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Apes (collectively Hominoidea ) are a superfamily of
Old World The "Old World" () is a term for Afro-Eurasia coined by Europeans after 1493, when they became aware of the existence of the Americas. It is used to contrast the continents of Africa, Europe, and Asia in the Eastern Hemisphere, previously ...
simian The simians, anthropoids, or higher primates are an infraorder (Simiiformes ) of primates containing all animals traditionally called monkeys and apes. More precisely, they consist of the parvorders New World monkey, Platyrrhini (New World mon ...
s native to
sub-Saharan Africa Sub-Saharan Africa is the area and regions of the continent of Africa that lie south of the Sahara. These include Central Africa, East Africa, Southern Africa, and West Africa. Geopolitically, in addition to the list of sovereign states and ...
and
Southeast Asia Southeast Asia is the geographical United Nations geoscheme for Asia#South-eastern Asia, southeastern region of Asia, consisting of the regions that are situated south of China, east of the Indian subcontinent, and northwest of the Mainland Au ...
(though they were more widespread in Africa, most of Asia, and Europe in prehistory, and counting humans are found globally). Apes are more closely related to
Old World monkey Old World monkeys are primates in the family Cercopithecidae (). Twenty-four genera and 138 species are recognized, making it the largest primate family. Old World monkey genera include baboons (genus '' Papio''), red colobus (genus '' Piliocolob ...
s (family Cercopithecidae) than to the
New World monkey New World monkeys are the five families of primates that are found in the tropical regions of Mexico, Central and South America: Callitrichidae, Cebidae, Aotidae, Pitheciidae, and Atelidae. The five families are ranked together as the Ceboi ...
s (Platyrrhini) with both Old World monkeys and apes placed in the
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
Catarrhini The parvorder Catarrhini (known commonly as catarrhine monkeys, Old World anthropoids, or Old World monkeys) consists of the Cercopithecoidea and apes (Hominoidea). In 1812, Geoffroy grouped those two groups together and established the name ...
. Apes do not have tails due to a mutation of the TBXT gene. In traditional and non-scientific use, the term ''ape'' can include tailless primates taxonomically considered Cercopithecidae (such as the
Barbary ape The Barbary macaque (''Macaca sylvanus''), also known as Barbary ape, is a macaque species native to the Atlas Mountains of Algeria, Tunisia and Morocco, along with a small introduced population in Gibraltar. It is the type species of the genus ' ...
and black ape), and is thus not equivalent to the scientific
taxon In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
Hominoidea. There are two
extant Extant or Least-concern species, least concern is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Exta ...
branches of the superfamily Hominoidea: the
gibbon Gibbons () are apes in the family Hylobatidae (). The family historically contained one genus, but now is split into four extant genera and 20 species. Gibbons live in subtropical and tropical forests from eastern Bangladesh and Northeast Indi ...
s, or lesser apes; and the
hominid The Hominidae (), whose members are known as the great apes or hominids (), are a taxonomic family of primates that includes eight extant species in four genera: '' Pongo'' (the Bornean, Sumatran and Tapanuli orangutan); '' Gorilla'' (the ...
s, or great apes. * The family
Hylobatidae Gibbons () are apes in the Family (biology), family Hylobatidae (). The family historically contained one genus, but now is split into four extant genera and 20 species. Gibbons live in subtropical and tropical forests from eastern Bangladesh an ...
, the lesser apes, include four genera and a total of 20 species of gibbon, including the lar gibbon and the
siamang The siamang (, ; ''Symphalangus syndactylus'') is an endangered arboreal, black-furred gibbon native to the forests of Indonesia, Malaysia, and Thailand. The largest of the gibbons, the siamang can be twice the size of other gibbons, reaching i ...
, all native to
Asia Asia ( , ) is the largest continent in the world by both land area and population. It covers an area of more than 44 million square kilometres, about 30% of Earth's total land area and 8% of Earth's total surface area. The continent, which ...
. They are highly
arboreal Arboreal locomotion is the locomotion of animals in trees. In habitats in which trees are present, animals have evolved to move in them. Some animals may scale trees only occasionally (scansorial), but others are exclusively arboreal. The hab ...
and
bipedal Bipedalism is a form of terrestrial locomotion where an animal moves by means of its two rear (or lower) limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning 'two feet' (from Latin ''bis'' ...
on the ground. They have lighter bodies and smaller social groups than great apes. * The family
Hominidae The Hominidae (), whose members are known as the great apes or hominids (), are a taxonomic Family (biology), family of primates that includes eight Neontology#Extant taxa versus extinct taxa, extant species in four Genus, genera: ''Orangutan ...
(''hominids''), the great apes, include four genera comprising three extant species of
orangutan Orangutans are great apes native to the rainforests of Indonesia and Malaysia. They are now found only in parts of Borneo and Sumatra, but during the Pleistocene they ranged throughout Southeast Asia and South China. Classified in the genus ...
s and their subspecies, two extant species of
gorilla Gorillas are primarily herbivorous, terrestrial great apes that inhabit the tropical forests of equatorial Africa. The genus ''Gorilla'' is divided into two species: the eastern gorilla and the western gorilla, and either four or five su ...
s and their subspecies, two extant species of
chimpanzees The chimpanzee (; ''Pan troglodytes''), also simply known as the chimp, is a species of great ape native to the forests and savannahs of tropical Africa. It has four confirmed subspecies and a fifth proposed one. When its close relative the ...
and their subspecies, and
human Humans (''Homo sapiens'') or modern humans are the most common and widespread species of primate, and the last surviving species of the genus ''Homo''. They are Hominidae, great apes characterized by their Prehistory of nakedness and clothing ...
s in a single extant subspecies. Except for gorillas and humans, hominoids are agile climbers of trees. Apes eat a variety of plant and animal foods, with the majority of food being plant foods, which can include fruits, leaves, stalks, roots and seeds, including nuts and grass seeds. Human diets are sometimes substantially different from that of other hominoids due in part to the development of technology and a wide range of habitation. All extant non-human hominoids are rare and threatened with
extinction Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
. The main threat is
habitat loss Habitat destruction (also termed habitat loss or habitat reduction) occurs when a natural habitat is no longer able to support its native species. The organisms once living there have either moved elsewhere, or are dead, leading to a decrease ...
, though some populations are further imperiled by hunting. The great apes of Africa are also facing threat from the
Ebola virus ''Orthoebolavirus zairense'' or Zaire ebolavirus, more commonly known as Ebola virus (; EBOV), is one of six known species within the genus ''Ebolavirus''. Four of the six known ebolaviruses, including EBOV, cause a severe and often fatal vira ...
.


