Hominoidea
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Apes (collectively Hominoidea ) are a
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyly, monophyletic – that is, composed of a common ancestor and all its lineage (evolution), lineal descendants – on a phylogenetic tree. ...
of
Old World The "Old World" is a term for Afro-Eurasia that originated in Europe , after Europeans became aware of the existence of the Americas. It is used to contrast the continents of Africa, Europe, and Asia, which were previously thought of by thei ...
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 monkeys (Platyrrhini) and Catarrhi ...
s native to sub-Saharan
Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...
and
Southeast Asia Southeast Asia, also spelled South East Asia and South-East Asia, and also known as Southeastern Asia, South-eastern Asia or SEA, is the geographical United Nations geoscheme for Asia#South-eastern Asia, south-eastern region of Asia, consistin ...
(though they were more widespread in Africa, most of Asia, and as well as Europe in prehistory), which together with its sister group
Cercopithecidae Old World monkey is the common English name for a family (biology), family of primates known wikt:taxonomy, taxonomically as the Cercopithecidae (). Twenty-four genus, genera and 138 species are recognized, making it the largest primate family. ...
form the catarrhine
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyly, monophyletic – that is, composed of a common ancestor and all its lineage (evolution), lineal descendants – on a phylogenetic tree. ...
, cladistically making them
monkey Monkey is a common name that may refer to most mammals of the infraorder Simiiformes, also known as the simians. Traditionally, all animals in the group now known as simians are counted as monkeys except the apes, which constitutes an incomple ...
s (though this is the subject of much debate). 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 and black ape), and is thus not equivalent to the scientific
taxon In biology, a taxon (back-formation from ''Taxonomy (biology), taxonomy''; plural 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 ...
Hominoidea. There are two extant branches of the superfamily Hominoidea: the
gibbon 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 rainforest from eastern Bangladesh ...
s, or lesser apes; and the
hominid 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, ...
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 rainforest from eastern Bangladesh ...
, 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 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 in height, a ...
, all native to
Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa Africa is ...
. They are highly arboreal and
bipedal Bipedalism is a form of terrestrial locomotion Terrestrial locomotion has evolved as animal Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material ...
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 Hominidae, 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 ...
s and their subspecies, two extant species of
gorilla Gorillas are herbivorous, predominantly ground-dwelling 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 fi ...
s and their subspecies, two extant species of panins ( bonobos and
chimpanzees The chimpanzee (''Pan troglodytes''), also known as simply the chimp, is a species of Hominidae, great ape native to the forest and savannah of tropical Africa. It has four confirmed subspecies and a fifth proposed subspecies. When its close r ...
) and their subspecies, and
human Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedality, bipedalism and exceptional cognitive skills due to a large and complex Human brain, brain. This has enabled the development of ad ...
s in a single extant subspecies. Except for gorillas and humans, hominoids are agile climbers of
tree In botany, a tree is a perennial plant with an elongated Plant stem, stem, or trunk (botany), trunk, usually supporting branches and leaves. In some usages, the definition of a tree may be narrower, including only woody plants with secondar ...
s. 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. Humans are by far the most numerous of the hominoid species, in fact outnumbering all other primates by a factor of several thousand to one. All non-human hominoids are rare and threatened with
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the Endling, last individual of the species, although the Functional ext ...
ion. The eastern hoolock gibbon is the least threatened, only being vulnerable to extinction. Five gibbon species are critically endangered, as are all species of orangutan and gorilla. The remaining species of gibbon, the bonobo, and all four species 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 and invas ...
. The chief threat to most of the endangered species is loss of tropical
rainforest Rainforests are characterized by a closed and continuous tree Canopy (biology), canopy, moisture-dependent vegetation, the presence of epiphytes and lianas and the absence of wildfire. Rainforest can be classified as tropical rainforest or tem ...
habitat, though some populations are further imperiled by hunting for
bushmeat Bushmeat is meat from wildlife species that are Hunting, hunted for human consumption, most often referring to the meat of Game (hunting), game in Africa. Bushmeat represents a primary source of animal protein and a cash-earning commodity for in ...
. The great apes of Africa are also facing threat from the
Ebola virus ''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 viral hemorrhagic fever, hemorrhag ...
. 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 becom ...
infection is responsible for the death of at least one third of all gorillas and chimpanzees since 1990.


