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| range_map_caption = Distribution of great ape species
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The '''Hominidae''' ({{IPAc-en|h|ɒ|ˈ|m|ɪ|n|ᵻ|d|iː}}), whose members are known as the '''great apes'''{{NoteTag|"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"<ref>{{cite book |last = Dawkins |first = R. |author-link = Richard Dawkins |year = 2005 |title = The Ancestor's Tale: A Pilgrimage to the Dawn of Life |title-link = The Ancestor's Tale |edition = p/b |location = London, England |publisher = Phoenix (Orion Books) |isbn = 978-0-7538-1996-8 |page = [https://archive.org/details/ancestorstalepil0000dawk/page/114 114]}}</ref> and "gibbons are faithfully monogamous, unlike the great apes which are our closer relatives."{{sfnp|Dawkins|2005|p=126}}}} or '''hominids''' ({{IPAc-en|ˈ|h|ɒ|m|ᵻ|n|ɪ|d|z}}), are a taxonomic [[Family (biology)|family]] of [[primate]]s that includes eight [[Neontology#Extant taxa versus extinct taxa|extant]] species in four [[Genus|genera]]: ''[[Orangutan|Pongo]]'' (the [[Bornean orangutan|Bornean]], [[Sumatran orangutan|Sumatran]] and [[Tapanuli orangutan]]); ''[[Gorilla]]'' (the [[Eastern gorilla|eastern]] and [[western gorilla]]); ''[[Pan (genus)|Pan]]'' (the [[chimpanzee]] and the [[bonobo]]); and ''[[Homo]]'', of which only [[Human|modern humans (''Homo sapiens'')]] remain.<ref name="MSW3" />
 
The '''Hominidae''' ({{IPAc-en|h|ɒ|ˈ|m|ɪ|n|ᵻ|d|iː}}), whose members are known as the '''great apes'''{{NoteTag|"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"<ref>{{cite book |last = Dawkins |first = R. |author-link = Richard Dawkins |year = 2005 |title = The Ancestor's Tale: A Pilgrimage to the Dawn of Life |title-link = The Ancestor's Tale |edition = p/b |location = London, England |publisher = Phoenix (Orion Books) |isbn = 978-0-7538-1996-8 |page = [https://archive.org/details/ancestorstalepil0000dawk/page/114 114]}}</ref>{{Better source needed | date = September 2024 | reason = When hypernym is mentioned together with its hyponym, the hyponym is often understood to be excluded from the hypernym in that context. This may be contextual and generally is applicable to all hypernym/hyponym usage. These aren't good examples as a general usage of 'Great ape' that excludes human.}} and "gibbons are faithfully monogamous, unlike the great apes which are our closer relatives."{{sfnp|Dawkins|2005|p=126}}{{Better source needed | date = September 2024 | reason = See previous BSN tag.}}}} or '''hominids''' ({{IPAc-en|ˈ|h|ɒ|m|ᵻ|n|ɪ|d|z}}), are a taxonomic [[Family (biology)|family]] of [[primate]]s that includes eight [[Neontology#Extant taxa versus extinct taxa|extant]] species in four [[Genus|genera]]: ''[[Orangutan|Pongo]]'' (the [[Bornean orangutan|Bornean]], [[Sumatran orangutan|Sumatran]] and [[Tapanuli orangutan]]); ''[[Gorilla]]'' (the [[Eastern gorilla|eastern]] and [[western gorilla]]); ''[[Pan (genus)|Pan]]'' (the [[chimpanzee]] and the [[bonobo]]); and ''[[Homo]]'', of which only [[Human|modern humans (''Homo sapiens'')]] remain.<ref name="MSW3" />
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".
 
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 and other apes were considered to be "hominids".
 