Name and terminology

"Ape", from Old English ''apa'', is a word of uncertain origin. The term has a history of rather imprecise usage—and of comedic or punning usage in the vernacular. Its earliest meaning was generally of any non-human anthropoid primate, as is still the case for its cognates in other Germanic languages. Later, after the term "
monkey Monkey is a common name that may refer to most mammals of the infraorder Simiiformes, also known as simians. Traditionally, all animals in the group now known as simians are counted as monkeys except the apes. Thus monkeys, in that sense, co ...
" had been introduced into English, "ape" was specialized to refer to a tailless (therefore exceptionally human-like) primate. Thus, the term "ape" obtained two different meanings, as shown in the 1911 ''Encyclopædia Britannica'' entry: it could be used as a synonym for "monkey" and it could denote the tailless human-like primate in particular. Some, or recently all, hominoids are also called "apes", but the term is used broadly and has several different senses within both popular and scientific settings. "Ape" has been used as a synonym for "monkey" or for naming any primate with a human-like appearance, particularly those without a tail. Biologists have traditionally used the term "ape" to mean a member of the superfamily Hominoidea ''other than'' humans, but more recently to mean ''all'' members of Hominoidea. So "ape"—not to be confused with "great ape"—now becomes another word for hominoid ''including'' humans. The taxonomic term ''hominoid'' is derived from, and intended as encompassing, the ''
hominid The Hominidae (), whose members are known as the great apes or hominids (), are a taxonomic family of primates that includes eight extant species in four genera: '' Pongo'' (the Bornean, Sumatran and Tapanuli orangutan); '' Gorilla'' (the ...
s,'' the family of
great ape The Hominidae (), whose members are known as the great apes or hominids (), are a taxonomic family of primates that includes eight extant species in four genera: '' Pongo'' (the Bornean, Sumatran and Tapanuli orangutan); '' Gorilla'' (the ...
s. Both terms were introduced by Gray (1825). The term ''
hominin The Hominini (hominins) form a taxonomic tribe of the subfamily Homininae (hominines). They comprise two extant genera: ''Homo'' (humans) and '' Pan'' (chimpanzees and bonobos), and in standard usage exclude the genus '' Gorilla'' ( gorillas) ...
s'' is also due to Gray (1824), intended as including the human lineage (see also Hominidae#Terminology, Human taxonomy). The distinction between apes and monkeys is complicated by the traditional
paraphyly Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
of monkeys: Apes emerged as a sister group of Old World Monkeys in the
catarrhines The parvorder Catarrhini (known commonly as catarrhine monkeys, Old World anthropoids, or Old World monkeys) consists of the Cercopithecoidea and apes (Hominoidea). In 1812, Geoffroy grouped those two groups together and established the name ...
, which are a sister group of New World Monkeys. Therefore, cladistically, apes, catarrhines and related contemporary extinct groups such as
Parapithecidae Parapithecidae is an now extinct family of primates which lived in the Eocene and Oligocene periods in Egypt. Eocene fossils from Myanmar are sometimes included in the family in addition. They showed certain similarities in dentition to Condylar ...
are monkeys as well, for any consistent definition of "monkey". "Old World monkey" may also legitimately be taken to be meant to include all the catarrhines, including apes and extinct species such as ''
Aegyptopithecus ''Aegyptopithecus'' ("Egyptian ape", from Greek ''Αίγυπτος'' "Egypt" and ''πίθηκος'' "ape") is an early fossil Catarrhini, catarrhine that predates the divergence between hominoids (apes) and Cercopithecidae, cercopithecids (Old Wo ...
'', in which case the apes, Cercopithecoidea and ''Aegyptopithecus'' emerged within the Old World monkeys. The primates called "apes" today became known to Europeans after the 18th century. As zoological knowledge developed, it became clear that taillessness occurred in a number of different and otherwise distantly related
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
. Sir Wilfrid Le Gros Clark was one of those primatologists who developed the idea that there were trends in primate evolution and that the extant members of the order could be arranged in an "ascending series", leading from "monkeys" to "apes" to humans. Within this tradition "ape" came to refer to all members of the superfamily Hominoidea except humans. As such, this use of "apes" represented a
paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
grouping, meaning that, even though all species of apes were descended from a common ancestor, this grouping did ''not'' include all the descendant species, because humans were excluded from being among the apes. Traditionally, the English-language
vernacular name Vernacular is the ordinary, informal, spoken form of language, particularly when perceived as having lower social status or less prestige than standard language, which is more codified, institutionally promoted, literary, or formal. More n ...
"apes" does not include humans, but phylogenetically, humans (''
Homo ''Homo'' () is a genus of great ape (family Hominidae) that emerged from the genus ''Australopithecus'' and encompasses only a single extant species, ''Homo sapiens'' (modern humans), along with a number of extinct species (collectively called ...
'') form part of the family Hominidae within Hominoidea. Thus, there are at least three common, or traditional, uses of the term "ape": non-specialists may not distinguish between "monkeys" and "apes", that is, they may use the two terms interchangeably; or they may use "ape" for any tailless monkey or non-human hominoid; or they may use the term "ape" to just mean the non-human hominoids. Modern taxonomy aims for the use of
monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...
groups for taxonomic classification; Some literature may now use the common name "ape" to mean all members of the superfamily Hominoidea, including humans. For example, in his 2005 book, Benton wrote "The apes, Hominoidea, today include the gibbons and orang-utan ... the gorilla and chimpanzee ... and humans"., p. 371 Modern biologists and primatologists refer to apes that are not human as "non-human" apes. Scientists broadly, other than paleoanthropologists, may use the term "
hominin The Hominini (hominins) form a taxonomic tribe of the subfamily Homininae (hominines). They comprise two extant genera: ''Homo'' (humans) and '' Pan'' (chimpanzees and bonobos), and in standard usage exclude the genus '' Gorilla'' ( gorillas) ...
" to identify the human clade, replacing the term "
hominid The Hominidae (), whose members are known as the great apes or hominids (), are a taxonomic family of primates that includes eight extant species in four genera: '' Pongo'' (the Bornean, Sumatran and Tapanuli orangutan); '' Gorilla'' (the ...
". See terminology of primate names. See below, History of hominoid taxonomy, for a discussion of changes in scientific classification and terminology regarding hominoids.


