Primate

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(Redirected from
Non human primates
)

Primates
Temporal range: 65.9–0 
Ma
Early Paleocene to Present
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Mirorder: Primatomorpha
Order: Primates
Linnaeus, 1758[1]
Suborders

sister:

Dermoptera

Range and density of non-human primates.
Synonyms

cladistically including crown primates[2]
)

Primates are the members of a diverse

simians (monkeys and apes). Primates arose 85–55 million years ago first from small terrestrial mammals, which adapted to living in the trees of tropical forests: many primate characteristics represent adaptations to life in this challenging environment, including large brains, visual acuity, color vision, a shoulder girdle allowing a large degree of movement in the shoulder joint, and dexterous hands. Primates range in size from Madame Berthe's mouse lemur, which weighs 30 g (1 oz), to the eastern gorilla, weighing over 200 kg (440 lb). There are 376–524 species 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 brains (relative to body size) compared to other mammals, as well as an increased reliance on visual acuity at the expense of the sense of

bipedalism) and modified walking on four limbs (knuckle-walking
).

Primates are among the most social of animals, forming pairs or family groups, uni-male harems, and multi-male/multi-female groups. Non-human primates have at least four types of

use 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

, 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 is derived from Old French or French primat, from a noun use of Latin primat-, from primus ('prime, first rank').[3] The name was given by Carl Linnaeus because he thought this the "highest" order of animals.[4] 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.[5][6]

Sir

paraphyletic, or rather they do not include all the descendants of a common ancestor.[8]

In contrast with Clark's methodology, modern classifications typically identify (or name) only those groupings that are

monophyletic; that is, such a named group includes all the descendants of the group's common ancestor.[9]

The cladogram below shows one possible classification sequence of the living primates:[10][11] groups that use common (traditional) names are shown on the right.

Primatomorpha

Dermoptera

Primates
Strepsirrhini
Lemuriformes[a]

lemurs (superfamily Lemuroidea)

lorises and allies (superfamily Lorisoidea)

Haplorhini
Tarsiiformes

tarsiers (superfamily Tarsioidea)

Simiiformes

New World monkeys (parvorder Platyrrhini)

Catarrhini

Old World monkeys (superfamily Cercopithecoidea)

Hominoidea

gibbons (family Hylobatidae)

Hominidae

orangutans (subfamily Ponginae)

Homininae

gorillas (tribe Gorillini)

Hominini

humans (g. Homo)

chimpanzees, bonobos (g. Pan)

prosimians
monkeys
lesser apes
great apes

All groups with scientific names are clades, 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 apes
    , 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

palaeontologist Benton wrote, "The apes, Hominoidea, today include the gibbons and orangutan ... the gorilla and chimpanzee ... and humans";[12]
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.

As of 2021[update], 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".[13]

Classification of living primates

A 1927 drawing of chimpanzees, a gibbon (top right) and two orangutans (center and bottom center): The chimpanzee in the upper left is brachiating; the orangutan at the bottom center is knuckle-walking.
Homo sapiens
is the only living primate species that is fully bipedal
Nilgiri langur (Trachypithecus johnii), an Old World monkey

A list of the families of the living primates is given below, together with one possible classification into ranks between order and family.[1][10][14][15] Other classifications are also used. For example, an alternative classification of the living Strepsirrhini divides them into two infraorders, Lemuriformes and Lorisiformes.[16]

  • Order Primates
    • Suborder Strepsirrhini: lemurs, galagos and lorisids
      • Infraorder Lemuriformes[a]
        • Superfamily
          Lemuroidea
          • Family Cheirogaleidae: dwarf lemurs and mouse-lemurs (41 species)
          • Family Daubentoniidae: aye-aye (1 species)
          • Family Lemuridae: ring-tailed lemur and allies (21 species)
          • Family
            Lepilemuridae
            : sportive lemurs (26 species)
          • Family Indriidae: woolly lemurs and allies (19 species)
        • Superfamily Lorisoidea
          • Family Lorisidae: lorisids (16 species)
          • Family
            Galagidae
            : galagos (23 species)
    • Suborder Haplorhini: tarsiers, monkeys and apes
      • Infraorder Tarsiiformes
      • Infraorder
        Simiiformes
        (or Anthropoidea)
        • Parvorder
          Platyrrhini
          : New World monkeys
          • Family Callitrichidae: marmosets and tamarins (49 species)
          • Family Cebidae: capuchins and squirrel monkeys (29 species)
          • Family
            Aotidae
            : night or owl monkeys (douroucoulis) (11 species)
          • Family Pitheciidae: titis, sakis and uakaris (56 species)
          • Family Atelidae: howler, spider, woolly spider and woolly monkeys (26 species)
        • Parvorder Catarrhini
          • Superfamily Cercopithecoidea
          • Superfamily
            Hominoidea
            • Family
              Hylobatidae
              : gibbons or "lesser apes" (20 species)
            • Family Hominidae: great apes, including humans (8 species)

