and Bibliographic database, bibliographic data for the known species of mammals. It is now in its third edition, ...
'' (MSW3). However, publications since the taxonomy in MSW3 was compiled in 2003 have pushed the number to 522 species, or 708 including subspecies.


Hybrids

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.


Clones

On 24 January 2018, scientists in China reported in the journal '' Cell'' the creation of two crab-eating macaque clones, named '' Zhong Zhong'' and '' Hua Hua'', using the complex DNA transfer method that produced ''Dolly'' the sheep, for the first time.


Anatomy and physiology


Head

The primate skull has a large, domed cranium, which is particularly prominent in
anthropoids The simians, anthropoids, or higher primates are an infraorder (Simiiformes ) of primates containing all animals traditionally called monkeys and apes. More precisely, they consist of the parvorders Platyrrhini (New World monkeys) and Catar ...
. 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
human Humans (''Homo sapiens'') or modern humans are the most common and widespread species of primate, and the last surviving species of the genus ''Homo''. They are Hominidae, great apes characterized by their Prehistory of nakedness and clothing ...
s than in the greatest nonhuman primate, reflecting a larger brain size. The mean endocranial volume is 1,201 cubic centimeters in humans, 469 cm3 in
gorilla Gorillas are primarily herbivorous, terrestrial great apes that inhabit the tropical forests of equatorial Africa. The genus ''Gorilla'' is divided into two species: the eastern gorilla and the western gorilla, and either four or five su ...
s, 400 cm3 in
chimpanzee The chimpanzee (; ''Pan troglodytes''), also simply known as the chimp, is a species of Hominidae, great ape native to the forests and savannahs of tropical Africa. It has four confirmed subspecies and a fifth proposed one. When its close rel ...
s and 397 cm3 in
orangutan Orangutans are great apes native to the rainforests of Indonesia and Malaysia. They are now found only in parts of Borneo and Sumatra, but during the Pleistocene they ranged throughout Southeast Asia and South China. Classified in the genus ...
s. The primary evolutionary trend of primates has been the elaboration of the brain, in particular the
neocortex The neocortex, also called the neopallium, isocortex, or the six-layered cortex, is a set of layers of the mammalian cerebral cortex involved in higher-order brain functions such as sensory perception, cognition, generation of motor commands, ...
(a part of the
cerebral cortex The cerebral cortex, also known as the cerebral mantle, is the outer layer of neural tissue of the cerebrum of the brain in humans and other mammals. It is the largest site of Neuron, neural integration in the central nervous system, and plays ...
), which is involved with sensory perception, generation of motor commands, spatial reasoning, conscious thought and, in humans,
language Language is a structured system of communication that consists of grammar and vocabulary. It is the primary means by which humans convey meaning, both in spoken and signed language, signed forms, and may also be conveyed through writing syste ...
. 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. The visual acuity of
human Humans (''Homo sapiens'') or modern humans are the most common and widespread species of primate, and the last surviving species of the genus ''Homo''. They are Hominidae, great apes characterized by their Prehistory of nakedness and clothing ...
s and other hominids is exceptional; they have the most acute vision known among all vertebrates, with the exception of certain species of predatory birds. Primates have forward-facing eyes on the front of the skull;
binocular vision Binocular vision is seeing with two eyes. The Field_of_view, field of view that can be surveyed with two eyes is greater than with one eye. To the extent that the visual fields of the two eyes overlap, #Depth, binocular depth can be perceived. Th ...
allows accurate distance perception, useful for the brachiating ancestors of all great apes. A bony ridge above the eye sockets reinforces weaker bones in the face, which are put under strain during chewing. Strepsirrhines have a postorbital bar, a bone around the eye socket, to protect their eyes; in contrast, the higher primates, haplorhines, have evolved fully enclosed sockets. Primates show an evolutionary trend towards a reduced
snout A snout is the protruding portion of an animal's face, consisting of its nose, mouth, and jaw. In many animals, the structure is called a muzzle, Rostrum (anatomy), rostrum, beak or proboscis. The wet furless surface around the nostrils of the n ...
. 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. Dental pattern in primates vary considerably; although some have lost most of their incisors, all retain at least one lower incisor. In most strepsirrhines, the lower incisors form a toothcomb, which is used in grooming and sometimes foraging. Old World monkeys have eight
premolar The premolars, also called premolar Tooth (human), teeth, or bicuspids, are transitional teeth located between the Canine tooth, canine and Molar (tooth), molar teeth. In humans, there are two premolars per dental terminology#Quadrant, quadrant in ...
s, compared with 12 in New World monkeys. The Old World species are divided into apes and monkeys depending on the number of cusps on their
molars The molars or molar teeth are large, flat tooth, teeth at the back of the mouth. They are more developed in mammal, mammals. They are used primarily to comminution, grind food during mastication, chewing. The name ''molar'' derives from Latin, '' ...
: monkeys have four, apes have five - although humans may have four or five. The main hominid molar cusp (
hypocone A cusp is a pointed, projecting, or elevated feature. In animals, it is usually used to refer to raised points on the crowns of teeth. The concept is also used with regard to the leaflets of the four heart valves. The mitral valve, which has ...
) 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 noses and forward-facing lower front teeth.


Body

Primates generally have five digits on each limb (
pentadactyly In biology, dactyly is the arrangement of wikt:digit, digits (fingers and toes) on the hands, feet, or sometimes wings of a tetrapod animal. The term is derived from the Greek (language), Greek word () meaning "finger." Sometimes the suffix "- ...
), with a characteristic type of keratin fingernail on the end of each finger and toe. The bottom sides of the hands and feet have sensitive pads on the fingertips. Most have opposable thumbs, a characteristic primate feature most developed in
human Humans (''Homo sapiens'') or modern humans are the most common and widespread species of primate, and the last surviving species of the genus ''Homo''. They are Hominidae, great apes characterized by their Prehistory of nakedness and clothing ...
s, though not limited to this order (
opossum Opossums () are members of the marsupial order Didelphimorphia () endemic to the Americas. The largest order of marsupials in the Western Hemisphere, it comprises 126 species in 18 genera. Opossums originated in South America and entered North A ...
s and
koala The koala (''Phascolarctos cinereus''), sometimes inaccurately called the koala bear, is an arboreal herbivorous marsupial native to Australia. It is the only Extant taxon, extant representative of the Family (biology), family ''Phascolar ...
s, for example, also have them). Thumbs allow some species to use
tools A tool is an object that can extend an individual's ability to modify features of the surrounding environment or help them accomplish a particular task. Although many animals use simple tools, only human beings, whose use of stone tools dates ...
. 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 locomotion. Prosimians have clawlike nails on the second toe of each foot, called toilet-claws, which they use for grooming. The primate collar bone is a prominent element of the pectoral girdle; this allows the shoulder joint broad mobility. Compared to Old World monkeys, apes have more mobile shoulder joints and arms due to the dorsal position of the
scapula The scapula (: scapulae or scapulas), also known as the shoulder blade, is the bone that connects the humerus (upper arm bone) with the clavicle (collar bone). Like their connected bones, the scapulae are paired, with each scapula on either side ...
, broad ribcages that are flatter front-to-back, a shorter, less mobile spine, and with lower
vertebra Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spina ...
e greatly reduced - resulting in tail loss in some species. Prehensile tails are found in the New World atelids, including the howler,
spider Spiders (order (biology), order Araneae) are air-breathing arthropods that have eight limbs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude spider silk, silk. They are the largest order of arachnids and ran ...
, woolly spider, woolly monkeys; and in capuchins. Male primates have a low-hanging penis and testes descended into a scrotum.


