View source for Ape

{{Short description|Branch of primates}}
{{Distinguish-redirect|Hominoid|Hominid}}
{{About|the branch of primates}}
{{pp-vandalism|small=yes}}
{{pp-move-indef}}
{{Use dmy dates|date=December 2019}}
{{automatic taxobox
| name = Hominoids<br/>Apes
| fossil_range = [[Miocene]]-[[Holocene]]
| image = 013 Alpha male chimpanzee at Kibale forest National Park Photo by Giles Laurent.jpg
| image_caption = Male [[chimpanzee]] (''Pan troglodytes'')
| taxon = Hominoidea
| authority = [[John Edward Gray|Gray]], 1825<ref>{{Cite journal |last=Gray |first=J. E. |title=An outline of an attempt at the disposition of Mammalia into tribes and families, with a list of the genera apparently appertaining to each tribe. |url=http://www.rhinoresourcecenter.com/index.php?s=1&act=pdfviewer&id=1223371163&folder=122 |journal=Annals of Philosophy |series=New Series |volume=10 |pages=337–344 |access-date=27 April 2022 |archive-date=27 April 2022 |archive-url=https://web.archive.org/web/20220427185350/http://www.rhinoresourcecenter.com/index.php?s=1&act=pdfviewer&id=1223371163&folder=122 |url-status=live }}</ref>
| type_species = ''[[Human|Homo sapiens]]''
| type_species_authority = [[Carl Linnaeus|Linnaeus]], [[10th edition of Systema Naturae|1758]]
| subdivision_ranks = Families
| subdivision = * †[[Proconsulidae]]
* †[[Afropithecidae]]
* †[[Pliobates|Pliobatidae]]
* †[[Dendropithecidae]]
* [[Gibbon|Hylobatidae]]
* [[Hominidae]]
}}

'''Apes''' (collectively '''Hominoidea''' {{IPAc-en|h|ɒ|m|ᵻ|ˈ|n|ɔɪ|d|i|.|ə}}) are a [[clade]] of [[Old World]] [[simian]]s native to [[sub-Saharan Africa]] and [[Southeast Asia]] (though they were more widespread in Africa, most of Asia, and Europe in prehistory, and counting humans are found globally). Apes are more closely related to [[Old World monkey]]s (family Cercopithecidae) than to the [[New World monkey]]s (Platyrrhini) with both Old World monkeys and apes placed in the [[clade]] [[Catarrhini]]. Apes do not have tails due to a mutation of the [[T-box transcription factor T|TBXT gene]].<ref>{{Cite bioRxiv |last1=Xia |first1=Bo |last2=Zhang |first2=Weimin |last3=Wudzinska |first3=Aleksandra |last4=Huang |first4=Emily |last5=Brosh |first5=Ran |last6=Pour |first6=Maayan |last7=Miller |first7=Alexander |last8=Dasen |first8=Jeremy S. |last9=Maurano |first9=Matthew T. |last10=Kim |first10=Sang Y. |last11=Boeke |first11=Jef D. |date=2021-09-16 |title=The genetic basis of tail-loss evolution in humans and apes |biorxiv=10.1101/2021.09.14.460388}}</ref><ref name="CNN-20240323">{{cite news |last=Weisberger |first=Mindy |title=Why don't humans have tails? Scientists find answers in an unlikely place |url=https://www.cnn.com/2024/03/23/world/humans-tails-genetic-mutation-junk-dna-scn/index.html |date=March 23, 2024 |work=[[CNN]] |url-status=live |archiveurl=https://archive.today/20240324031927/https://www.cnn.com/2024/03/23/world/humans-tails-genetic-mutation-junk-dna-scn/index.html |archivedate=March 24, 2024 |accessdate=March 24, 2024 }}</ref> In traditional and non-scientific use, the term ''ape'' can include tailless primates taxonomically considered Cercopithecidae (such as the [[Barbary ape]] and [[Celebes crested macaque|black ape]]), and is thus not equivalent to the scientific [[taxon]] Hominoidea. There are two extant branches of the superfamily Hominoidea: the [[gibbon]]s, or lesser apes; and the [[hominid]]s, or [[great apes]].
* The family [[Hylobatidae]], the lesser apes, include four genera and a total of 20 species of gibbon, including the [[lar gibbon]] and the [[siamang]], all native to [[Asia]]. They are highly arboreal and [[bipedal]] on the ground. They have lighter bodies and smaller social groups than great apes.
* The family [[Hominidae]] (''hominids''), the great apes, include four genera comprising three extant species of [[orangutan]]s and their subspecies, two extant species of [[gorilla]]s and their subspecies, two extant species of [[Pan (genus)|chimpanzees]] and their subspecies, and [[human]]s in a single extant subspecies.{{efn|name=DawkinsUse1|Although Dawkins is clear that he uses "apes" for Hominoidea, he also uses "great apes" in ways which exclude humans. Thus in {{Harvnb|Dawkins|2005}}: "Long before people thought in terms of evolution ... great apes were often confused with humans" (p. 114); "gibbons are faithfully monogamous, unlike the great apes which are our closer relatives" (p. 126).}}{{Sfn|Dixson|1981|pp=13}}<ref>{{Cite journal |last=Grehan |first=J. R. |year=2006 |title=Mona Lisa smile: the morphological enigma of human and great ape evolution |journal=Anatomical Record |volume=289B |pages=139–157 |doi=10.1002/ar.b.20107 |pmid=16865704 |issue=4 |doi-access=free}}</ref><ref name=Benton2005p371/>