Order Primates was established by

Ruminantia),[21]
but these new taxa were not accepted.

Before Anderson and Jones introduced the classification of Strepsirrhini and Haplorhini in 1984,

Anthropoidea.[24] Prosimii included all of the prosimians: Strepsirrhini plus the tarsiers. Anthropoidea contained all of the simians
.

Phylogeny and genetics

Euarchontoglires  
Glires 

Rodentia (rodents)

Lagomorpha (rabbits, hares, pikas)

 Euarchonta 

Scandentia (treeshrews)

Primatomorpha

Dermoptera (colugos)

Primates

Plesiadapiformes

crown primates

Order Primates is part of the clade Euarchontoglires, which is nested within the clade Eutheria of Class Mammalia. Recent molecular genetic research on primates, colugos, and treeshrews has shown that the two species of colugos are more closely related to primates than to treeshrews,[25] even though treeshrews were at one time considered primates.[26] These three orders make up the clade Euarchonta. The combination of this clade with the clade Glires (composed of Rodentia and Lagomorpha) forms the clade Euarchontoglires. Variously, both Euarchonta and Euarchontoglires are ranked as superorders. Some scientists consider Dermoptera to be a suborder of Primates and use the suborder Euprimates for the "true" primates.[27]

Evolutionary history

The primate lineage is thought to go back at least near the

mya).[28][29][30][31][32] The earliest possible primate/proto-primate may be Purgatorius, which dates back to Early Paleocene of North America ~66mya.[33][34] The oldest known primates from the fossil record date to the Late Paleocene of Africa, c.57 mya (Altiatlasius)[35] or the Paleocene-Eocene transition in the northern continents, c. 55 mya (Cantius, Donrussellia, Altanius, Plesiadapis and Teilhardina).[36][37][33] 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.[38][39][40]

By modern

omomyids.[43][44] 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.[45] 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,[44] while adapiforms survived until about 10 mya.[46]

According to genetic studies, the lemurs of Madagascar diverged from the lorisoids approximately 75 mya.

rafting event between 50 and 80 mya.[42][47][48] Other colonization options have been suggested, such as multiple colonizations from Africa and India,[43] but none are supported by the genetic and molecular evidence.[42]

Common brown lemur, a Strepsirrhine primate

Until recently, the aye-aye has been difficult to place within Strepsirrhini.[1] 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.[42]

Suborder

eosimiids, developed in Asia, but became extinct millions of years ago.[54]

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.

island hopping, facilitated by Atlantic Ocean ridges and a lowered sea level.[43] Alternatively, a single rafting event may explain this transoceanic colonization. Due to continental drift, the Atlantic Ocean was not nearly as wide at the time as it is today.[43] Research suggests that a small 1 kg (2.2 lb) primate could have survived 13 days on a raft of vegetation.[56]
Given estimated current and wind speeds, this would have provided enough time to make the voyage between the continents.

Emperor tamarin, a New World monkey

Apes and monkeys spread from Africa into Europe and Asia starting in the Miocene.[57] 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.[44] Old World monkeys disappeared from Europe about 1.8 mya.[58] Molecular and fossil studies generally show that modern humans originated in Africa 100,000–200,000 years ago.[59]

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;[11] the author, Colin Groves, increased that number to 376 for his contribution to the third edition of Mammal Species of the World (MSW3).[1] However, publications since the taxonomy in MSW3 was compiled in 2003 have pushed the number to 522 species, or 708 including subspecies.[60]

Hybrids

Primate hybrids usually arise in captivity,[61] but there have also been examples in the wild.[62][63] 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.[62] 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.[64]

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.[65][66][67][68][69]

Anatomy and physiology

Head

Primate skulls showing postorbital bar, and increasing brain sizes

The primate skull has a large, domed

hominids is exceptional; they have the most acute vision known among all vertebrates, with the exception of certain species of predatory birds.[74][75]

Primates have forward-facing eyes on the front of the skull;

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.[76]

An 1893 drawing of the hands and feet of various primates

Primates show an evolutionary trend towards a reduced

hypocone
) evolved in early primate history, while the cusp of the corresponding primitive lower molar (paraconid) was lost. Prosimians are distinguished by their immobilized upper lips, the moist tip of their noses and forward-facing lower front teeth.

