Primate

This article is about the type of animal. For other uses, 2010.
see Primate (disambiguation).
Considered generalist mammals, primates exhibit a wide
range of characteristics. Some primates (including some
A primate ( i /pramet/ PRY-mayt) is a mammal of great apes and baboons) are primarily terrestrial rather
the order Primates (Latin: prime, rst rank).[2][3] than arboreal, but all species possess adaptations for
In taxonomy, primates include two distinct lineages, climbing trees. Locomotion techniques used include
strepsirrhines and haplorhines.[1] Primates arose from an- leaping from tree to tree, walking on two or four limbs,
cestors that lived in the trees of tropical forests; many pri- knuckle-walking, and swinging between branches of trees
mate characteristics represent adaptations to life in this (brachiation).
challenging three-dimensional environment. Most pri- Primates are characterized by large brains relative to
mate species remain at least partly arboreal.
other mammals, as well as an increased reliance on
With the exception of humans, who inhabit every continent except for Antarctica,[4] most primates live in tropical or subtropical regions of the Americas, Africa and
Asia.[5] They range in size from Madame Berthes mouse
lemur, which weighs only 30 g (1 oz), to the eastern
gorilla, weighing over 200 kg (440 lb). Based on fossil evidence, the earliest known true primates, represented by the genus Teilhardina, date to 55.8 million
years old.[6] An early close primate relative known from
abundant remains is the Late Paleocene Plesiadapis, c.
5558 million years old.[7] Molecular clock studies suggest that the primate branch may be even older, originating near the CretaceousPaleogene boundary or around
6374 mya.[8][9][10][11]

stereoscopic vision at the expense of smell, the dominant sensory system in most mammals. These features are
more developed in monkeys and apes and noticeably less
so in lorises and lemurs. Three-color vision has developed in some primates. Most also have opposable thumbs
and some have prehensile tails. Many species are sexually
dimorphic; dierences include body mass, canine tooth
size, and coloration. Primates have slower rates of development than other similarly sized mammals and reach
maturity later, but have longer lifespans. Depending on
the species, adults may live in solitude, in mated pairs, or
in groups of up to hundreds of members.

The order Primates was traditionally divided into two

main groupings: prosimians and anthropoids (simians).
Prosimians have characteristics more like those of the
earliest primates, and include the lemurs of Madagascar,
lorisoids, and tarsiers. Simians include monkeys, apes
and hominins. More recently, taxonomists have preferred
to split primates into the suborder Strepsirrhini, or wetnosed primates, consisting of non-tarsier prosimians, and
the suborder Haplorhini, or dry-nosed primates, consisting of tarsiers and the simians.

1 Historical and modern terminology

The relationships among the dierent 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.[12]

Simians are divided into two groups: catarrhine (narrownosed) monkeys and apes of Africa and southeastern Asia
and platyrrhine (at-nosed) or New World monkeys
of South and Middle America. Catarrhines consist of
Old World monkeys (such as baboons and macaques),
gibbons and great apes; New World monkeys include the
capuchin, howler and squirrel monkeys. Humans are the
only extant catarrhines to have spread successfully outside
of Africa, South Asia, and East Asia, although fossil evidence shows many other species were formerly present in
Europe. New primate species are still being discovered.
More than 25 species were taxonomically described in the
decade of the 2000s and eleven have been described since

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.[13]
Commonly used names for groups of primates such as
"prosimians", "monkeys", "lesser apes", and "great apes"
reect this methodology. According to our current understanding of the evolutionary history of the primates,
several of these groups are paraphyletic: a paraphyletic
group is one which does not include all the descendants
of the groups common ancestor.[14]
In contrast with Clarks methodology, modern classications typically identify (or name) only those group1

2 CLASSIFICATION OF LIVING PRIMATES

ings that are monophyletic; that is, such a named group

includes all the descendants of the groups common
ancestor.[15]

2 Classication of living primates

The cladogram below shows one possible classication

sequence of the living primates,[16][17] with groups that
use common (traditional) names are shown on the right.
All groups with scientic names are monophyletic (that
is, they are clades), and the sequence of scientic classication reects 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 in particular,
are paraphyletic because they exclude humans.
Thus, the members of the two sets of groups, and hence
names, do not match, which causes problems in relating
scientic 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
palaeontologist Benton wrote, The apes, Hominoidea,
today include the gibbons and orang-utan ... the gorilla
and chimpanzee ... and humans";[18] thereby Benton was
using apes to mean hominoids. In that case, the group
heretofore called apes must now be identied as the
non-human apes.

A 1927 drawing of chimpanzees, a gibbon (top right) and two

orangutans (center and bottom center): The chimp in the upper
left is brachiating; the orangutan at the bottom center is knucklewalking.

As of 2005, there is no consensus as to which methodology will rule, 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 will be
found in biological sources, often in the same work, and
sometimes by the same author. Thus, Benton denes
apes to include humans, then he repeatedly uses apelike 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.[19]

Infraorder Lemuriformes[lower-alpha 1]
Superfamily Lemuroidea
Family Cheirogaleidae:
dwarf
lemurs and mouse-lemurs (34
species)
Family Daubentoniidae: aye-aye
(one species)
Family Lemuridae:
ring-tailed
lemur and allies (21 species)
Family Lepilemuridae: sportive
lemurs (26 species)

A list of the families of the living primates is given below,

together with one possible classication into ranks between order and family.[1][16][20][21] Other classications
are also used. For example, an alternative classication of
the living Strepsirrhini divides them into two infraorders,
Lemuriformes and Lorisiformes.[22]
Order Primates
Suborder Strepsirrhini: lemurs, galagos and
lorisids

Nilgiri langur (Trachypithecus johnii), an Old World monkey

Homo sapiens is the only living primate species that is fully

bipedal

Family Indriidae: woolly lemurs

and allies (19 species)
Superfamily Lorisoidea
Family Lorisidae: lorisids (14
species)
Family Galagidae: galagos (19
species)

Family Atelidae: howler, spider,

woolly spider and woolly monkeys
(29 species)
Parvorder Catarrhini
Superfamily Cercopithecoidea
Family Cercopithecidae:
Old
World monkeys (138 species)
Superfamily Hominoidea
Family Hylobatidae: gibbons or
lesser apes (17 species)
Family Hominidae: great apes, including humans (7 species)

