Hominidae (/hɒˈmɪnɪdiː/), 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 chimpanzee and the bonobo; and Homo, which
includes modern humans and its extinct relatives (e.g., the
Neanderthal), and ancestors, such as
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 extinct 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). 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
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
Ponginae (see classification graphic below). The most recent
common ancestor of all
Hominidae lived roughly 14 million years
ago, when the ancestors of the orangutans speciated from the
ancestral line of the other three genera. Those ancestors of the
Hominidae had already speciated from the family Hylobatidae
(the gibbons), perhaps 15 million to 20 million years ago.
1 Evolution and taxonomy
1.1 Taxonomic history
1.1.2 Extant and fossil relatives of humans
2 Physical description
3 Legal status
5 See also
8 External links
Evolution and taxonomy
Sumatran orangutan (Pongo abelii)
In the early Miocene, about 22 million years ago, there were many
species of arboreally adapted primitive catarrhines from East Africa;
the variety suggests a long history of prior diversification. Fossils
at 20 million years ago include fragments attributed to
Victoriapithecus, the earliest Old World 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.
At sites far distant from East Africa, the presence of other
generalized non-cercopithecids, that is, non-monkey primates, of
Miocene age—Otavipithecus from cave deposits in Namibia, and
Dryopithecus from France, Spain and Austria—is
further evidence of a wide diversity of ancestral ape forms across
Africa and the Mediterranean basin during the relatively warm and
equable climatic regimes of the early and 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.
Molecular evidence indicates that the lineage of gibbons (family
Hylobatidae), the lesser apes, diverged from that of the great apes
some 18–12 million 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, dated to around 10
million years ago, may be represented by
Sivapithecus from India and
Griphopithecus from Turkey.
A reconstruction of a female
Australopithecus afarensis (National
Museum of Natural History)
Species close to the last common ancestor of gorillas, chimpanzees and
humans may be represented by
Nakalipithecus fossils found in Kenya and
Ouranopithecus found in Greece. Molecular evidence suggests that
between 8 and 4 million years ago, first the gorillas (genus Gorilla),
and then the chimpanzees (genus Pan) split off from the line leading
to the humans.
DNA is approximately 98.4% identical to that of
chimpanzees when comparing single nucleotide polymorphisms (see human
evolutionary genetics). The fossil record, however, of gorillas and
chimpanzees is limited; both poor preservation—rain forest soils
tend to be acidic and dissolve bone—and sampling bias probably
contribute most to this problem.
Other hominins probably adapted to the drier environments outside the
African equatorial belt; and there they encountered antelope, hyenas,
elephants and other forms becoming adapted to surviving in the East
African savannas, particularly the regions of the
Sahel and the
Serengeti. The wet equatorial belt contracted after about 8 million
years ago, and there is very little fossil evidence for the divergence
of the hominin lineage from that of gorillas and chimpanzees—which
split was thought to have occurred around that time. The earliest
fossils argued by some to belong to the human lineage are
Sahelanthropus tchadensis (7 Ma) and
Orrorin tugenensis (6 Ma),
Ardipithecus (5.5–4.4 Ma), with species Ar. kadabba and
Humans are one of the four extant hominid genera.
The classification of the great apes has been revised several times in
the last few decades; these revisions have led to a varied use of the
word "hominid" over time. The original meaning of the term referred to
only humans and their closest relatives—what is now the modern
meaning of the term "hominin". The meaning of the taxon Hominidae
changed gradually, leading to a different (modern) usage of "hominid"
that today includes all the great apes including humans.
The term hominid is easily confused with a number of very similar
A hominoid, commonly called an ape, is a member of the superfamily
Hominoidea: extant members are the gibbons (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 subfamily Homininae: gorillas,
chimpanzees, and humans (excludes orangutans).
A hominin is a member of the tribe Hominini: chimpanzees and humans.
A homininan, following a suggestion by Wood and Richmond (2000), would
be a member of the subtribe
Hominina of the tribe Hominini: that is,
modern humans and their closest relatives, including
Australopithecina, but excluding chimpanzees.
A human is a member of the genus Homo, of which
Homo sapiens is the
only extant species, and within that
Homo sapiens sapiens is the only
A cladogram indicating common names (c.f. more detailed cladogram
Hominoidea (hominoids, apes)
Hominidae (hominids, great apes)
Extant and fossil relatives of humans
A model of a modern human hominid skull (or hominin skull)
A fossil hominid exhibit at The Museum of Osteology, Oklahoma City,
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
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 groups—this 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
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.)  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.
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.
The exact criteria for membership in the tribe
Hominini under the
current understanding of human origins are not clear, but the taxon
generally includes 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 concept—including 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 disproven) that gorillas or
chimpanzees ever develop a theory of mind. This is also the case
for some New World monkeys outside the family of great apes, as, for
example, the capuchin monkeys.
