Ape
Encyclopedia
Apes are Old World anthropoid
Anthropoid
Anthropoid may refer to:*Simian, monkeys and apes *Anthropoides, a genus of cranes*Operation Anthropoid, the codename for the assassination of SS-Obergruppenführer and Reichsprotektor of Bohemia and Moravia Reinhard Heydrich...

 mammal
Mammal
Mammals are members of a class of air-breathing vertebrate animals characterised by the possession of endothermy, hair, three middle ear bones, and mammary glands functional in mothers with young...

s, more specifically a clade of tailless catarrhine primate
Primate
A primate is a mammal of the order Primates , which contains prosimians and simians. Primates arose from ancestors that lived in the trees of tropical forests; many primate characteristics represent adaptations to life in this challenging three-dimensional environment...

s, belonging to the biological superfamily Hominoidea. The apes are native to Africa
Africa
Africa is the world's second largest and second most populous continent, after Asia. At about 30.2 million km² including adjacent islands, it covers 6% of the Earth's total surface area and 20.4% of the total land area...

 and South-east Asia, although in relatively recent times human
Human
Humans are the only living species in the Homo genus...

s have spread all over the world. Apes are the world's largest primates; the orangutan
Orangutan
Orangutans are the only exclusively Asian genus of extant great ape. The largest living arboreal animals, they have proportionally longer arms than the other, more terrestrial, great apes. They are among the most intelligent primates and use a variety of sophisticated tools, also making sleeping...

, an ape, is the world's largest living arboreal animal. Hominoids are traditionally forest dwellers, although chimpanzee
Chimpanzee
Chimpanzee, sometimes colloquially chimp, is the common name for the two extant species of ape in the genus Pan. The Congo River forms the boundary between the native habitat of the two species:...

s may range into savanna, and the extinct australopithecine
Australopithecine
The term australopithecine refers generally to any species in the related genera Australopithecus or Paranthropus. These species occurred in the Plio-Pleistocene era, and were bipedal and dentally similar to humans, but with a brain size not much larger than modern apes, lacking the...

s are famous for being savanna inhabitants, inferred from their morphology. Humans inhabit almost every terrestrial habitat.

Hominoidea contains two families
Family (biology)
In biological classification, family is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, class, order, genus, and species, with family fitting between order and genus. As for the other well-known ranks, there is the option of an immediately lower rank, indicated by the...

 of living (extant) species:
  • Hylobatidae
    Gibbon
    Gibbons are apes in the family Hylobatidae . The family is divided into four genera based on their diploid chromosome number: Hylobates , Hoolock , Nomascus , and Symphalangus . The extinct Bunopithecus sericus is a gibbon or gibbon-like ape which, until recently, was thought to be closely related...

     consists of four genera and sixteen species of gibbon
    Gibbon
    Gibbons are apes in the family Hylobatidae . The family is divided into four genera based on their diploid chromosome number: Hylobates , Hoolock , Nomascus , and Symphalangus . The extinct Bunopithecus sericus is a gibbon or gibbon-like ape which, until recently, was thought to be closely related...

    , including the lar gibbon
    Lar Gibbon
    The lar gibbon , also known as the white-handed gibbon, is a primate in the Hylobatidae or gibbon family. It is one of the better-known gibbons and is often seen in zoos.-Range:...

     and the siamang
    Siamang
    The siamang is a tailless, arboreal, black-furred gibbon native to the forests of Malaysia, Thailand, and Sumatra. The largest of the lesser apes, the siamang can be twice the size of other gibbons, reaching 1 m in height, and weighing up to 14 kg...

    . They are commonly referred to as lesser apes.
  • Hominidae
    Hominidae
    The Hominidae or include them .), as the term is used here, form a taxonomic family, including four extant genera: chimpanzees , gorillas , humans , and orangutans ....

     consists of orangutan
    Orangutan
    Orangutans are the only exclusively Asian genus of extant great ape. The largest living arboreal animals, they have proportionally longer arms than the other, more terrestrial, great apes. They are among the most intelligent primates and use a variety of sophisticated tools, also making sleeping...

    s, gorilla
    Gorilla
    Gorillas are the largest extant species of primates. They are ground-dwelling, predominantly herbivorous apes that inhabit the forests of central Africa. Gorillas are divided into two species and either four or five subspecies...

    s, chimpanzee
    Chimpanzee
    Chimpanzee, sometimes colloquially chimp, is the common name for the two extant species of ape in the genus Pan. The Congo River forms the boundary between the native habitat of the two species:...

    s, bonobo
    Bonobo
    The bonobo , Pan paniscus, previously called the pygmy chimpanzee and less often, the dwarf or gracile chimpanzee, is a great ape and one of the two species making up the genus Pan. The other species in genus Pan is Pan troglodytes, or the common chimpanzee...

    s and human
    Human
    Humans are the only living species in the Homo genus...

    s. Alternatively, the hominidae family are collectively described as the great apes.

Members of the superfamily are called hominoids (not to be confused with "hominids" or "hominins").

Some or all hominoids are also called "apes". However, the term "ape" is used in several different senses. It has been used as a synonym for "monkey" or for any tailless primate with a humanlike appearance. Thus the Barbary macaque
Barbary Macaque
The Barbary Macaque , or Common macaque, is a macaque with no tail. Found in the Atlas Mountains of Algeria and Morocco with a small population, of unknown origin, in Gibraltar, the Barbary Macaque is one of the best-known Old World monkey species. Besides humans, they are the only primates that...

, a kind of monkey, is popularly called the "Barbary ape" to indicate its lack of a tail. Biologists have used the term "ape" to mean a member of the superfamily Hominoidea other than humans, or more recently to mean all members of the superfamily Hominoidea, so that "ape" becomes another word for "hominoid". See also Primate: Historical and modern terminology.

