Hominidae

The Hominidae (/hmndi/), whose members are known as great apes[note 1] 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 eastern and western gorilla; Pan, the common
Hominidae[1]
chimpanzee and the bonobo; and Homo, which includes modern Humans and its extinct relatives (e.g., the Neanderthal), and ancestors, such as Homo Temporal range: Miocene -
erectus.[1] Present , 23.030 Ma

Pre OS D C P T J K Pg N
Several revisions in classifying the great apes have caused the use of the term
"hominid" to vary over time. Its original meaning referred only to humans
(Homo) and their closest non-extant relatives. That restrictive meaning has now been largely assumed by the term "hominin", which comprises all
members of the human clade after the split from the chimpanzees (Pan). (See below, for a fuller discussion of related and very similar terms, at
Terminology.) The current, 21st-century meaning of "hominid" includes all the great apes including humans. Usage still varies, however, and some
scientists and laypersons still use "hominid" in the original restrictive sense; the scholarly literature generally shows the traditional usage until around the
turn of the 21st century.[4]

Within the taxon Hominidae, a number of extant and known extinct, that is, fossil, genera are grouped with the humans, chimpanzees, and gorillas in the
subfamily Homininae; others with orangutans in the subfamily Ponginae (see classification graphic below). The most recent common ancestor of all
Hominidae lived roughly 14 million years ago,[5] when the ancestors of the orangutans speciated from the ancestral line of the other three genera.[6]
Those ancestors of the family Hominidae had already speciated from the family Hylobatidae (the gibbons), perhaps 15 million to 20 million years
ago.[6][7]

Contents
1 History Two hominids: A human (Homo
1.1 Taxonomic history
1.1.1 Terminology sapiens) and a chimpanzee (Pan
1.1.2 Extant and fossil relatives of humans troglodytes)
1.2 Classification
Scientific classification
1.2.1 Evolutionary tree
1.2.2 Extant Kingdom: Animalia
1.2.3 Fossil
Phylum: Chordata
2 Physical description
Class: Mammalia
3 Legal status
4 Conservation Order: Primates
5 See also Suborder: Haplorhini
6 Notes
Infraorder: Simiiformes
7 References
Superfamily: Hominoidea
8 External links
Family: Hominidae
Gray, 1825
History Type genus
In the early Miocene, about 22 million years ago, there were many species of arboreally adapted primitive catarrhines from East Africa; the variety Homo
suggests a long history of prior diversification. Fossils at 20 million years ago include fragments attributed to Victoriapithecus, the earliest Old World
Linnaeus, 1758
monkey. Among the genera thought to be in the ape lineage leading up to 13 million years ago are Proconsul, Rangwapithecus, Dendropithecus,
Limnopithecus, Nacholapithecus, Equatorius, Nyanzapithecus, Afropithecus, Heliopithecus, and Kenyapithecus, all from East Africa. Subfamily

At sites far distant from East Africa, the presence of other generalized non-cercopithecids, that is, non-monkey primates, of middle Miocene age Ponginae
Otavipithecus from cave deposits in Namibia, and Pierolapithecus and Dryopithecus from France, Spain and Austriais further evidence of a wide
Homininae
diversity of ancestral ape forms across Africa and the Mediterranean basin during the relatively warm and equable climatic regimes of the early and
Ouranopithecus
middle Miocene. The most recent of these far-flung Miocene apes (hominoids) is Oreopithecus, from the fossil-rich coal beds in northern Italy and dated
to 9 million years ago. Kenyapithecinae