Suborder Haplorhini: tarsiers, monkeys and

Order Primates was established by Carl Linnaeus in
apes
1758, in the tenth edition of his book Systema Nat Infraorder Tarsiiformes
urae,[25] for the genera Homo (humans), Simia (other
Family Tarsiidae:
tarsiers (11
apes and monkeys), Lemur (prosimians) and Vespertilio
species)
(bats). In the rst edition of the same book (1735),
Infraorder Simiiformes (or Anthro- he had used the name Anthropomorpha for Homo,
poidea)
Simia and Bradypus (sloths).[26] In 1839, Henri Marie
Parvorder Platyrrhini: New World Ducrotay de Blainville, following Linnaeus and imitatmonkeys
ing his nomenclature, established the orders Secundates
Family Callitrichidae: marmosets (including the suborders Chiroptera, Insectivora and
and tamarins (42 species)
Carnivora), Tertiates (or Glires) and Quaternates (includ Family Cebidae: capuchins and ing Gravigrada, Pachydermata and Ruminantia),[27] but
these new taxa were not accepted.
squirrel monkeys (14 species)
Family Aotidae: night or owl monkeys (douroucoulis) (11 species)
Family Pitheciidae: titis, sakis and
uakaris (43 species)

Before Anderson and Jones introduced the classication of Strepsirrhini and Haplorhini in 1984,[28] (followed by McKenna and Bells 1997 work Classication
of Mammals: Above the species level),[29] the Pri-

3 EVOLUTIONARY HISTORY

mates were divided into two superfamilies: Prosimii and

Anthropoidea.[30] Prosimii included all of the prosimians:
Strepsirrhini plus the tarsiers. Anthropoidea contained all
of the simians.

Evolutionary history

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,[31] even though treeshrews were at
one time considered primates.[32] 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.[33]

3.1

Evolution

Human timeline
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Human-like
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Nakalipithecus
Ouranopithecus
Sahelanthropus
Orrorin
Ardipithecus
Australopithecus

Homo habilis

Homo erectus
Neanderthal
Homo sapiens

Earlier apes

Earliest bipedal

Early bipedal

Earliest stone tools

Earliest exit
from Africa

Earliest re use

Earliest cooking

Earliest clothes

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Evolution

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Axis scale: millions of years.

Also see: Life timeline & Nature timeline

Life timeline
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Single-celled
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Earliest humans

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Eukaryotes
Multicellular
life
Land life
Dinosaurs
Mammals
Flowers

Earliest Earth (4540)

Earliest water

Earliest life
(4100)

LHB meteorites

Earliest oxygen

Atmospheric oxygen

Oxygen Crisis

Earliest sexual reproduction

Cambrian explosion

H
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Axis scale: millions of years.
Also see: Human timeline & Nature timeline

Further information: Evolution of primates

The primate lineage is thought to go back at least 65 million years ago (mya),[34] even though the oldest known
primates from the fossil record date to the Late Paleocene of Africa (Altiatlasius)[35] or the Paleocene-Eocene
transition in the northern continents, c. 55 mya (Cantius,
Donrussellia, Altanius, and Teilhardina).[36] Other studies, including molecular clock studies, have estimated the
origin of the primate branch to have been in the midCretaceous period, around 85 mya.[37][38][39]
By modern cladistic reckoning, the order Primates is
monophyletic. The suborder Strepsirrhini, the "wetnosed" primates, is generally thought to have split o
from the primitive primate line about 63 mya,[40] although earlier dates are also supported.[41] The seven
strepsirrhine families are the ve related lemur families

3 EVOLUTIONARY HISTORY

and the two remaining families that include the lorisids

and the galagos.[1][20] Older classication schemes wrap
Lepilemuridae into Lemuridae and Galagidae into
Lorisidae, yielding a four-one family distribution instead
of ve-two as presented here.[1] During the Eocene,
most of the northern continents were dominated by two
groups, the adapiforms and the omomyids.[42][43] The former are considered members of Strepsirrhini, but did
not have a toothcomb like modern lemurs; recent analysis has demonstrated that Darwinius masillae ts into
this grouping.[44] 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,[43] while adapiforms survived until about
10 mya.[45]

vision, originating about 58 mya.[48][49] The earliest

known haplorhine skeleton, that of 55 MA old tarsierlike Archicebus, was found in central China,[50] supporting an already suspected Asian origin for the group.[51]
The infraorder Simiiformes (simian primates, consisting
of monkeys and apes) emerged about 40 mya,[43] possibly also in Asia; if so, they dispersed across the Tethys
Sea from Asia to Africa soon afterwards.[52] There are
two simian clades, both parvorders: Catarrhini, 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.[1] A third clade, which included the eosimiids, developed in Asia, but became extinct millions of years
ago.[53]

According to genetic studies, the lemurs of Madagascar diverged from the lorisoids approximately 75 mya.[41]
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.[41][46]
However, Madagascar split from Africa 160 mya and
from India 90 mya.[47] 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.[41][46][47] Other colonization options have been examined, such as multiple colonizations from Africa and India, but none are supported
by the genetic and molecular evidence.[42]

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.[7][54] The anthropoid primates possibly traversed
the Atlantic Ocean from Africa to South America during the Eocene by island hopping, facilitated by Atlantic
Ocean ridges and a lowered sea level.[42] 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.[42] Research suggests that a small 1 kg (2.2 lb) primate could
have survived 13 days on a raft of vegetation.[55] Given
estimated current and wind speeds, this would have provided enough time to make the voyage between the continents.