Except for gorillas and humans, hominoids are agile climbers of trees. Their diet is best described as vegetarian or omnivorous, consisting of leaves, nuts, seeds and fruits, including grass seeds, and in most cases other animals, either hunted or scavenged (or farmed in the case of humans), along with anything else available and easily digested.

Most nonhuman hominoids are rare or endangered
Endangered species
An endangered species is a population of organisms which is at risk of becoming extinct because it is either few in numbers, or threatened by changing environmental or predation parameters...

. The chief threat to most of the endangered species is loss of tropical rainforest
Rainforest
Rainforests are forests characterized by high rainfall, with definitions based on a minimum normal annual rainfall of 1750-2000 mm...

 habitat, though some populations are further imperiled by hunting for bushmeat
Bushmeat
Bushmeat initially referred to the hunting of wild animals in West and Central Africa and is a calque from the French viande de brousse. Today the term is commonly used for meat of terrestrial wild animals, killed for subsistence or commercial purposes throughout the humid tropics of the Americas,...

.

Historical and modern terminology

"Ape", from Old English apa, is possibly an onomatopoetic imitation of animal chatter. The term has a history of rather imprecise usage. Its earliest meaning was a tailless (and therefore exceptionally human-like) nonhuman primate. The original usage of "ape" in English might have referred to the baboon
Baboon
Baboons are African and Arabian Old World monkeys belonging to the genus Papio, part of the subfamily Cercopithecinae. There are five species, which are some of the largest non-hominoid members of the primate order; only the mandrill and the drill are larger...

, an Old World monkey
Old World monkey
The Old World monkeys or Cercopithecidae are a group of primates, falling in the superfamily Cercopithecoidea in the clade Catarrhini. The Old World monkeys are native to Africa and Asia today, inhabiting a range of environments from tropical rain forest to savanna, shrubland and mountainous...

. Two tailless species of macaque
Macaque
The macaques constitute a genus of Old World monkeys of the subfamily Cercopithecinae. - Description :Aside from humans , the macaques are the most widespread primate genus, ranging from Japan to Afghanistan and, in the case of the barbary macaque, to North Africa...

 have common names including "ape": the Barbary ape of North Africa (introduced into Gibraltar
Gibraltar
Gibraltar is a British overseas territory located on the southern end of the Iberian Peninsula at the entrance of the Mediterranean. A peninsula with an area of , it has a northern border with Andalusia, Spain. The Rock of Gibraltar is the major landmark of the region...

), Macaca sylvanus, and the Sulawesi black ape or Celebes crested macaque
Celebes Crested Macaque
The Celebes crested macaque , also known as the crested black macaque, Sulawesi crested macaque, or the black ape, is an Old World monkey that lives in the northeast of the Indonesian island of Sulawesi as well as on smaller neighboring islands.-Description:Its skin and hairless face is, with the...

, M. nigra.

As zoological
Zoology
Zoology |zoölogy]]), is the branch of biology that relates to the animal kingdom, including the structure, embryology, evolution, classification, habits, and distribution of all animals, both living and extinct...

 knowledge developed, it became clear that taillessness occurred in a number of different and otherwise unrelated species
Species
In biology, a species is one of the basic units of biological classification and a taxonomic rank. A species is often defined as a group of organisms capable of interbreeding and producing fertile offspring. While in many cases this definition is adequate, more precise or differing measures are...

. The term "ape" was then used in two different senses, as shown in the 1910 Encyclopaedia Britannica
Encyclopædia Britannica Eleventh Edition
The Encyclopædia Britannica Eleventh Edition is a 29-volume reference work, an edition of the Encyclopædia Britannica. It was developed during the encyclopaedia's transition from a British to an American publication. Some of its articles were written by the best-known scholars of the time...

 entry. Either "ape" was still used for a tailless humanlike primate or it became a synonym for "monkey".

Sir Wilfrid Le Gros Clark
Wilfrid Le Gros Clark
Sir Wilfrid Edward Le Gros Clark was a British anatomist surgeon, primatologist and palaeoanthropologist, today best remembered for his contribution to the study of human evolution....

 was one of the primatologists who developed the idea that there were "trends" in primate evolution and that the living members of the order could be arranged in a series, leading through "monkeys" and "apes" to humans. Within this tradition, "ape" refers to all the members of the superfamily Hominoidea, except humans. Thus "apes" are a paraphyletic
Paraphyly
A group of taxa is said to be paraphyletic if the group consists of all the descendants of a hypothetical closest common ancestor minus one or more monophyletic groups of descendants...

 group, meaning that although all the species of apes descend from a common ancestor, the group does not include all the descendants of that ancestor, because humans are excluded. The diagram below shows the currently accepted evolutionary relationships of the Hominoidea, with the apes marked by a bracket.

The "apes" are traditionally divided further into the "lesser apes" and the "great apes":

In summary, there are three common uses of the term "ape": non-biologists may not distinguish between "monkeys" and "apes", or may use "ape" for any tailless monkey or nonhuman hominoid, whereas biologists traditionally used the term "ape" for all non-human hominoids as shown above.

In recent years biologists have generally preferred to use only monophyletic
Monophyly
In common cladistic usage, a monophyletic group is a taxon which forms a clade, meaning that it contains all the descendants of the possibly hypothetical closest common ancestor of the members of the group. The term is synonymous with the uncommon term holophyly...

 groups in classifications, that is only groups which include all the descendants of a common ancestor. The superfamily Hominoidea is one such group (or "clade
Clade
A clade is a group consisting of a species and all its descendants. In the terms of biological systematics, a clade is a single "branch" on the "tree of life". The idea that such a "natural group" of organisms should be grouped together and given a taxonomic name is central to biological...

"). Some then use the term "ape" to mean all the members of the superfamily Hominoidea. For example, in a 2005 book, Benton wrote "The apes, Hominoidea, today include the gibbons and orang-utan ... the gorilla and chimpanzee ... and humans". The group traditionally called "apes" by biologists is then called the "nonhuman apes".