Molecular evidence indicates that the lineage of gibbons (family Hylobatidae), the lesser apes, diverged from that of the great apes some 1812 million Synonyms
years ago, and that of orangutans (subfamily Ponginae) diverged from the other great apes at about 12 million years. There are no fossils that clearly
document the ancestry of gibbons, which may have originated in a still-unknown South East Asian hominoid population; but fossil proto-orangutans, Pongidae Elliot, 1913
dated to around 10 million years ago, may be represented bySivapithecus from
India and Griphopithecus from Turkey.[8]
Human timeline
Species close to the last common ancestor of gorillas, chimpanzees and view discuss
humans may be represented by Nakalipithecus fossils found in Kenya and 0 Homo sapiens Modern humans
P Modern speech
Ouranopithecus found in Greece. Molecular evidence suggests that between 8
l Neanderthal Earliest clothes
and 4 million years ago, first the gorillas (genus Gorilla), and then the e Earliest cooking
chimpanzees (genus Pan) split off from the line leading to the humans. Human -1 i Earliest in Europe
s Homo erectus
DNA is approximately 98.4% identical to that of chimpanzees when t Earliest fire use
o Earliest exit from
comparing single nucleotide polymorphisms (see human evolutionary
-2 c Africa
genetics).[9] The fossil record, however, of gorillas and chimpanzees is limited; e Homo habilis
both poor preservationrain forest soils tend to be acidic and dissolve bone n
e
Sumatran orangutan (Pongo abelii) and sampling bias probably contribute most to this problem. -3
Earliest stone tools
Other hominins probably adapted to the drier environments outside the African Australopithecus
P
equatorial belt; and there they encountered antelope, hyenas, elephants and other forms becoming adapted to surviving in the -4 l Earliest bipedal
East African savannas, particularly the regions of the Sahel and the Serengeti. The wet equatorial belt contracted after about 8 i
o Ardipithecus H
c
Human-like
 c
million years ago, and there is very little fossil evidence for the divergence of the hominin -5 e Human-like o
n apes
lineage from that of gorillas and chimpanzeeswhich split was thought to have occurred m
e
around that time. The earliest fossils argued by some to belong to the human lineage are Separation from
-6 Orrorin i chimpanzees
Sahelanthropus tchadensis (7 Ma) and Orrorin tugenensis (6 Ma), followed by Ardipithecus
(5.54.4 Ma), with speciesAr. kadabba and Ar. ramidus. n

-7 M Sahelanthropus i Possibly bipedal

i
Taxonomic history o d
c
s
-8 e
Terminology n
e
A reconstruction of a The classification of the great apes has been revised several times in the last few decades;
female Australopithecus these revisions have led to a varied use of the word "hominid" over time. The original -9 Ouranopithecus Separation from
afarensis (National gorillas
meaning of the term referred to only humans and their closest relativeswhat is now the
Museum of Natural
modern meaning of the term "hominin". And the meaning of the taxon Hominidae changed Nakalipithecus
History) -10 Earlier apes
gradually, leading to a different (modern) usage of "hominid" that today includes all the Axis scale: millions of years.
great apes including humans. Also see: Life timeline and Nature timeline

The term hominid is easily confused with a number of very similar words:

A hominoid, commonly called an ape, is a member of the superfamilyHominoidea: extant members are thegibbons (lesser apes,
family Hylobatidae) and the hominids.
A hominid is a member of the family Hominidae, the great apes:orangutans, gorillas, chimpanzees, and humans.
A hominine is a member of the subfamilyHomininae: gorillas, chimpanzees, and humans (excludesorangutans).
A hominin is a member of the tribeHominini: Chimpanzees and humans.
A homininan is a member of the subtribeHominina of the tribe Hominini: that is, modern humans and their closest relativesafter their
split from chimpanzees.
A human is a member of the genusHomo, of which Homo sapiens is the only extant species, and within that Homo sapiens sapiensis
the only surviving subspecies.
(See below, a graphic of the taxonomic classifications atEvolutionary tree of the Hominoidea.)

Humans are one of the four extant

hominid genera.

Hominoidea (hominoids) (20.4 Mya) Hylobatidae

Hominidae (hominids) (15.7 Mya) Ponginae

Homininae (hominines) (8.8 Mya)

Gorillinae

Panina
Hominini (hominins) (6.3 Mya)
Hominina (homininans)

For each clade it is indicated approximately when newer extant clades emerged. Some (older) texts will refer to Homonini as the Hominina branch. Many scientists, including paleoanthropologists,
continue to use the term "hominid" to mean humans and their direct and near
-direct bipedal ancestors.

Extant and fossil relatives of humans

As mentioned, Hominidae was originally the name given to the family of humans and their (extinct) close relatives, with the other great apes (that is, the
orangutans, gorillas, and chimpanzees) all being placed in a separate family, the Pongidae. However, that definition eventually made Pongidae
paraphyletic because at least one great ape species (the chimpanzees) proved to be more closely related to humans than to other great apes. Most
taxonomists today encourage monophyletic groupsthis would require, in this case, the use of Pongidae to be restricted to just one closely related
grouping. Thus, many biologists now assign Pongo (as the subfamily Ponginae) to the family Hominidae. The taxonomy shown here follows the
monophyletic groupings according to the modern understanding of human and great ape relationships.