However, even without the ability to test whether early members of the
Hominini (such as
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, and
some[who?] say the orangutan may also satisfy those criteria for the
theory of mind concept. These scientific debates take on political
significance for advocates of great ape personhood.
Below is a cladogram with extinct species. It is indicated
approximately how many million years ago (Mya) the clades diverged
into newer clades.
Kenyapithecus (†13 Mya)
Crown Homininae (10)
Hominoidea (emphasis on family Hominidae): after an
initial separation from the main line by the
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).
Model of the phylogeny of Hominidae, with adjacent branches of
Hominoidea, over the past 20 million years.
All extant species
There are eight living species of great ape which are classified in
four genera. The following classification is commonly accepted:
Family Hominidae: humans and other great apes; extinct genera and
Bornean orangutan, Pongo pygmaeus
Pongo pygmaeus pygmaeus
Pongo pygmaeus morio
Pongo pygmaeus wurmbii
Sumatran orangutan, Pongo abelii
Tapanuli orangutan, Pongo tapanuliensis
Western lowland gorilla,
Gorilla gorilla gorilla
Cross River gorilla,
Gorilla gorilla diehli
Gorilla beringei beringei
Eastern lowland gorilla,
Gorilla beringei graueri
Chimpanzee (common chimpanzee), Pan troglodytes
Pan troglodytes troglodytes
Pan troglodytes verus
Pan troglodytes ellioti
Pan troglodytes schweinfurthii
Bonobo (pygmy chimpanzee), Pan paniscus
Anatomically modern human,
Homo sapiens sapiens
A reconstruction of
Replica of the skull sometimes known as "Nutcracker Man", found by
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
Tribe Lufengpithecini †
Oreopithecus (placement disputed)
† (placement debated)
Homo – close relatives of modern humans
† (thought by some to be an early subspecies of Homo
† (considered by some to be an early subspecies of Homo
† (sometimes called Archaic
Denisovans (scientific name not yet assigned)†
† (sometimes called
Homo sapiens idaltu†
Red Deer Cave people
Red Deer Cave people
† (scientific name has not yet been assigned;
perhaps a race of modern humans or a hybrid of modern humans and
The great apes are large, tailless primates, with the smallest living
species being the bonobo at 30–40 kilograms in weight, and the
largest being the eastern gorillas, with males weighing 140–180
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
predominantly quadrupedal, they are all able to use their hands for
gathering food or nesting materials, and, in some cases, for tool
Most species are omnivorous, but fruit is the
preferred food among all but some human groups.
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
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.
Gestation in great apes lasts 8–9 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.
The gorillas and the common chimpanzee live in family groups of around
five to ten individuals, although much larger groups are sometimes
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
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
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 a research
ban to protect great apes from any kind of scientific testing.
On June 25, 2008, the Spanish parliament supported a new law that
would make "keeping apes for circuses, television commercials or
On September 8, 2010, the
European Union banned the testing of great
The following table lists the estimated number of great ape
individuals living outside zoos.
Dawn of Humanity (2015 PBS film)
Great ape language
Great ape research ban
Great Apes Survival Partnership
Kinshasa Declaration on Great Apes
List of human evolution fossils
List of individual apes
Prehistoric Autopsy (2012 BBC documentary)
The Mind of an Ape
Timeline of human evolution
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Wikimedia Commons has media related to Hominidae.
Wikispecies has information related to Hominidae
The Wikibook Dichotomous Key has a page on the topic of: Hominidae
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: Toumaï the
Species at TalkOrigins Archive
For more details on Hominid species, including excellent photos of
New Scientist 19 May 2003 – Chimps are human, gene study implies
Scientific American magazine (April 2006 Issue) Why Are Some Animals
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
Human Timeline (Interactive) – Smithsonian, National Museum of
Natural History (August 2016).
Extant primate families
Extant species of family
Hominidae (great apes)
Bornean orangutan (P. pygmaeus)
Sumatran orangutan (P. abelii)
Tapanuli orangutan (P. tapanuliensis)
Western gorilla (G. gorilla)
Eastern gorilla (G. beringei)
Common chimpanzee (P. troglodytes)
Bonobo (P. paniscus)
H. s. sapiens
Extant ape species
Chimpanzee (Pan spp.)
Bonobo (Pan paniscus)
Common chimpanzee (Pan troglodytes)
Western gorilla (
Eastern gorilla (
Orangutan (Pongo spp.)
Bornean orangutan (Pongo abelii)
Sumatran orangutan (Pongo pygmaeus)
Tapanuli orangutan (Pongo tapanuliensis)
Gibbon (family: Hylobatidae)
Study of apes
Chimpanzee genome project
Human genome project
Neanderthal genome project
Lone Drøscher Nielsen
Primate Learning Sanctuary
Legal and social status
List of individual apes
List of individual apes (non-human)
Apes in space (non-human)
Chimpanzee–human last common ancestor
List of fictional primates (non-human)
Evolutionary biology portal