See the section History of hominoid taxonomy below for a discussion of changes in scientific classification and terminology.

Biology

The "lesser apes" are the gibbon family, Hylobatidae with sixteen medium-sized species. Their major differentiating characteristic is their long arms, which they use to brachiate
Brachiation
Brachiation is a form of arboreal locomotion in which primates swing from tree limb to tree limb using only their arms.- Brachiators :...

 through the trees. As an evolutionary adaptation to this arboreal lifestyle, their wrists are ball and socket joint
Ball and socket joint
A ball and socket joint is a joint in which the distal bone is capable of motion around an indefinite number of axes, which have one common center...

s. The largest of the gibbons, the siamang
Siamang
The siamang is a tailless, arboreal, black-furred gibbon native to the forests of Malaysia, Thailand, and Sumatra. The largest of the lesser apes, the siamang can be twice the size of other gibbons, reaching 1 m in height, and weighing up to 14 kg...

, weighs up to 14 kg (31 lb). In comparison, the smallest "great ape" is the common chimpanzee
Common Chimpanzee
The common chimpanzee , also known as the robust chimpanzee, is a great ape. Colloquially, the common chimpanzee is often called the chimpanzee , though technically this term refers to both species in the genus Pan: the common chimpanzee and the closely related bonobo, formerly called the pygmy...

 at 40 to 65 kg (88.2 to 143.3 lb).

The "great apes" were formerly treated as the family Pongidae
Pongidae
Pongidae is a taxonomical family which is no longer in use.*Originally, until the last few years of the 20th century, Pongidae comprised all the extant species in the current family Hominidae, excluding Homo sapiens and sometimes including the family Hylobatidae .*Currently pongid may refer to the...

. As noted above, this definition makes the Pongidae paraphyletic, and does not show that orangutans, gorillas, chimpanzees and humans are all more closely related to one another than any of these four groups are to gibbons. Further, current evidence implies that humans share a common extinct ancestor with the chimpanzee line, from which we separated more recently than the gorilla line.

The superfamily Hominoidea falls within Catarrhini
Catarrhini
Catarrhini is one of the two subdivisions of the higher primates . It contains the Old World monkeys and the apes, which in turn are further divided into the lesser apes or gibbons and the great apes, consisting of the orangutans, gorillas, chimpanzees, bonobos, and humans...

, which also includes the Old World monkeys of Africa and Eurasia. Within this group, both families (Hylobatidae and Hominidae) can be distinguished from Old World monkeys by the number of cusps
Cusp (dentistry)
A cusp is an occlusal or incisal eminence on a tooth.Canine teeth, otherwise known as cuspids, each possess a single cusp, while premolars, otherwise known as bicuspids, possess two each. Molars normally possess either four or five cusps...

 on their molar
Molar (tooth)
Molars are the rearmost and most complicated kind of tooth in most mammals. In many mammals they grind food; hence the Latin name mola, "millstone"....

s (hominoids have five—the "Y-5" molar pattern, Old World monkeys have only four in a bilophodont pattern). Hominoids have more mobile shoulder joints and arms due to the dorsal position of the scapula
Scapula
In anatomy, the scapula , omo, or shoulder blade, is the bone that connects the humerus with the clavicle ....

, broad ribcages that are flatter front-to-back, and a shorter, less mobile spine compared to Old World monkeys, with caudal (tail) vertebrae greatly reduced, resulting in complete tail loss in living species. These are all anatomical adaptations to vertical hanging and swinging locomotion (brachiation
Brachiation
Brachiation is a form of arboreal locomotion in which primates swing from tree limb to tree limb using only their arms.- Brachiators :...

), as well as better balance in a bipedal pose. However, there are also primates in other families that lack tails, and at least one (the pig-tailed langur
Pig-tailed Langur
The pig-tailed langur is a large, rather heavily built Old World monkey, which is adapted to climbing with its long arms. Its fur is black-brown, and its hairless face is also black. It is the only monkey in the subfamily Colobinae to have a relatively short tail; the tail is only slightly furred...

) that has been known to walk significant distances bipedally. The front skull is characterised by its sinuses, fusion of the frontal bone and post-orbital constriction
Post-Orbital Constriction
In physical anthropology, post-orbital constriction is a narrowing of the cranium just behind the eye sockets , in primates — including primitive hominids. This constriction is very noticeable in non-human primates, slightly less so in Australopithecines, even less in Homo erectus and the...

.

Although the hominoid fossil record is far from complete, and the evidence is often fragmentary, there is enough to give a good outline of the evolutionary history of humans. The time of the split between humans and other living hominoids used to be thought to have occurred 15 to 20 million years ago. Some species occurring within that time period, such as Ramapithecus, used to be considered as hominins, and possible ancestors of humans. Later fossil finds indicated that Ramapithecus was more closely related to the orangutan, and new biochemical evidence indicated that the last common ancestor of humans and other hominins occurred between 5 and 10 million years ago, and probably in the lower end of that range.

Behaviour and cognition

Although there had been earlier studies, the scientific investigation of behaviour and cognition in nonhuman members of the superfamily Hominoidea expanded enormously during the latter half of the twentieth century. Major studies of behaviour in the field were completed on the three better-known "great apes", for example by Jane Goodall
Jane Goodall
Dame Jane Morris Goodall, DBE , is a British primatologist, ethologist, anthropologist, and UN Messenger of Peace. Considered to be the world's foremost expert on chimpanzees, Goodall is best known for her 45-year study of social and family interactions of wild chimpanzees in Gombe Stream National...