Except for gorillas and humans, hominoids are agile climbers of trees. Apes eat a variety of plant and animal foods, with the majority of food being plant foods, which can include fruits, leaves, stalks, roots and seeds, including nuts and grass seeds. Human diets are sometimes substantially different from that of other hominoids due in part to the development of technology and a wide range of habitation.

All non-human hominoids are rare and threatened with [[extinct]]ion. The main threat is [[habitat loss]], though some populations are further imperiled by hunting. The great apes of Africa are also facing threat from the [[Ebola virus]].<ref>{{cite news |url=https://www.independent.co.uk/news/world/africa/ebola-virus-has-killed-a-third-of-the-worlds-gorillas-and-chimpanzees--and-could-pose-greatest-threat-to-their-survival-conservationists-warn-9998386.html |title=Ebola virus 'has killed a third of world's gorillas and chimpanzees' – and could pose greatest threat to their survival, conservationists warn |last=Rush |first=J. |date=23 January 2015 |work=The Independent |access-date=26 March 2015 |archive-url=https://web.archive.org/web/20150330035716/http://www.independent.co.uk/news/world/africa/ebola-virus-has-killed-a-third-of-the-worlds-gorillas-and-chimpanzees--and-could-pose-greatest-threat-to-their-survival-conservationists-warn-9998386.html |archive-date=30 March 2015 |url-status=dead}}</ref>

== Name and terminology ==
{{Human timeline}}
"Ape", from Old English ''apa'', is a word of uncertain origin.<!---->{{efn|reference=The hypothetical Proto-Germanic form is given as ''*apōn'' (F. Kluge, ''Etymologisches Wörterbuch der Deutschen Sprache'' (2002), online version, s.v. "[http://etymology_de.deacademic.com/279/Affe Affe]"; V. Orel, ''A handbook of Germanic etymology'' (2003), s.v. "[https://archive.org/stream/Orel-AHandbookOfGermanicEtymology/2003OrelV.-AHandbookOfGermanicEtymology#page/n59/mode/2up *apōn]" or as *''apa(n)'' (''Online Etymology Dictionary'' (2001–2014), s.v. "[http://www.etymonline.com/index.php?l=a&p=36 ape]"; M. Philippa, F. Debrabandere, A. Quak, T. Schoonheim & N. van der Sijs, ''Etymologisch woordenboek van het Nederlands'' (2003–2009), s.v. "[http://www.etymologiebank.nl/trefwoord/aap aap]"). Perhaps ultimately derived from a non-Indo-European language, the word might be a direct borrowing from Celtic, or perhaps from Slavic, although in both cases it is also argued that the borrowing, if it took place, went in the opposite direction.}} The term has a history of rather imprecise usage—and of comedic or punning usage in the vernacular. Its earliest meaning was generally of any non-human anthropoid primate, as is still the case for its cognates in other Germanic languages.{{efn|reference="Any simian known on the Mediterranean during the Middle Ages; monkey or ape"; cf. ''ape-ward'': "a juggler who keeps a trained monkey for the amusement of the crowd." (Middle English Dictionary, s.v. "[http://quod.lib.umich.edu/cgi/m/mec/med-idx?size=First+100&type=headword&q1=ape&rgxp=constrained ape]").}}{{Sfn|Terry|1977|pp=3}}
Later, after the term "[[monkey]]" had been introduced into English, "ape" was specialized to refer to a tailless (therefore exceptionally human-like) primate.{{Sfn|Terry|1977|pp=3–4}} Thus, the term "ape" obtained two different meanings, as shown in the 1911 [[Encyclopædia Britannica Eleventh Edition|''Encyclopædia Britannica'']] entry: it could be used as a synonym for "monkey" and it could denote the tailless human-like primate in particular.<ref name="EB1911">{{Cite EB1911|wstitle=Ape|volume=2|page=160}}</ref>