Sexual dimorphism

Sexual dimorphism Sexual dimorphism is the condition where sexes of the same species exhibit different Morphology (biology), morphological characteristics, including characteristics not directly involved in reproduction. The condition occurs in most dioecy, di ...
is often exhibited in
simian The simians, anthropoids, or higher primates are an infraorder (Simiiformes ) of primates containing all animals traditionally called monkeys and apes. More precisely, they consist of the parvorders New World monkey, Platyrrhini (New World mon ...
s, 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 Sexual selection is a mechanism of evolution in which members of one sex mate choice, choose mates of the other sex to mating, mate with (intersexual selection), and compete with members of the same sex for access to members of the opposite sex ...
,
natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the Heredity, heritable traits characteristic of a population over generation ...
, 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 Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last comm ...
of dimorphism, and some extinct hominids probably had greater dimorphism than any living primate.


Locomotion

Primate species move by brachiation,
bipedalism Bipedalism is a form of terrestrial locomotion where an animal moves by means of its two rear (or lower) Limb (anatomy), limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning 'two feet' (from ...
, leaping, arboreal and terrestrial
quadruped Quadrupedalism is a form of locomotion in which animals have four legs that are used to bear weight and move around. An animal or machine that usually maintains a four-legged posture and moves using all four legs is said to be a quadruped (fr ...
alism, 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
indri The indri (; ''Indri indri''), also called the babakoto, is one of the largest living lemurs, with a head-body length of about and a weight of between . It has a black and white coat and maintains an upright posture when climbing or clinging. ...
s), sportive lemurs, and all
tarsier Tarsiers ( ) are haplorhine primates of the family Tarsiidae, which is the lone extant family within the infraorder Tarsiiformes. Although the group was prehistorically more globally widespread, all of the existing species are restricted to M ...
s. 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.
Gibbon Gibbons () are apes in the family Hylobatidae (). The family historically contained one genus, but now is split into four extant genera and 20 species. Gibbons live in subtropical and tropical forests from eastern Bangladesh and Northeast Indi ...
s, muriquis and spider monkeys all brachiate extensively, with gibbons sometimes doing so in remarkably acrobatic fashion. Woolly monkeys also brachiate at times.
Orangutan Orangutans are great apes native to the rainforests of Indonesia and Malaysia. They are now found only in parts of Borneo and Sumatra, but during the Pleistocene they ranged throughout Southeast Asia and South China. Classified in the genus ...
s 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.
Chimpanzee The chimpanzee (; ''Pan troglodytes''), also simply known as the chimp, is a species of Hominidae, great ape native to the forests and savannahs of tropical Africa. It has four confirmed subspecies and a fifth proposed one. When its close rel ...
s and
gorilla Gorillas are primarily herbivorous, terrestrial great apes that inhabit the tropical forests of equatorial Africa. The genus ''Gorilla'' is divided into two species: the eastern gorilla and the western gorilla, and either four or five su ...
s 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.


Vision

The evolution of color vision in primates is unique among most
eutheria Eutheria (from Greek , 'good, right' and , 'beast'; ), also called Pan-Placentalia, is the clade consisting of Placentalia, placental mammals and all therian mammals that are more closely related to placentals than to marsupials. Eutherians ...
n mammals. While the remote
vertebrate Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...
ancestors of the primates possessed three color vision (trichromaticism), the
nocturnal Nocturnality is a ethology, behavior in some non-human animals characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnality, diurnal meaning the opposite. Nocturnal creatur ...
,
warm-blooded Warm-blooded is a term referring to animal species whose bodies maintain a temperature higher than that of their environment. In particular, homeothermic species (including birds and mammals) maintain a stable body temperature by regulating ...
, mammalian ancestors lost one of three cones in the
retina The retina (; or retinas) is the innermost, photosensitivity, light-sensitive layer of tissue (biology), tissue of the eye of most vertebrates and some Mollusca, molluscs. The optics of the eye create a focus (optics), focused two-dimensional ...
during the
Mesozoic The Mesozoic Era is the Era (geology), era of Earth's Geologic time scale, geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceous Period (geology), Periods. It is characterized by the dominance of archosaurian r ...
era. Fish,
reptile Reptiles, as commonly defined, are a group of tetrapods with an ectothermic metabolism and Amniotic egg, amniotic development. Living traditional reptiles comprise four Order (biology), orders: Testudines, Crocodilia, Squamata, and Rhynchocepha ...
s and birds are therefore trichromatic or tetrachromatic, while all mammals, with the exception of some primates and
marsupial Marsupials are a diverse group of mammals belonging to the infraclass Marsupialia. They are natively found in Australasia, Wallacea, and the Americas. One of marsupials' unique features is their reproductive strategy: the young are born in a r ...
s, 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 Gene duplication (or chromosomal duplication or gene amplification) is a major mechanism through which new genetic material is generated during molecular evolution. It can be defined as any duplication of a region of DNA that contains a gene ...
of the red-green
opsin Animal opsins are G-protein-coupled receptors and a group of proteins made light-sensitive via a chromophore, typically retinal. When bound to retinal, opsins become retinylidene proteins, but are usually still called opsins regardless. Most pro ...
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 Zygosity (the noun, zygote, is from the Greek "yoked," from "yoke") () is the degree to which both copies of a chromosome or gene have the same genetic sequence. In other words, it is the degree of similarity of the alleles in an organism. Mos ...
for two
allele An allele is a variant of the sequence of nucleotides at a particular location, or Locus (genetics), locus, on a DNA molecule. Alleles can differ at a single position through Single-nucleotide polymorphism, single nucleotide polymorphisms (SNP), ...
s 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 Gene duplication (or chromosomal duplication or gene amplification) is a major mechanism through which new genetic material is generated during molecular evolution. It can be defined as any duplication of a region of DNA that contains a gene ...
. Howler monkeys are one of the most specialized leaf-eaters of the New World monkeys; fruits are not a major part of their diets, 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 selected by environment.