, Dian Fossey
Dian Fossey
Dian Fossey was an American zoologist who undertook an extensive study of gorilla groups over a period of 18 years. She studied them daily in the mountain forests of Rwanda, initially encouraged to work there by famous anthropologist Louis Leakey...

 and Birute Galdikas
Birute Galdikas
Birutė Marija Filomena Galdikas, OC , is a primatologist, conservationist, ethologist, and author of several books relating to the endangered orangutan, particularly the Bornean orangutan. Well known in the field of modern primatology, Galdikas is recognized as a leading authority on orangutans...

 (field work on gibbons and the bonobo
Bonobo
The bonobo , Pan paniscus, previously called the pygmy chimpanzee and less often, the dwarf or gracile chimpanzee, is a great ape and one of the two species making up the genus Pan. The other species in genus Pan is Pan troglodytes, or the common chimpanzee...

 is still relatively underdeveloped). These studies have shown that in their natural environments, the nonhuman hominoids show sharply varying social structure: gibbons are monogamous, territorial pair-bonders, orangutans are solitary, gorillas live in small troops with a single adult male leader, while chimpanzees live in larger troops with bonobos exhibiting promiscuous sexual behaviour. Their diets also vary; gorillas are foliovores while the others are all primarily frugivore
Frugivore
A frugivore is a fruit eater. It can be any type of herbivore or omnivore where fruit is a preferred food type. Because approximately 20% of all mammalian herbivores also eat fruit, frugivory is considered to be common among mammals. Since frugivores eat a lot of fruit they are highly dependent...

s, although the common chimpanzee does some hunting for meat. Foraging behaviour is correspondingly variable.

All the nonhuman hominoids are generally thought of as highly intelligent, and scientific study has broadly confirmed that they perform outstandingly well on a wide range of cognitive tests - though there is relatively little data on gibbon cognition. The early studies by Wolfgang Köhler
Wolfgang Köhler
Wolfgang Köhler was a German psychologist and phenomenologist who, like Max Wertheimer, and Kurt Koffka, contributed to the creation of Gestalt psychology.-Early life:...

 demonstrated exceptional problem-solving abilities in chimpanzees, which Köhler attributed to insight
Insight
Insight is the understanding of a specific cause and effect in a specific context. Insight can be used with several related meanings:*a piece of information...

. The use of tools
Tool use by animals
Tools are used by some animals, particularly primates, to perform simple tasks such as the acquisition of food, or grooming. Originally thought to be a skill only possessed by humans, tool use requires some level of intelligence. Primates have been observed exploiting sticks and stones to...

 has been repeatedly demonstrated; more recently, the manufacture of tools has been documented, both in the wild and in laboratory tests. Imitation
Imitation
Imitation is an advanced behavior whereby an individual observes and replicates another's. The word can be applied in many contexts, ranging from animal training to international politics.-Anthropology and social sciences:...

 is much more easily demonstrated in "great apes" than in other primate species. Almost all the studies in animal language acquisition have been completed with "great apes", and though there is continuing dispute as to whether they demonstrate real language abilities, there is no doubt that they involve significant feats of learning. Chimpanzees in different parts of Africa have developed tools that are used in food acquisition, demonstrating a form of animal culture.

Distinction from monkeys

Apes do not possess a tail, unlike most monkey
Monkey
A monkey is a primate, either an Old World monkey or a New World monkey. There are about 260 known living species of monkey. Many are arboreal, although there are species that live primarily on the ground, such as baboons. Monkeys are generally considered to be intelligent. Unlike apes, monkeys...

s. Monkeys are more likely to be in trees and use their tails for balance. Apes are considerably larger than monkeys, with the exception of gibbons
Gibbons
Notable people named Gibbons include:* Alan Gibbons, a British author* Beth Gibbons , a British singer* Billy Gibbons, a guitarist for ZZ Top* Carroll Gibbons , an American-born British bandleader...

, which are smaller than some monkeys. Apes are considered to be more intelligent than monkeys, which are considered to have more primitive brains. Unlike female monkeys which go through the estrous cycle
Estrous cycle
The estrous cycle comprises the recurring physiologic changes that are induced by reproductive hormones in most mammalian placental females. Estrous cycles start after puberty in sexually mature females and are interrupted by anestrous phases or pregnancies...

, great apes
Great Apes
Great Apes may refer to*Great apes, species in the biological family Hominidae, including humans, chimpanzees, gorillas, and orangutans*Great Apes , a 1997 novel by Will Self...

, including humans, go through a menstrual cycle.

History of hominoid taxonomy

The history of hominoid taxonomy is somewhat confusing and complex. The names of subgroups have changed their meaning over time as new evidence, from fossil discoveries and comparisons of anatomy and DNA sequences, has changed understanding of the relationships between hominoids. There has been a gradual demotion of humans from a special position in the taxonomy to being one branch among many. This history illustrates the growing influence of cladistics
Cladistics
Cladistics is a method of classifying species of organisms into groups called clades, which consist of an ancestor organism and all its descendants . For example, birds, dinosaurs, crocodiles, and all descendants of their most recent common ancestor form a clade...

 (the science of classifying living things by strict descent) on taxonomy.

As of 2006, there are eight extant genera
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...

 of hominoids. They are the four genera in the family Hominidae (Homo
Homo (genus)
Homo is the genus that includes modern humans and species closely related to them. The genus is estimated to be about 2.3 to 2.4 million years old, evolving from australopithecine ancestors with the appearance of Homo habilis....

– humans, Pan
Chimpanzee
Chimpanzee, sometimes colloquially chimp, is the common name for the two extant species of ape in the genus Pan. The Congo River forms the boundary between the native habitat of the two species:...

– chimpanzees and bonobos, Gorilla
Gorilla
Gorillas are the largest extant species of primates. They are ground-dwelling, predominantly herbivorous apes that inhabit the forests of central Africa. Gorillas are divided into two species and either four or five subspecies...

, and Pongo
Orangutan
Orangutans are the only exclusively Asian genus of extant great ape. The largest living arboreal animals, they have proportionally longer arms than the other, more terrestrial, great apes. They are among the most intelligent primates and use a variety of sophisticated tools, also making sleeping...

