Great ape
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Twas Now
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{{redirect|Hominidae|an explanation of similar terms|Hominid}}
{{Taxobox
| name = Hominids<ref name=MSW3>{{MSW3 Groves|pages=181-184|id=12100786}}</ref>
| fossil range = [[Miocene]] to Recent
| image = Austrolopithecus africanus.jpg
| image_width = 250px
| image_caption = ''[[Australopithecus|Australopithecus africanus]]'' reconstruction
| regnum = [[Animal]]ia
| phylum = [[Chordata]]
| classis = [[Mammal]]ia
| ordo = [[Primate]]s
| subordo = [[Haplorrhini]]
| infraordo = [[Simiiformes]]
| parvordo = [[Catarrhini]]
| superfamilia = [[Hominoidea]]
| familia = '''Hominidae'''
| familia_authority = [[John Edward Gray|Gray]], 1825
| subdivision_ranks = [[Genus|Genera]]
| subdivision =
*Subfamily [[Ponginae]]
**''[[Orangutan|Pongo]]'' – [[orangutan]]s
*Subfamily [[Homininae]]
**''[[Gorilla]]'' – [[gorilla]]s
**''[[chimpanzee|Pan]]'' – [[chimpanzee]]s
**''[[Homo (genus)|Homo]]'' – [[human]]s
}}
The '''great apes''' are the members of the [[biological family]] '''Hominidae''' which includes [[human]]s, [[chimpanzee]]s, [[gorilla]]s, and [[orangutan]]s.<!--These are the common names for the four living genera in Hominidae. Please don't change it to include a mix of species and genera common names. See the classification below.--><ref name=MSW3/><ref>"Great ape" is a common name rather than a taxonomic label and there are differences in usage. Subtly, it may seem to exclude human beings ("humans and the great apes") or to include them ("humans and non-human great apes"). ''Homo sapiens'' is not at any especial remove from other members of the biological family, and humans are therefore described here as great apes.</ref>
==Characteristics==
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 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 use.<ref name=EoM>{{cite book |editor=Macdonald, D.|author= Harcourt, A.H., MacKinnon, J. & Wrangham, R.W.|year=1984 |title= The Encyclopedia of Mammals|publisher= Facts on File|location=New York|pages= 422-439|isbn= 0-87196-871-1}}</ref>
Most species are [[omnivore|omnivorous]], but fruit is the preferred food among all but humans. 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. In contrast, humans consume a large proportion of highly processed, low fiber foods, unusual proportions of grains and vertebrate flesh, as well as a wide variety of other foodstuffs. The teeth are similar to those of the [[Old World monkey]]s and [[gibbon]]s, although they are especially large in the gorilla. The [[dentition|dental formula]] is:{{dentition2|2.1.2.3|2.1.2.3}} Human teeth and jaws are markedly smaller for our size than those of other apes. Some scholars{{Fact|date=March 2008}} have surmised that humans' small teeth and jaws are adaptations to eating cooked food for possibly as long as a million years or more. This idea remains controversial.
[[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 the mother must care for them for long periods of time. Compared with most other mammals, great apes have a remarkably long adolescence, not being [[weaning|weaned]] for several years, and not becoming fully mature for 8-13 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.<ref name = EoM/>
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 male(s) can no longer control them and the females often mate with other subordinate males, whether by choice or not. 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 maturity. By contrast, orangutans are generally solitary. The social structure of humans is complex and highly variable. DNA studies suggest that, as in other apes, female humans are the sex that leaves the group at maturity, or at least has done so over evolutionary time.
==Classification==
[[Image:Gorilla 019.jpg|thumb|left|[[Gorilla]]]]
This classification of the great apes has been [[ape#History of hominoid taxonomy|revised several times]] in the last few decades. Also see [[hominid]] for a helpful clarification of terms. Originally, the group was restricted to humans and their [[extinct]] relatives, with the other great [[ape]]s being placed in a separate family, the '''Pongidae'''. This definition is still used by many [[anthropologist]]s and by [[laity|lay persons]]. However, that definition makes Pongidae [[paraphyletic]], whereas most taxonomists nowadays encourage [[monophyletic]] groups. Thus many [[biologist]]s consider Hominidae to include Pongidae as the subfamily [[Ponginae]], or restrict the latter to the orangutans and their extinct relatives like ''[[Gigantopithecus]]''. The taxonomy shown here follows the monophyletic groupings.