Body

Vervet hindfoot showing fingerprint ridges on the sole

Primates generally have five digits on each limb (

toilet-claws, which they use for grooming.[70]

The primate

woolly spider, woolly monkeys; and in capuchins.[80][81] Male primates have a low-hanging penis and testes descended into a scrotum.[82][78]

Sexual dimorphism

Distinct sexual size dimorphism can be seen between the male and female mountain gorilla.

pelage and skin color.[87] The dimorphism can be attributed to and affected by different factors, including mating system,[88] size,[88] habitat and diet.[89]

Comparative analyses have generated a more complete understanding of the relationship between

hominids probably had greater dimorphism than any living primate.[90]

Locomotion

Diademed sifaka, a lemur that is a vertical clinger and leaper

Primate species move by

avahis and indris), sportive lemurs, and all tarsiers.[92] Other prosimians are arboreal quadrupeds and climbers. Some are also terrestrial quadrupeds, while some are leapers. Most monkeys are both arboreal and terrestrial quadrupeds and climbers. Gibbons, muriquis and spider monkeys all brachiate extensively,[58] with gibbons sometimes doing so in remarkably acrobatic fashion. Woolly monkeys also brachiate at times.[93] Orangutans use a similar form of locomotion called quadramanous climbing, in which they use their arms and legs to carry their heavy bodies through the trees.[58] Chimpanzees and gorillas knuckle walk,[58] 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.[94]

Vision

The tapetum lucidum of a northern greater galago, typical of prosimians, reflects the light of the photographer's flash.

The evolution of color vision in primates is unique among most eutherian mammals. While the remote vertebrate ancestors of the primates possessed three color vision (trichromaticism), the nocturnal, warm-blooded, mammalian ancestors lost one of three cones in the retina during the Mesozoic era. Fish, reptiles and birds are therefore trichromatic or tetrachromatic, while all mammals, with the exception of some primates and marsupials,[95] are dichromats or monochromats (totally color blind).[78] Nocturnal primates, such as the night monkeys and bush babies, are often monochromatic. Catarrhines are routinely trichromatic due to a gene duplication of the red-green opsin gene at the base of their lineage, 30 to 40 million years ago.[78][96] Platyrrhines, on the other hand, are trichromatic in a few cases only.[97] Specifically, individual females must be heterozygous for two alleles of the opsin gene (red and green) located on the same locus of the X chromosome.[78] 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.[78]

Like catarrhines, howler monkeys (a family of platyrrhines) show routine trichromatism that has been traced to an evolutionarily recent gene duplication.[98] Howler monkeys are one of the most specialized leaf-eaters of the New World monkeys; fruits are not a major part of their diets,[93] 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.[97]

Behavior

Social systems

Richard Wrangham stated that social systems of primates are best classified by the amount of movement by females occurring between groups.[99] 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.[99] This organization can be seen in chimpanzees, where the males, who are typically related, will cooperate in defense of the group's territory.[100] Evidence of this social system has also been found among Neanderthal remains in Spain[101] and in remains of Australopithecus and Paranthropus robustus groups in southern Africa.[102][103] Among New World Monkeys, spider monkeys and muriquis use this system.[104]
A social huddle of ring-tailed lemurs. The two individuals on the right exposing their white ventral surface are sunning themselves.

Other systems are known to occur as well. For example, with

callitrichid monkeys also use this system.[58]

The transfer of females or males from their native group is likely an adaptation for avoiding inbreeding.[109] 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.[109][110] This effect of inbreeding on infant mortality is probably largely a result of increased expression of deleterious recessive alleles (see Inbreeding depression).

Chimpanzees are social great apes.