Humans and close relatives including the tribes Hominini and Gorillini form the subfamily Homininae (see classification graphic below). (A few
researchers go so far as to refer the chimpanzees and the gorillas to the genus Homo along with humans.) [10][11][12] But, it is those fossil relatives more
closely related to humans than the chimpanzees that represent the especially close members of the human family, and without necessarily assigning
subfamily or tribal categories.[13]

Many extinct hominids have been studied to help understand the relationship between modern humans and the other extant hominids. Some of the extinct
members of this family include Gigantopithecus, Orrorin, Ardipithecus, Kenyanthropus, and the australopithecines Australopithecus and
Paranthropus.[14]
A model of a modern human hominid
The exact criteria for membership in the tribe Hominini under the current understanding of human origins are not clear, but the taxon generally includes
skull (or hominin skull)
those species that share more than 97% of their DNA with the modern human genome, and exhibit a capacity for language or for simple cultures beyond
their 'local family' or band. The theory of mind conceptincluding such faculties as empathy, attribution of mental state, and even empathetic deception
is a controversial criterion; it distinguishes the adult human alone among the hominids. Humans acquire this capacity after about four years of age, whereas it has not been proven (nor has it been
[15] This is also the case for someNew World monkeys outside the family of great apes, as, for example, thecapuchin monkeys.
disproven) that gorillas or chimpanzees ever develop a theory of mind.
However, even without the ability to test whether early members of the Hominini (such as Homo erectus, Homo neanderthalensis, or even the
australopithecines) had a theory of mind, it is difficult to ignore similarities seen in their living cousins. Orangutans have shown the development of
culture comparable to that of chimpanzees,[16] and some say the orangutan may also satisfy those criteria for the theory of mind concept. These scientific
debates take on political significance for advocates ofgreat ape personhood.

Below is a cladogram with extinct species.[17] It is indicated approximately how many million years ago (Mya) the clades diver
ged into newer clades.[18]

Kenyapithecus (13 Mya)

Sivapithecus (9)

Ponginae (14) Crown Ponginae

A fossil hominid exhibit at The
Ankarapithecus (9) Museum of Osteology, Oklahoma
City, Oklahoma
Giganthopithecus (0.1)

Khoratpithecus (7)

Pierolapithecus (11)
Hominidae (17.61) (12)
(13)
Hispanopithecus (10)

Lufengpithecus (7)

Ouranopithecus (8)

Hominini

Crown Homininae (9.68)

Homininae (10) Crown Gorilla
Gorilla
Chororapithecus ()

Nakalipithecus (10)

Samburupithecus (9)

Classification

Evolutionary tree

Evolutionary tree of theHominoidea (emphasis on family

Hominidae): after an initial separation from the main line by the
Hylobatidae (gibbons) some 18 million years ago, the line of
Ponginae broke away, leading to the orangutan; later, the Homininae
split into the tribes Hominini (led to humans and chimpanzees) and
Gorillini (led to gorillas).

Extant
[1]
There are eight living species of great ape which are classified in fourgenera. The following classification is commonly accepted:

[1]
Family Hominidae: humans and other great apes; extinct genera and species excluded

Subfamily Ponginae

Tribe Pongini

Genus Pongo
Skulls of an orangutan and a gorilla
Bornean orangutan, Pongo pygmaeus

Pongo pygmaeus pygmaeus

Pongo pygmaeus morio
Pongo pygmaeus wurmbii
Sumatran orangutan, Pongo abelii
Tapanuli orangutan, Pongo tapanuliensis[19]
Subfamily Homininae

Tribe Gorillini

Genus Gorilla

Western gorilla, Gorilla gorilla

Western lowland gorilla, Gorilla gorilla gorilla

 Cross River gorilla, Gorilla gorilla diehli
Eastern gorilla, Gorilla beringei

Mountain gorilla, Gorilla beringei beringei

Eastern lowland gorilla, Gorilla beringei graueri
Tribe Hominini

Subtribe Panina

Genus Pan

Chimpanzee (common chimpanzee),Pan troglodytes

Central chimpanzee, Pan troglodytes troglodytes

Western chimpanzee, Pan troglodytes verus Human and chimp skulls and brains
Nigeria-Cameroon chimpanzee, Pan troglodytes ellioti (not to scale), as illustrated in
Eastern chimpanzee, Pan troglodytes schweinfurthii Gervais' Histoire naturelle des
Bonobo (pygmy chimpanzee),Pan paniscus mammifres
Subtribe Hominina