The earlier restrictive meaning has now been largely assumed by the term ''[[Hominini|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.<ref>{{cite web|last1=Morton|first1=Mary|title=Hominid vs. hominin|url=https://www.earthmagazine.org/article/hominid-vs-hominin|website=Earth Magazine|access-date=17 July 2017}}</ref>
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At sites far distant from East Africa, the presence of other generalized non-[[Old World monkey|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.<ref name="Srivastava2009">{{cite book|author=Srivastava| first=R. P.|title=Morphology of the Primates And Human Evolution|url=https://books.google.com/books?id=kCerOsM8XMwC&pg=PA87|access-date=6 November 2011|date=2009|publisher=PHI Learning Pvt. Ltd.|isbn=978-81-203-3656-8|page=87}}</ref> 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]]).<ref>{{Cite journal|last1=Chen|first1=Feng-Chi|last2=Li|first2=Wen-Hsiung|date=2001-01-15|title=Genomic Divergences between Humans and Other Hominoids and the Effective Population Size of the Common Ancestor of Humans and Chimpanzees|journal=American Journal of Human Genetics|volume=68|issue=2|pages=444–456|issn=0002-9297|pmc=1235277|pmid=11170892|doi=10.1086/318206}}</ref> 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''.
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* 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.<ref name=Wood2010>{{cite journal | author = B. Wood | year = 2010 | title = Reconstructing human evolution: Achievements, challenges, and opportunities | journal = Proceedings of the National Academy of Sciences | volume = 107 | issue = Suppl 2 | pages = 8902–8909 | doi = 10.1073/pnas.1001649107 | bibcode=2010PNAS..107.8902W | pmid=20445105 | pmc=3024019| doi-access = free}}</ref>
* 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.<ref name=WoodRichmond>{{cite journal | last1=Wood | first1=Bernard Wood| last2=Richmond | year first2=Brian 2000G. | title = Human evolution: taxonomy and paleobiology | journal = Journal of Anatomy | volume = 197 | pages issue= 19–601 |pmid date= 109992702000 |last2 issn= Richmond0021-8782 |first2 pmid= B. G.10999270 | pmc = 1468107 | doi = 10.1046/j.1469-7580.2000.19710019.x | issue pages= Pt 119–60}}. In this suggestion, the new subtribe of ''Hominina'' was to be designated as including the genus ''Homo'' exclusively, so that ''Hominini'' would have two subtribes, ''Australopithecina'' and ''Hominina'', with the only known genus in ''Hominina'' being ''Homo''. ''[[Orrorin]]'' (2001) has been proposed as a possible ancestor of ''Hominina'' but not ''Australopithecina''.{{cite book |url = https://books.google.com/books?id=PrJ1lmjMakoC&pg=PA116|title=African Genesis: Perspectives on Hominin Evolution |isbn = 9781107019959 |last1=Reynolds |first1=Sally C. |last2=Gallagher|first2=Andrew |date=2012-03-29|publisher=Cambridge University Press}}. Designations alternative to ''Hominina'' have been proposed: ''Australopithecinae'' (Gregory & Hellman 1939) and ''Preanthropinae'' (Cela-Conde & Altaba 2002); {{cite journal | last1 = Brunet | first1 = M. | display-authors = etal | year = 2002 | title = A new hominid from the upper Miocene of Chad, central Africa | journal = Nature | volume = 418 | issue = 6894| pages = 145–151 | doi=10.1038/nature00879 | pmid=12110880 | bibcode = 2002Natur.418..145B| s2cid = 1316969| url = http://doc.rero.ch/record/13388/files/PAL_E190.pdf}}
{{cite journal | last1 = Cela-Conde | first1 = C.J. | last2 = Ayala | first2 = F.J. | year = 2003 | title = Genera of the human lineage | journal = PNAS | volume = 100 | issue = 13| pages = 7684–7689 | doi=10.1073/pnas.0832372100 | pmid=12794185 | pmc=164648 | bibcode = 2003PNAS..100.7684C| doi-access = free}}
{{cite journal | last1 = Wood | first1 = B. | last2 = Lonergan | first2 = N. | year = 2008 | title = The hominin fossil record: taxa, grades and clades | url = http://www.gwu.edu/~hogwash/BW_PDFs/RP156.pdf | journal = J. Anat. | volume = 212 | issue = 4| pages = 354–376 | doi=10.1111/j.1469-7580.2008.00871.x | pmid=18380861 | pmc=2409102}}</ref><ref>{{Cite web|url=https://www.geol.umd.edu/~tholtz/G204/lectures/204scatterlings.html|title=GEOL 204 The Fossil Record: The Scatterlings of Africa: The Origins of Humanity|website=www.geol.umd.edu|access-date=2019-11-07}}</ref>
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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 [[biologist]]s 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.)<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 | archive-url= https://web.archive.org/web/20030601115459/http://news.nationalgeographic.com/news/2003/05/0520_030520_chimpanzees.html | url-status= dead | archive-date= 1 June 2003 | date=20 May 2003 | access-date= 4 August 2007 | work= [[National Geographic Society]]}}</ref><ref>[{{cite web | title=Relationship Humans-Gorillas | website=berggorilla.de | date=2007-10-21 | url=http://www.berggorilla.de/english/gjournal/texte/32mensch-gorilla-groves.html Relationship| Humansarchive-Gorillas] {{Webarchive|url=https://web.archive.org/web/20071130023024/http://www.berggorilla.de/english/gjournal/texte/32mensch-gorilla-groves.html | archive-date=2007-11-30 November| 2007url-status=dead | access-date=2024-09-15}}.</ref><ref>{{cite book | author=Watson, E. E. | display-authors=etal | date=2001 | chapter=Homo genus: a review of the classification of humans and the great apes | pages= 311–323 | title= Humanity from African Naissance to Coming Millennia | editor= Tobias, P. V. | display-editors= etal | location= Florence | publisher= Firenze Univ. Press}}</ref> But, those fossil relatives more closely related to humans than the chimpanzees represent the especially close members of the human family, {{Clarify | text =and without necessarily assigning subfamily or tribal categories.<ref>Schwartz,| J.Hdate = September 2024 | reason = This seems to conflict with the cladogram above: chimp and Australopithecine et al. (1986)in Primatethe systematicstribe Homonini, and aAustralopithecine classificationet ofal. in the ordersubtribe Homonina.}}<ref>{{cite book | editor-last=Swindler | editor-first=Daris R. | editor-last2=Erwin | editor-first2=J. | title=Comparative primatePrimate biologyBiology | volume = 1: Systematics, evolution, and anatomy (ed. by| D.R.chapter Swindler,= Primate systematics and J.a Erwin),classification pp.of 1–41,the Alanorder R| last1 = Schwartz | first1 = J.H. | publisher=Wiley-Liss, | location = New York. | date=1986 | isbn=978-0-471-62644-2 | page=1–41}}</ref>
 