Behavior


Social systems

Richard Wrangham Richard Walter Wrangham (born 1948) is an English anthropologist and primatologist; he is Professor of Biological Anthropology at Harvard University. His research and writing have involved ape behavior, human evolution, violence, and cooking. ...
stated that
social systems In sociology, a social system is the patterned network of relationships constituting a coherent whole that exist between individuals, groups, and institutions. It is the formal Social structure, structure of role and status that can form in a smal ...
of primates are best classified by the amount of movement by females occurring between groups. He proposed four categories: * Female transfer systems – females move away from the group in which they were born. Females of a group will not be closely related whereas males will have remained with their natal groups, and this close association may be influential in social behavior. The groups formed are generally quite small. This organization can be seen in
chimpanzee The chimpanzee (; ''Pan troglodytes''), also simply known as the chimp, is a species of Hominidae, great ape native to the forests and savannahs of tropical Africa. It has four confirmed subspecies and a fifth proposed one. When its close rel ...
s, where the males, who are typically related, will cooperate in defense of the group's territory. Evidence of this social system (called
patrilocal residence In social anthropology, patrilocal residence or patrilocality, also known as virilocal residence or virilocality, are terms referring to the social system in which a married couple resides with or near the husband's parents. The concept of loca ...
when used by the
Anthropology Anthropology is the scientific study of humanity, concerned with human behavior, human biology, cultures, society, societies, and linguistics, in both the present and past, including archaic humans. Social anthropology studies patterns of behav ...
) has also been found among
Neanderthal Neanderthals ( ; ''Homo neanderthalensis'' or sometimes ''H. sapiens neanderthalensis'') are an extinction, extinct group of archaic humans who inhabited Europe and Western and Central Asia during the Middle Pleistocene, Middle to Late Plei ...
remains in
Spain Spain, or the Kingdom of Spain, is a country in Southern Europe, Southern and Western Europe with territories in North Africa. Featuring the Punta de Tarifa, southernmost point of continental Europe, it is the largest country in Southern Eur ...
and in remains of ''
Australopithecus ''Australopithecus'' (, ; or (, ) is a genus of early hominins that existed in Africa during the Pliocene and Early Pleistocene. The genera ''Homo'' (which includes modern humans), ''Paranthropus'', and ''Kenyanthropus'' evolved from some ''Aus ...
'' and ''
Paranthropus robustus ''Paranthropus robustus'' is a species of robust australopithecine from the Early and possibly Middle Pleistocene of the Cradle of Humankind, South Africa, about 2.27 to 0.87 (or, more conservatively, 2 to 1) million years ago. It has been ide ...
'' groups in southern Africa. 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. Group sizes are usually larger. This system is common among the
ring-tailed lemur The ring-tailed lemur (''Lemur catta'') is a medium- to larger-sized strepsirrhine (wet-nosed) primate and the most internationally recognized lemur species, owing to its long, black-and-white, ringed tail. It belongs to Lemuridae, one of fiv ...
, 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.
Indri The indri (; ''Indri indri''), also called the babakoto, is one of the largest living lemurs, with a head-body length of about and a weight of between . It has a black and white coat and maintains an upright posture when climbing or clinging. ...
, lariang tarsiers, Callitrichidae monkeys and gibbons use this system, although "monogamy" in this context does not necessarily mean absolute sexual fidelity. These species do not live in larger groups. * Solitary species – males and females live in overlapping home ranges. This type of organization is found in lorises, galagos, mouse lemurs, aye-ayes and orangutans. Other systems are known to occur as well. For example, with howler monkeys and
gorilla Gorillas are primarily herbivorous, terrestrial great apes that inhabit the tropical forests of equatorial Africa. The genus ''Gorilla'' is divided into two species: the eastern gorilla and the western gorilla, and either four or five su ...
s 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 also use this system. The transfer of females or males from their native group is likely an adaptation for avoiding inbreeding. An analysis of breeding records of captive primate colonies representing numerous different species indicates that the infant mortality of inbred young is generally higher than that of non-inbred young. This effect of inbreeding on infant mortality is probably largely a result of increased expression of deleterious recessive alleles (see
Inbreeding depression Inbreeding depression is the reduced biological fitness caused by loss of genetic diversity as a consequence of inbreeding, the breeding of individuals closely related genetically. This loss of genetic diversity results from small population siz ...
). Primatologist Jane Goodall, who studied in the
Gombe Stream National Park Gombe National Park () is a national park in Tanzania, located in the Kigoma District of the Kigoma Region. It was formerly called Gombe Stream National Park.Tanzania National Parks“Gombe Stream National Park”, 2008. Overview Established in ...
, noted fission-fusion societies in chimpanzees. There is ''fission'' when 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 bonobo (; ''Pan paniscus''), also historically called the pygmy chimpanzee (less often the dwarf chimpanzee or gracile chimpanzee), is an endangered great ape and one of the two species making up the genus ''Pan (genus), Pan'' (the other bei ...
. The gelada has a similar social structure in which many smaller groups come together to form temporary herds of up to 600 monkeys.
Human Humans (''Homo sapiens'') or modern humans are the most common and widespread species of primate, and the last surviving species of the genus ''Homo''. They are Hominidae, great apes characterized by their Prehistory of nakedness and clothing ...
s also form fission-fusion societies. In hunter-gatherer societies, humans form groups which are made up of several individuals that may split up to obtain different resources. These social systems are affected by three main ecological factors: distribution of resources, group size, and
predation Predation is a biological interaction in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common List of feeding behaviours, feeding behaviours that includes parasitism and micropredation ...
. Within a social group there is a balance between cooperation and competition. Cooperative behaviors in many primates species include
social grooming Social grooming is a behavior in which social animals, including humans, clean or maintain one another's bodies or appearances. A related term, allogrooming, indicates social grooming between members of the same species. Grooming is a major s ...
(removing skin parasites and cleaning wounds), food sharing, and collective defense against predators or of a territory. Aggressive behaviors often signal competition for food, sleeping sites or mates. Aggression is also used in establishing dominance hierarchies. In November 2023, scientists reported, for the first time, evidence that groups of primates, particularly
bonobo The bonobo (; ''Pan paniscus''), also historically called the pygmy chimpanzee (less often the dwarf chimpanzee or gracile chimpanzee), is an endangered great ape and one of the two species making up the genus ''Pan (genus), Pan'' (the other bei ...
s, are capable of cooperating with each other.


Interspecific associations

Several species of primates are known to associate in the wild. Some of these associations have been extensively studied. In the Tai Forest of Africa, several species coordinate anti-predator behavior. These include the Diana monkey,
Campbell's mona monkey Campbell's mona monkey (''Cercopithecus campbelli''), also known as Campbell's guenon and Campbell's monkey, is a species of primate in the family Cercopithecidae found in the Ivory Coast, Gambia, Ghana, Guinea, Guinea-Bissau, Liberia, Senegal, ...
, lesser spot-nosed monkey, western red colobus, king colobus (western black and white colobus), and sooty mangabey, which coordinate anti-predator alarm calls. Among the predators of these monkeys is the
common chimpanzee The chimpanzee (; ''Pan troglodytes''), also simply known as the chimp, is a species of great ape native to the forests and savannahs of tropical Africa. It has four confirmed subspecies and a fifth proposed one. When its close relative the ...
. 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. In South America, squirrel monkeys associate with capuchin monkeys. This may have more to do with foraging benefits to the squirrel monkeys than anti-predation benefits.


