Family of primates
The
Hominidae
(
), whose members are known as the
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
chimpanzee
and the
bonobo
); and
Homo
, of which only
modern humans (
Homo sapiens
)
remain.
[1]
Numerous revisions in classifying the great apes have caused the use of the term
hominid
to change over time. The original meaning of "hominid" referred only to humans (
Homo
) and their closest extinct relatives. However, by the 1990s humans, apes, and their ancestors were considered to be "hominids".
The earlier 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 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 the turn of the 21st century.
[5]
Within the taxon Hominidae, a number of extant and extinct 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,
[6]
when the ancestors of the orangutans speciated from the ancestral line of the other three genera.
[7]
Those ancestors of the family Hominidae had already speciated from the family
Hylobatidae
(the
gibbons
), perhaps 15 to 20 million years ago.
[7]
[8]
Due to the close genetic relationship between humans and the 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
. Twenty-nine countries have instituted
research bans
to protect great apes from any kind of scientific testing.
[9]
Evolution
In the early
Miocene
, about 22 million years ago, there were many species of
tree-adapted
primitive
catarrhines
from East Africa; the variety suggests a long history of prior diversification. Fossils from 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 middle Miocene age?
Otavipithecus
from cave deposits in Namibia, and
Pierolapithecus
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.
[10]
Species close to the last common ancestor of gorillas, chimpanzees and humans may be represented by
Nakalipithecus
fossils found in Kenya and
Ouranopithecus
fossils 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 humans. Human
DNA
is approximately 98.4% identical to that of chimpanzees when comparing single nucleotide polymorphisms (see
human evolutionary genetics
).
[11]
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), followed by
Ardipithecus
(5.5?4.4 Ma), with species
Ar. kadabba
and
Ar. ramidus
.
Taxonomy
Terminology
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 modern usage of "hominid" that includes all the great apes including humans.
A number of very similar words apply to related classifications:
- A
hominoid
, sometimes 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.
[12]
- 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.
[13]
[14]
- 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 surviving
subspecies
.
A cladogram indicating common names (cf.
more detailed cladogram below
):
Hominoidea
|
|
Hylobatidae
gibbons
|
|
Hominidae
|
|
hominids, great apes
|
|
hominoids, apes
|
Extant and fossil relatives of humans
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
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
sub
family
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.)
[15]
[16]
[17]
But, those fossil relatives more closely related to humans than the chimpanzees represent the especially close members of the human family, and without necessarily assigning subfamily or tribal categories.
[18]
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
.
[19]
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.
[20]
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 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,
[21]
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
.
Phylogeny
Below is a cladogram with extinct species.
[22]
[23]
[24]
[
failed verification
]
It is indicated approximately how many million years ago (Mya) the clades diverged into newer clades.
[25]
Taxonomy of
Hominoidea
(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
There are eight living species of great ape which are classified in four
genera
. The following classification is commonly accepted:
[1]
- Family Hominidae
: humans and other great apes; extinct genera and species excluded
[1]
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.
[27]
|
|
−10 —
–
−9 —
–
−8 —
–
−7 —
–
−6 —
–
−5 —
–
−4 —
–
−3 —
–
−2 —
–
−1 —
–
0 —
| | |
|
Family Hominidae
Description
The great apes are tailless primates, with the smallest living species being the bonobo at 30 to 40 kilograms (66 to 88 lb) in weight, and the largest being the eastern gorillas, with males weighing 140 to 180 kilograms (310 to 400 lb). 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. 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 not only having supplanted with extensive tool use the role of jaws in hunting and fighting, but also eating cooked food since the end of the
Pleistocene
.
[32]
[33]
Behavior
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 use.
[34]
They build complex sleeping platforms, also called nests, in trees to sleep in at night, but chimpanzees and gorillas also build terrestrial nests, and gorillas can also sleep on the bare ground.
[35]
All species are
omnivorous
,
[36]
although 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
, have consumed mostly
cereals
and other
starchy
foods, including increasingly highly
processed foods
, as well as many other
domesticated plants
(including fruits) and
meat
.
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,
[37]
and not becoming fully mature for eight to thirteen years in most species (longer in orangutans and humans). As a result, females typically give birth only once every few years. There is no distinct breeding season.
[34]
Gorillas and chimpanzees 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.
This fact is related to gorillas' greater sexual dimorphism relative to that of chimpanzees; that is, the difference in size between male and female gorillas is much greater than that between male and female chimpanzees. This enables gorilla males to physically dominate female gorillas more easily. In both chimpanzees and gorillas, the groups include at least one dominant male, and young males leave the group at maturity.
Legal status
Due to the close genetic relationship between humans and the other great apes, certain
animal rights
organizations, such as the
Great Ape Project
, argue that nonhuman
great apes are persons
and, per the
Declaration on Great Apes
, should be given basic
human rights
. In 1999, New Zealand was the first country to ban any great ape experimentation, and now 29 countries have currently instituted a
research ban
to protect great apes from any kind of scientific testing.
On 25 June 2008, the Spanish parliament supported a new law that would make "keeping apes for circuses, television commercials or filming" illegal.
[38]
On 8 September 2010, the
European Union
banned the testing of great apes
.
[39]
Conservation
The following table lists the estimated number of great ape individuals living outside zoos.
See also
Notes
- ^
"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 Dawkins writes "Long before people thought in terms of evolution ... great apes were often confused with humans"
[3]
and "gibbons are faithfully monogamous, unlike the great apes which are our closer relatives."
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External links
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