– orangutans), and the four genera in the family Hylobatidae or gibbon
Gibbon
Gibbons are apes in the family Hylobatidae . The family is divided into four genera based on their diploid chromosome number: Hylobates , Hoolock , Nomascus , and Symphalangus . The extinct Bunopithecus sericus is a gibbon or gibbon-like ape which, until recently, was thought to be closely related...

s (Hylobates
Hylobates
The genus Hylobates is one of the four genera of gibbons. It was once considered the only genus, but recently its subgenera have been elevated to the genus level. Hylobates remains the most speciose and widespread of gibbon genera, ranging from southern China to western and central Java...

, Hoolock, Nomascus
Nomascus
Nomascus is the second most speciose genus of gibbons . Originally this genus was a subgenus of Hylobates, and all individuals were considered one species, Hylobates concolor. Species within Nomascus are characterized by 52 chromosomes...

and Symphalangus). (The genus for the hoolock gibbon
Hoolock gibbon
The hoolock gibbons , also known as hoolocks, are two primate species from the family of the gibbons .Hoolocks are the second largest of the gibbons, after the Siamang. They reach a size of 60 to 90 cm and weigh 6 to 9 kg...

s was recently changed from Bunopithecus to Hoolock.)

In 1758, Carolus Linnaeus
Carolus Linnaeus
Carl Linnaeus , also known after his ennoblement as , was a Swedish botanist, physician, and zoologist, who laid the foundations for the modern scheme of binomial nomenclature. He is known as the father of modern taxonomy, and is also considered one of the fathers of modern ecology...

, relying on second- or third-hand accounts, placed a second species in Homo along with H. sapiens: Homo troglodytes ("cave-dwelling man"). It is not clear to which animal this name refers, as Linnaeus had no specimen to refer to, hence no precise description. Linnaeus named the orangutan Simia satyrus ("satyr monkey"). He placed the three genera Homo, Simia
Simia
In his Systema Naturae of 1758, Carolus Linnaeus divided the Order Primates into four genera: Homo, Simia, Lemur, and Vespertilio. His Vespertilio included all bats, and has since been moved from Primates to Chiroptera...

and Lemur in the order of Primates.

The troglodytes name was used for the chimpanzee by Blumenbach
Johann Friedrich Blumenbach
Johann Friedrich Blumenbach was a German physician, physiologist and anthropologist, one of the first to explore the study of mankind as an aspect of natural history, whose teachings in comparative anatomy were applied to classification of what he called human races, of which he determined...

 in 1775 but moved to the genus Simia. The orangutan was moved to the genus Pongo
Orangutan
Orangutans are the only exclusively Asian genus of extant great ape. The largest living arboreal animals, they have proportionally longer arms than the other, more terrestrial, great apes. They are among the most intelligent primates and use a variety of sophisticated tools, also making sleeping...

in 1799 by Lacépède.

Linnaeus's inclusion of humans in the primates with monkeys and apes was troubling for people who denied a close relationship between humans and the rest of the animal kingdom. Linnaeus's Lutheran archbishop had accused him of "impiety." In a letter to Johann Georg Gmelin
Johann Georg Gmelin
Johann Georg Gmelin was a German naturalist, botanist and geographer.- Early life and education :Gmelin was born in Tübingen, the son of an professor at the University of Tübingen. He was a gifted child and begun attending university lectures at the age of 14. In 1727, he graduated with a medical...

 dated 25 February 1747, Linnaeus wrote:
It is not pleasing to me that I must place humans among the primates, but man is intimately familiar with himself. Let's not quibble over words. It will be the same to me whatever name is applied. But I desperately seek from you and from the whole world a general difference between men and simians from the principles of Natural History. I certainly know of none. If only someone might tell me one! If I called man a simian or vice versa I would bring together all the theologians against me. Perhaps I ought to, in accordance with the law of Natural History.


Accordingly, Johann Friedrich Blumenbach
Johann Friedrich Blumenbach
Johann Friedrich Blumenbach was a German physician, physiologist and anthropologist, one of the first to explore the study of mankind as an aspect of natural history, whose teachings in comparative anatomy were applied to classification of what he called human races, of which he determined...

 in the first edition of his Manual of Natural History (1779), proposed that the primates be divided into the Quadrumana
Quadrumana
Quadrumana and Bimana form an obsolete division of the primates: the Quadrumana are primates with four hands , and the Bimana being those with two hands and two feet...

 (four-handed, i.e. apes and monkeys) and Bimana (two-handed, i.e. humans). This distinction was taken up by other naturalists, most notably Georges Cuvier
Georges Cuvier
Georges Chrétien Léopold Dagobert Cuvier or Jean Léopold Nicolas Frédéric Cuvier , known as Georges Cuvier, was a French naturalist and zoologist...

. Some elevated the distinction to the level of order
Order (biology)
In scientific classification used in biology, the order is# a taxonomic rank used in the classification of organisms. Other well-known ranks are life, domain, kingdom, phylum, class, family, genus, and species, with order fitting in between class and family...

.

However, the many affinities between humans and other primates — and especially the "great apes" — made it clear that the distinction made no scientific sense. Charles Darwin
Charles Darwin
Charles Robert Darwin FRS was an English naturalist. He established that all species of life have descended over time from common ancestry, and proposed the scientific theory that this branching pattern of evolution resulted from a process that he called natural selection.He published his theory...

 wrote, in The Descent of Man
The Descent of Man, and Selection in Relation to Sex
The Descent of Man, and Selection in Relation to Sex is a book on evolutionary theory by English naturalist Charles Darwin, first published in 1871. It was Darwin's second great book on evolutionary theory, following his 1859 work, On The Origin of Species. In The Descent of Man, Darwin applies...