Mating systems

The mating systems of primates vary between
monogamy Monogamy ( ) is a social relation, relationship of Dyad (sociology), two individuals in which they form a mutual and exclusive intimate Significant other, partnership. Having only one partner at any one time, whether for life or #Serial monogamy ...
,
polyandry Polyandry (; ) is a form of polygamy in which a woman takes two or more husbands at the same time. Polyandry is contrasted with polygyny, involving one male and two or more females. If a marriage involves a plural number of "husbands and wives ...
,
polygyny Polygyny () is a form of polygamy entailing the marriage of a man to several women. The term polygyny is from Neoclassical Greek πολυγυνία (); . Incidence Polygyny is more widespread in Africa than in any other continent. Some scholar ...
and polygynandry. In monogamous species, adult males and females form long-lasting
pair bond In biology, a pair bond is the strong affinity that develops in some species between a mating pair, often leading to the production and rearing of young and potentially a lifelong bond. Pair-bonding is a term coined in the 1940s that is frequently ...
s. Compared to other systems, there is little competition for mating rights and males and females tend to be similar in size. Polyandry, which involves groups consisting of single females mating with multiple males, may arise as a secondary mating system in monogamous species. In the brown-mantled tamarin, a female may breeding with one or two males. Polyandry may have developed due to the high frequency of twin births, which require more help in raising. Polygynous species include gorillas, Hanuman langurs, geladas, hamadryas baboons, proboscis monkeys, and golden snub-nosed monkeys, and consists of one male mating with multiple females within a
harem A harem is a domestic space that is reserved for the women of the house in a Muslim family. A harem may house a man's wife or wives, their pre-pubescent male children, unmarried daughters, female domestic Domestic worker, servants, and other un ...
or one-male unit. Sexual dimorphism tends to be higher in these species and males may also develop prominent
secondary sex characteristic A secondary sex characteristic is a physical characteristic of an organism that is related to or derived from its sex, but not directly part of its reproductive system. In humans, these characteristics typically start to appear during pubert ...
s. In the patriarchal hamadryas baboon, the males aggressively herd females into their groups and violently discipline those that wander. By contrast, in gelada society, which is based on female kinship, a male is dependent on the support of the females in his unit and cannot impose on them. Polygynous males must defend their harems from rivals, who may try to take over. In some species, such as ring-tailed lemurs, sifakas, macaques, most
baboon Baboons are primates comprising the biology, genus ''Papio'', one of the 23 genera of Old World monkeys, in the family Cercopithecidae. There are six species of baboon: the hamadryas baboon, the Guinea baboon, the olive baboon, the yellow ba ...
s, mangabeys, squirrel monkeys, woolly monkeys, spider monkeys, woolly spider monkeys, chimpanzees and bonobos, both males and females mate with multiple partners. Polygynandry occurs in multimale-multifemale groups, and since females mate many times before conception, males have large testicles for
sperm competition Sperm competition is the competitive process between Spermatozoon, spermatozoa of two or more different males to fertility, fertilize the same Egg cell, egg during sexual reproduction. Competition can occur when females have multiple potential m ...
. Males may exist in a
dominance hierarchy In the zoological field of ethology, a dominance hierarchy (formerly and colloquially called a pecking order) is a type of social hierarchy that arises when members of animal social animal, social groups interact, creating a ranking system. Dif ...
and those at the top will try to monopolize access to the females. Consortships may occur in some species but these are short-term. In solitary-living species, males and females mate with partners whose home ranges they overlap with. This is known as a 'dispersed' mating system. Genetic evidence indicates that humans were predominantly polygynous for most of their existence as a species, but that this began to shift during the Neolithic, when
monogamy Monogamy ( ) is a social relation, relationship of Dyad (sociology), two individuals in which they form a mutual and exclusive intimate Significant other, partnership. Having only one partner at any one time, whether for life or #Serial monogamy ...
started becoming widespread concomitantly with the transition from nomadic to sedentary societies. Most modern human societies consist of monogamous
marriage Marriage, also called matrimony or wedlock, is a culturally and often legally recognised union between people called spouses. It establishes rights and obligations between them, as well as between them and their children (if any), and b ...
s, but allow for polygyny, particularly for those of a high status.


Sexual behavior

Female primates may signal to the male their receptiveness though various displays including eye-contact, tongue-clicking and presenting of the rump. Female lemurs, lorises and galagos will position themselves in the lordosis pose while female chimpanzees, bonobos and some Old World monkeys develop sexual swellings on the rump. Copulation in primates typically involves the males mounting the females from behind, as with most mammals. Belly-to-belly copulation has been recorded in apes, both gibbons and the great apes. Human sex positions are modifications of these two positions. Primates may engage in sexual activity as part of social bonding; including homosexual behaviour. Such behavior play an important role in bonobo society in particular. female bonobos engage in mutual genital-rubbing behavior, possibly to bond socially with each other, thus forming a female nucleus of bonobo society. The bonding among females enables them to dominate most of the males.


Life history

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 Ovulation is an important part of the menstrual cycle in female vertebrates where the egg cells are released from the ovaries as part of the ovarian cycle. In female humans ovulation typically occurs near the midpoint in the menstrual cycle and ...
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.
Infanticide Infanticide (or infant homicide) is the intentional killing of infants or offspring. Infanticide was a widespread practice throughout human history that was mainly used to dispose of unwanted children, its main purpose being the prevention of re ...
is common in polygynous species such as gray langurs and gorillas. Adult males may kill dependent offspring that are not theirs so the female will return to estrus and thus they can sire offspring of their own. Social monogamy in some species may have evolved to combat this behavior. Polygynandry may also lessen the risk of infanticide since paternity becomes uncertain. Primates have a longer juvenile period between weaning and sexual maturity than other mammals of similar size. Some primates such as galagos and New World monkeys use tree-holes for nesting, and park juveniles in leafy patches while foraging. Other primates follow a strategy of "riding", i.e. carrying individuals on the body while feeding. Adults may construct or use nesting sites, sometimes accompanied by juveniles, for the purpose of resting, a behavior which has developed secondarily in the great apes. During the juvenile period, primates are more susceptible than adults to
predation Predation is a biological interaction in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common List of feeding behaviours, feeding behaviours that includes parasitism and micropredation ...
and
starvation Starvation is a severe deficiency in caloric energy intake, below the level needed to maintain an organism's life. It is the most extreme form of malnutrition. In humans, prolonged starvation can cause permanent organ damage and eventually, de ...
; 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, this may be partially due to their slower metabolisms. Late in life, female catarrhine primates appear to undergo a cessation of reproductive function known as
menopause Menopause, also known as the climacteric, is the time when Menstruation, menstrual periods permanently stop, marking the end of the Human reproduction, reproductive stage for the female human. It typically occurs between the ages of 45 and 5 ...
; 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 nutrients including
carbohydrate A carbohydrate () is a biomolecule composed of carbon (C), hydrogen (H), and oxygen (O) atoms. The typical hydrogen-to-oxygen atomic ratio is 2:1, analogous to that of water, and is represented by the empirical formula (where ''m'' and ''n'' ...
s and
lipid Lipids are a broad group of organic compounds which include fats, waxes, sterols, fat-soluble vitamins (such as vitamins A, D, E and K), monoglycerides, diglycerides, phospholipids, and others. The functions of lipids include storing ...
s for energy. Primates in the suborder
Strepsirrhini Strepsirrhini or Strepsirhini (; ) is a Order (biology), suborder of primates that includes the Lemuriformes, lemuriform primates, which consist of the lemurs of Fauna of Madagascar, Madagascar, galagos ("bushbabies") and pottos from Fauna of A ...
(non-tarsier prosimians) are able to synthesize
vitamin C Vitamin C (also known as ascorbic acid and ascorbate) is a water-soluble vitamin found in citrus and other fruits, berries and vegetables. It is also a generic prescription medication and in some countries is sold as a non-prescription di ...
, like most other mammals, while primates of the suborder
Haplorhini Haplorhini (), the haplorhines (Greek language, Greek for "simple-nosed") or the "dry-nosed" primates is a suborder of primates containing the Tarsiiformes, tarsiers and the simians (Simiiformes or anthropoids), as sister of the Strepsirrhini ("m ...
(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
insect Insects (from Latin ') are Hexapoda, hexapod invertebrates of the class (biology), class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (Insect morphology#Head, head, ...
s. 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 The aye-aye (''Daubentonia madagascariensis'') is a long-fingered lemur, a Strepsirrhini, strepsirrhine primate native to Madagascar with rodent-like teeth that perpetually grow and a special thin middle finger that they can use to catch grubs ...
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.