Genus Homo

Human, Homo sapiens

Anatomically modern human, Homo sapiens sapiens

Fossil
In addition to the extant species and subspecies, archaeologists, paleontologists, and anthropologists have discovered and classified numerous extinct
great ape species as below, based on the taxonomy shown.[20]

Family Hominidae

Ouranopithecus)

Ouranopithecus macedoniensis
Otavipithecus

Otavipithecus namibiensis
Morotopithecus

Morotopithecus bishopi
Subfamily Ponginae[21]

Tribe Lufengpithecini

Lufengpithecus
A reconstruction of Pierolapithecus
Lufengpithecus lufengensis catalaunicus
Lufengpithecus keiyuanensis
Lufengpithecus hudienensis
Tribe Sivapithecini

Ankarapithecus

Ankarapithecus meteai
Sivapithecus

Sivapithecus brevirostris
Sivapithecus punjabicus
Sivapithecus parvada
Sivapithecus sivalensis
Sivapithecus indicus
Gigantopithecus

Gigantopithecus bilaspurensis
Gigantopithecus blacki
Gigantopithecus giganteus
Tribe Pongini

Khoratpithecus

Khoratpithecus ayeyarwadyensis Replica of the skull sometimes

Khoratpithecus piriyai known as "Nutcracker Man", found
Khoratpithecus chiangmuanensis by Mary Leakey.
Pongo (orangutans)

Pongo hooijeri

Subfamily Homininae[22][23]

Pierolapithecus

Pierolapithecus catalaunicus
Udabnopithecus

Udabnopithecus garedziensis
Tribe Dryopithecini

Oreopithecus (placement disputed)

Oreopithecus bambolii
Nakalipithecus

Nakalipithecus nakayamai
Anoiapithecus
 Anoiapithecus brevirostris
Hispanopithecus

Hispanopithecus laietanus
Hispanopithecus crusafonti
Dryopithecus

Dryopithecus wuduensis
Dryopithecus fontani
Dryopithecus brancoi
Dryopithecus laietanus
Dryopithecus crusafonti
Rudapithecus

Rudapithecus hungaricus
Samburupithecus

Samburupithecus kiptalami
Tribe Gorillini

Chororapithecus (placement debated)

Chororapithecus abyssinicus
Tribe Hominini

Graecopithecus [24]

Graecopithecus freybergi
Sahelanthropus

Sahelanthropus tchadensis
Orrorin

Orrorin tugenensis
Subtribe Hominina

Ardipithecus

Ardipithecus ramidus
Ardipithecus kadabba
Kenyanthropus

Kenyanthropus platyops
Praeanthropus[25]

Praeanthropus bahrelghazali
Praeanthropus anamensis
Praeanthropus afarensis
Australopithecus

Australopithecus africanus
Australopithecus garhi
Australopithecus sediba
Australopithecus deyiremeda
Paranthropus

Paranthropus aethiopicus
Paranthropus robustus
Paranthropus boisei
Homo immediate ancestors of modernhumans

Homo gautengensis
Homo rudolfensis
Homo naledi
Homo habilis
Homo floresiensis
Homo erectus
Homo ergaster
Homo antecessor
Homo heidelbergensis
Homo cepranensis
Homo helmei
Homo palaeojavanicus
Homo tsaichangensis
Denisovans (scientific name has not yet been assigned)
Homo neanderthalensis
Homo rhodesiensis
Homo sapiens

Homo sapiens idaltu

Archaic Homo sapiens
[26] of modern humans and Denisovans[27] )
Red Deer Cave people (scientific name has not yet been assigned; perhaps a race of modern humans or a hybrid