Some, or recently all, hominoids are also called "apes", but the term is used broadly and has several different senses within both popular and scientific settings. "Ape" has been used as a synonym for "monkey" or for naming any primate with a human-like appearance, particularly those without a tail.<ref name="EB1911"/> Biologists have traditionally used the term "ape" to mean a member of the superfamily Hominoidea ''other than'' humans,{{Sfn|Dixson|1981|pp=13}} but more recently to mean ''all'' members of Hominoidea. So "ape"—not to be confused with "great ape"—now becomes another word for '''hominoid''' ''including'' humans.<ref name=Benton2005p371/>{{efn|name=DawkinsUse2|{{Harvnb|Dawkins|2005}}; for example "[a]ll apes except humans are hairy" (p. 99), "[a]mong the apes, gibbons are second only to humans" (p. 126).}}

The taxonomic term ''hominoid'' is derived from, and intended as encompassing, the ''[[hominid]]s,'' the family of [[great ape]]s. Both terms were introduced by Gray (1825).<ref>{{Cite journal |last=Gray |first=JE |title=An outline of an attempt at the disposition of Mammalia into tribes and families, with a list of the genera apparently appertaining to each tribe. |url=http://www.rhinoresourcecenter.com/index.php?s=1&act=pdfviewer&id=1223371163&folder=122 |journal=Annals of Philosophy |series=New Series |volume=10 |pages=337–344 |access-date=27 April 2022 |archive-date=27 April 2022 |archive-url=https://web.archive.org/web/20220427185350/http://www.rhinoresourcecenter.com/index.php?s=1&act=pdfviewer&id=1223371163&folder=122 |url-status=live }}</ref> The term ''[[hominin]]s'' is also due to Gray (1824), intended as including the human lineage (see also [[Hominidae#Terminology]], [[Human taxonomy]]).

The distinction between apes and monkeys is complicated by the traditional [[paraphyly]] of monkeys: Apes emerged as a sister group of Old World Monkeys in the [[Catarrhini|catarrhines]], which are a sister group of New World Monkeys. Therefore, [[Cladistics|cladistically]], apes, catarrhines and related contemporary extinct groups such as [[Parapithecidae]] are monkeys as well, for any consistent definition of "monkey". "Old World monkey" may also legitimately be taken to be meant to include all the catarrhines, including apes and extinct species such as ''[[Aegyptopithecus]]'',<ref name="OsmanHill1953">{{Cite book |title=Primates Comparative Anatomy and Taxonomy I—Strepsirhini |last=Osman Hill |first=W. C. |publisher=Edinburgh University Press |year=1953 |series=Edinburgh Univ Pubs Science & Maths, No 3 |pages=53 |oclc=500576914 |author-link=William Charles Osman Hill}}</ref><ref>{{Cite book |url=https://archive.org/details/generalintroduct00martrich |title=A General Introduction to the Natural History of Mammiferous Animals, With a Particular View of the Physical History of man, and the More Closely Allied Genera of the Order Quadrumana, or Monkeys |last=Martin |first=W. C. L. |publisher=Wright and Co. printers |year=1841 |location=London |pages=340, 361}}</ref><ref>{{Cite journal |last=Geoffroy Saint-Hilaire |first=M. É. |date=1812 |title=Tableau des quadrumanes, ou des animaux composant le premier ordre de la classe des Mammifères |url=https://www.biodiversitylibrary.org/item/23270#page/107/mode/1up |journal=Annales du Muséum d'Histoire Naturelle |location=Paris |volume=19 |pages=85–122 |access-date=16 July 2019 |archive-date=27 March 2019 |archive-url=https://web.archive.org/web/20190327091031/https://www.biodiversitylibrary.org/item/23270#page/107/mode/1up |url-status=live }}</ref><ref name="Bugge1974">{{Cite journal |last=Bugge |first=J. |date=1974 |title=Chapter 4 |journal=Cells Tissues Organs |volume=87 |issue=Suppl. 62 |pages=32–43 |doi=10.1159/000144209 |issn=1422-6405}}</ref>{{citation needed|date=October 2016}} in which case the apes, Cercopithecoidea and ''Aegyptopithecus'' emerged within the Old World monkeys.