Evolution

Although the hominoid fossil record is still incomplete and fragmentary, there is now enough evidence to provide an outline of the evolutionary history of humans. Previously, the divergence between humans and other extant hominoids was thought to have occurred 15 to 20 million years ago, and several species of that time period, such as ''
Ramapithecus ''Sivapithecus'' () (syn: ''Ramapithecus)'' is a genus of extinct apes. Fossil remains of animals now assigned to this genus, dated from 12.2 million years old in the Miocene, have been found since the 19th century in the Sivalik Hills of the Ind ...
'', were once thought to be
hominin The Hominini (hominins) form a taxonomic tribe of the subfamily Homininae (hominines). They comprise two extant genera: ''Homo'' (humans) and '' Pan'' (chimpanzees and bonobos), and in standard usage exclude the genus '' Gorilla'' ( gorillas) ...
s and possible ancestors of humans. But, later fossil finds indicated that ''Ramapithecus'' was more closely related to the orangutan; and new biochemical evidence indicates that the last common ancestor of humans and non-hominins (that is, the chimpanzees) occurred between 5 and 10 million years ago, and probably nearer the lower end of that range (more recent); see
Chimpanzee–human last common ancestor The chimpanzee–human last common ancestor (CHLCA) is the last common ancestor shared by the extant ''Homo'' (human) and '' Pan'' (chimpanzee and bonobo) genera of Hominini. Estimates of the divergence date vary widely from thirteen to five mill ...
(CHLCA).