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]]''.<ref>{{cite journal | last1 = Schwartz, | first1 = J.H. (2004b)| year = 2004 | title = Issues in hominid systematics | journal = Paleoantropología | url = https://www.academia.edu/8420749 Zona| Arqueologíaissue = 4, | location = ALCALA DE HENARES | pages = 360–371. }}</ref>
 
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 [[culture]]s 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.<ref>{{cite journal |author last1 = Heyes, | first1 = C. M. | date = 1998 | title = Theory of Mind in Nonhuman Primates | journal = Behavioral and Brain Sciences | issue = 1 | id = bbs00000546 | volume = 21 | doi = 10.1017/S0140525X98000703 | pmid = 10097012 | pages = 101–14 | s2cid = 6469633 | url = http://discovery.ucl.ac.uk/117063/1/download.14pdf.pdf}}</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, even without the ability to test whether early members of the Hominini (such as ''[[Homo erectus]]'', ''[[Neanderthal|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,<ref>{{cite journal | author = Van Schaik C.P. | date = 2003 | title = Orangutan cultures and the evolution of material culture | journal = Science | volume = 299 | issue = 5603| pages = 102–105 | doi = 10.1126/science.1078004 | pmid = 12511649 | last2 = Ancrenaz | first2 = M | last3 = Borgen | first3 = G | last4 = Galdikas | first4 = B | last5 = Knott | first5 = CD | last6 = Singleton | first6 = I | last7 = Suzuki | first7 = A | last8 = Utami | first8 = SS | last9 = Merrill | first9 = M | bibcode = 2003Sci...299..102V| s2cid = 25139547}}</ref> and some{{who|date=September 2012}} 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=Description==
[[File:Gorilla 019.jpg|thumb|Gorilla]]
The great apes are tailless primates, with the smallest living species being the bonobo at {{Convert|30 to 40|kg|lb}} in weight, and the largest being the eastern gorillas, with males weighing {{Convert|140 to 180|kg|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 monkey]]s and gibbons, although they are especially large in gorillas. The [[dentition|dental formula]] is {{DentalFormula|upper=2.1.2.3|lower=2.1.2.3}}. Human teeth and jaws are markedly smaller relative to body size compared to those of other apes. This may be an adaptation not only to the extensive use of tools, which has supplanted the role of jaws in hunting and fighting, but also to eating cooked food since the end of the [[Pleistocene]].<ref name=Brace&Mahler>{{Cite journal|date=1971|last1=Brace |first1=C. Loring |author-link=C. Loring Brace |last2=Mahler |first2=Paul Emil |title=Post-Pleistocene changes in the human dentition|journal=[[American Journal of Physical Anthropology]]|volume=34|issue=2 |pages=191–203|doi=10.1002/ajpa.1330340205 |pmid=5572603 |hdl=2027.42/37509 |url = https://deepblue.lib.umich.edu/bitstream/2027.42/37509/1/1330340205_ftp.pdf|hdl-access=free}}</ref><ref name=Wrangham2007>{{cite book |year = 2007 |first = Richard |last = Wrangham | chapter=Chapter 12: The Cooking Enigma | editor=Charles Pasternak | title=What Makes Us Human? | location=Oxford | publisher=Oneworld Press |isbn=978-1-85168-519-6}}</ref>
 