Primatologist

fission-fusion societies in chimpanzees.[111] 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,[112] spider monkeys[93] and the bonobo.[112] The gelada has a similar social structure in which many smaller groups come together to form temporary herds of up to 600 monkeys.[112] Humans 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.[113]

These social systems are affected by three main ecological factors: distribution of resources, group size, and predation.[114] Within a social group there is a balance between cooperation and competition. Cooperative behaviors in many primates species include social grooming (removing skin parasites and cleaning wounds), food sharing, and collective defense against predators or of a territory. Aggressive behaviors often signal competition for food, sleeping sites or mates. Aggression is also used in establishing dominance hierarchies.[114][115]

In November 2023, scientists reported, for the first time, evidence that groups of primates, particularly bonobos, are capable of cooperating with each other.[116][117]

Interspecific associations

Several species of primates are known to associate in the wild. Some of these associations have been extensively studied. In the

common chimpanzee.[119]

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.[112] Several of these species are preyed upon by the common chimpanzee.[120]

In South America, squirrel monkeys associate with capuchin monkeys.[121] This may have more to do with foraging benefits to the squirrel monkeys than anti-predation benefits.[121]

Communication

mark territories with pheromones, which are detected by the vomeronasal organ; this process forms a large part of the communication behavior of these primates.[72] In Old World monkeys and apes this ability is mostly vestigial, having regressed as trichromatic eyes evolved to become the main sensory organ.[122] Primates also use vocalizations, gestures, and facial expressions to convey psychological state.[123][124] 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.[125] Hand and arm gestures are also important forms of communication for great apes and a single gesture can have multiple functions.[124]

Primates are a particularly vocal group of mammals.

alarm calls.[126] 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.[127] Male howler monkeys are among the loudest land mammals as their roars can be heard up to 4.8 km (3.0 mi), and relate to intergroup spacing, territorial protection and possibly mate-guarding.[128][129] Roars are produced by a modified larynx and enlarged hyoid bone which contains an air sac.[130] 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.[131] Male and female siamangs both possess inflatable pouches in the throat with which pair -bonds use to sing "duets" to each other.[132]

Many non-human primates have the vocal anatomy to produce human speech but lack the proper brain wiring.

great apes in general.[136]

Life history

A crab-eating macaque breastfeeding her baby

Primates have slower rates of development than other mammals.[58] 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.[58] In some species, infants are protected and transported by males in the group, particularly males who may be their fathers.[58] Other relatives of the infant, such as siblings and aunts, may participate in its care as well.[58] Most primate mothers cease ovulation while breastfeeding an infant; once the infant is weaned the mother can reproduce again.[58] This often leads to weaning conflict with infants who attempt to continue breastfeeding.[58]

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.[137] Promiscuity may also lessen the risk of infanticide since paternity becomes uncertain.[138]

Primates have a longer

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.[139][140] During the juvenile period, primates are more susceptible than adults to predation and starvation; they gain experience in feeding and avoiding predators during this time.[58] They learn social and fighting skills, often through playing.[58] Primates, especially females, have longer lifespans than other similarly sized mammals,[58] this may be partially due to their slower metabolisms.[141] Late in life, female catarrhine primates appear to undergo a cessation of reproductive function known as menopause; other groups are less studied.[142]

Diet and feeding

Leaf eating mantled guereza, a species of black-and-white colobus
A mouse lemur holds a cut piece of fruit in its hands and eats.

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.[143] Most primates include fruit in their diets to obtain easily digested nutrients including carbohydrates and lipids for energy.[58] Primates in the suborder Strepsirrhini (non-tarsier prosimians) are able to synthesize vitamin C, like most other mammals, while primates of the suborder Haplorhini (tarsiers, monkeys and apes) have lost this ability, and require the vitamin in their diet.[144]

Many primates have anatomical specializations that enable them to exploit particular foods, such as fruit, leaves, gum or insects.[58] 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.[58] 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.[58] The aye-aye combines rodent-like teeth with a long, thin middle finger to fill the same ecological niche as a woodpecker. It taps on trees to find insect larvae, then gnaws holes in the wood and inserts its elongated middle finger to pull the larvae out.[145] 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.[58] The gelada is the only primate species that feeds primarily on grass.[146]

Hunting

Portrait of a Dayak hunter in Borneo with a boar over his shoulder
Humans have traditionally hunted prey for subsistence.