:
The greater number of naturalists who have taken into consideration the whole structure of man, including his mental faculties, have followed Blumenbach and Cuvier, and have placed man in a separate Order, under the title of the Bimana, and therefore on an equality with the orders of the Quadrumana, Carnivora, etc. Recently many of our best naturalists have recurred to the view first propounded by Linnaeus, so remarkable for his sagacity, and have placed man in the same Order with the Quadrumana, under the title of the Primates. The justice of this conclusion will be admitted: for in the first place, we must bear in mind the comparative insignificance for classification of the great development of the brain in man, and that the strongly marked differences between the skulls of man and the Quadrumana (lately insisted upon by Bischoff, Aeby, and others) apparently follow from their differently developed brains. In the second place, we must remember that nearly all the other and more important differences between man and the Quadrumana are manifestly adaptive in their nature, and relate chiefly to the erect position of man; such as the structure of his hand, foot, and pelvis, the curvature of his spine, and the position of his head.


Changes in taxonomy

Until about 1960, the hominoids were usually divided into two families: humans and their extinct relatives in Hominidae
Hominidae
The Hominidae or include them .), as the term is used here, form a taxonomic family, including four extant genera: chimpanzees , gorillas , humans , and orangutans ....

, all other hominoids in Pongidae
Pongidae
Pongidae is a taxonomical family which is no longer in use.*Originally, until the last few years of the 20th century, Pongidae comprised all the extant species in the current family Hominidae, excluding Homo sapiens and sometimes including the family Hylobatidae .*Currently pongid may refer to the...

.
The 1960s saw the application of techniques from molecular biology
Molecular biology
Molecular biology is the branch of biology that deals with the molecular basis of biological activity. This field overlaps with other areas of biology and chemistry, particularly genetics and biochemistry...

 to primate taxonomy. Goodman used his 1964 immunological study of serum proteins to propose a division of the hominoids into three families, with the "great apes" in Pongidae and the "lesser apes" (gibbons) in Hylobatidae
Gibbon
Gibbons are apes in the family Hylobatidae . The family is divided into four genera based on their diploid chromosome number: Hylobates , Hoolock , Nomascus , and Symphalangus . The extinct Bunopithecus sericus is a gibbon or gibbon-like ape which, until recently, was thought to be closely related...

. The trichotomy of hominoid families, however, prompted scientists to ask which family speciated
Speciation
Speciation is the evolutionary process by which new biological species arise. The biologist Orator F. Cook seems to have been the first to coin the term 'speciation' for the splitting of lineages or 'cladogenesis,' as opposed to 'anagenesis' or 'phyletic evolution' occurring within lineages...

 first from the common hominoid ancestor.
Within the superfamily Hominoidea, gibbons are the outgroup: this means that the rest of the hominoids are more closely related to each other than any of them are to gibbons. This led to the placing of the "great apes" into the family Hominidae along with humans, by demoting the Pongidae to a subfamily; the Hominidae family now contained the subfamilies Homininae
Homininae
Homininae is a subfamily of Hominidae, which includes humans, gorillas and chimpanzees, and some extinct relatives; it comprises all those hominids, such as Australopithecus, that arose after the split from orangutans . Our family tree, which has 3 main branches leading to chimpanzees, humans and...

 and Ponginae
Ponginae
Ponginae is a subfamily in the hominidae family. It contains a number of genera, all but one extinct:*Pongo *†Gigantopithecus*†Sivapithecus*†Lufengpithecus*†Ankarapithecus*†Ouranopithecus*†Griphopithecus...

. Again, the three-way split in Ponginae led scientists to ask which of the three genera is least related to the others.
Investigation showed orangutan
Orangutan
Orangutans are the only exclusively Asian genus of extant great ape. The largest living arboreal animals, they have proportionally longer arms than the other, more terrestrial, great apes. They are among the most intelligent primates and use a variety of sophisticated tools, also making sleeping...

s to be the outgroup, but comparing humans to all three other hominid genera showed that African "apes" (chimpanzees and gorillas) and humans are more closely related to each other than any of them are to orangutans. This led to the placing of the African hominoids in the subfamily Homininae, forming another three-way split. This classification was first proposed by M. Goodman in 1974.
To try to resolve the hominine trichotomy, some authors proposed the division of the subfamily Homininae into the tribes Gorillini (African "apes") and Hominini (humans).
However, DNA comparisons provide convincing evidence that within the subfamily Homininae, gorillas are the outgroup. This suggests that chimpanzees should be in Hominini along with humans. This classification was first proposed (though one rank lower) by M. Goodman et al. in 1990. See Human evolutionary genetics
Human evolutionary genetics
Human evolutionary genetics studies how one human genome differs from the other, the evolutionary past that gave rise to it, and its current effects. Differences between genomes have anthropological, medical and forensic implications and applications...

 for more information on the speciation of humans and "great apes".
Later DNA comparisons split the gibbon genus Hylobates into four genera: Hylobates
Hylobates
The genus Hylobates is one of the four genera of gibbons. It was once considered the only genus, but recently its subgenera have been elevated to the genus level. Hylobates remains the most speciose and widespread of gibbon genera, ranging from southern China to western and central Java...

, Hoolock, Nomascus
Nomascus
Nomascus is the second most speciose genus of gibbons . Originally this genus was a subgenus of Hylobates, and all individuals were considered one species, Hylobates concolor. Species within Nomascus are characterized by 52 chromosomes...

, and Symphalangus.

Classification and evolution

As discussed above, hominoid taxonomy has undergone several changes. Genetic analysis shows that hominoids diverged from the Old World monkey
Old World monkey
The Old World monkeys or Cercopithecidae are a group of primates, falling in the superfamily Cercopithecoidea in the clade Catarrhini. The Old World monkeys are native to Africa and Asia today, inhabiting a range of environments from tropical rain forest to savanna, shrubland and mountainous...

s between 29 million and 34.5 million years ago. The gibbons split from the rest about 18 mya, and the hominid splits happened 14 mya (Pongo), 7 mya (Gorilla), and 3-5 mya (Homo & Pan).