Especially close human relatives form a [[subfamily]], the [[Homininae]]. Some researchers go so far as to include chimpanzees<ref>{{cite web | author = Pickrell, John | title = Chimps Belong on Human Branch of Family Tree, Study Says | url = http://news.nationalgeographic.com/news/2003/05/0520_030520_chimpanzees.html | date = [[2003-05-20]] | accessdate = 2007-08-04 | work = [http://www.nationalgeographic.com/index.html National Geographic]}}</ref> and gorillas<ref>[http://www.berggorilla.de/english/gjournal/texte/32mensch-gorilla-groves.html Relationship Humans-Gorillas<!-- Bot generated title -->]</ref><ref>Watson, E. E. et al. (2001) Homo genus: a review of the classification of humans and the great apes. Pp. 311-323 in: Humanity from African Naissance to Coming Millennia (eds. Tobias, P. V. et al.). Florence: Firenze Univ. Press</ref> in the genus ''[[Homo (genus)|Homo]]'' along with humans, but it is more commonly accepted to describe the relationships as shown here.
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 [[australopithecine]]s ''[[Australopithecus]]'' and ''[[Paranthropus]]''.
The exact criteria for membership in the Homininae are not clear, but the subfamily 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 [[culture]]s beyond the family or band. The [[theory of mind]] including such faculties as mental state attribution, empathy and even empathetic deception is a controversial criterion distinguishing the adult human alone among the hominids. Humans acquire this capacity at about four and a half years of age, whereas it has neither been proven nor disproven that gorillas and chimpanzees develop a theory of mind.<ref>{{cite journal | author = Heyes, C. M. | year = 1998 | title = Theory of Mind in Nonhuman Primates | journal = Behavioral and Brain Sciences | id = bbs00000546 | url = http://www.bbsonline.org/documents/a/00/00/05/46/index.html | volume = 21 | doi = 10.1017/S0140525X98000703 <!--Retrieved from CrossRef by DOI bot-->}}</ref> This is also the case for some [[new world monkey]]s outside the family of great apes, as, for example, the [[Capuchin monkey#Theory of mind|capuchin monkeys]].
However, without the ability to test whether early members of the Homininae (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. Despite an apparent lack of real culture and significant physiological and psychological differences, some say that the orangutan may also satisfy these criteria. These scientific debates take on political significance for advocates of [[Great Ape personhood]].
==Evolution==
[[Image:Man of the woods.jpg|thumb|right|Orangutan]]
On July 19, 2001,a 7 million year old [[fossil]] [[skull]] nicknamed "Toumaï" by its discoverers, and formally classified as ''[[Sahelanthropus tchadensis]]'', was discovered in [[Chad]] and is possibly the earliest hominid [[fossil]] ever found. In addition to its age, Toumaï, unlike the 3 – 4 million year younger [[gracile australopithecine]] dubbed "[[Australopithecus afarensis|Lucy]]", has a relatively flat face without the prominent snout seen on other pre-''[[Homo (genus)|Homo]]'' hominids. Some researchers have made the suggestion that this previously unknown species may in fact be a direct ancestor of modern humans (or at least closely related to a direct ancestor). Others contend that one fossil is not enough to make such a claim because it would overturn the conclusions of over 100 years of [[anthropology|anthropological]] study. A report on this finding was published in the journal ''[[Nature (journal)|Nature]]'' on [[July 11]], [[2002]]. While some scientists claim that it is merely the skull of a female gorilla ancestor, others have called it the most important hominin fossil since ''Australopithecus''.
In addition to the Toumai fossil, some experts use evidence from the genome to argue that the species associated with the chimpanzees and proto-humans split interbred over a long period of time, swapping genes, before making a final separation. A paper, whose authors include [[David Reich]] and [[Eric Lander]] (Harvard and the Massachusetts Institute of Technology (MIT)), was published in journal ''[[Nature (journal)|Nature]]'' in May 2006.
It is generally believed that the ''Pan/Homo'' split occurred about 6.5–7.4 million years ago, but the [[molecular clock]] (a method of calculating evolution based on the speed at which genes mutate) suggests the genera split 5.4–6.3 million years ago. Previous studies looked at average genetic differences between human and chimp. The new study compares the ages of key sequences of genes of modern humans and modern chimps. Some sequences are younger than others, indicating that chimps and humans gradually split apart over a period of 4 million years. The youngest human chromosome is the X sex chromosome which is about 1.2 million years more recent than the 22 autosomes. The X chromosome is known to be vulnerable to selective pressure. Its age suggests there was an initial split between the two species, followed by gradual divergence and interbreeding that resulted in younger genes, and then a final separation.