==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.<ref name=EoM>{{cite book|editor-last= Macdonald|editor-first= D.|last1= Harcourt|first1= A.H.|last2= MacKinnon|first2= J.|last3= Wrangham |first3= R.W.|date= 1984|title= The Encyclopedia of Mammals|publisher= Facts on File|location= New York|pages= [https://archive.org/details/encyclopediaofma00mals_0/page/422 422–439]|isbn= 978-0-87196-871-5|url-access= registration|url= https://archive.org/details/encyclopediaofma00mals_0/page/422}}</ref> 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.<ref>{{cite journal | doi=10.1007/s10764-020-00186-z | title=Deciding Where to Sleep: Spatial Levels of Nesting Selection in Chimpanzees (Pan troglodytes) Living in Savanna at Issa, Tanzania | date=2020 | last1=Hernandez-Aguilar | first1=R. Adriana | last2=Reitan | first2=Trond | journal=International Journal of Primatology | volume=41 | issue=6 | pages=870–900 | doi-access=free }}</ref>
 
All species are [[omnivore|omnivorous]],<ref>{{Cite web |last=Alina |first=Bradford |date=29 May 2015 |title=Facts About Apes |url=https://www.livescience.com/51017-ape-facts.html |access-date=2021-01-25 |website=livescience.com |language=en}}</ref> 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 [[cereal]]s and other [[starch]]y foods, including increasingly highly [[Food processing|processed foods]], as well as many other [[Domestication#Plants|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 [[weaning|weaned]] for several years,<ref name="Hamilton 2017">{{cite web |last=Hamilton |first=Jon |title=Orangutan Moms Are The Primate Champs Of Breast-Feeding |website=NPR |date=2017-05-17 |url=https://www.npr.org/sections/health-shots/2017/05/17/528776636/orangutan-moms-are-the-primate-champs-of-breast-feeding |access-date=2024-05-20}}</ref> 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.<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 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==
{{Main|Great ape personhood|Great Ape Project|Countries banning non-human ape experimentation}}
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 ape personhood|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 [[Great ape research ban|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.<ref>{{cite news |url = https://www.reuters.com/article/scienceNews/idUSL256586320080625 |title = Spanish parliament to extend rights to apes |access-date = 11 July 2008 |date = 25 June 2008 |work = Reuters}}</ref> On 8 September 2010, the [[European Union]] [[great ape research ban|banned the testing of great apes]].<ref>{{cite web |url = https://www.independent.co.uk/life-style/health-and-families/new-eu-rules-on-animal-testing-ban-use-of-apes-2077443.html |title = New EU rules on animal testing ban use of apes |website = [[Independent.co.uk]] |date = 12 September 2010}}</ref>
 