Hunting

Tarsier Tarsiers ( ) are haplorhine primates of the family Tarsiidae, which is the lone extant family within the infraorder Tarsiiformes. Although the group was prehistorically more globally widespread, all of the existing species are restricted to M ...
s are the only
extant Extant or Least-concern species, least concern is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Exta ...
obligate carnivorous primates, exclusively eating insects, crustaceans, small vertebrates and snakes (including venomous species). Capuchin monkeys can exploit many different types of plant matter, including fruit, leaves, flowers, buds, nectar and seeds, but also eat insects and other
invertebrate Invertebrates are animals that neither develop nor retain a vertebral column (commonly known as a ''spine'' or ''backbone''), which evolved from the notochord. It is a paraphyletic grouping including all animals excluding the chordata, chordate s ...
s, bird eggs, and small vertebrates such as birds,
lizard Lizard is the common name used for all Squamata, squamate reptiles other than snakes (and to a lesser extent amphisbaenians), encompassing over 7,000 species, ranging across all continents except Antarctica, as well as most Island#Oceanic isla ...
s,
squirrel Squirrels are members of the family Sciuridae (), a family that includes small or medium-sized rodents. The squirrel family includes tree squirrels, ground squirrels (including chipmunks and prairie dogs, among others), and flying squirrel ...
s and bats. The
common chimpanzee The chimpanzee (; ''Pan troglodytes''), also simply known as the chimp, is a species of great ape native to the forests and savannahs of tropical Africa. It has four confirmed subspecies and a fifth proposed one. When its close relative the ...
eats an
omnivorous An omnivore () is an animal that regularly consumes significant quantities of both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize ...
frugivorous diet. It prefers fruit above all other food items and even seeks out and eats them when they are not abundant. It also eats leaves and leaf buds, seeds, blossoms, stems, pith, bark and resin. Insects and meat make up a small proportion of their diet, estimated as 2%. The meat consumption includes predation on other primate species, such as the western red colobus monkey. The
bonobo The bonobo (; ''Pan paniscus''), also historically called the pygmy chimpanzee (less often the dwarf chimpanzee or gracile chimpanzee), is an endangered great ape and one of the two species making up the genus ''Pan (genus), Pan'' (the other bei ...
is an
omnivorous An omnivore () is an animal that regularly consumes significant quantities of both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize ...
frugivore – the majority of its diet is fruit, but it supplements this with leaves, meat from small
vertebrate Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...
s, such as anomalures,
flying squirrel Flying squirrels (scientifically known as Pteromyini or Petauristini) are a tribe (biology), tribe of 50 species of squirrels in the family (biology), family Squirrel, Sciuridae. Despite their name, they are not in fact capable of full flight i ...
s and
duiker A duiker is a small to medium-sized brown antelope native to sub-Saharan Africa, found in heavily wooded areas. The 22 extant species, including three sometimes considered to be subspecies of the other species, form the subfamily Cephalophinae ...
s, and
invertebrate Invertebrates are animals that neither develop nor retain a vertebral column (commonly known as a ''spine'' or ''backbone''), which evolved from the notochord. It is a paraphyletic grouping including all animals excluding the chordata, chordate s ...
s. In some instances, bonobos have been shown to consume lower-order primates. Until the development of agriculture approximately 10,000 years ago, ''Homo sapiens'' employed a hunter-gatherer method as their sole means of food collection. This involved combining stationary food sources (such as fruits, grains, tubers, and mushrooms, insect larvae and aquatic mollusks) with wild game, which must be hunted and killed in order to be consumed. It has been proposed that humans have used fire to prepare and cook food since the time of ''
Homo erectus ''Homo erectus'' ( ) is an extinction, extinct species of Homo, archaic human from the Pleistocene, spanning nearly 2 million years. It is the first human species to evolve a humanlike body plan and human gait, gait, to early expansions of h ...
''. Around ten thousand years ago, humans developed agriculture, which substantially altered their diet. This change in diet may also have altered human biology; with the spread of
dairy farming Dairy farming is a class of agriculture for the long-term production of milk, which is processed (either on the farm or at a dairy plant, either of which may be called a dairy) for the eventual sale of a dairy product. Dairy farming has a h ...
providing a new and rich source of food, leading to the evolution of the ability to digest
lactose Lactose is a disaccharide composed of galactose and glucose and has the molecular formula C12H22O11. Lactose makes up around 2–8% of milk (by mass). The name comes from (Genitive case, gen. ), the Latin word for milk, plus the suffix ''-o ...
in some adults.


As prey

Predators of primates include various species of
carnivora Carnivora ( ) is an order of placental mammals specialized primarily in eating flesh, whose members are formally referred to as carnivorans. The order Carnivora is the sixth largest order of mammals, comprising at least 279 species. Carnivor ...
ns,
birds of prey Birds of prey or predatory birds, also known as (although not the same as) raptors, are hypercarnivorous bird species that actively predation, hunt and feed on other vertebrates (mainly mammals, reptiles and smaller birds). In addition to speed ...
,
reptile Reptiles, as commonly defined, are a group of tetrapods with an ectothermic metabolism and Amniotic egg, amniotic development. Living traditional reptiles comprise four Order (biology), orders: Testudines, Crocodilia, Squamata, and Rhynchocepha ...
s, and other primates. Even gorillas have been recorded as prey. Predators of primates have diverse hunting strategies and as such, primates have evolved several different antipredator adaptations including
crypsis In ecology, crypsis is the ability of an animal or a plant to avoid observation or detection by other animals. It may be part of a predation strategy or an antipredator adaptation. Methods include camouflage, nocturnality, subterranean life ...
, alarm calls and mobbing. Several species have separate alarm calls for different predators such as air-borne or ground-dwelling predators. Predation may have shaped group size in primates as species exposed to higher predation pressures appear to live in larger groups.