Taxonomic classification and phylogeny

Genetic analysis combined with fossil evidence indicates that hominoids diverged from the
Old World monkey Old World monkeys are primates in the family Cercopithecidae (). Twenty-four genera and 138 species are recognized, making it the largest primate family. Old World monkey genera include baboons (genus '' Papio''), red colobus (genus '' Piliocolob ...
s about 25 million years ago (mya), near the Oligocene–Miocene boundary. 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''). In 2015, a new genus and species were described, '' Pliobates cataloniae'', which lived 11.6 mya, and appears to predate the split between Hominidae and Hylobatidae. The families, and extant genera and species of hominoids are: * Superfamily Hominoidea ** Family
Hominidae The Hominidae (), whose members are known as the great apes or hominids (), are a taxonomic Family (biology), family of primates that includes eight Neontology#Extant taxa versus extinct taxa, extant species in four Genus, genera: ''Orangutan ...
: hominids ("great apes") *** Genus '' Pongo'': orangutans ****
Bornean orangutan The Bornean orangutan (''Pongo pygmaeus'') is an orangutan species endemic to the island of Borneo. It belongs to the only genus of great apes native to Asia and is the largest of the three ''Pongo'' species. It has a coarse, reddish coat and up ...
, ''P. pygmaeus'' ****
Sumatran orangutan The Sumatran orangutan (''Pongo abelii'') is one of the three species of orangutans. Critically endangered, and found only in the north of the Indonesian island of Sumatra, it is rarer than the Bornean orangutan but more common than the recently ...
, ''P. abelii'' **** Tapanuli orangutan, ''P. tapanuliensis'' *** Genus ''
Gorilla Gorillas are primarily herbivorous, terrestrial great apes that inhabit the tropical forests of equatorial Africa. The genus ''Gorilla'' is divided into two species: the eastern gorilla and the western gorilla, and either four or five su ...
'': gorillas ****
Western gorilla The western gorilla (''Gorilla gorilla'') is a great ape found in Africa, one of two species of the hominid genus ''Gorilla''. Large and robust with males weighing around , the species is found in a region of midwest Africa, geographically iso ...
, ''G. gorilla'' ****
Eastern gorilla The eastern gorilla (''Gorilla beringei'') is a critically endangered species of the genus ''Gorilla'' and the largest living primate. At present, the species is subdivided into two subspecies. There are 6,800 eastern lowland gorillas or Grauer ...
, ''G. beringei'' *** Genus ''
Homo ''Homo'' () is a genus of great ape (family Hominidae) that emerged from the genus ''Australopithecus'' and encompasses only a single extant species, ''Homo sapiens'' (modern humans), along with a number of extinct species (collectively called ...
'': humans ****
Human Humans (''Homo sapiens'') or modern humans are the most common and widespread species of primate, and the last surviving species of the genus ''Homo''. They are Hominidae, great apes characterized by their Prehistory of nakedness and clothing ...
, ''H. sapiens'' *** Genus '' Pan'': chimpanzees ****
Chimpanzee The chimpanzee (; ''Pan troglodytes''), also simply known as the chimp, is a species of Hominidae, great ape native to the forests and savannahs of tropical Africa. It has four confirmed subspecies and a fifth proposed one. When its close rel ...
, ''P. troglodytes'' ****
Bonobo The bonobo (; ''Pan paniscus''), also historically called the pygmy chimpanzee (less often the dwarf chimpanzee or gracile chimpanzee), is an endangered great ape and one of the two species making up the genus ''Pan (genus), Pan'' (the other bei ...
, ''P. paniscus'' ** Family
Hylobatidae Gibbons () are apes in the Family (biology), family Hylobatidae (). The family historically contained one genus, but now is split into four extant genera and 20 species. Gibbons live in subtropical and tropical forests from eastern Bangladesh an ...
: gibbons ("lesser apes") *** Genus ''
Hylobates The genus ''Hylobates'' is one of the four genera of gibbons. Its name means "forest walker", from the Greek (, "forest") and (, "one who treads"). It was once considered the only genus, but recently its subgenera (''Hoolock'' ormerly ''Bun ...
'' **** Lar gibbon or white-handed gibbon, ''H. lar'' **** Bornean white-bearded gibbon, ''H. albibarbis'' **** Agile gibbon or black-handed gibbon, ''H. agilis'' **** Western grey gibbon or Abbott's grey gibbon, ''H. abbotti'' **** Eastern grey gibbon or northern grey gibbon, ''H. funereus'' ****
Müller's gibbon Müller's gibbon (''Hylobates muelleri''), also known as the southern grey gibbon, is a primate in the gibbon family, Hylobatidae. Taxonomy Formerly, the western grey gibbon (''H. abbotti'') and eastern grey gibbon (''H. funereus'') were consi ...
or southern grey gibbon, ''H. muelleri'' **** Silvery gibbon, ''H. moloch'' ****
Pileated gibbon The pileated gibbon (''Hylobates pileatus'') is a primate in the gibbon family, Hylobatidae. The pileated gibbon has sexual dimorphism in fur coloration: males have purely black fur, while females have white-grey colored fur with only the belly ...
or capped gibbon, ''H. pileatus'' **** Kloss's gibbon or Mentawai gibbon or bilou, ''H. klossii'' *** Genus ''
Hoolock The hoolock gibbons are three primate species of genus ''Hoolock'' in the gibbon family, Hylobatidae, native to eastern Bangladesh, Northeast India, Myanmar, and Southwest China. Description Hoolocks are the second-largest of the gibbons, after ...
'' ****
Western hoolock gibbon The western hoolock gibbon (''Hoolock hoolock'') is a primate from the gibbon family, Hylobatidae. The species is found in Assam, Mizoram, and Meghalaya in India, Bangladesh, and Myanmar west of the Chindwin River. Classification Mootnick and ...
, ''H. hoolock'' **** Eastern hoolock gibbon, ''H. leuconedys'' **** Skywalker hoolock gibbon, ''H. tianxing'' *** Genus ''
Symphalangus The siamang (, ; ''Symphalangus syndactylus'') is an endangered arboreal, black-furred gibbon native to the forests of Indonesia, Malaysia, and Thailand. The largest of the gibbons, the siamang can be twice the size of other gibbons, reaching i ...
'' ****
Siamang The siamang (, ; ''Symphalangus syndactylus'') is an endangered arboreal, black-furred gibbon native to the forests of Indonesia, Malaysia, and Thailand. The largest of the gibbons, the siamang can be twice the size of other gibbons, reaching i ...
, ''S. syndactylus'' *** Genus ''
Nomascus ''Nomascus'' is the second-most speciose genus of the gibbon family, Hylobatidae. Originally, this genus was a subgenus of ''Hylobates'', with all individuals considered to be one species, ''H. concolor''. Species within ''Nomascus'' are charact ...
'' ****
Northern buffed-cheeked gibbon The northern buffed-cheeked gibbon (''Nomascus annamensis'') is a newly discovered species of crested gibbon which is found in Vietnam, Cambodia, and Laos. Its habitat is in the humid subtropical and seasonal tropical forests of these countries. ...
, ''N. annamensis'' ****
Black crested gibbon The black crested gibbon (''Nomascus concolor'') is a Critically Endangered species of gibbon found in China, Laos, and northern Vietnam, with four subspecies. Taxonomy The taxonomy of the species is confused. Previously grouped in the gen ...
, ''N. concolor'' **** Eastern black crested gibbon, ''N. nasutus'' **** Hainan black crested gibbon, ''N. hainanus'' **** Southern white-cheeked gibbon ''N. siki'' **** White-cheeked crested gibbon, ''N. leucogenys'' **** Yellow-cheeked gibbon, ''N. gabriellae''