The families, genera and extant species of hominoids are:
  • Superfamily Hominoidea
    • Family Hylobatidae: Gibbons ("Lesser apes")
      • Genus Hylobates
        Hylobates
        The genus Hylobates is one of the four genera of gibbons. It was once considered the only genus, but recently its subgenera have been elevated to the genus level. Hylobates remains the most speciose and widespread of gibbon genera, ranging from southern China to western and central Java...

        • Lar Gibbon
          Lar Gibbon
          The lar gibbon , also known as the white-handed gibbon, is a primate in the Hylobatidae or gibbon family. It is one of the better-known gibbons and is often seen in zoos.-Range:...

           or White-handed Gibbon, H. lar
        • Bornean White-bearded Gibbon
          Bornean white-bearded gibbon
          The Bornean white-bearded Gibbon, , also known as the Bornean agile Gibbon or Southern Gibbon, is an endangered species of gibbon endemic to southern Borneo, between the Kapuas and Barito rivers....

          , H. albibarbis
        • Agile Gibbon
          Agile Gibbon
          The agile gibbon , also known as the black-handed gibbon, is an Old World primate in the Hylobatidae or gibbon family, a group also collectively referred to as the "lesser apes"...

           or Black-handed Gibbon, H. agilis
        • Müller's Bornean Gibbon
          Müller's Bornean Gibbon
          Müller's Bornean gibbon , also known as the grey gibbon, is a primate in the Hylobatidae or gibbon family.Unlike other gibbon species, Müller's Bornean gibbon does not show sexual dimorphism in its fur coloration. Its fur is grey or brown colored with a ring of bright fur around its face. On the...

           or Grey Gibbon, H. muelleri
        • Silvery Gibbon
          Silvery Gibbon
          The silvery gibbon is a primate in the Hylobatidae or gibbon family. Its coat is bluish-grey in colour, with a dark grey or black cap. Like all gibbons, Silvery Gibbons lack external tails, have dorsally-placed scapula and reduced flexibility in the lumbar region. They have long, curved fingers...

          , H. moloch
        • Pileated Gibbon
          Pileated Gibbon
          The pileated gibbon is a primate in the Hylobatidae or gibbon family.The pileated gibbon has sexual dimorphism in fur coloration: males have a purely black fur, while the females have a white-grey colored fur with only the belly and head black...

           or Capped Gibbon, H. pileatus
        • Kloss's Gibbon
          Kloss's Gibbon
          Kloss's gibbon , also known as the Mentawai gibbon or the Bilou, is a primate in the Hylobatidae or gibbon family. It resembles the Siamang with its black fur, but is considerably smaller and lacks the Siamang's distinctive throat pouch. Kloss's gibbon reaches a size 44 to 63 cm and weigh at most 6...

           or Mentawai Gibbon or Bilou, H. klossii
      • Genus Hoolock
        • Western Hoolock Gibbon
          Western Hoolock Gibbon
          The western hoolock gibbon is a primate from the Hylobatidae family. The species is found in Assam, Bangladesh and in Myanmar west of the Chindwin River.-Classification:...

          , H. hoolock
        • Eastern Hoolock Gibbon
          Eastern Hoolock Gibbon
          The eastern hoolock gibbon is a primate from the Hylobatidae family. The species is found in extreme eastern corner of Assam and parts of Arunachal Pradesh, Myanmar east of the Chindwin River, and in south west Yunnan of China....

          , H. leuconedys
      • Genus Symphalangus
        • Siamang
          Siamang
          The siamang is a tailless, arboreal, black-furred gibbon native to the forests of Malaysia, Thailand, and Sumatra. The largest of the lesser apes, the siamang can be twice the size of other gibbons, reaching 1 m in height, and weighing up to 14 kg...

          , S. syndactylus
      • Genus Nomascus
        Nomascus
        Nomascus is the second most speciose genus of gibbons . Originally this genus was a subgenus of Hylobates, and all individuals were considered one species, Hylobates concolor. Species within Nomascus are characterized by 52 chromosomes...

        • Black Crested Gibbon, N. concolor
        • Eastern Black Crested Gibbon
          Eastern Black Crested Gibbon
          The eastern black crested gibbon is a species of gibbon that was once widespread in China and Vietnam. There are two subspecies....

          , N. nasutus
        • Hainan Black-crested Gibbon
          Hainan Black-Crested Gibbon
          The Hainan Black Crested Gibbon or Hainan Gibbon , is found only on Hainan Island, China. It was formerly considered a subspecies of the Eastern Black Crested Gibbon from Hoa Binh and Cao Bang provinces of Vietnam and Jingxi County in Guangxi Zhuang Autonomous Region, China...

          , N. hainanus
        • Southern White-cheeked Gibbon
          Southern White-cheeked Gibbon
          The southern white-cheeked gibbon is a species of gibbon native to Vietnam and Laos. It is closely related to the northern white-cheeked gibbon and the yellow-cheeked gibbon ; it has previously been identified as a subspecies of each of these, and may potentially be a hybrid of the two rather...

           N. siki
        • White-cheeked Crested Gibbon
          White-cheeked Crested Gibbon
          The northern white-cheeked gibbon is a species of gibbon native to Vietnam, Laos, and the Yunnan province of China. It is closely related to the southern white-cheeked gibbon , with which it was previously considered conspecific. The females of the two species are virtually indistinguishable in...

          , N. leucogenys
        • Yellow-cheeked Gibbon
          Yellow-cheeked Gibbon
          The yellow-cheeked gibbon , also called the yellow-cheeked crested gibbon, the golden-cheeked crested gibbon or the buffed-cheeked gibbon, is a species of gibbon native to Vietnam, Laos and Cambodia....