Communication

Lemur Lemurs ( ; from Latin ) are Strepsirrhini, wet-nosed primates of the Superfamily (biology), superfamily Lemuroidea ( ), divided into 8 Family (biology), families and consisting of 15 genera and around 100 existing species. They are Endemism, ...
s,
loris Loris is the common name for the strepsirrhine mammals of the subfamily Lorinae (sometimes spelled Lorisinae) in the family Lorisidae. ''Loris'' is one genus in this subfamily and includes the slender lorises, ''Nycticebus'' is the genus cont ...
es,
tarsier Tarsiers ( ) are haplorhine primates of the family Tarsiidae, which is the lone extant family within the infraorder Tarsiiformes. Although the group was prehistorically more globally widespread, all of the existing species are restricted to M ...
s, and New World monkeys rely on olfactory signals for many aspects of social and reproductive behavior. Specialized glands are used to mark territories with
pheromone A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
s, 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. Facial musculature is very developed in primates, particularly in monkeys and apes, allowing for complex facial communication. Like humans, chimpanzees can distinguish the faces of familiar and unfamiliar individuals. Hand and arm gestures are also important forms of communication for great apes and a single gesture can have multiple functions. Chest-beating in male gorillas is a form of visual and non-vocal sound communication that serves to show fitness to both rivals and females. Primates are a particularly vocal group of mammals.
Indri The indri (; ''Indri indri''), also called the babakoto, is one of the largest living lemurs, with a head-body length of about and a weight of between . It has a black and white coat and maintains an upright posture when climbing or clinging. ...
s and
black-and-white ruffed lemur The black-and-white ruffed lemur (''Varecia variegata'') is an endangered species of ruffed lemur, ruffed lemur, one of two which are endemic to the island of Madagascar. Despite having a larger range than the red ruffed lemur, it has a much sma ...
s make distinctive, loud songs and choruses which maintain territories and act as alarm calls. The Philippine tarsier, has a high-frequency limit of auditory sensitivity of approximately 91 kHz with a dominant frequency of 70 kHz, among the highest recorded for any terrestrial mammal. For Philippine tarsiers, these ultrasonic vocalizations might represent a private channel of communication that subverts detection by predators, prey and competitors, enhances energetic efficiency, or improves detection against low-frequency background noise. Male howler monkeys are among the loudest land mammals as their roars can be heard up to , and relate to intergroup spacing, territorial protection and possibly mate-guarding. Male and female siamangs both possess inflatable pouches in the throat with which pair -bonds use to sing "duets" to each other. The vervet monkey gives a distinct alarm call for each of at least four different predators, and the reactions of other monkeys vary according to the call. Furthermore, many primate species including
chimpanzee The chimpanzee (; ''Pan troglodytes''), also simply known as the chimp, is a species of Hominidae, great ape native to the forests and savannahs of tropical Africa. It has four confirmed subspecies and a fifth proposed one. When its close rel ...
s,
Campbell's mona monkey Campbell's mona monkey (''Cercopithecus campbelli''), also known as Campbell's guenon and Campbell's monkey, is a species of primate in the family Cercopithecidae found in the Ivory Coast, Gambia, Ghana, Guinea, Guinea-Bissau, Liberia, Senegal, ...
s or Diana monkeys have been shown to combine vocalizations in sequences, suggesting
syntax In linguistics, syntax ( ) is the study of how words and morphemes combine to form larger units such as phrases and sentences. Central concerns of syntax include word order, grammatical relations, hierarchical sentence structure (constituenc ...
may not be uniquely humans as previously thought but rather evolutionary ancient, and its origins may be deeply rooted in the primate lineage. Consonant- and vowel-like sounds exist in some orangutan calls and they maintain their meaning over great distances. The time range for the evolution of human language and/or its anatomical prerequisites extends, at least in principle, from the phylogenetic divergence of ''
Homo ''Homo'' () is a genus of great ape (family Hominidae) that emerged from the genus ''Australopithecus'' and encompasses only a single extant species, ''Homo sapiens'' (modern humans), along with a number of extinct species (collectively called ...
'' (2.3 to 2.4 million years ago) from '' Pan'' (5 to 6 million years ago) to the emergence of full behavioral modernity some 50,000–150,000 years ago. Few dispute that ''
Australopithecus ''Australopithecus'' (, ; or (, ) is a genus of early hominins that existed in Africa during the Pliocene and Early Pleistocene. The genera ''Homo'' (which includes modern humans), ''Paranthropus'', and ''Kenyanthropus'' evolved from some ''Aus ...
'' probably lacked vocal communication significantly more sophisticated than that of
great ape The Hominidae (), whose members are known as the great apes or hominids (), are a taxonomic family of primates that includes eight extant species in four genera: '' Pongo'' (the Bornean, Sumatran and Tapanuli orangutan); '' Gorilla'' (the ...
s in general.


Intelligence and cognition

Primates have advanced cognitive abilities: some make tools and use them to acquire food and for social displays; some can perform tasks 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 In psychology and philosophy, theory of mind (often abbreviated to ToM) refers to the capacity to understand other individuals by ascribing mental states to them. A theory of mind includes the understanding that others' beliefs, desires, intent ...
, 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 Spiders (order (biology), order Araneae) are air-breathing arthropods that have eight limbs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude spider silk, silk. They are the largest order of arachnids and ran ...
and capuchin monkeys have scored highly by some measures), as well as in the results of different studies.


Tool use and manufacture

In 1960, Jane Goodall observed a
chimpanzee The chimpanzee (; ''Pan troglodytes''), also simply known as the chimp, is a species of Hominidae, great ape native to the forests and savannahs of tropical Africa. It has four confirmed subspecies and a fifth proposed one. When its close rel ...
poking pieces of grass into a
termite Termites are a group of detritivore, detritophagous Eusociality, eusocial cockroaches which consume a variety of Detritus, decaying plant material, generally in the form of wood, Plant litter, leaf litter, and Humus, soil humus. They are dist ...
mound and then raising the grass to his mouth. After he left, Goodall approached the mound and repeated the behaviour because she was unsure what the chimpanzee was doing. She found that the termites bit onto the grass with their jaws. The chimpanzee had been using the grass as a tool to "fish" or "dip" for termites. There are more limited reports of the closely related
bonobo The bonobo (; ''Pan paniscus''), also historically called the pygmy chimpanzee (less often the dwarf chimpanzee or gracile chimpanzee), is an endangered great ape and one of the two species making up the genus ''Pan (genus), Pan'' (the other bei ...
using tools in the wild; it has been claimed they rarely use tools in the wild although they use tools as readily as chimpanzees when in captivity. It has been reported that females, both chimpanzee and bonobo, use tools more avidly than males.
Orangutan Orangutans are great apes native to the rainforests of Indonesia and Malaysia. They are now found only in parts of Borneo and Sumatra, but during the Pleistocene they ranged throughout Southeast Asia and South China. Classified in the genus ...
s in
Borneo Borneo () is the List of islands by area, third-largest island in the world, with an area of , and population of 23,053,723 (2020 national censuses). Situated at the geographic centre of Maritime Southeast Asia, it is one of the Greater Sunda ...
scoop catfish out of small ponds. Over two years, anthropologist Anne Russon observed orangutans learning to jab sticks at catfish to scare them out of the ponds and in to their waiting hands. There are few reports of
gorilla Gorillas are primarily herbivorous, terrestrial great apes that inhabit the tropical forests of equatorial Africa. The genus ''Gorilla'' is divided into two species: the eastern gorilla and the western gorilla, and either four or five su ...
s using tools in the wild. An adult female western lowland gorilla used a branch as a walking stick apparently to test water depth and to aid her in crossing a pool of water. Another adult female used a detached trunk from a small shrub as a stabilizer during food gathering, and another used a log as a bridge. The first direct observation of a non-ape primate using a tool in a wild environment occurred in 1988. Primatologist Sue Boinski watched an adult male white-faced capuchin beat a fer-de-lance snake to death with a dead branch. The black-striped capuchin was the first non-ape primate for which routine tool use was documented in the wild; individuals were observed cracking nuts by placing them on a stone anvil and hitting them with another large stone. In Thailand and Myanmar, crab-eating macaques use stone tools to open nuts, oysters and other bivalves, and various types of sea snails. Chacma baboons use stones as weapons; stoning by these baboons is done from the rocky walls of the canyon where they sleep and retreat to when they are threatened. Stones are lifted with one hand and dropped over the side whereupon they tumble down the side of the cliff or fall directly to the canyon floor. Although they have not been observed to use tools in the wild, lemurs in controlled settings have been shown to be capable of understanding the functional properties of the objects they had been trained to use as tools, performing as well as tool-using haplorhines. Soon after her initial discovery of tool use, Goodall observed other chimpanzees picking up leafy twigs, stripping off the leaves and using the stems to fish for insects. This change of a leafy twig into a tool was a major discovery. Prior to this, scientists thought that only humans manufactured and used tools, and that this ability was what separated humans from other animals. Chimpanzees have also been observed making "sponges" out of leaves and moss that suck up water. Sumatran orangutans have been observed making and using tools. They will break off a tree branch that is about 30 cm long, snap off the twigs, fray one end and then use the stick to dig in tree holes for termites. In the wild, mandrills have been observed to clean their ears with modified tools. Scientists filmed a large male mandrill at
Chester Zoo Chester Zoo is a zoo in Upton-by-Chester, Cheshire, England. Chester Zoo was opened in 1931 by George Mottershead and his family. The zoo is one of the UK's largest zoos at and the zoo has a total land holding of approximately . Chester Zoo ...
(UK) stripping down a twig, apparently to make it narrower, and then using the modified stick to scrape dirt from underneath its toenails. Captive gorillas have made a variety of tools.