Common brown lemur, a strepsirrhine primate

Until recently, the aye-aye has been dicult 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 conrmed to be most closely related to the
other Malagasy lemurs, likely having descended from the
same ancestral population that colonized the island.[41]

Emperor tamarin, a New World monkey

Apes and monkeys spread from Africa into Europe and

Asia starting in the Miocene.[56] Soon after, the lorises
and tarsiers made the same journey. The rst hominin
fossils were discovered in northern Africa and date back
58 mya.[43] Old World monkeys disappeared from Eu[57]
Suborder Haplorhini, the simple-nosed or dry-nosed rope about 1.8 mya. Molecular and fossil studies genoriginated in Africa
primates, is composed of two sister clades.[1] Prosimian erally show that modern humans
[58]
100,000200,000
years
ago.
tarsiers in the family Tarsiidae (monotypic in its own
infraorder Tarsiiformes), represent the most basal di- Although primates are well studied in comparison to

7
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;[17] 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 424 species, or 658 including subspecies.[21]

tects the large brain, a distinguishing characteristic of this

group.[63] The endocranial volume (the volume within the
skull) is three times greater in humans than in the greatest nonhuman primate, reecting a larger brain size.[65]
The mean endocranial volume is 1,201 cubic centimeters
in humans, 469 cm3 in gorillas, 400 cm3 in chimpanzees
and 397 cm3 in orangutans.[65] The primary evolutionary
trend of primates has been the elaboration of the brain,
in particular the neocortex (a part of the cerebral cortex), which is involved with sensory perception, generation of motor commands, spatial reasoning, conscious
thought and, in humans, language.[5] While other mam3.2 Hybrids
mals rely heavily on their sense of smell, the arboreal life
of primates has led to a tactile, visually dominant sensory
Primate hybrids usually arise in captivity,[59] but there system,[5] a reduction in the olfactory region of the brain
have also been examples in the wild.[60][61] Hybridization and increasingly complex social behavior.[66]
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.[60] Intergeneric hybridizations, hybrids of
dierent 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.[62]

Anatomy, physiology, and morphology

Primates have forward-facing eyes on the front of the

skull; binocular vision allows accurate distance perception, useful for the brachiating ancestors of all great
apes.[63] 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.[64]

An 1893 drawing of the hands and feet of various primates

Primates generally have ve digits on each limb

(pentadactyly), with keratin nails on the end of each nger and toe. The bottom sides of the hands and feet have
sensitive pads on the ngertips. Most have opposable
thumbs, a characteristic primate feature, though not limited to this order, (opossums and koalas, for example, also
have them).[63] Thumbs allow some species to use tools.
In primates, the combination of opposing thumbs, short
Primate crania with brain masses indicated
ngernails (rather than claws) and long, inward-closing
The primate skull has a large, domed cranium, which is ngers is a relict of the ancestral practice of gripping
particularly prominent in anthropoids. The cranium pro- branches, and has, in part, allowed some species to de-

4 ANATOMY, PHYSIOLOGY, AND MORPHOLOGY

velop brachiation (swinging by the arms from tree limb to

tree limb) as a signicant means of locomotion. Prosimians have clawlike nails on the second toe of each foot,
called toilet-claws, which they use for grooming.[63]

The tapetum lucidum of a northern greater galago, typical of

prosimians, reects the light of the photographers ash

Vervet hindfoot showing ngerprint ridges on the sole

The primate collar bone is retained as prominent element

of the pectoral girdle; this allows the shoulder joint broad
mobility.[67] Apes have more mobile shoulder joints and
arms due to the dorsal position of the scapula, broad
ribcages that are atter front-to-back, and a shorter, less
mobile spine compared to Old World monkeys (with
lower vertebrae greatly reduced, resulting in tail loss in
some species). Old World monkeys are unlike apes in
that most have tails. New World atelids, including the
howler, spider, woolly spider and woolly monkeys, and
New World capuchins have prehensile tails.[68][69] Male
primates typically have a pendulous penis and scrotal
testes.[70][71]
Primates show an evolutionary trend towards a reduced
snout.[67] 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.[66]
In New World monkeys, the nostrils face sideways; in
Old World monkeys, they face downwards.[66] Dental pattern in primates vary considerably; although some have
lost most of their incisors, all retain at least one lower
incisor.[66] In most strepsirrhines, the lower incisors and
canines form a toothcomb, which is used in grooming
and sometimes foraging,[66][71] and the rst lower premolar is shaped like a canine.[71] Old World monkeys
have eight premolars, compared with 12 in New World
monkeys.[66] The Old World species are divided into apes
and monkeys depending on the number of cusps on their
molars; apes have ve, Old World monkeys have four,[66]
although humans may have four or ve.[72] The main hominid molar cusp (hypocone) evolved in early primate
history, while the cusp of the corresponding primitive
lower molar (paraconid) was lost. Prosimians are distinguished by their immobilized upper lips, the moist tip of
their noses and forward-facing lower front teeth.

(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,[73]
are dichromats or monochromats (totally color blind).[71]
Nocturnal primates, such as the night monkeys and bush
babies, are often monochromatic. Catarrhines are routinely trichromatic due to a gene duplication of the redgreen opsin gene at the base of their lineage, 30 to 40
million years ago.[71][74] Platyrrhines, on the other hand,
are trichromatic in a few cases only.[75] Specically, individual females must be heterozygous for two alleles of
the opsin gene (red and green) located on the same locus
of the X chromosome.[71] 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.[71]
Like catarrhines, howler monkeys (a family of
platyrrhines) show routine trichromatism that has
been traced to an evolutionarily recent gene duplication.[76] Howler monkeys are one of the most specialized
leaf-eaters of the New World monkeys; fruits are not
a major part of their diets,[77] 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.[75]

4.1 Sexual dimorphism

Main article: Sexual dimorphism in non-human primates

Sexual dimorphism is often exhibited in simians, though
to a greater degree in Old World species (apes and some
monkeys) than New World species. Recent studies involve comparing DNA to examine both the variation in
The evolution of color vision in primates is unique among the expression of the dimorphism among primates and
most eutherian mammals. While the remote vertebrate the fundamental causes of sexual dimorphism. Primates
ancestors of the primates possessed three color vision usually have dimorphism in body mass[78][79] and canine

Distinct sexual size dimorphism can be seen between the female

and two male hamadryas baboons.

tooth size[80][81] along with pelage and skin color.[82] The

dimorphism can be attributed to and aected by dierent
factors, including mating system,[83] size,[83] habitat and
diet.[84]
Comparative analyses have generated a more complete
understanding of the relationship between sexual selection, natural selection, and mating systems in primates.
Studies have shown that dimorphism is the product of
changes in both male and female traits.[85] Ontogenetic
scaling, where relative extension of a common growth
trajectory occurs, may give some insight into the relationship between sexual dimorphism and growth patterns.[86]
Some evidence from the fossil record suggests that there
was convergent evolution of dimorphism, and some extinct hominids probably had greater dimorphism than any
living primate.[85]