==Conservation==
The following table lists the estimated number of great ape individuals living outside zoos.
{| class="wikitable sortable" style="font-weight: bold;"
|-
! style="background-color:#efefef;" | Species
! style="background-color:#efefef;" | Estimated<br />number
! style="background-color:#efefef;" | Conservation<br />status
! style="background-color:#efefef;" class="unsortable" | Refs
|-
| style="background-color:#eeddbb;"|[[Bornean orangutan]]
| style="background-color:#eeddbb; text-align: right; padding-right: 1em;"|104,700
| style="background-color:#eeddbb; text-align: right; padding-right: 1em;"|[[Critically endangered]]
| style="background-color:#eeddbb; text-align: center;"|<ref>{{cite web | title=Bornean Orangutan, Pongo pygmaeus | website=New England Primate Conservancy| date=2021-10-29 | url=https://neprimateconservancy.org/bornean-orangutan/ | access-date=2024-09-18 | archiveurl = https://web.archive.org/web/20240529110845/https://neprimateconservancy.org/bornean-orangutan/ | archivedate = 2024-05-29 | url-status = live}}</ref>
|-
| style="background-color:#eeddbb;"|[[Sumatran orangutan]]
| style="background-color:#eeddbb; text-align: right; padding-right: 1em;"|6,667
| style="background-color:#eeddbb; text-align: right; padding-right: 1em;"|[[Critically endangered]]
| style="background-color:#eeddbb; text-align: center;"|<ref name="orangutans">An estimate of the number of wild [[orangutans]] in 2004: {{cite web |url = http://www.yorku.ca/arusson/Papers/GoI%20OU%20action%20plan%2007-17.pdf|title= Orangutan Action Plan 2007–2017 |year = 2007 |publisher= [[Government of Indonesia]] | archiveurl = https://web.archive.org/web/20240901105624/http://www.yorku.ca/arusson/Papers/GoI%20OU%20action%20plan%2007-17.pdf | archivedate = 2024-09-01 | url-status = live | quote = Pada IUCN Red List Edisi tahun 2002 orangutan sumatera dikategorikan Critically Endangered, artinya sudah sangat terancam kepunahan, sedangkan orangutan kalimantan dikategorikan Endangered atau langka.}}</ref>
|-
| style="background-color:#eeddbb;"|[[Tapanuli orangutan]]
| style="background-color:#eeddbb; text-align: right; padding-right: 1em;"|800
| style="background-color:#eeddbb; text-align: right; padding-right: 1em;"|[[Critically endangered]]
| style="background-color:#eeddbb; text-align: center;"|<ref>{{cite web | title=Tapanuli Orangutan, Pongo tapanuliensis | website=New England Primate Conservancy| date=2021-10-30 | url=https://neprimateconservancy.org/tapanuli-orangutan/ | access-date=2024-09-18 | archiveurl = https://web.archive.org/web/20240529123735/https://neprimateconservancy.org/tapanuli-orangutan/ | archivedate = 2024-05-29 | url-status = live}}</ref>
|-
| style="background-color:#cef2e0;"|[[Western gorilla]]
| style="background-color:#cef2e0; text-align: right; padding-right: 1em;"|200,000
| style="background-color:#cef2e0; text-align: right; padding-right: 1em;"|[[Critically endangered]]
| style="background-color:#cef2e0; text-align: center;"|<ref name="gorillas">{{cite web |url = http://www.unep.org/Documents.Multilingual/Default.asp?DocumentID=556&ArticleID=6033&l=en&t=long |title= Gorillas on Thin Ice |date = 15 January 2009 |publisher = [[United Nations Environment Programme]] |access-date= 19 May 2010|url-status= dead |archive-url = http://arquivo.pt/wayback/20160518164244/http://www.unep.org/Documents.Multilingual/Default.asp?DocumentID=556&ArticleID=6033&l=en&t=long |archive-date= 18 May 2016}}</ref>
|-
| style="background-color:#cef2e0;"|[[Eastern gorilla]]
| style="background-color:#cef2e0; text-align: right; padding-right: 1em;"|<5,000
| style="background-color:#cef2e0; text-align: right; padding-right: 1em;"|[[Critically endangered]]
| style="background-color:#cef2e0; text-align: center;"|<ref>{{cite report | author=IUCN | title=Eastern Gorilla: Gorilla beringei Plumptre, A., Robbins, M.M. & Williamson, E.A.: The IUCN Red List of Threatened Species 2019: e.T39994A115576640 | date=2018-08-02 | doi=10.2305/iucn.uk.2019-1.rlts.t39994a115576640.en | url = https://www.iucnredlist.org/species/39994/115576640 | archiveurl = https://web.archive.org/web/20240704163729/https://www.iucnredlist.org/species/39994/115576640 | archivedate = 2024-07-04 | url-status = live}}</ref>
|-
| style="background-color:#ddcef2;"|[[Chimpanzee]]
| style="background-color:#ddcef2; text-align: right; padding-right: 1em;"|200,000
| style="background-color:#ddcef2; text-align: right; padding-right: 1em;"|[[Endangered]]
| style="background-color:#ddcef2; text-align: center;"|<ref name="chimpanzees">{{cite journal |first = Linda |last = Vigilant |year = 2004 |title = Chimpanzees |journal = [[Current Biology]] |volume = 14 |issue = 10 |pages = R369–R371 |doi= 10.1016/j.cub.2004.05.006 |pmid = 15186757|doi-access = free|bibcode = 2004CBio...14.R369V}}</ref><ref>{{cite web | title = Chimpanzees | website = WWF | date = 2024-05-28 | url = https://wwf.panda.org/discover/knowledge_hub/endangered_species/great_apes/chimpanzees/ | access-date = 2024-09-17 | archiveurl = https://web.archive.org/web/20240814133459/https://wwf.panda.org/discover/knowledge_hub/endangered_species/great_apes/chimpanzees/ | archivedate = 2024-08-14 | url-status = live}}</ref>
|-
| style="background-color:#ddcef2;"|[[Bonobo]]
| style="background-color:#ddcef2; text-align: right; padding-right: 1em;"|10,000
| style="background-color:#ddcef2; text-align: right; padding-right: 1em;"|[[Endangered]]
| style="background-color:#ddcef2; text-align: center;"|<ref name="chimpanzees" />
|-
| style="background-color:#c3d52f;"|[[Human]]
| style="background-color:#c3d52f; text-align: right; padding-right: 1em;"|8,040,640,000
| style="background-color:#c3d52f; text-align: right; padding-right: 1em;"|N/A
| style="background-color:#c3d52f; text-align: center;"|<ref>{{cite web|url=https://www.census.gov/popclock/|title=U.S. and World Population Clock|publisher=United States Census Bureau|access-date=22 November 2022}}</ref>
|}
 