obligate carnivorous primates, exclusively eating insects, crustaceans, small vertebrates and snakes (including venomous species).[147] Capuchin monkeys can exploit many different types of plant matter, including fruit, leaves, flowers, buds, nectar and seeds, but also eat insects and other invertebrates, bird eggs, and small vertebrates such as birds, lizards, squirrels and bats.[93]

The

omnivorous frugivore – the majority of its diet is fruit, but it supplements this with leaves, meat from small vertebrates, such as anomalures, flying squirrels and duikers,[150] and invertebrates.[151] In some instances, bonobos have been shown to consume lower-order primates.[152][153]

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.[154] It has been proposed that humans have used fire to prepare and cook food since the time of Homo erectus.[155] Around ten thousand years ago, humans developed agriculture,[156] which substantially altered their diet. This change in diet may also have altered human biology; with the spread of dairy farming providing a new and rich source of food, leading to the evolution of the ability to digest lactose in some adults.[157][158]

As prey

Predators of primates include various species of

antipredator adaptations including crypsis, 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.[159]

Intelligence and cognition

Primates have advanced cognitive abilities: some make tools and use them to acquire food and for social displays;

conspecifics;[165][166] and they can learn to use symbols and understand aspects of human language including some relational syntax and concepts of number and numerical sequence.[167][168][169] Research in primate cognition explores problem solving, memory, social interaction, a theory of mind, and numerical, spatial, and abstract concepts.[170] 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.[171][172] However, there is a great deal of variation in each group (e.g., among New World monkeys, both spider[171] and capuchin monkeys[172] have scored highly by some measures), as well as in the results of different studies.[171][172]

Tool use and manufacture

Chimpanzees using twigs to dip for ants
Crab-eating macaques with stone tools

In 1960, Jane Goodall observed a chimpanzee poking pieces of grass into a termite 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.[173] There are more limited reports of the closely related bonobo 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.[174] It has been reported that females, both chimpanzee and bonobo, use tools more avidly than males.[175] Orangutans in Borneo 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.[176] There are few reports of gorillas 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.[177]

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.[178] 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.[179] In Thailand and Myanmar, crab-eating macaques use stone tools to open nuts, oysters and other bivalves, and various types of sea snails.[180] 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.[181]

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.[182]

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.[173] Chimpanzees have also been observed making "sponges" out of leaves and moss that suck up water.[183] 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.[184][185] In the wild, mandrills have been observed to clean their ears with modified tools. Scientists filmed a large male mandrill at 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.[186] Captive gorillas have made a variety of tools.[187]

Ecology

Rhesus macaque at Agra Fort, India

Non-human primates primarily live in the

Honshū and Hokkaido; the Barbary macaque which lives in North Africa and several species of langur which live in China. Primates tend to live in tropical rainforests but are also found in temperate forests, savannas, deserts, mountains and coastal areas.[188] The number of primate species within tropical areas has been shown to be positively correlated to the amount of rainfall and the amount of rain forest area.[189] Accounting for 25% to 40% of the fruit-eating animals (by weight) within tropical rainforests, primates play an important ecological role by dispersing seeds of many tree species.[190]

Primate habitats span a range of altitudes: the

black snub-nosed monkey has been found living in the Hengduan Mountains at altitudes of 4,700 meters (15,400 ft),[191] the mountain gorilla can be found at 4,200 meters (13,200 ft) crossing the Virunga Mountains,[192] and the gelada has been found at elevations of up to 5,000 m (16,000 ft) in the Ethiopian Highlands.[193] 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.[194] Some primates, such as the rhesus macaque and gray langurs, can exploit human-modified environments and even live in cities.[112][195]

Interactions between humans and other primates

Disease transmission

Close interactions between humans and non-human primates (NHPs) can create pathways for the transmission of

Herpes B Virus), Poxviridae, measles, ebola, rabies, the Marburg virus and viral hepatitis can be transmitted to humans; in some cases the viruses produce potentially fatal diseases in both humans and non-human primates.[196]

Legal and social status

Slow lorises are popular in the exotic pet trade, which threatens wild populations.

Only humans are recognized as persons and protected in law by the United Nations Universal Declaration of Human Rights.[b] The legal status of NHPs, on the other hand, is the subject of much debate, with organizations such as the Great Ape Project (GAP) campaigning to award at least some of them legal rights.[198] In June 2008, Spain became the first country in the world to recognize the rights of some NHPs, when its parliament's cross-party environmental committee urged the country to comply with GAP's recommendations, which are that chimpanzees, orangutans and gorillas are not to be used for animal experiments.[199][200]

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.