4.2

Locomotion

lemurs, and all tarsiers.[87] 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,[57] with gibbons sometimes doing so in
remarkably acrobatic fashion. Woolly monkeys also
brachiate at times.[77] 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.[57] Chimpanzees and gorillas knuckle
walk,[57] 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.[88]

5 Behavior
5.1 Social systems
Primates are among the most social of animals, forming pairs or family groups, uni-male harems, and multimale/multi-female groups.[89] Richard Wrangham stated
that social systems of non-human primates are best classied by the amount of movement by females occurring
between groups.[90] 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 inuential in social behavior.
The groups formed are generally quite small. This
organization can be seen in chimpanzees, where
the males, who are typically related, will cooperate in defense of the groups territory. Among New
World Monkeys, spider monkeys and muriquis use
this system.[91]

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

Primate species move by brachiation, bipedalism,

leaping, arboreal and terrestrial quadrupedalism,
climbing, knuckle-walking or by a combination of these
methods. Several prosimians are primarily vertical A social huddle of ring-tailed lemurs. The two individuals on the
clingers and leapers. These include many bushbabies, right exposing their white ventral surface are sunning themselves.
all indriids (i.e., sifakas, avahis and indris), sportive

10
Male transfer systems while the females remain in
their natal groups, the males will emigrate as adolescents. Polygynous and multi-male societies are
classed in this category. Group sizes are usually
larger. This system is common among the ringtailed lemur, capuchin monkeys and cercopithecine
monkeys.[57]
Monogamous species a malefemale bond, sometimes accompanied by a juvenile ospring. There is
shared responsibility of parental care and territorial
defense. The ospring leaves the parents territory
during adolescence. Gibbons essentially use this
system, although monogamy in this context does
not necessarily mean absolute sexual delity.[92]

5 BEHAVIOR
group returns at night to sleep as a group. This social structure can also be observed in the hamadryas
baboon,[97] spider monkeys[77] and the bonobo.[97] The
gelada has a similar social structure in which many
smaller groups come together to form temporary herds
of up to 600 monkeys.[97]
These social systems are aected by three main ecological factors: distribution of resources, group size, and
predation.[98] Within a social group there is a balance between cooperation and competition. Cooperative behaviors include social grooming (removing skin parasites and
cleaning wounds), food sharing, and collective defense
against predators or of a territory. Aggressive behaviors
often signal competition for food, sleeping sites or mates.
Aggression is also used in establishing dominance hierarchies.[98][99]

Solitary species often males who defend territories that include the home ranges of several females. This type of organization is found in the
prosimians such as the slow loris. Orangutans do 5.2 Interspecic associations
not defend their territory but eectively have this
organization.[93]
Several species of primates are known to associate in
the wild. Some of these associations have been extenOther systems are known to occur as well. For exam- sively studied. In the Tai Forest of Africa several species
ple, with howler monkeys both the males and females coordinate anti-predator behavior. These include the
typically transfer from their natal group on reaching sex- Diana monkey, Campbells mona monkey, lesser spotual maturity, resulting in groups in which neither the nosed monkey, western red colobus, king colobus and
males nor females are typically related.[77] Some prosimi- sooty mangabey, which coordinate anti-predator alarm
ans, colobine monkeys and callitrichid monkeys use this calls.[100] Among the predators of these monkeys is the
system.[57]
common chimpanzee.[101]
The transfer of females or males from their native group is The red-tailed monkey associates with several species,
likely an adaptation for avoiding inbreeding.[94] An analy- including the western red colobus, blue monkey, Wolfs
sis of breeding records of captive primate colonies repre- mona monkey, mantled guereza, black crested mangabey
senting numerous dierent species indicates that the in- and Allens swamp monkey.[97] Several of these species
fant mortality of inbred young is generally higher than are preyed upon by the common chimpanzee.[102]
that of non-inbred young.[94][95] This eect of inbreeding
on infant mortality is probably largely a result of increased In South America,[103]squirrel monkeys associate with
This may have more to do with
expression of deleterious recessive alleles (see Inbreeding capuchin monkeys.
foraging
benets
to
the
squirrel monkeys than antidepression).
predation benets.[103]

5.3 Cognition and communication

Chimpanzees are social animals.

Primatologist Jane Goodall, who studied in the Gombe

Stream National Park, noted ssion-fusion societies in
chimpanzees.[96] There is ssion when the main group
splits up to forage during the day, then fusion when the A pair of black howler monkeys vocalizing.

5.4

Life history

Main article: Primate cognition

Further information: Great ape language
Primates have advanced cognitive abilities: some make
tools and use them to acquire food and for social displays;[104][105] some have sophisticated hunting strategies requiring cooperation, inuence and
rank;[106] they are status conscious, manipulative and
capable of deception;[107] they can recognise kin and
conspecics;[108][109] and they can learn to use symbols
and understand aspects of human language including
some relational syntax and concepts of number and numerical sequence.[110][111][112] Research in primate cognition explores problem solving, memory, social interaction, a theory of mind, and numerical, spatial, and abstract concepts.[113] Comparative studies show a trend
towards higher intelligence going from prosimians to
New World monkeys to Old World monkeys, and signicantly higher average cognitive abilities in the great
apes.[114][115] However, there is a great deal of variation
in each group (e.g., among New World monkeys, both
spider[114] and capuchin monkeys[115] have scored highly
by some measures), as well as in the results of dierent
studies.[114][115]
Lemurs, lorises, tarsiers, and New World monkeys rely
on olfactory signals for many aspects of social and reproductive behavior.[5] Specialized glands are used to mark
territories with pheromones, which are detected by the
vomeronasal organ; this process forms a large part of
the communication behavior of these primates.[5] In Old
World monkeys and apes this ability is mostly vestigial,
having regressed as trichromatic eyes evolved to become
the main sensory organ.[116] Primates also use vocalizations, gestures, and facial expressions to convey psychological state.[117] Facial musculature is complex in primates, particularly in monkeys and apes. Like humans,
chimpanzees can distinguish the faces of familiar and unfamiliar individuals.[118]
The Philippine tarsier, has a high-frequency limit of auditory sensitivity of approximately 91 kHz with a dominant
frequency of 70 kHz. Such values are among the highest
recorded for any terrestrial mammal, and a relatively extreme example of ultrasonic communication. For Philippine tarsiers, ultrasonic vocalizations might represent a
private channel of communication that subverts detection
by predators, prey and competitors, enhances energetic
eciency, or improves detection against low-frequency
background noise.[119] Male howler monkeys are among
the loudest land mammals and their roars can be heard
up to 4.8 km (3.0 mi).[120] Roars are produced by modied larynx and enlarged hyoid bone which contains an
air sac.[121] These calls are thought to relate to intergroup
spacing and territoral protection as well as possibly mateguarding.[122]