History of hominoid taxonomy

The history of hominoid taxonomy is complex and somewhat confusing. Recent evidence has changed our understanding of the relationships between the hominoids, especially regarding the human lineage; and the traditionally used terms have become somewhat confused. Competing approaches to methodology and terminology are found among current scientific sources. Over time, authorities have changed the names and the meanings of names of groups and subgroups as new evidence — that is, new discoveries of fossils and tools and of observations in the field, plus continual comparisons of anatomy and
DNA sequences A nucleic acid sequence is a succession of bases within the nucleotides forming alleles within a DNA (using GACT) or RNA (GACU) molecule. This succession is denoted by a series of a set of five different letters that indicate the order of the ...
— has changed the understanding of relationships between hominoids. There has been a gradual demotion of humans from being 'special' in the taxonomy to being one branch among many. This recent turmoil (of history) illustrates the growing influence on all taxonomy of
cladistics Cladistics ( ; from Ancient Greek 'branch') is an approach to Taxonomy (biology), biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesiz ...
, the science of classifying living things strictly according to their lines of descent. Today, there are eight extant
genera Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial s ...
of hominoids. They are the four genera in the family Hominidae, namely ''
Homo ''Homo'' () is a genus of great ape (family Hominidae) that emerged from the genus ''Australopithecus'' and encompasses only a single extant species, ''Homo sapiens'' (modern humans), along with a number of extinct species (collectively called ...
'', '' Pan'', ''
Gorilla Gorillas are primarily herbivorous, terrestrial great apes that inhabit the tropical forests of equatorial Africa. The genus ''Gorilla'' is divided into two species: the eastern gorilla and the western gorilla, and either four or five su ...
'', and '' Pongo''; plus four genera in the family Hylobatidae (gibbons): ''
Hylobates The genus ''Hylobates'' is one of the four genera of gibbons. Its name means "forest walker", from the Greek (, "forest") and (, "one who treads"). It was once considered the only genus, but recently its subgenera (''Hoolock'' ormerly ''Bun ...
'', ''
Hoolock The hoolock gibbons are three primate species of genus ''Hoolock'' in the gibbon family, Hylobatidae, native to eastern Bangladesh, Northeast India, Myanmar, and Southwest China. Description Hoolocks are the second-largest of the gibbons, after ...
'', ''
Nomascus ''Nomascus'' is the second-most speciose genus of the gibbon family, Hylobatidae. Originally, this genus was a subgenus of ''Hylobates'', with all individuals considered to be one species, ''H. concolor''. Species within ''Nomascus'' are charact ...
'' and ''
Symphalangus The siamang (, ; ''Symphalangus syndactylus'') is an endangered arboreal, black-furred gibbon native to the forests of Indonesia, Malaysia, and Thailand. The largest of the gibbons, the siamang can be twice the size of other gibbons, reaching i ...
''. (The two subspecies of hoolock gibbons were recently moved from the genus ''
Bunopithecus ''Bunopithecus'' is an extinct genus of primate represented by one species, ''Bunopithecus sericus'', a gibbon or gibbon-like ape. Its remains were first discovered in Sichuan, China, in strata from the Middle Pleistocene. Although the three hoo ...
'' to the new genus ''Hoolock'' and re-ranked as species; a third species was described in January 2017). In 1758,
Carl Linnaeus Carl Linnaeus (23 May 1707 – 10 January 1778), also known after ennoblement in 1761 as Carl von Linné,#Blunt, Blunt (2004), p. 171. was a Swedish biologist and physician who formalised binomial nomenclature, the modern system of naming o ...
, relying on second- or third-hand accounts, placed a second species in ''Homo'' along with ''H. sapiens'': ''Homo troglodytes'' ("cave-dwelling man"). Although the term "Orang Outang" is listed as a variety – ''Homo sylvestris'' – under this species, it is nevertheless not clear to which animal this name refers, as Linnaeus had no specimen to refer to, hence no precise description. Linnaeus may have based ''Homo troglodytes'' on reports of mythical creatures, then-unidentified
simian The simians, anthropoids, or higher primates are an infraorder (Simiiformes ) of primates containing all animals traditionally called monkeys and apes. More precisely, they consist of the parvorders New World monkey, Platyrrhini (New World mon ...
s, or Asian natives dressed in animal skins. Linnaeus named the orangutan ''Simia satyrus'' ("satyr monkey"). He placed the three genera ''Homo'', ''
Simia In his ''Systema Naturae'' of 1758, Carl Linnaeus divided the Order Primates within ''Mammalia'' into four genera: ''Homo'', ''Simia'', ''Lemur'', and ''Vespertilio''. His ''Vespertilio'' included all bats, and has since been moved from Primates t ...
'' and ''Lemur'' in the order of Primates. The ''troglodytes'' name was used for the chimpanzee by Blumenbach in 1775, but moved to the genus ''Simia''. The orangutan was moved to the genus '' Pongo'' 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 (8 August 1709 – 20 May 1755) was a German naturalist, botanist and geographer. Early life and education Gmelin was born in Tübingen, the son of a professor at the University of Tübingen. He was a gifted child and began ...
dated 25 February 1747, Linnaeus wrote: Accordingly,
Johann Friedrich Blumenbach Johann Friedrich Blumenbach (11 May 1752 – 22 January 1840) was a German physician, naturalist, physiologist and anthropologist. He is considered to be a main founder of zoology and anthropology as comparative, scientific disciplines. He has be ...
in the first edition of his ''Manual of Natural History'' (1779), proposed that the primates be divided into the
Quadrumana Quadrumana is an outdated taxonomic division within the primates. The division of "Quadrumana" from "Bimana" was an attempt at distinguishing ''Homo sapiens'' from the rest of the great apes. For a century, modern science has considered humans as p ...
(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 Jean Léopold Nicolas Frédéric, baron Cuvier (23 August 1769 – 13 May 1832), known as Georges Cuvier (; ), was a French natural history, naturalist and zoology, zoologist, sometimes referred to as the "founding father of paleontology". Cuv ...
. Some elevated the distinction to the level of
order Order, ORDER or Orders may refer to: * A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica * Categorization, the process in which ideas and objects are recognized, differentiated, and understood ...
. However, the many affinities between humans and other primates – and especially the "great apes" – made it clear that the distinction made no scientific sense. In his 1871 book ''
The Descent of Man, and Selection in Relation to Sex ''The Descent of Man, and Selection in Relation to Sex'' is a book by English naturalist Charles Darwin, first published in 1871, which applies evolutionary theory to human evolution, and details his theory of sexual selection, a form of biolog ...
'',
Charles Darwin Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English Natural history#Before 1900, naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all speci ...
wrote:


Changes in taxonomy and terminology


Characteristics

The lesser apes are the gibbon family, Hylobatidae, of sixteen species; all are native to Asia. Their major differentiating characteristic is their long arms, which they use to
brachiate Brachiation (from "brachium", Latin for "arm"), or arm swinging, is a form of arboreal locomotion in which primates swing from tree limb to tree limb using only their arms. During brachiation, the body is alternately supported under each forelimb ...
through trees. Their wrists are
ball and socket joint The ball-and-socket joint (or spheroid joint) is a type of synovial joint in which the ball-shaped surface of one rounded bone fits into the cup-like depression of another bone. The distal bone is capable of motion around an indefinite number of ...
s as an evolutionary adaptation to their
arboreal Arboreal locomotion is the locomotion of animals in trees. In habitats in which trees are present, animals have evolved to move in them. Some animals may scale trees only occasionally (scansorial), but others are exclusively arboreal. The hab ...
lifestyle. Generally smaller than the African apes, the largest gibbon, the
siamang The siamang (, ; ''Symphalangus syndactylus'') is an endangered arboreal, black-furred gibbon native to the forests of Indonesia, Malaysia, and Thailand. The largest of the gibbons, the siamang can be twice the size of other gibbons, reaching i ...
, weighs up to ; in comparison, the smallest "great ape", the
bonobo The bonobo (; ''Pan paniscus''), also historically called the pygmy chimpanzee (less often the dwarf chimpanzee or gracile chimpanzee), is an endangered great ape and one of the two species making up the genus ''Pan (genus), Pan'' (the other bei ...
, is . The superfamily Hominoidea falls within the
parvorder Order () is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and recognized ...
Catarrhini The parvorder Catarrhini (known commonly as catarrhine monkeys, Old World anthropoids, or Old World monkeys) consists of the Cercopithecoidea and apes (Hominoidea). In 1812, Geoffroy grouped those two groups together and established the name ...
, which also includes the Old World monkeys of Africa and Eurasia. Within this grouping, the two families Hylobatidae and Hominidae can be distinguished from Old World monkeys by the number of cusps on their
molars The molars or molar teeth are large, flat tooth, teeth at the back of the mouth. They are more developed in mammal, mammals. They are used primarily to comminution, grind food during mastication, chewing. The name ''molar'' derives from Latin, '' ...
; hominoids have five in the "Y-5" molar pattern, whereas Old World monkeys have only four in a
bilophodont The molars or molar teeth are large, flat teeth at the back of the mouth. They are more developed in mammals. They are used primarily to grind food during chewing. The name ''molar'' derives from Latin, ''molaris dens'', meaning "millstone toot ...
pattern. Further, in comparison with Old World monkeys, hominoids are noted for: more mobile shoulder joints and arms due to the dorsal position of the
scapula The scapula (: scapulae or scapulas), also known as the shoulder blade, is the bone that connects the humerus (upper arm bone) with the clavicle (collar bone). Like their connected bones, the scapulae are paired, with each scapula on either side ...
; broader ribcages that are flatter front-to-back; and a shorter, less mobile spine, with greatly reduced caudal (tail) vertebrae—resulting in complete loss of the tail in extant hominoid species. These are anatomical adaptations, first, to vertical hanging and swinging locomotion (
brachiation Brachiation (from "brachium", Latin for "arm"), or arm swinging, is a form of arboreal locomotion in which primates swing from tree limb to tree limb using only their arms. During brachiation, the body is alternately supported under each forelimb ...
) and, later, to developing balance in a
bipedal Bipedalism is a form of terrestrial locomotion where an animal moves by means of its two rear (or lower) limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning 'two feet' (from Latin ''bis'' ...
pose. Note there are primates in other families that also lack tails, and at least one, the
pig-tailed langur The pig-tailed langur (''Simias concolor''), monotypic in genus ''Simias'', is a large Old World monkey, endemic to several small islands off the coast of Sumatra in Indonesia. Its face is black, its fur is black-brown and it has a relatively sho ...
, is known to walk significant distances bipedally. The front of the ape skull is characterised by its sinuses, fusion of the frontal bone, and by
post-orbital constriction In physical anthropology, post-orbital constriction is the narrowing of the cranium (skull) just behind the eye sockets (the orbits, hence the name) found in most non-human primates and early hominins. This constriction is very noticeable in non-hu ...
.