==Phylogeny==
{{more citations needed|section|date=July 2019}}
{{anchor|Classification}}
Line 124 ⟶ 196:
</ref>
{{Clade|{{Clade
|label1=''[[Ponginae]] (14)''
|1={{Clade
|1=''[[Kenyapithecus]]'' (†13 Mya)
|2=''[[Sivapithecus]]'' (†9)
|3=''Crown [[Ponginae|Crown Ponginae]]''
|state4=dashed
|4=''[[Ankarapithecus]]'' (†9)
|state5=dashed
|5=''[[Giganthopithecus]]'' (†0.1)
|state6=dashed
|6=''[[Khoratpithecus]]'' (†7)
}}
|label2=(13)
Line 140 ⟶ 212:
|label1=(12)
|1={{Clade
|1=''[[Pierolapithecus]]'' (†11)
|2=''[[Hispanopithecus]]'' (†10)
}}
|2={{Clade
|1=''[[Lufengpithecus]]'' (†7)
|2=''[[Khoratpithecus]]'' (†9)
}}
}}
|label3=''[[Homininae]]'' (13)
|3={{Clade
|label2=Crown ''[[Homininae|Crown Homininae]]'' (10)
|1={{Clade
|label1=''[[Hominini]]'' (7)|1={{clade
|1={{clade
|1=''[[Ardipithecus]]'' (incl. ''[[Homo]])''|2=''[[Pan (genus)|Pan]]''}}
|2=[[Graecopithecus]] (†8)
}}
|2= ''[[Ouranopithecus]]'' (†7)
|label3=''[[Gorillini]]''
|3={{Clade
|1=''Crown [[Gorillini|Crown Gorillini]]''
|2=''[[Chororapithecus]]'' (†)
}}
}}
|3=''[[Nakalipithecus]] ''(†10)
|state4=dashed
|4=''[[Samburupithecus]]'' (†9)
}}
}}|label1='''Hominidae (18)'''}}
Line 181 ⟶ 253:
{{anchor|Foo}}
 
====Extant====
{{see also|List of hominoids}}
There are eight living species of great ape which are classified in four [[genus|genera]]. The following classification is commonly accepted:<ref name=MSW3 />
Line 217 ⟶ 289:
******* [[Anatomically modern human]], ''Homo sapiens sapiens''
 
====Fossil====
[[File:Paranthropus boisei skull.jpg|thumb|upright|Replica of [[OH 5|the skull sometimes known as "Nutcracker Man"]], found by [[Mary Leakey]]]]
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.<ref>{{cite web |url=http://www.helsinki.fi/~mhaaramo/metazoa/deuterostoma/chordata/synapsida/eutheria/primates/hominoidea/hominoidea.html | title = Hominoidea | publisher = Mikko's Phylogeny Archive | date = 14 January 2005 | first = Mikko | last = Haaramo}}</ref>
Line 338 ⟶ 410:
****** ''[[Homo sapiens idaltu]]''[[†]]
****** [[Archaic humans|Archaic ''Homo sapiens'']][[†]]
 
==Description==
[[File:Gorilla 019.jpg|thumb|Gorilla]]
The great apes are tailless primates, with the smallest living species being the bonobo at {{Convert|30 to 40|kg|lb}} in weight, and the largest being the eastern gorillas, with males weighing {{Convert|140 to 180|kg|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 monkey]]s and gibbons, although they are especially large in gorillas. The [[dentition|dental formula]] is {{DentalFormula|upper=2.1.2.3|lower=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]].<ref name=Brace&Mahler>{{Cite journal|date=1971|last1=Brace |first1=C. Loring |author-link=C. Loring Brace |last2=Mahler |first2=Paul Emil |title=Post-Pleistocene changes in the human dentition|journal=[[American Journal of Physical Anthropology]]|volume=34|issue=2 |pages=191–203|doi=10.1002/ajpa.1330340205 |pmid=5572603 |hdl=2027.42/37509 |url = https://deepblue.lib.umich.edu/bitstream/2027.42/37509/1/1330340205_ftp.pdf|hdl-access=free}}</ref><ref name=Wrangham2007>{{cite book |year = 2007 |first = Richard |last = Wrangham | chapter=Chapter 12: The Cooking Enigma | editor=Charles Pasternak | title=What Makes Us Human? | location=Oxford | publisher=Oneworld Press |isbn=978-1-85168-519-6}}</ref>
 