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 the simians. Traditionally, all animals in the group now known as simians are counted as monkeys except the apes, which constitutes an incomple ...
" 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 (biology), family of primates that includes eight Neontology#Extant taxa versus extinct taxa, extant species in four Genus, genera: ''Orangutan, ...
s,'' the family of
great ape 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, ...
s. Both terms were introduced by Gray (1825). The term ''
hominin The Hominini form a Tribe (biology), taxonomic tribe of the subfamily Homininae ("hominines"). Hominini includes the extant genera ''Homo'' (humans) and ''Pan (genus), Pan'' (chimpanzees and bonobos) and in standard usage excludes the genus '' ...
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 In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...
of monkeys: Apes emerged as a sister group of Old World Monkeys in the catarrhines, which are a sister group of New World Monkeys. Therefore, cladistically, apes, catarrhines and related contemporary extinct groups such as Parapithecidaea 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 ...
, 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 In biology, a species is the basic unit of Taxonomy (biology), classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of ...
. 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 In taxonomy, a group is paraphyletic if it consists of the group's last common ancestor and most of its descendants, excluding a few monophyletic subgroups. The group is said to be paraphyletic ''with respect to'' the excluded subgroups. In ...
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 A vernacular or vernacular language is in contrast with a "standard language". It refers to the language or dialect that is spoken by people that are inhabiting a particular country or region. The vernacular is typically the native language, n ...
"apes" does not include humans, but phylogenetically, humans (''
Homo ''Homo'' () is the genus that emerged in the (otherwise extinct) genus ''Australopithecus'' that encompasses the extant species ''Homo sapiens'' (Human, modern humans), plus several extinct species classified as either ancestral to or closely ...
'') form part of the family Hominidae within Hominoidaea. 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 cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population Population typically refers to the number of ...
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 form a Tribe (biology), taxonomic tribe of the subfamily Homininae ("hominines"). Hominini includes the extant genera ''Homo'' (humans) and ''Pan (genus), Pan'' (chimpanzees and bonobos) and in standard usage excludes the genus '' ...
" 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 (biology), family of primates that includes eight Neontology#Extant taxa versus extinct taxa, extant species in four Genus, genera: ''Orangutan, ...
". 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'', were once thought to be
hominin The Hominini form a Tribe (biology), taxonomic tribe of the subfamily Homininae ("hominines"). Hominini includes the extant genera ''Homo'' (humans) and ''Pan (genus), Pan'' (chimpanzees and bonobos) and in standard usage excludes the genus '' ...
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 (genus), Pan'' (chimpanzee and bonobo) genera of Hominini. Due to complex hybrid speciation, it is not currently possi ...
(CHLCA).