Physical description
The great apes are large, tailless primates, with the smallest living species being the bonobo at 3040 kilograms in weight, and the largest being the eastern gorillas, with males weighing 140180
kilograms. In all great apes, the males are, on average, larger and stronger than the females, although the degree of sexual dimorphism varies greatly among species. Although most living species are
[28]
predominantly quadrupedal, they are all able to use their hands for gathering food or nesting materials, and, in some cases, for tool use.
Most species are omnivorous, but fruit is the preferred food among all but some human groups. Chimpanzees and orangutans primarily eat fruit. When gorillas run short of fruit at certain times of the
year or in certain regions, they resort to eating shoots and leaves, often of bamboo, a type of grass. Gorillas have extreme adaptations for chewing and digesting such low-quality forage, but they still
prefer fruit when it is available, often going miles out of their way to find especially preferred fruits. Humans, since the neolithic revolution, consume mostly cereals and other starchy foods, including
increasingly highly processed foods, as well as many other domesticated plants (including fruits) and meat. Hominid teeth are similar to those of the Old World monkeys and gibbons, although they are
especially large in gorillas. The dental formula is 2.1.2.3
2.1.2.3
. Human teeth and jaws are markedly smaller for their size than those of other apes, which may be an adaptation to eating cooked food since the
end of the Pleistocene.[29][30]

Gestation in great apes lasts 89 months, and results in the birth of a single offspring, or, rarely, twins. The young are born helpless, and require care for
long periods of time. Compared with most other mammals, great apes have a remarkably long adolescence, not being weaned for several years, and not
becoming fully mature for eight to thirteen years in most species (longer in humans). As a result, females typically give birth only once every few years.
There is no distinct breeding season.[28]

The gorillas and the common chimpanzee live in family groups of around five to ten individuals, although much larger groups are sometimes noted.
Chimpanzees live in larger groups that break up into smaller groups when fruit becomes less available. When small groups of female chimpanzees go off
in separate directions to forage for fruit, the dominant males can no longer control them and the females often mate with other subordinate males. In
contrast, groups of gorillas stay together regardless of the availability of fruit. When fruit is hard to find, they resort to eating leaves and shoots. Because Gorilla
gorilla groups stay together, the male is able to monopolize the females in his group. This fact is related to gorillas' greater sexual dimorphism than
chimpanzees'. In both chimpanzees and gorillas, the groups include at least one dominant male, and females leave the group at maturity
.

Legal status
Due to the close genetic relationship between humans and other great apes, certain animal rights organizations, such as the Great Ape Project, argue that nonhuman great apes are persons and should be
given basic human rights. Some countries have instituted aresearch ban to protect great apes from any kind of scientific testing.

[31]
On June 25, 2008, the Spanish parliament supported a new law that would make "keeping apes for circuses, television commercials or filming" illegal.

On September 8, 2010, theEuropean Union banned the testing of great apes.[32]

Conservation
The following table lists the estimated number of great ape individuals living outside zoos.

Estimated
Species Refs
number

Sumatran orangutan 6,667 [33]

Bornean orangutan 61,234 [33]

Tapanuli orangutan 800 [34]

Western gorilla 200,000 [35]

Eastern gorilla 6,000 [35]

Common chimpanzee 100,000 [36]

Bonobo 10,000 [36]

Human 7,405,745,000 [37]

See also
Bili ape List of individual apes
Dawn of Humanity (2015 PBS film) Oldest apes
Great ape language Prehistoric Autopsy (2012 BBC documentary)
Great ape research ban Primate cognition
Great Apes Survival Partnership The Mind of an Ape
Kinshasa Declaration on Great Apes Timeline of human evolution
List of human evolution fossils

Notes
1. "Great ape" is a common name rather than a taxonomic label, and there are differences in usage, even by the same author . The term may or may not include humans, as when
[2] and "gibbons are faithfully monogamous, unlike the great
Dawkins writes "Long before people thought in terms of evolution ... great apes were often confused with humans"
apes which are our closer relatives."[3]

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External links
The Animal Legal and Historical Center at Michigan State University College of Law, Great Apes and the Law
Renderings of Hominid Exemplars at the Smithsonian
Additional information on great apes
NPR News: Touma the Human Ancestor
Hominid Species at TalkOrigins Archive
For more details on Hominid species, including excellent photos of fossil hominids
New Scientist 19 May 2003 Chimps are human, gene study implies
Scientific American magazine (April 2006 Issue) Why Are Some Animals So Smart?
A new mediterranean hominoid-hominid link discovered,Anoiapithecus brevirostris, "Lluc": A unique Middle Miocene European hominoid and the origins of the great ape and
human clade Link to graphical reconstruction
Human Timeline (Interactive) Smithsonian, National Museum of Natural History(August 2016).

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