==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.<ref name=EoM>{{cite book|editor-last= Macdonald|editor-first= D.|last1= Harcourt|first1= A.H.|last2= MacKinnon|first2= J.|last3= Wrangham |first3= R.W.|date= 1984|title= The Encyclopedia of Mammals|publisher= Facts on File|location= New York|pages= [https://archive.org/details/encyclopediaofma00mals_0/page/422 422–439]|isbn= 978-0-87196-871-5|url-access= registration|url= https://archive.org/details/encyclopediaofma00mals_0/page/422}}</ref> 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.<ref>{{cite journal | url=https://link.springer.com/article/10.1007/s10764-020-00186-z | doi=10.1007/s10764-020-00186-z | title=Deciding Where to Sleep: Spatial Levels of Nesting Selection in Chimpanzees (Pan troglodytes) Living in Savanna at Issa, Tanzania | date=2020 | last1=Hernandez-Aguilar | first1=R. Adriana | last2=Reitan | first2=Trond | journal=International Journal of Primatology | volume=41 | issue=6 | pages=870–900 | doi-access=free }}</ref>
 
All species are [[omnivore|omnivorous]],<ref>{{Cite web |last=Alina |first=Bradford |date=29 May 2015 |title=Facts About Apes |url=https://www.livescience.com/51017-ape-facts.html |access-date=2021-01-25 |website=livescience.com |language=en}}</ref> 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 [[cereal]]s and other [[starch]]y foods, including increasingly highly [[Food processing|processed foods]], as well as many other [[Domestication#Plants|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 [[weaning|weaned]] for several years,<ref name="Hamilton 2017">{{cite web |last=Hamilton |first=Jon |title=Orangutan Moms Are The Primate Champs Of Breast-Feeding |website=NPR |date=2017-05-17 |url=https://www.npr.org/sections/health-shots/2017/05/17/528776636/orangutan-moms-are-the-primate-champs-of-breast-feeding |access-date=2024-05-20}}</ref> 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.<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 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==
{{Main|Great ape personhood|Great Ape Project|Countries banning non-human ape experimentation}}
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 ape personhood|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 [[Great ape research ban|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.<ref>{{cite news |url = https://www.reuters.com/article/scienceNews/idUSL256586320080625 |title = Spanish parliament to extend rights to apes |access-date = 11 July 2008 |date = 25 June 2008 |work = Reuters}}</ref> On 8 September 2010, the [[European Union]] [[great ape research ban|banned the testing of great apes]].<ref>{{cite web |url = https://www.independent.co.uk/life-style/health-and-families/new-eu-rules-on-animal-testing-ban-use-of-apes-2077443.html |title = New EU rules on animal testing ban use of apes |website = [[Independent.co.uk]] |date = 12 September 2010}}</ref>
 