Taxonomic classification and phylogeny

Genetic analysis combined with fossil evidence indicates that hominoids diverged from the
Old World monkey Old World monkey is the common English name for a family (biology), family of primates known wikt:taxonomy, taxonomically as the Cercopithecidae (). Twenty-four genus, genera and 138 species are recognized, making it the largest primate family. ...
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 a species of orangutan endemic to the island of Borneo Borneo (; id, Kalimantan) is the List of islands by area, third-largest island in the world and the largest in Asia. At the geographic ce ...
, ''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 The Tapanuli orangutan (''Pongo tapanuliensis'') is a species of orangutan restricted to South Tapanuli Regency, South Tapanuli in the island of Sumatra in Indonesia. It is one of three known species of orangutan, alongside the Sumatran orangutan ...
, ''P. tapanuliensis'' *** Genus ''
Gorilla Gorillas are herbivorous, predominantly ground-dwelling 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 fi ...
'': gorillas ****
Western gorilla The western gorilla (''Gorilla gorilla'') is a great ape found in Africa, one of two species of the Hominidae, hominid genus ''Gorilla''. Large and robust with males weighing around , the hair is significantly lighter in color than the eastern g ...
, ''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 3,800 eastern lowland gorillas or Grauer ...
, ''G. beringei'' *** Genus ''
Homo ''Homo'' () is the genus that emerged in the (otherwise extinct) genus ''Australopithecus'' that encompasses the extant species ''Homo sapiens'' (Human, modern humans), plus several extinct species classified as either ancestral to or closely ...
'': humans ****
Human Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedality, bipedalism and exceptional cognitive skills due to a large and complex Human brain, brain. This has enabled the development of ad ...
, ''H. sapiens'' *** Genus '' Pan'': chimpanzees ****
Chimpanzee The chimpanzee (''Pan troglodytes''), also known as simply the chimp, is a species of great ape native to the forest and savannah of tropical Africa Although tropical Africa is mostly familiar to the Western world, West for its rainfore ...
, ''P. troglodytes'' ****
Bonobo The bonobo (; ''Pan paniscus''), also historically called the pygmy chimpanzee and 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 be ...
, ''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 rainforest from eastern Bangladesh ...
: 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 ''Bu ...
'' **** 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 or southern grey gibbon, ''H. muelleri'' ****
Silvery gibbon The silvery gibbon (''Hylobates moloch''), also known as the Javan gibbon, is a primate in the gibbon family Hylobatidae. It is endemic to the Indonesian island of Java, where it inhabits undisturbed rainforests up to an altitude of . It is liste ...
, ''H. moloch'' ****
Pileated gibbon The pileated gibbon (''Hylobates pileatus'') is a primate Primates are a diverse order (biology), order of mammals. They are divided into the Strepsirrhini, strepsirrhines, which include the lemurs, galagos, and lorisids, and the Haplorhin ...
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 C ...
, ''H. hoolock'' **** Eastern hoolock gibbon, ''H. leuconedys'' **** Skywalker hoolock gibbon, ''H. tianxing'' *** Genus ''
Symphalangus The siamang (, ; ''Symphalangus syndactylus'') is an 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 in height, a ...
'' ****
Siamang The siamang (, ; ''Symphalangus syndactylus'') is an 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 in height, a ...
, ''S. syndactylus'' *** Genus ''
Nomascus ''Nomascus'' is the second-most speciose genus of gibbons (family Hylobatidae). Originally, this genus was a subgenus of ''Hylobates'', and all individuals were considered one species, ''Hylobates concolor''. Species within ''Nomascus'' are char ...
'' **** Northern buffed-cheeked gibbon, ''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 genus ...
, ''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 The yellow-cheeked gibbon (''Nomascus gabriellae''), also called the golden-cheeked gibbon, the yellow-cheeked crested gibbon, the golden-cheeked crested gibbon, the red-cheeked gibbon, or the buffed-cheeked gibbon, is a species of gibbon native ...
, ''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 — 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 (; ) 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 hypothesized relationships is typically ...
, the science of classifying living things strictly according to their lines of descent. Today, there are eight extant
genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of hominoids. They are the four genera in the family Hominidae, namely ''
Homo ''Homo'' () is the genus that emerged in the (otherwise extinct) genus ''Australopithecus'' that encompasses the extant species ''Homo sapiens'' (Human, modern humans), plus several extinct species classified as either ancestral to or closely ...
'', '' Pan'', ''
Gorilla Gorillas are herbivorous, predominantly ground-dwelling 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 fi ...
'', 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'' [formerly ''Bu ...
'', ''
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 gibbons (family Hylobatidae). Originally, this genus was a subgenus of ''Hylobates'', and all individuals were considered one species, ''Hylobates concolor''. Species within ''Nomascus'' are char ...
'' and ''
Symphalangus The siamang (, ; ''Symphalangus syndactylus'') is an 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 in height, a ...
''. (The two subspecies of hoolock gibbons were recently moved from the genus ''Bunopithecus'' 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 his Nobility#Ennoblement, ennoblement in 1761 as Carl von Linné#Blunt, Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalise ...
, 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 monkeys (Platyrrhini) and Catarrhi ...
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 ' (originally in Latin language, Latin written ' with the Orthographic ligature, ligature æ) is one of the major works of the Sweden, Swedish botanist, zoologist and physician Carl Linnaeus (1707–1778) and introduce ...
'' 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 natural history, 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 ...
dated 25 February 1747, Linnaeus wrote: Accordingly,
Johann Friedrich Blumenbach Johann Friedrich Blumenbach (11 May 1752 – 22 January 1840) was a German physician A physician (American English), medical practitioner (English in the Commonwealth of Nations, Commonwealth English), medical doctor, or simply doctor, ...
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 primate Primates are a diverse order (biology), order of mammals. They are divided into the Strepsirrhini, strepsirrhines, which include the lemurs, galagos, and lorisids, and the Haplo ...
(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". Cuvier ...
. Some elevated the distinction to the level of order. 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 natural history, naturalist Charles Darwin, first published in 1871, which applies evolutionary theory to human evolution, and details his theory of sexual selection, ...
'',
Charles Darwin Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all species of life have descended ...
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 Arboreal locomotion is the locomotion of animals in trees. In habitats in which trees are present, animals have evolved to move in them. Some ...
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 Animal locomotion, 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, but others are exclusively arboreal. Th ...
lifestyle. Generally smaller than the African apes, the largest gibbon, the
siamang The siamang (, ; ''Symphalangus syndactylus'') is an 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 in height, a ...
, weighs up to ; in comparison, the smallest "great ape", the
bonobo The bonobo (; ''Pan paniscus''), also historically called the pygmy chimpanzee and 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 be ...
, is . The superfamily Hominoidea falls within the parvorder
Catarrhini The parvorder Catarrhini , catarrhine monkeys, Old World anthropoids, or Old World monkeys, consisting of the Cercopithecoidea and apes (Hominoidea). In 1812, Geoffroy grouped those two groups together and established the name Catarrhini, "Ol ...
, 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; 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 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, ...
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 (plural 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 ...
; 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 primate Primates are a diverse order (biology), order of mammals. They are divided into the Strepsirrhini, strepsirrhines, which inc ...
) and, later, to developing balance in a
bipedal Bipedalism is a form of terrestrial locomotion Terrestrial locomotion has evolved as animal Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material ...
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.