Ecology

Human Humans (''Homo sapiens'') or modern humans are the most common and widespread species of primate, and the last surviving species of the genus ''Homo''. They are Hominidae, great apes characterized by their Prehistory of nakedness and clothing ...
s are the most adaptable primate species, despite having a low or narrow tolerance for many of the earth's extreme environments. Currently the species is present in all eight biogeographical realms, although their presence in the Antarctic realm is very limited to research stations and annually there is a population decline in the winter months of this realm. Non-human primates primarily live in the
tropical The tropics are the regions of Earth surrounding the equator, where the sun may shine directly overhead. This contrasts with the temperate or polar regions of Earth, where the Sun can never be directly overhead. This is because of Earth's ax ...
latitudes of Africa, Asia, and the Americas. Species that live outside of the tropics include the
Japanese macaque The Japanese macaque (''Macaca fuscata''), also known as the snow monkey, is a terrestrial Old World monkey species that is native to Japan. Colloquially, they are referred to as "snow monkeys" because some live in areas where snow covers the g ...
which lives in the Japanese islands of
Honshū , historically known as , is the largest of the four main islands of Japan. It lies between the Pacific Ocean (east) and the Sea of Japan (west). It is the seventh-largest island in the world, and the second-most populous after the Indonesian ...
and
Hokkaido is the list of islands of Japan by area, second-largest island of Japan and comprises the largest and northernmost prefectures of Japan, prefecture, making up its own list of regions of Japan, region. The Tsugaru Strait separates Hokkaidō fr ...
; the
Barbary macaque The Barbary macaque (''Macaca sylvanus''), also known as Barbary ape, is a macaque species native to the Atlas Mountains of Algeria, Tunisia and Morocco, along with a small introduced population in Gibraltar. It is the type species of the genus ' ...
which lives in North Africa and several species of langur which live in China. Primates tend to live in
tropical rainforest Tropical rainforests are dense and warm rainforests with high rainfall typically found between 10° north and south of the Equator. They are a subset of the tropical forest biome that occurs roughly within the 28° latitudes (in the torrid zo ...
s but are also found in
temperate forest A temperate forest is a forest found between the tropical and boreal regions, located in the temperate zone. It is the second largest terrestrial biome, covering 25% of the world's forest area, only behind the boreal forest, which covers about 3 ...
s,
savanna A savanna or savannah is a mixed woodland-grassland (i.e. grassy woodland) biome and ecosystem characterised by the trees being sufficiently widely spaced so that the canopy does not close. The open canopy allows sufficient light to reach th ...
s,
desert A desert is a landscape where little precipitation occurs and, consequently, living conditions create unique biomes and ecosystems. The lack of vegetation exposes the unprotected surface of the ground to denudation. About one-third of the la ...
s, mountains and coastal areas. The number of primate species within tropical areas has been shown to be positively
correlated In statistics, correlation or dependence is any statistical relationship, whether causal or not, between two random variables or bivariate data. Although in the broadest sense, "correlation" may indicate any type of association, in statistic ...
to the amount of rainfall and the amount of rain forest area. Accounting for 25% to 40% of the fruit-eating animals (by
weight In science and engineering, the weight of an object is a quantity associated with the gravitational force exerted on the object by other objects in its environment, although there is some variation and debate as to the exact definition. Some sta ...
) within tropical rainforests, primates play an important ecological role by dispersing seeds of many tree species. Primate habitats span a range of altitudes: the black snub-nosed monkey has been found living in the
Hengduan Mountains The Hengduan Mountains () are a group of mountain ranges in southwest China, southwest China that connect the southeast portions of the Tibetan Plateau with the Yunnan–Guizhou Plateau. The Hengduan Mountains are primarily large north-south ...
at altitudes of 4,700 meters (15,400 ft), the
mountain gorilla The mountain gorilla (''Gorilla beringei beringei'') is one of the two subspecies of the eastern gorilla. It is listed as endangered by the IUCN . There are two populations: One is found in the Virunga Mountains, Virunga volcanic mountains of C ...
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 in the
Ethiopian Highlands The Ethiopian Highlands (also called the Abyssinian Highlands) is a rugged mass of mountains in Ethiopia in Northeast Africa. It forms the largest continuous area of its elevation in the continent, with little of its surface falling below , whil ...
. Some species interact with aquatic environments and may swim or even dive, including the proboscis monkey, De Brazza's monkey and Allen's swamp monkey. Some primates, such as the
rhesus macaque The rhesus macaque (''Macaca mulatta''), colloquially rhesus monkey, is a species of Old World monkey. There are between six and nine recognised subspecies split between two groups, the Chinese-derived and the Indian-derived. Generally brown or g ...
and gray langurs, can exploit human-modified environments and even live in cities.


Interactions between humans and other primates


Disease transmission

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 ''Poxviridae'' is a family of double-stranded DNA viruses. Vertebrates and arthropods serve as natural hosts. The family contains 22 genera that are assigned to two subfamilies: ''Chordopoxvirinae'' and ''Entomopoxvirinae''. ''Entomopoxvirinae'' ...
'',
measles Measles (probably from Middle Dutch or Middle High German ''masel(e)'', meaning "blemish, blood blister") is a highly contagious, Vaccine-preventable diseases, vaccine-preventable infectious disease caused by Measles morbillivirus, measles v ...
,
ebola Ebola, also known as Ebola virus disease (EVD) and Ebola hemorrhagic fever (EHF), is a viral hemorrhagic fever in humans and other primates, caused by ebolaviruses. Symptoms typically start anywhere between two days and three weeks after in ...
,
rabies Rabies is a viral disease that causes encephalitis in humans and other mammals. It was historically referred to as hydrophobia ("fear of water") because its victims panic when offered liquids to drink. Early symptoms can include fever and abn ...
, 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.