          , N. gabriellae
    • Family Hominidae
      Hominidae
      The Hominidae or include them .), as the term is used here, form a taxonomic family, including four extant genera: chimpanzees , gorillas , humans , and orangutans ....

      : Hominids ("Great apes", including humans)
      • Genus Pongo
        Orangutan
        Orangutans are the only exclusively Asian genus of extant great ape. The largest living arboreal animals, they have proportionally longer arms than the other, more terrestrial, great apes. They are among the most intelligent primates and use a variety of sophisticated tools, also making sleeping...

        : Orangutans
        • Bornean orangutan
          Bornean Orangutan
          The Bornean orangutan, Pongo pygmaeus, is a species of orangutan native to the island of Borneo. Together with the slightly smaller Sumatran orangutan, it belongs to the only genus of great apes native to Asia....

          , P. pygmaeus
        • Sumatran orangutan
          Sumatran Orangutan
          The Sumatran orangutan is one of the two species of orangutans. Found only on the island of Sumatra, in Indonesia, it is rarer and smaller than the Bornean orangutan. The Sumatran orangutan grows to about tall and in males...

          , P. abelii
      • Genus Gorilla
        Gorilla
        Gorillas are the largest extant species of primates. They are ground-dwelling, predominantly herbivorous apes that inhabit the forests of central Africa. Gorillas are divided into two species and either four or five subspecies...

        : Gorillas
        • Western Gorilla
          Western Gorilla
          The western gorilla is a great ape and the most populous species of the genus Gorilla.-Taxonomy:Nearly all of the individuals of this taxon belong to the western lowland gorilla subspecies whose population is approximately 95,000 individuals...

          , G. gorilla
        • Eastern Gorilla
          Eastern Gorilla
          The eastern gorilla is a species of the genus Gorilla and the largest living primate. At present, the species is subdivided into two subspecies. The eastern lowland gorilla is the most populous, at about 5,000 individuals. The mountain gorilla has only about 700 individuals...

          , G. beringei
      • Genus Homo
        Homo (genus)
        Homo is the genus that includes modern humans and species closely related to them. The genus is estimated to be about 2.3 to 2.4 million years old, evolving from australopithecine ancestors with the appearance of Homo habilis....

        : Humans
        • Human
          Human
          Humans are the only living species in the Homo genus...

          , H. sapiens
      • Genus Pan
        Chimpanzee
        Chimpanzee, sometimes colloquially chimp, is the common name for the two extant species of ape in the genus Pan. The Congo River forms the boundary between the native habitat of the two species:...

        : Chimpanzees
        • Common Chimpanzee
          Common Chimpanzee
          The common chimpanzee , also known as the robust chimpanzee, is a great ape. Colloquially, the common chimpanzee is often called the chimpanzee , though technically this term refers to both species in the genus Pan: the common chimpanzee and the closely related bonobo, formerly called the pygmy...

          , P. troglodytes
        • Bonobo
          Bonobo
          The bonobo , Pan paniscus, previously called the pygmy chimpanzee and less often, the dwarf or gracile chimpanzee, is a great ape and one of the two species making up the genus Pan. The other species in genus Pan is Pan troglodytes, or the common chimpanzee...

          , P. paniscus
    • Family †Parapithecidae
      Parapithecidae
      Parapithecidae is an extinct family of primates which lived in the Eocene and Oligocene periods in Egypt. Eocene fossils from Burma are sometimes included in the family in addition. They showed certain similarities in dentition to Condylarthra, but had short faces and jaws shaped like those of...

       
      • Genus Apidium
        Apidium
        The genus Apidium is that of at least three extinct primates living in the early Oligocene, roughly 36 to 32 millions years ago. Apidium fossils are common in the Fayoum deposits of Egypt...

      • Genus Arsinoea
        Arsinoea
        Arsinoea is an extinct genus of primates of which there is one known species....

      • Genus Biretia
        Biretia
        Biretia is an extinct genus of Old World monkey, of the extinct family Parapithecidae. Fossils are found from Late Eocene strata of Egypt....

      • Genus Parapithecus
        Parapithecus
        Parapithecus is an extinct genus of primates. There are two known species.-Species:*†Parapithecus fraasi Schlosser 1910*†Parapithecus grangeri Simons 1974...

      • Genus Qatratia
        Qatratia
        Qatratia is an extinct genus of primates. There are two known species.-Species:*†Qatratia fleaglei Simons & Kay, 1983*†Qatratia wingi Simons & Kay, 1983...


Cultural aspects of apes

Often, "apes" (nonhuman hominoids) are said to be the result of a curse—a Jewish folktale claims that one of the races who built the Tower of Babel
Tower of Babel
The Tower of Babel , according to the Book of Genesis, was an enormous tower built in the plain of Shinar .According to the biblical account, a united humanity of the generations following the Great Flood, speaking a single language and migrating from the east, came to the land of Shinar, where...

 became apes as punishment, while Muslim lore says that the Jews of Eilat became apes as punishment for fishing on the Sabbath. Some sects of Christianity have folklore that claims that these apes are a symbol of lust and were created by Satan in response to God's creation of humans. It is uncertain whether any of these references are to any specific apes. All of these concepts date from a period when neither the distinction between apes and monkeys, nor the fact that humans are closely related to chimpanzees, was widely understood, if understood at all.

See also

  • Chimp Haven
  • Declaration on Great Apes from the Great Ape Project
    Great Ape Project
    The Great Ape Project , founded in 1994, is an international organization of primatologists, anthropologists, ethicists, and other experts who advocate a United Nations Declaration of the Rights of Great Apes that would confer basic legal rights on non-human great apes: chimpanzees, bonobos,...

  • Human
    Human
    Humans are the only living species in the Homo genus...

  • List of human evolution fossils
  • List of apes (for notable non-fictional apes)
  • List of fictional apes

External links

Agreement between cladograms based on molecular and anatomical data.
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