11

5.4 Life history

A crab-eating macaque breastfeeding her baby

Primates have slower rates of development than other

mammals.[57] 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.[57] In some
species, infants are protected and transported by males in
the group, particularly males who may be their fathers.[57]
Other relatives of the infant, such as siblings and aunts,
may participate in its care as well.[57] Most primate mothers cease ovulation while breastfeeding an infant; once the
infant is weaned the mother can reproduce again.[57] This
often leads to weaning conict with infants who attempt
to continue breastfeeding.[57]
Infanticide is common in polygynous species such as gray
langurs and gorillas. Adult males may kill dependent ospring that are not theirs so the female will return to estrus and thus they can sire ospring of their own. Social
monogamy in some species may have evolved to combat
this behavior.[123] Promiscuity may also lessen the risk of
infanticide since paternity becomes uncertain.[124]
Primates have a longer juvenile period between weaning and sexual maturity than other mammals of similar
size.[57] 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.[125][126] 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.[57] They learn social and ghting skills, often through playing.[57] Primates, especially
females, have longer lifespans than other similarly sized
mammals,[57] this may be partially due to their slower
metabolisms.[127] Late in life, female catarrhine primates
appear to undergo a cessation of reproductive function
known as menopause; other groups are less studied.[128]

12

5 BEHAVIOR
ecological niche as a woodpecker. It taps on trees to nd
insect larvae, then gnaws holes in the wood and inserts its
elongated middle nger to pull the larvae out.[131] 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.[57] The gelada is the only primate species
that feeds primarily on grass.[132]
5.5.1 Hunting

Leaf eating mantled guereza, a species of black-and-white

colobus

A tiny mouse lemur holds a cut piece of fruit in its hands and eats

5.5

Diet, feeding and hunting

Primates exploit a variety of food sources. It has been

said that many characteristics of modern primates, including humans, derive from an early ancestors practice
of taking most of its food from the tropical canopy.[129]
Most primates include fruit in their diets to obtain easily digested carbohydrates and lipids for energy.[57] However, they require other foods, such as leaves or insects,
for amino acids, vitamins and minerals. Primates in the
suborder Strepsirrhini (non-tarsier prosimians) are able
to synthesize vitamin C, like most other mammals, while
primates of the suborder Haplorrhini (tarsiers, monkeys
and apes) have lost this ability, and require the vitamin in
their diet.[130]
Many primates have anatomical specializations that enable them to exploit particular foods, such as fruit, leaves,
gum or insects.[57] 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 dicult to
digest.[57] 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.[57] The aye-aye combines rodentlike teeth with a long, thin middle nger to ll the same

Humans have traditionally hunted prey for subsistence.

Tarsiers are the only extant obligate carnivorous primates,

exclusively eating insects, crustaceans, small vertebrates
and snakes (including venomous species).[133] Capuchin
monkeys can exploit many dierent types of plant matter,
including fruit, leaves, owers, buds, nectar and seeds,
but also eat insects and other invertebrates, bird eggs,
and small vertebrates such as birds, lizards, squirrels and
bats.[77]
The common chimpanzee has a varied diet that includes
predation on other primate species, such as the western
red colobus monkey.[101] This sometimes involves tool
use. Common chimpanzees sharpen sticks to use as
weapons when hunting mammals. This is considered the
rst evidence of systematic use of weapons in a species
other than humans. Researchers documented 22 occasions where wild chimpanzees fashioned sticks into
spears to hunt lesser bush babies (Galago senegalensis).
In each case, a chimpanzee modied a branch by breaking
o one or both ends and, frequently using its teeth, sharp-

5.7

Tool use and manufacture

13

ened the stick. The tools, on average, were about 60 cm

(24 in) long and 1.1 cm (0.4 in) in circumference. The
chimpanzees then jabbed their spears into hollows in tree
trunks where bush babies slept. There was a single case
in which a chimpanzee successfully extracted a bush baby
with the tool. The bonobo is an omnivorous frugivore
the majority of its diet is fruit, but it supplements this with
leaves, meat from small vertebrates, such as anomalures,
ying squirrels and duikers,[134] and invertebrates.[135] In
some instances, bonobos have been shown to consume
lower-order primates.[136][137]

5.6

As prey

Predators of primates include various species of

carnivorans, birds of prey, reptiles and other primates.
Even gorillas have been recorded as prey. Predators of
primates have diverse hunting strategies and as such,
primates have evolved several dierent antipredator
adaptations including crypsis, alarm calls and mobbing.
Several species have separate alarm calls for dierent
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.[138] With their technology and
increased intelligence, modern humans are nearly free
of threats from predators and are themselves apex
predators.

5.7

Tool use and manufacture

Main article: Tool use by animals

5.7.1

Tool use

A Western lowland gorilla using a stick possibly to gauge the depth

of water

There are many reports of non-human primates using

tools, both in the wild or when captive. The use of tools
by primates is varied and includes hunting (mammals, invertebrates, sh), collecting honey, processing food (nuts,

Crab-eating macaques with stone tools

fruits, vegetables and seeds), collecting water, weapons

and shelter.
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 sh or
dip for termites.[139] 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,[140] It has been reported that both female
chimpanzees and bonobos use tools more avidly than
males.[141] Orangutans in Borneo scoop catsh out of
small ponds. Anthropologist Anne Russon saw several
animals on these forested islands learn on their own to jab
at catsh with sticks, so that the panicked prey would op
out of ponds and into the orangutans waiting hands[142]
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.[143]
The black-striped capuchin was the rst non-ape primate
for which 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.[144]
In Thailand and Myanmar, crab-eating macaques use
stone tools to open nuts, oysters and other bivalves, and
various types of sea snails.[145] 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 cli or fall directly to the canyon
oor.[146]