United States – Mexico border, with prices ranging from US$3000 for monkeys to $30,000 for apes.[203]

Primates are used as

quadriplegic humans; their intelligence, memory, and manual dexterity make them ideal helpers.[205]

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.[206]

Role in scientific research

Sam, a rhesus macaque, was flown to the edge of space by NASA in the 1959 Little Joe 2 flight of Project Mercury.

Thousands of non-human primates are used around the world in research because of their psychological and physiological similarity to humans.[207][208] In particular, the brains and eyes of NHPs more closely parallel human anatomy than those of any other animals. NHPs are commonly used in preclinical trials, neuroscience, ophthalmology studies, and toxicity studies. Rhesus macaques are often used, as are other macaques, African green monkeys, chimpanzees, baboons, squirrel monkeys, and marmosets, both wild-caught and purpose-bred.[207][209]

In 2005, GAP reported that 1,280 of the 3,100 NHPs living in captivity in the United States were used for experiments.[198] In 2004, the European Union used around 10,000 NHPs in such experiments; in 2005 in Great Britain, 4,652 experiments were conducted on 3,115 NHPs.[210] 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.[211] 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.[212]

Extinction threats

Humans are known to hunt other primates for food, called bushmeat. Pictured are two men who have killed a number of silky sifaka and white-headed brown lemurs.

The International Union for Conservation of Nature (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.[213] Additionally, 75% of primate species have decreasing populations.[213] 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.[214][215]

White-fronted Brown Lemur (Eulemur albifrons) killed in northeast Madagascar for bushmeat.
White-headed lemur (Eulemur albifrons) killed in northeast Madagascar for bushmeat.

Common threats to primate species include

forest fragmentation, monkey drives (resulting from primate crop raiding),[216] 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.[217][218][219] More than 90% of primate species occur in tropical forests.[218][220] 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.[220] 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.[221]

The critically endangered silky sifaka

Primates with a large body size (over 5 kg) are at increased extinction risk due to their greater profitability to

drill are hunted at levels that far exceed sustainable levels.[223] This is due to their large body size, ease of transport and profitability per animal.[223] As farming encroaches on forest habitats, primates feed on the crops, causing the farmers large economic losses.[224] Primate crop raiding gives locals a negative impression of primates, hindering conservation efforts.[225]

critically endangered
Sumatran orangutan

In Asia, Hinduism, Buddhism, and Islam prohibit eating primate meat; however, primates are still hunted for food.[220] Some smaller traditional religions allow the consumption of primate meat.[226][227] The pet trade and traditional medicine also increase demand for illegal hunting.[202][228][229] The rhesus macaque, a model organism, was protected after excessive trapping threatened its numbers in the 1960s; the program was so effective that they are now viewed as a pest throughout their range.[219]

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.

edge effects such as farmland encroachment, lower levels of humidity and a change in plant life.[231][232] 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.[233][234]

There are 21 critically endangered primates, 7 of which have remained on the IUCN's "The World's 25 Most Endangered Primates" list since the year 2000: the silky sifaka, Delacour's langur, the white-headed langur, the gray-shanked douc, the Tonkin snub-nosed monkey, the Cross River gorilla and the Sumatran orangutan.[235] Miss Waldron's red colobus was recently declared extinct when no trace of the subspecies could be found from 1993 to 1999.[236] A few hunters have found and killed individuals since then, but the subspecies' prospects remain bleak.[237]

See also

Footnotes

  1. ^ a b Although the monophyletic relationship between lemurs and lorisoids is widely accepted, their clade name is not. The term "lemuriform" is used here because it derives from one popular taxonomy that clumps the clade of toothcombed primates into one infraorder and the extinct, non-toothcombed adapiforms into another, both within the suborder Strepsirrhini.[17][18] However, another popular alternative taxonomy places the lorisoids in their own infraorder, Lorisiformes.[16]
  2. ^ Article 6: Everyone has the right to recognition everywhere as a person before the law.[197]

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Literature cited

Further reading

  • David J. Chivers; Bernard A. Wood; Alan Bilsborough, eds. (1984). Food Acquisition and Processing in Primates. New York & London: Plenum Press. .

External links