Legal and social status

Only
human Humans (''Homo sapiens'') or modern humans are the most common and widespread species of primate, and the last surviving species of the genus ''Homo''. They are Hominidae, great apes characterized by their Prehistory of nakedness and clothing ...
s are recognized as persons and protected in law by the
United Nations The United Nations (UN) is the Earth, global intergovernmental organization established by the signing of the Charter of the United Nations, UN Charter on 26 June 1945 with the stated purpose of maintaining international peace and internationa ...
Universal Declaration of Human Rights The Universal Declaration of Human Rights (UDHR) is an international document adopted by the United Nations General Assembly that enshrines the Human rights, rights and freedoms of all human beings. Drafted by a UN Drafting of the Universal D ...
. 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
chimpanzee The chimpanzee (; ''Pan troglodytes''), also simply known as the chimp, is a species of Hominidae, great ape native to the forests and savannahs of tropical Africa. It has four confirmed subspecies and a fifth proposed one. When its close rel ...
s,
orangutan Orangutans are great apes native to the rainforests of Indonesia and Malaysia. They are now found only in parts of Borneo and Sumatra, but during the Pleistocene they ranged throughout Southeast Asia and South China. Classified in the genus ...
s and
gorilla Gorillas are primarily herbivorous, terrestrial great apes that inhabit the tropical forests of equatorial Africa. The genus ''Gorilla'' is divided into two species: the eastern gorilla and the western gorilla, and either four or five su ...
s are not to be used for animal experiments. Many species of NHP are kept as pets by humans. The Allied Effort to Save Other Primates (AESOP) estimates that around 15,000 NHPs live as exotic pets in the United States. 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$ The United States dollar (Currency symbol, symbol: Dollar sign, $; ISO 4217, currency code: USD) is the official currency of the United States and International use of the U.S. dollar, several other countries. The Coinage Act of 1792 introdu ...
3000 for monkeys to $30,000 for apes. Primates are used as
model organism A model organism is a non-human species that is extensively studied to understand particular biological phenomena, with the expectation that discoveries made in the model organism will provide insight into the workings of other organisms. Mo ...
s in laboratories and have been used in space missions. They serve as
service animal Service animals are Working animal, working animals that have been trained to perform tasks that assist disabled people. Service animals may also be referred to as assistance animals or helper animals depending on the country and the animal's fu ...
s for disabled humans. Capuchin monkeys can be trained to assist
quadriplegic Tetraplegia, also known as quadriplegia, is defined as the dysfunction or loss of Motor control, motor and/or Sense, sensory function in the Cervical vertebrae, cervical area of the spinal cord. A loss of motor function can present as either weak ...
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 towards conservation, education and research. GAP does not insist that all NHPs should be released from zoos, primarily because captive-born primates lack the knowledge and experience to survive in the wild if released.


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 Neuroscience is the scientific study of the nervous system (the brain, spinal cord, and peripheral nervous system), its functions, and its disorders. It is a multidisciplinary science that combines physiology, anatomy, molecular biology, ...
,
ophthalmology Ophthalmology (, ) is the branch of medicine that deals with the diagnosis, treatment, and surgery of eye diseases and disorders. An ophthalmologist is a physician who undergoes subspecialty training in medical and surgical eye care. Following a ...
studies, and toxicity studies.
Rhesus macaque The rhesus macaque (''Macaca mulatta''), colloquially rhesus monkey, is a species of Old World monkey. There are between six and nine recognised subspecies split between two groups, the Chinese-derived and the Indian-derived. Generally brown or g ...
s are often used, as are other macaques, African green monkeys, chimpanzees,
baboon Baboons are primates comprising the biology, genus ''Papio'', one of the 23 genera of Old World monkeys, in the family Cercopithecidae. There are six species of baboon: the hamadryas baboon, the Guinea baboon, the olive baboon, the yellow ba ...
s, 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 The European Union (EU) is a supranational union, supranational political union, political and economic union of Member state of the European Union, member states that are Geography of the European Union, located primarily in Europe. The u ...
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.


Extinction threats

The
International Union for Conservation of Nature The International Union for Conservation of Nature (IUCN) is an international organization working in the field of nature conservation and sustainable use of natural resources. Founded in 1948, IUCN has become the global authority on the stat ...
(IUCN) lists more than a third of primates as critically endangered or vulnerable. About 60% of primate species are threatened with extinction, including: 87% of species in Madagascar, 73% in Asia, 37% in Africa, and 36% in South and Central America. Additionally, 75% of primate species have decreasing populations. 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 Deforestation or forest clearance is the removal and destruction of a forest or stand of trees from land that is then converted to non-forest use. Deforestation can involve conversion of forest land to farms, ranches, or urban use. Ab ...
, 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 Bushmeat is meat from wildlife species that are Hunting, hunted for human consumption. Bushmeat represents a primary source of animal protein and a cash-earning commodity in poor and rural communities of humid tropical forest regions of the worl ...
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 Madagascar, officially the Republic of Madagascar, is an island country that includes the island of Madagascar and numerous smaller peripheral islands. Lying off the southeastern coast of Africa, it is the world's List of islands by area, f ...
, 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 Hinduism () is an Hypernymy and hyponymy, umbrella term for a range of Indian religions, Indian List of religions and spiritual traditions#Indian religions, religious and spiritual traditions (Sampradaya, ''sampradaya''s) that are unified ...
,
Buddhism Buddhism, also known as Buddhadharma and Dharmavinaya, is an Indian religion and List of philosophies, philosophical tradition based on Pre-sectarian Buddhism, teachings attributed to the Buddha, a wandering teacher who lived in the 6th or ...
, and
Islam Islam is an Abrahamic religions, Abrahamic monotheistic religion based on the Quran, and the teachings of Muhammad. Adherents of Islam are called Muslims, who are estimated to number Islam by country, 2 billion worldwide and are the world ...
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 The rhesus macaque (''Macaca mulatta''), colloquially rhesus monkey, is a species of Old World monkey. There are between six and nine recognised subspecies split between two groups, the Chinese-derived and the Indian-derived. Generally brown or g ...
, a
model organism A model organism is a non-human species that is extensively studied to understand particular biological phenomena, with the expectation that discoveries made in the model organism will provide insight into the workings of other organisms. Mo ...
, 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, seven 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.


See also

* Arboreal theory * Great Ape Project * Human evolution * International Primate Day * List of primates * List of fossil primates * Monkey Day * Primatology


Footnotes


References


Literature cited

* * * *


Further reading

*


External links


Primate Info Net


at Animal Diversity Web
Primate Research Institute
Kyoto University *
EUPRIM-Net: European Primate Network

PrimateImages: Natural History Collection

Interactive views
of various primate skeletons at eSkeletons.org (associated with the University of Texas at Austin)
Tree of Life web project
{{Featured article Extant Thanetian first appearances Primates, Taxa named by Carl Linnaeus