Distinction from monkeys

Cladistically, apes, catarrhines, and extinct species such as
Aegyptopithecus ''Aegyptopithecus'' ("Egyptian ape", from Greek ''Αίγυπτος'' "Egypt" and ''πίθηκος'' "ape") is an early fossil Catarrhini, catarrhine that predates the divergence between hominoids (apes) and Cercopithecidae, cercopithecids (Old Wo ...
and
Parapithecidae Parapithecidae is an now extinct family of primates which lived in the Eocene and Oligocene periods in Egypt. Eocene fossils from Myanmar are sometimes included in the family in addition. They showed certain similarities in dentition to Condylar ...
a, are monkeys, so one can only specify ape features not present in other monkeys. Unlike most
monkey Monkey is a common name that may refer to most mammals of the infraorder Simiiformes, also known as simians. Traditionally, all animals in the group now known as simians are counted as monkeys except the apes. Thus monkeys, in that sense, co ...
s, apes do not possess a tail. Monkeys are more likely to be in trees and use their tails for balance. While the great apes are considerably larger than monkeys,
gibbon Gibbons () are apes in the family Hylobatidae (). The family historically contained one genus, but now is split into four extant genera and 20 species. Gibbons live in subtropical and tropical forests from eastern Bangladesh and Northeast Indi ...
s (lesser apes) are smaller than some monkeys. Apes are considered to be more intelligent than monkeys, which are considered to have more primitive brains. The enzyme
urate oxidase The enzyme urate oxidase (UO), uricase or factor-independent urate hydroxylase, absent in humans, catalyzes the oxidation of uric acid to 5-hydroxyisourate: :Uric acid + O2 + H2O → 5-hydroxyisourate + H2O2 : 5-hydroxyisourate + H2O → 2- ...
has become inactive in all apes, its function having been lost in two
primate Primates is an order (biology), order of mammals, which is further divided into the Strepsirrhini, strepsirrhines, which include lemurs, galagos, and Lorisidae, lorisids; and the Haplorhini, haplorhines, which include Tarsiiformes, tarsiers a ...
lineages during the middle Miocene; first in the common ancestors of Hominidae, and later in the common ancestor of Hylobatidae. It has been hypothesized that in both incidents it was a mutation that occurred in apes living in Europe when the climate was getting colder, leading to starvation during winter. The mutation changed the biochemistry of the apes and made it easier to accumulate fat, which allowed the animals to survive longer periods of starvation. When they migrated to Asia and Africa, this genetic trait remained.


Behaviour

Major studies of behaviour in the field were completed on the three better-known "great apes", for example by
Jane Goodall Dame Jane Morris Goodall (; born Valerie Jane Morris-Goodall; 3 April 1934), formerly Baroness Jane van Lawick-Goodall, is an English zoologist, Primatology, primatologist and Anthropology, anthropologist. She is considered the world's foremo ...
,
Dian Fossey Dian Fossey ( ; January 16, 1932 – ) was an American primatologist and conservationist known for undertaking an extensive study of mountain gorilla groups from 1966 until her murder in 1985. She studied them daily in the mountain forests of ...
and
Birutė Galdikas Birutė Marija Filomena Galdikas or Birutė Mary Galdikas, OC (born 10 May 1946), is a Lithuanian-Canadian anthropologist, primatologist, conservationist, ethologist, and author. She is a professor at Simon Fraser University. In the field of ...
. These studies have shown that in their natural environments, the non-human 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
foliovore In zoology, a folivore is a herbivore that specializes in eating leaves. Mature leaves contain a high proportion of hard-to-digest cellulose, less energy than other types of foods, and often toxic compounds.Jones, S., Martin, R., & Pilbeam, D. (1 ...
s, while the others are all primarily
frugivore A frugivore ( ) is an animal that thrives mostly on raw fruits or succulent fruit-like produce of plants such as roots, shoots, nuts and seeds. Approximately 20% of mammalian herbivores eat fruit. Frugivores are highly dependent on the abundance ...
s, although the common chimpanzee hunts for meat. Foraging behaviour is correspondingly variable. In November 2023, scientists reported, for the first time, evidence that groups of
primate Primates is an order (biology), order of mammals, which is further divided into the Strepsirrhini, strepsirrhines, which include lemurs, galagos, and Lorisidae, lorisids; and the Haplorhini, haplorhines, which include Tarsiiformes, tarsiers a ...
s, including apes, and, particularly
bonobo The bonobo (; ''Pan paniscus''), also historically called the pygmy chimpanzee (less often the dwarf chimpanzee or gracile chimpanzee), is an endangered great ape and one of the two species making up the genus ''Pan (genus), Pan'' (the other bei ...
s, are capable of cooperating with each other.


Diet

Apart from humans and gorillas, apes eat a predominantly
frugivorous A frugivore ( ) is an animal that thrives mostly on raw fruits or succulent fruit-like produce of plants such as roots, shoots, nuts and seeds. Approximately 20% of mammalian herbivores eat fruit. Frugivores are highly dependent on the abundance ...
diet, mostly fruit, but supplemented with a variety of other foods. Gorillas are predominantly folivorous, eating mostly stalks, shoots, roots and leaves with some fruit and other foods. Non-human apes usually eat a small amount of raw animal foods such as insects or eggs. In the case of humans, migration and the invention of hunting tools and cooking has led to an even wider variety of foods and diets, with many human diets including large amounts of cooked
tuber Tubers are a type of enlarged structure that plants use as storage organs for nutrients, derived from stems or roots. Tubers help plants perennate (survive winter or dry months), provide energy and nutrients, and are a means of asexual reproduc ...
s (
root In vascular plants, the roots are the plant organ, organs of a plant that are modified to provide anchorage for the plant and take in water and nutrients into the plant body, which allows plants to grow taller and faster. They are most often bel ...
s) or
legume Legumes are plants in the pea family Fabaceae (or Leguminosae), or the fruit or seeds of such plants. When used as a dry grain for human consumption, the seeds are also called pulses. Legumes are grown agriculturally, primarily for human consum ...
s. Other food production and processing methods including animal husbandry and industrial refining and processing have further changed human diets. Humans and other apes occasionally eat other primates. Some of these primates are now close to extinction with habitat loss being the underlying cause.