14

INTERACTIONS BETWEEN HUMANS AND OTHER PRIMATES

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.[147]
5.7.2

Tool manufacture

Tool manufacture is much rarer than simple tool use and

probably represents higher cognitive functioning. Soon
after her initial discovery of tool use, Goodall observed
other chimpanzees picking up leafy twigs, stripping o
the leaves and using the stems to sh 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.[139] Both bonobos and
chimpanzees have also been observed making sponges
out of leaves and moss that suck up water and are used
as grooming tools. Sumatran orangutans have been observed making and using tools. They will break o a
tree branch that is about 30 cm long, snap o the twigs,
fray one end and then use the stick to dig in tree holes
for termites.[148][149] In the wild, mandrills have been observed to clean their ears with modied tools. Scientists
lmed a large male mandrill at Chester Zoo (UK) stripping down a twig, apparently to make it narrower, and
then using the modied stick to scrape dirt from underneath its toenails.[150] Captive gorillas have made a variety
of tools.[151]

Rhesus macaque at Agra Fort, India

ing in the Hengduan Mountains at altitudes of 4,700 meters (15,400 ft),[154] the mountain gorilla can be found
at 4,200 meters (13,200 ft) crossing the Virunga Mountains,[155] and the gelada has been found at elevations
of up to 5,000 m (16,000 ft) in the Ethiopian Highlands. Although most species are generally shy of water,
a few are good swimmers and are comfortable in swamps
and watery areas, including the proboscis monkey, De
6 Habitat and distribution
Brazzas monkey and Allens swamp monkey, which has
developed small webbing between its ngers. Some priSee also: List of primates by population
mates, such as the rhesus macaque and gray langurs, can
Primates evolved from arboreal animals, and many exploit human-modied environments and even live in
species live most of their lives in trees. Most primate cities.[97][156]
species live in tropical rain forests. The number of primate species within tropical areas has been shown to be
positively correlated to the amount of rainfall and the
amount of rain forest area.[152] Accounting for 25% to 7 Interactions between humans
40% of the fruit-eating animals (by weight) within tropiand other primates
cal rainforests, primates play an important ecological role
by dispersing seeds of many tree species.[153]
Close interactions between humans and non-human priSome species are partially terrestrial, such as baboons
and patas monkeys, and a few species are fully terrestrial, such as geladas and humans. Non-human primates
live in a diverse number of forested habitats in the tropical latitudes of Africa, India, Southeast Asia, and South
America, including rainforests, mangrove forests, and
montane forests. There are some examples of non-human
primates that live outside of the tropics; the mountaindwelling Japanese macaque lives in the north of Honsh
where there is snow-cover eight months of the year; the
Barbary macaque lives in the Atlas Mountains of Algeria and Morocco. Primate habitats span a range of altitudes: the black snub-nosed monkey has been found liv-

mates (NHPs) can create pathways for the transmission of

zoonotic diseases. Viruses such as Herpesviridae (most
notably Herpes B Virus), Poxviridae, measles, ebola,
rabies, the Marburg virus and viral hepatitis can be transmitted to humans; in some cases the viruses produce potentially fatal diseases in both humans and non-human
primates.[157]

7.1 Legal and social status

Further information: Great ape personhood

7.2

Role in scientic research

15
Many species of NHP are kept as pets by humans, the
Allied Eort to Save Other Primates (AESOP) estimates that around 15,000 NHPs live as exotic pets in the
United States.[162] The expanding Chinese middle class
has increased demand for NHPs as exotic pets in recent
years.[163] Although NHP import for the pet trade was
banned in the U.S. in 1975, smuggling still occurs along
the United States Mexico border, with prices ranging
from US$3000 for monkeys to $30,000 for apes.[164]

Primates are used as model organisms in laboratories

and have been used in space missions.[165] They serve as
service animals for disabled humans. Capuchin monkeys
can be trained to assist quadriplegic humans; their intelligence, memory, and manual dexterity make them ideal
[166]
Slow lorises are popular in the exotic pet trade, which threatens helpers.
wild populations.
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, edOnly humans are recognized as persons and protected in ucation and research. GAP does not insist that all NHPs
law by the United Nations Universal Declaration of Hu- should be released from zoos, primarily because captiveman Rights.[lower-alpha 2] The legal status of NHPs, on the born primates lack the knowledge and experience to surother hand, is the subject of much debate, with organiza- vive in the wild if released.[167]
tions such as the Great Ape Project (GAP) campaigning
to award at least some of them legal rights.[159] In June
2008, Spain became the rst country in the world to recognize the rights of some NHPs, when its parliaments
cross-party environmental committee urged the country
to comply with GAPs recommendations, which are that
chimpanzees, bonobos, orangutans, and gorillas are not
7.2 Role in scientic research
to be used for animal experiments.[160][161]
Further information: Animal testing on non-human
primates and International trade in primates

Capuchin monkeys' manual dexterity is one reason they can assist

quadriplegic humans.

Thousands of non-human primates are used around the

world in research because of their psychological and
physiological similarity to humans.[168][169] 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 purposebred.[168][170] In 2005, GAP reported that 1,280 of the
3,100 NHPs living in captivity in the United States were
used for experiments.[159] 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.[171] 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.[172]
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 Unions review of
animal testing legislation.[173]

16

INTERACTIONS BETWEEN HUMANS AND OTHER PRIMATES

Humans are known to hunt other primates for food, so-called

bushmeat. Pictured are two men who have killed a number of
silky sifaka and white-headed brown lemurs.

7.3

Conservation

The International Union for Conservation of Nature

(IUCN) lists more than a third of primates as critically endangered or vulnerable. Trade is regulated, as
all species are listed by CITES in Appendix II, except
50 species and subspecies listed in Appendix I, which
gain full protection from trade.[174][175] Common threats
to primate species include deforestation, forest fragmentation, monkey drives (resulting from primate crop
raiding),[176] 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.[177][178][179] More than 90% of primate species
occur in tropical forests.[178][180] 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.[180] 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.[181]
Primates with a large body size (over 5 kg) are at increased extinction risk due to their greater protability to
poachers compared to smaller primates.[180] 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.[182] Data
for some African cities show that half of all protein consumed in urban areas comes from the bushmeat trade.[183]
Endangered primates such as guenons and the drill are
hunted at levels that far exceed sustainable levels.[183]
This is due to their large body size, ease of transport and
protability per animal.[183] As farming encroaches on
forest habitats, primates feed on the crops, causing the
farmers large economic losses.[184] Primate crop raiding
gives locals a negative impression of primates, hindering
conservation eorts.[185]