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 ...
and Parapithecidaea, 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 the simians. Traditionally, all animals in the group now known as simians are counted as monkeys except the apes, which constitutes an incomple ...
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 (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 rainforest from eastern Bangladesh ...
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.


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 on 3 April 1934), formerly Baroness Jane van Lawick-Goodall, is an English Primatology, primatologist and Anthropology, anthropologist. Seen as the world's foremost expert on Common ...
,
Dian Fossey Dian Fossey (, January 16, 1932 – ) was an American primatologist and Conservationist movement, conservationist known for undertaking an extensive study of mountain gorilla groups from 1966 until her murder in 1985. She studied them daily in t ...
and Birute Galdikas. 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 foliovores, 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 and ...
s, although the common chimpanzee hunts for meat. Foraging behaviour is correspondingly variable.


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 and ...
diet, mostly fruit, but supplemented with a variety of other foods. Gorillas are predominately 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 used as storage organs for nutrients in some plants. They are used for the plant's perennation (survival of the winter or dry months), to provide energy and nutrients for regrowth during the next growing s ...
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 b ...
s) or
legume A legume () is a plant in the family Fabaceae (or Leguminosae), or the fruit or seed of such a plant. When used as a dry grain, the seed is also called a pulse. Legumes are grown agriculturally, primarily for human consumption, for livestock for ...
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 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 that leads to the "development of traditions, and ultimately our culture. I ...
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 pe ...
.


See also

* ''Dawn of Humanity'' (2015 PBS film) * World Declaration on Great Apes from the
Great Ape Project The Great Ape Project (GAP), founded in 1993, is an international organization of primatologists, anthropologist An anthropologist is a person engaged in the practice of anthropology. Anthropology is the study of aspects of humans within past ...
* International Primate Day *
List of individual apes This is a list of non-human Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedality, bipedalism and exceptional cognitive skills due to a large and complex Human brain, brain. This has ...
(for notable non-fictional non-human apes) * List of fictional primates * 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 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. In 2021, with 7. ...
(August 2016). {{Authority control Extant Chattian first appearances Taxa named by John Edward Gray