Cognition

All the non-human hominoids are generally thought of as highly intelligent, and scientific study has broadly confirmed that they perform very 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 (; 21 January 1887 – 11 June 1967) was a German psychologist and phenomenologist who, like Max Wertheimer and Kurt Koffka, contributed to the creation of Gestalt psychology. During the Nazi regime in Germany, he pro ...
demonstrated exceptional
problem-solving Problem solving is the process of achieving a goal by overcoming obstacles, a frequent part of most activities. Problems in need of solutions range from simple personal tasks (e.g. how to turn on an appliance) to complex issues in business an ...
abilities in chimpanzees, which Köhler attributed to
insight Insight is the understanding of a specific causality, cause and effect within a particular context. The term insight can have several related meanings: *a piece of information *the act or result of understanding the inner nature of things or of se ...
. The use of tools has been repeatedly demonstrated; more recently, the manufacture of tools has been documented, both in the wild and in laboratory tests.
Imitation Imitation (from Latin ''imitatio'', "a copying, imitation") is a behavior whereby an individual observes and replicates another's behavior. Imitation is also a form of learning that leads to the "development of traditions, and ultimately our cu ...
is much more easily demonstrated in "great apes" than in other primate species. Almost all the studies in animal language acquisition have been done 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 Animal culture can be defined as the ability of non-human animals to learn and transmit behaviors through processes of social or cultural learning. Culture is increasingly seen as a process, involving the social transmittance of behavior among p ...
.


Threats and conservation

All non-human hominoids are rare and threatened with
extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
ion. The eastern hoolock gibbon is the least threatened, only being vulnerable to extinction. Five gibbon species are
critically endangered An IUCN Red List critically endangered (CR or sometimes CE) species is one that has been categorized by the International Union for Conservation of Nature as facing an extremely high risk of extinction in the wild. As of December 2023, of t ...
, as are all species of orangutan and gorilla. The remaining species of gibbon, the bonobo, and all four subspecies of chimpanzees are
endangered An endangered species is a species that is very likely to become extinct in the near future, either worldwide or in a particular political jurisdiction. Endangered species may be at risk due to factors such as habitat loss, poaching, inv ...
. The chief threat to most of the endangered species is loss of tropical
rainforest Rainforests are forests characterized by a closed and continuous tree Canopy (biology), canopy, moisture-dependent vegetation, the presence of epiphytes and lianas and the absence of wildfire. Rainforests can be generally classified as tropi ...
habitat, though some populations are further imperiled by hunting for
bushmeat Bushmeat is meat from wildlife species that are Hunting, hunted for human consumption. Bushmeat represents a primary source of animal protein and a cash-earning commodity in poor and rural communities of humid tropical forest regions of the worl ...
. The great apes of Africa are also facing threat from the
Ebola virus ''Orthoebolavirus zairense'' or Zaire ebolavirus, more commonly known as Ebola virus (; EBOV), is one of six known species within the genus ''Ebolavirus''. Four of the six known ebolaviruses, including EBOV, cause a severe and often fatal vira ...
. Currently considered to be the greatest threat to survival of African apes,
Ebola Ebola, also known as Ebola virus disease (EVD) and Ebola hemorrhagic fever (EHF), is a viral hemorrhagic fever in humans and other primates, caused by ebolaviruses. Symptoms typically start anywhere between two days and three weeks after in ...
infection is responsible for the death of at least one third of all gorillas and chimpanzees since 1990. All the species of
great apes The Hominidae (), whose members are known as the great apes or hominids (), are a taxonomic family of primates that includes eight extant species in four genera: '' Pongo'' (the Bornean, Sumatran and Tapanuli orangutan); '' Gorilla'' (the ...
in Africa are considered endangered. Hunting, logging, agricultural expansion and mining are among the main threats.


See also

* *
Great Ape Project The Great Ape Project (GAP), founded in 1993, is an international organization of primatologists, anthropologists, ethicists, and others who advocate a United Nations Declaration of the Rights of Great Apes that would confer basic legal rights ...
* Great Apes Survival Partnership *
International Primate Day International Primate Day, September 1, is an annual educational observance event organized since 2005 largely by British-based Animal Defenders International (ADI) and supported annually by various primate-oriented advocacy organizations, speak ...
* Kinshasa Declaration on Great Apes *
List of individual apes This is a list of non-human apes of encyclopedic interest. It includes individual chimpanzees, gorillas, orangutans, bonobos, and gibbons that are in some way famous or notable. Actors * J. Fred Muggs (a chimpanzee born 1952) was a "co-host" ...
(for notable non-fictional non-human apes) * List of primates by population


Notes


References


Literature cited

* * * * *


External links

* * * Agreement between cladograms based on molecular and anatomical data.
Human Timeline (Interactive)
Smithsonian,
National Museum of Natural History The National Museum of Natural History (NMNH) is a natural history museum administered by the Smithsonian Institution, located on the National Mall in Washington, D.C., United States. It has free admission and is open 364 days a year. With 4.4 ...
(August 2016). {{Authority control Extant Chattian first appearances Taxa named by John Edward Gray