The critically endangered Sumatran orangutan

Madagascar, home to ve endemic primate families,

has experienced the greatest extinction of the recent
past; since human settlement 1,500 years ago, at least
eight classes and fteen of the larger species have become extinct due to hunting and habitat destruction.[5]
Among the primates wiped out were Archaeoindris (a
lemur larger than a silverback gorilla) and the families
Palaeopropithecidae and Archaeolemuridae.[5]
In Asia, Hinduism, Buddhism, and Islam prohibit eating primate meat; however, primates are still hunted
for food.[180] Some smaller traditional religions allow
the consumption of primate meat.[186][187] The pet trade
and traditional medicine also increase demand for illegal
hunting.[163][188][189] The rhesus macaque, a model organism, was protected after excessive trapping threatened its
numbers in the 1960s; the program was so eective that
they are now viewed as a pest throughout their range.[179]

The critically endangered silky sifaka

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.[177][190]
This increases the amount of forest vulnerable to edge ef-

17
fects such as farmland encroachment, lower levels of humidity and a change in plant life.[191][192] Movement restriction results in a greater amount of inbreeding, which
can cause deleterious eects leading to a population bottleneck, whereby a signicant percentage of the population is lost.[193][194]
There are 21 critically endangered primates, 7 of which
have remained on the IUCNs "The Worlds 25 Most Endangered Primates" list since the year 2000: the silky
sifaka, Delacours langur, the white-headed langur, the
gray-shanked douc, the Tonkin snub-nosed monkey, the
Cross River gorilla and the Sumatran orangutan.[195] Miss
Waldrons red colobus was recently declared extinct when
no trace of the subspecies could be found from 1993
to 1999.[196] A few hunters have found and killed individuals since then, but the subspecies prospects remain
bleak.[197]

See also
Arboreal theory
Human evolution
List of primates
List of fossil primates
Nest-building in primates
Primatology
Simian shelf

Footnotes

[1] 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.[23][24] However, another popular alternative taxonomy places the lorisoids in their own
infraorder, Lorisiformes.[22]
[2] Article 6: Everyone has the right to recognition everywhere as a person before the law.[158]

10

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0-801-88221-4.

[2] Primate.
Merriam-Webster Online Dictionary.
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11

Further reading

David J. Chivers; Bernard A. Wood; Alan Bilsborough, eds. (1984). Food Acquisition and Processing
in Primates. New York & London: Plenum Press.
ISBN 0-306-41701-4.

12

External links

Primate Info Net

Primates at Animal Diversity Web
Primate Research Institute, Kyoto University
High-Resolution Cytoarchitectural Primate Brain
Atlases
EUPRIM-Net: European Primate Network
PrimateImages: Natural History Collection
Tree of Life web project

26

13

13
13.1

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Images

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Images

File:Akha_cropped.png Source: https://upload.wikimedia.org/wikipedia/commons/c/c5/Akha_cropped.png License: CC BY-SA 2.0 de

Contributors: Extracted from File:Akha_couple.JPG, transferred from en.wikipedia Original artist: Manuel Jobi Weltenbummler84.
File:Brown_Lemur_in_Andasibe.jpg Source: https://upload.wikimedia.org/wikipedia/commons/b/b5/Brown_Lemur_in_Andasibe.
jpg License: CC BY-SA 2.0 Contributors: originally posted to Flickr as Brown Lemur in Andasibe Original artist: David Dennis
File:COLLECTIE_TROPENMUSEUM_Portret_van_een_Dajak_jager_op_Borneo_met_een_gevangen_zwijn_over_de_
schouder_TMnr_60043389.jpg Source: https://upload.wikimedia.org/wikipedia/commons/6/62/COLLECTIE_TROPENMUSEUM_
Portret_van_een_Dajak_jager_op_Borneo_met_een_gevangen_zwijn_over_de_schouder_TMnr_60043389.jpg License:
CC BYSA 3.0 Contributors: Tropenmuseum Original artist: Unknown<a href='//www.wikidata.org/wiki/Q4233718' title='wikidata:
Q4233718'><img
alt='wikidata:Q4233718'
src='https://upload.wikimedia.org/wikipedia/commons/thumb/f/ff/Wikidata-logo.
svg/20px-Wikidata-logo.svg.png'
width='20'
height='11'
srcset='https://upload.wikimedia.org/wikipedia/commons/thumb/f/ff/
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File:Cebus_albifrons_edit.jpg Source: https://upload.wikimedia.org/wikipedia/commons/6/64/Cebus_albifrons_edit.jpg License: CC
BY 2.5 Contributors: Own work Original artist: Photo by Whaldener Endo (Edit: Noise reduction and sharpening by User:Diliff.
File:Chimpanzee_male_white_background.jpg Source: https://upload.wikimedia.org/wikipedia/commons/e/e1/Chimpanzee_male_
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File:Chlorocebus_pygerythrus01.jpg Source: https://upload.wikimedia.org/wikipedia/commons/9/96/Chlorocebus_pygerythrus01.jpg
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File:Colubusmonkey.JPG Source: https://upload.wikimedia.org/wikipedia/commons/6/6b/Colubusmonkey.JPG License: CC-BY-SA3.0 Contributors: ? Original artist: ?
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File:Crab-eating_Macaque_nursing.jpg Source:
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nursing.jpg License: GFDL 1.2 Contributors: Own work Original artist: Muhammad Mahdi Karim Facebook
File:Diademed_ready_to_push_off.jpg Source: https://upload.wikimedia.org/wikipedia/commons/6/6c/Diademed_ready_to_push_
off.jpg License: Public domain Contributors: en:Image:Diademed ready to push o.jpg uploaded by en:User:Anlace on English Wikipedia
on 9th June 2006. Original artist: C. Michael Hogan - all rights released to Wikipedia
File:GarnettsGalago_CincinnatiZoo.jpg
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File:Gnome-mime-sound-openclipart.svg
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File:Gorilla_gorilla_white_background.jpg Source: https://upload.wikimedia.org/wikipedia/commons/f/f1/Gorilla_gorilla_white_
background.jpg License: CC BY-SA 2.5 Contributors: [1] Original artist: Pierre Fidenci
File:Gorilla_tool_use.png Source: https://upload.wikimedia.org/wikipedia/commons/c/c7/Gorilla_tool_use.png License: CC BY 2.5
Contributors: ? Original artist: See Source
File:Hamadryas_Baboon.jpg Source: https://upload.wikimedia.org/wikipedia/commons/e/e2/Hamadryas_Baboon.jpg License: CCBY-SA-3.0 Contributors: ? Original artist: ?
File:Howler_monkey.jpg Source: https://upload.wikimedia.org/wikipedia/commons/2/2f/Howler_monkey.jpg License: CC BY-SA 2.0
Contributors: howler monkees doing their thing Original artist: Steve from washington, dc, usa
File:Howler_monkey.ogg Source: https://upload.wikimedia.org/wikipedia/commons/b/b8/Howler_monkey.ogg License: CC BY 3.0
Contributors: Video recording Original artist: David O'Hara
File:Hunted_Silky_Sifakas.jpg Source: https://upload.wikimedia.org/wikipedia/commons/2/26/Hunted_Silky_Sifakas.jpg License:
Public domain Contributors: Own work Original artist: Simponafotsy
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File:Lock-green.svg Source: https://upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg License: CC0 Contributors: en:File:
Free-to-read_lock_75.svg Original artist: User:Trappist the monk
File:Macaca_fascicularis_aurea_stone_tools_-_journal.pone.0072872.g002.png Source: https://upload.wikimedia.org/wikipedia/
commons/8/87/Macaca_fascicularis_aurea_stone_tools_-_journal.pone.0072872.g002.png License: CC BY 2.5 Contributors: Haslam M,
Gumert MD, Biro D, Carvalho S, Malaivijitnond S (2013) Use-Wear Patterns on Wild Macaque Stone Tools Reveal Their Behavioural
History. PLoS ONE 8(8): e72872. doi:10.1371/journal.pone.0072872 Original artist: Haslam M, Gumert MD, Biro D, Carvalho S,
Malaivijitnond S (2013)
File:Macaque_India_3.jpg Source: https://upload.wikimedia.org/wikipedia/commons/9/96/Macaque_India_3.jpg License: CC BY-SA
3.0 Contributors: own work at http://www.retas.de/thomas/travel/india2007/index.html Original artist: Thomas Schoch
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background.jpg License: Public domain Contributors: ? Original artist: ?