== Species ==
[[Image:Hominidae.PNG|300px|thumb|right|Hominoid family tree]]
[[Image:Orang.gorilla.skulls.jpg|250px|thumb|right|Skulls of an orangutan and a gorilla]]
The seven living species of great ape are classified in four [[genus|genera]].
* '''Family Hominidae''': humans and other great apes; extinct genera and species excluded.<ref name=MSW3/>
** Subfamily [[Orangutan|Ponginae]]
*** Genus ''[[Orangutan|Pongo]]''
**** [[Bornean Orangutan]], ''Pongo pygmaeus''
***** ''Pongo pygmaeus pygmaeus''
***** ''Pongo pygmaeus morio''
***** ''Pongo pygmaeus wurmbii''
**** [[Sumatran Orangutan]], ''Pongo abelii''
** Subfamily [[Homininae]]
*** Tribe [[Gorilla|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]]
**** Genus ''[[chimpanzee|Pan]]''
***** [[Common Chimpanzee]], ''Pan troglodytes''
****** Central Chimpanzee, ''Pan troglodytes troglodytes''
****** West African Chimpanzee, ''Pan troglodytes verus''
****** Nigerian Chimpanzee, ''Pan troglodytes vellerosus''
****** Eastern Chimpanzee, ''Pan troglodytes schweinfurthii''
***** [[Bonobo]] (Pygmy Chimpanzee), ''Pan paniscus''
**** Genus ''[[Homo (genus)|Homo]]''
***** [[Human]], ''Homo sapiens sapiens''
In addition to the extant species and subspecies above, [[archaeologists]], [[paleontologists]], and [[anthropologists]] have discovered numerous extinct species. The list below are some of the genera of those discoveries.
*Subfamily [[Ponginae]]
**''[[Gigantopithecus]]''†
**''[[Sivapithecus]]''†
**''[[Lufengpithecus]]''†
**''[[Ankarapithecus]]''†
**''[[Ouranopithecus]]''†
*Subfamily [[Homininae]]
**''[[Homo (genus)|Homo]]'' – [[Human]]s and their immediate ancestors
**''[[Oreopithecus]]''†
**''[[Paranthropus]]''†
**''[[Australopithecus]]''†
**''[[Sahelanthropus]]''†
**''[[Orrorin]]''†
**''[[Ardipithecus]]''†
**''[[Kenyanthropus]]''†
== 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 non-human [[great ape personhood|great apes are persons]] and should be given basic [[human rights]]. Some countries have instituted a [[Great ape research ban|research ban]] to protect great apes from any kind of scientific testing.
On [[25 June]] [[2008]], the Spanish parliament gave support for a new law that would make "keeping apes for circuses, television commercials or filming" illegal under Spain's penal code.<ref>{{cite web |url=http://www.reuters.com/article/scienceNews/idUSL256586320080625 |title=Spanish parliament to extend rights to apes |accessdate=2008-07-11 |date=2008-06-25 |publisher=Reuters}}</ref>
== See also ==
{{Commonscat|Hominidae}}
{{3d commons}}
{{Wikispecies|Hominidae}}
{{Wikibookspar|Dichotomous Key|Hominidae}}
* [[Great ape language]]
** ''[[The Mind of an Ape]]''
* [[List of non-human apes]] – notable individual apes
* [[Ape extinction]]
** [[Great Apes Survival Project]]
** [[Declaration on Great Apes]]
** [[Kinshasa Declaration on Great Apes]]
* [[Evolution of Homo sapiens]]
** [[Graphical timeline of human evolution]]
== References ==
{{reflist}}
== External links ==
*[http://www.unep.org/grasp/Resources/index.asp Additional information on great apes]
*[http://www.npr.org/programs/atc/features/2002/july/toumai/index.html NPR News: Toumaï the Human Ancestor]
*[http://www.talkorigins.org/faqs/homs/species.html Hominid Species] at talkorigins.org
*[http://www.modernhumanorigins.net/ For more details on Hominid species, including excellent photos of fossil hominids]
*[http://www.newscientist.com/article.ns?id=dn3744 New Scientist 19 May 2003 – Chimps are human, gene study implies]
*[http://www.sciam.com/article.cfm?chanID=sa006&articleID=000C1E5D-B9BA-1422-B9BA83414B7F0103 Scientific American Magazine (April 2006 Issue) Why Are Some Animals So Smart?]
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