==Conservation==
The following table lists the estimated number of great ape individuals living outside zoos.
{| class="wikitable sortable" style="font-weight: bold;"
|-
! style="background-color:#efefef;" | Species
! style="background-color:#efefef;" | Estimated<br />number
! style="background-color:#efefef;" | Conservation<br />status
! style="background-color:#efefef;" class="unsortable" | Refs
|-
| style="background-color:#eeddbb;"|[[Bornean orangutan]]
| style="background-color:#eeddbb; text-align: right; padding-right: 1em;"|61,234
| style="background-color:#eeddbb; text-align: right; padding-right: 1em;"|[[Critically endangered]]
| style="background-color:#eeddbb; text-align: center;"|<ref name="orangutans2">{{cite web |url= http://www.yorku.ca/arusson/Papers/GoI%20OU%20action%20plan%2007-17.pdf |title= Orangutan Action Plan 2007–2017|date= 2007 |publisher= [[Government of Indonesia]] |page = 5 |language = id |access-date= 1 May 2010}}\</ref>
|-
| style="background-color:#eeddbb;"|[[Sumatran orangutan]]
| style="background-color:#eeddbb; text-align: right; padding-right: 1em;"|6,667
| style="background-color:#eeddbb; text-align: right; padding-right: 1em;"|[[Critically endangered]]
| style="background-color:#eeddbb; text-align: center;"|<ref name="orangutans">An estimate of the number of wild [[orangutans]] in 2004: {{cite web |url = http://www.yorku.ca/arusson/Papers/GoI%20OU%20action%20plan%2007-17.pdf|title= Orangutan Action Plan 2007–2017 |year = 2007 |publisher= [[Government of Indonesia]]}}</ref>
|-
| style="background-color:#eeddbb;"|[[Tapanuli orangutan]]
| style="background-color:#eeddbb; text-align: right; padding-right: 1em;"|800
| style="background-color:#eeddbb; text-align: right; padding-right: 1em;"|[[Critically endangered]]
| style="background-color:#eeddbb; text-align: center;"|<ref name=":4">{{Cite news |url = https://www.theguardian.com/science/2017/nov/02/new-species-of-orangutan-discovered-in-northern-sumatra-tapanuli-pongo-tapanuliensis |title = New species of orangutan discovered in Sumatra – and is already endangered |last=Davis |first=Nicola |date=2017-11-02 |newspaper = The Guardian |access-date=2017-11-03 |language=en-GB |issn = 0261-3077}}</ref>
|-
| style="background-color:#cef2e0;"|[[Western gorilla]]
| style="background-color:#cef2e0; text-align: right; padding-right: 1em;"|200,000
| style="background-color:#cef2e0; text-align: right; padding-right: 1em;"|[[Critically endangered]]
| style="background-color:#cef2e0; text-align: center;"|<ref name="gorillas">{{cite web |url = http://www.unep.org/Documents.Multilingual/Default.asp?DocumentID=556&ArticleID=6033&l=en&t=long |title= Gorillas on Thin Ice |date = 15 January 2009 |publisher = [[United Nations Environment Programme]] |access-date= 19 May 2010|url-status= dead |archive-url = http://arquivo.pt/wayback/20160518164244/http://www.unep.org/Documents.Multilingual/Default.asp?DocumentID=556&ArticleID=6033&l=en&t=long |archive-date= 18 May 2016}}</ref>
|-
| style="background-color:#cef2e0;"|[[Eastern gorilla]]
| style="background-color:#cef2e0; text-align: right; padding-right: 1em;"|6,000
| style="background-color:#cef2e0; text-align: right; padding-right: 1em;"|[[Critically endangered]]
| style="background-color:#cef2e0; text-align: center;"|<ref name="gorillas" />
|-
| style="background-color:#ddcef2;"|[[Chimpanzee]]
| style="background-color:#ddcef2; text-align: right; padding-right: 1em;"|200,000
| style="background-color:#ddcef2; text-align: right; padding-right: 1em;"|[[Endangered]]
| style="background-color:#ddcef2; text-align: center;"|<ref name="chimpanzees">{{cite journal |first = Linda |last = Vigilant |year = 2004 |title = Chimpanzees |journal = [[Current Biology]] |volume = 14 |issue = 10 |pages = R369–R371 |doi= 10.1016/j.cub.2004.05.006 |pmid = 15186757|doi-access = free|bibcode = 2004CBio...14.R369V}}</ref><ref>{{cite web | url=https://wwf.panda.org/discover/knowledge_hub/endangered_species/great_apes/chimpanzees/ | title=Chimpanzees}}</ref>
|-
| style="background-color:#ddcef2;"|[[Bonobo]]
| style="background-color:#ddcef2; text-align: right; padding-right: 1em;"|10,000
| style="background-color:#ddcef2; text-align: right; padding-right: 1em;"|[[Endangered]]
| style="background-color:#ddcef2; text-align: center;"|<ref name="chimpanzees" />
|-
| style="background-color:#c3d52f;"|[[Human]]
| style="background-color:#c3d52f; text-align: right; padding-right: 1em;"|8,040,640,000
| style="background-color:#c3d52f; text-align: right; padding-right: 1em;"|N/A
| style="background-color:#c3d52f; text-align: center;"|<ref>{{cite web|url=https://www.census.gov/popclock/|title=U.S. and World Population Clock|publisher=United States Census Bureau|access-date=22 November 2022}}</ref>
|}
 
==See also==
Line 456 ⟶ 457:
{{Taxonbar |from = Q635162}}
{{Portal bar|Evolutionary biology|Science}}
 
 
{{Authority control}}