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13

TEXT AND IMAGE SOURCES, CONTRIBUTORS, AND LICENSES

File:Microcebus_murinus_-Artis_Zoo,_Amsterdam,_Netherlands-8c.jpg Source:
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commons/e/ed/Microcebus_murinus_-Artis_Zoo%2C_Amsterdam%2C_Netherlands-8c.jpg License: CC BY 2.0 Contributors: originally
posted to Flickr as 2007-07-15-12h53m31.IMG_1008e Original artist: Arjan Haverkamp
File:NILGIRI_LANGUR_(Trachypithecus_johnii).jpg Source: https://upload.wikimedia.org/wikipedia/commons/0/0c/NILGIRI_
LANGUR_%28Trachypithecus_johnii%29.jpg License: CC BY-SA 4.0 Contributors: Own work Original artist: Anand2202
File:Nyticebus_white_background.jpg Source: https://upload.wikimedia.org/wikipedia/commons/1/11/Nyticebus_white_background.
jpg License: CC BY-SA 3.0 Contributors: File:Nycticebus pygmaeus 004.jpg Original artist: David Haring / Duke Lemur Center
File:Pongo_abelli_white_background.jpg Source:
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background.jpg License: CC BY-SA 3.0 Contributors: Own work Original artist: Ezhuttukari
File:PrimateFeet.jpg Source: https://upload.wikimedia.org/wikipedia/commons/b/bf/PrimateFeet.jpg License: Public domain Contributors: Lydekker, R. (1893). The Royal Natural History 1: 15. Original artist: Richard Lydekker (1849-1915)
File:Primate_skull_series_with_legend_cropped.png Source: https://upload.wikimedia.org/wikipedia/commons/d/db/Primate_skull_
series_with_legend_cropped.png License: CC BY 2.5 Contributors: Primate_skull_series_with_legend.png Molecular Insights into Human
Brain Evolution, Jane Bradbury, PLoS Biology Vol. 3, No. 3, e50 doi:10.1371/journal.pbio.0030050
Original artist: Primate_skull_series_with_legend.png: Christopher Walsh, Harvard Medical School
File:Primates-drawing.jpg Source: https://upload.wikimedia.org/wikipedia/commons/5/54/Primates-drawing.jpg License: Public domain Contributors: ? Original artist: ?
File:Red_Pencil_Icon.png Source: https://upload.wikimedia.org/wikipedia/commons/7/74/Red_Pencil_Icon.png License: CC0 Contributors: Own work Original artist: Peter coxhead
File:Ringstaartmakis_-_Ring-tailed_Lemur.jpg Source: https://upload.wikimedia.org/wikipedia/commons/7/79/Ringstaartmakis_-_
Ring-tailed_Lemur.jpg License: CC BY 2.5 Contributors: ? Original artist: ?
File:Sapajus_white_background.jpg Source: https://upload.wikimedia.org/wikipedia/commons/e/ef/Sapajus_white_background.jpg
License: CC BY-SA 2.0 de Contributors: Self-photographed Original artist: Masteraah
File:Silky_Sifaka_Pink_Face_Closeup.JPG Source:
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Face_Closeup.JPG License: Public domain Contributors: Own work Original artist: Simponafotsy
File:Slow_Loris_Female.jpg Source: https://upload.wikimedia.org/wikipedia/commons/9/9b/Slow_Loris_Female.jpg License: Public
domain Contributors: Own work (Original text: self-made) Original artist: Lionel Mauritson
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File:Tamarin_portrait_2_edit3.jpg Source: https://upload.wikimedia.org/wikipedia/commons/8/85/Tamarin_portrait_2_edit3.jpg License: GFDL Contributors: Own work Original artist: Brocken Inaglory, edited by Fir0002, edited by Brocken Inaglory
File:Tarsier_white_background.jpg Source: https://upload.wikimedia.org/wikipedia/commons/4/46/Tarsier_white_background.jpg
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apple.jpg License: CC BY-SA 2.0 Contributors: originally posted to Flickr as Whatcha got there?" Original artist: Matthew Hoelscher
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