The primates called "apes" today became known to Europeans after the 18th century. As zoological knowledge developed, it became clear that taillessness occurred in a number of different and otherwise distantly related [[species]]. Sir [[Wilfrid Le Gros Clark]] was one of those primatologists who developed the idea that there were trends in primate evolution and that the extant members of the order could be arranged in an "ascending series", leading from "monkeys" to "apes" to humans. Within this tradition "ape" came to refer to all members of the superfamily Hominoidea except humans.{{Sfn|Dixson|1981|pp=13}} As such, this use of "apes" represented a [[paraphyly|paraphyletic]] grouping, meaning that, even though all species of apes were descended from a common ancestor, this grouping did ''not'' include all the descendant species, because humans were excluded from being among the apes.{{efn|Definitions of [[paraphyly]] vary; for the one used here see e.g. {{Harvnb|Stace|2010|pp=106}}}}

Traditionally, the English-language [[Common name|vernacular name]] "apes" does not include humans, but phylogenetically, humans (''[[Homo]]'') form part of the family Hominidae within Hominoidea. Thus, there are at least three common, or traditional, uses of the term "ape": non-specialists may not distinguish between "monkeys" and "apes", that is, they may use the two terms interchangeably; or they may use "ape" for any tailless monkey or non-human hominoid; or they may use the term "ape" to just mean the non-human hominoids.

Modern taxonomy aims for the use of [[monophyly|monophyletic]] groups for taxonomic classification;<ref name="SpringerHolley2011">{{cite book |author1=Springer |author2=D. H. |title=An Introduction to Zoology: Investigating the Animal World |date=1 July 2011 |publisher=Jones & Bartlett Publishers |isbn=978-0-7637-5286-6 |pages=536 |quote=Through careful study taxonomists today struggle to eliminate polyphyletic and paraphyletic groups and taxons, reclassifying their members into appropriate monophyletic taxa}}</ref>{{efn|Definitions of [[monophyly]] vary; for the one used here see e.g. {{Harvnb|Mishler|2009|pp=114}}}}
Some literature may now use the common name "ape" to mean all members of the superfamily Hominoidea, including humans. For example, in his 2005 book, Benton wrote "The apes, Hominoidea, today include the gibbons and orang-utan ... the gorilla and chimpanzee ... and humans".<ref name=Benton2005p371>{{Cite book |last=Benton |first=M. J. |year=2005 |title=Vertebrate Palaeontology |publisher=Wiley-Blackwell |isbn=978-0-632-05637-8 |url=https://books.google.com/books?id=SyJO3vpCk8AC |access-date=10 July 2011 |archive-date=3 April 2023 |archive-url=https://web.archive.org/web/20230403152612/https://books.google.com/books?id=SyJO3vpCk8AC |url-status=live }}, p. 371</ref> Modern biologists and primatologists refer to apes that are not human as "non-human" apes. Scientists broadly, other than paleoanthropologists, may use the term "[[hominin]]" to identify [[the human clade]], replacing the term "[[hominid]]". See [[Hominidae#Terminology|terminology of primate names]].

See below, [[#History of hominoid taxonomy|History of hominoid taxonomy]], for a discussion of changes in scientific classification and terminology regarding hominoids.

== Evolution ==
{{Unreferenced section|date=September 2021}}

Although the hominoid fossil record is still incomplete and fragmentary, there is now enough evidence to provide an outline of the [[Timeline of human evolution|evolutionary history of humans]]. Previously, the divergence between humans and other extant hominoids was thought to have occurred 15 to 20 million years ago, and several species of that time period, such as ''[[Ramapithecus]]'', were once thought to be [[hominin]]s and possible ancestors of humans. But, later fossil finds indicated that ''Ramapithecus'' was more closely related to the orangutan; and new biochemical evidence indicates that the last common ancestor of humans and non-hominins (that is, the chimpanzees) occurred between 5 and 10 million years ago, and probably nearer the lower end of that range (more recent); see [[Chimpanzee–human last common ancestor]] (CHLCA).

== Taxonomic classification and phylogeny ==
{{main|List of hominoids}}

Genetic analysis combined with fossil evidence indicates that hominoids diverged from the [[Old World monkey]]s about 25 million years ago (mya), near the [[Oligocene–Miocene boundary]].<ref>{{cite web |url=https://www.science.org/content/article/fossils-may-pinpoint-critical-split-between-apes-and-monkeys |title=Fossils may pinpoint critical split between apes and monkeys |date=15 May 2013 |access-date=30 June 2022 |archive-date=16 December 2022 |archive-url=https://web.archive.org/web/20221216065413/https://www.science.org/content/article/fossils-may-pinpoint-critical-split-between-apes-and-monkeys |url-status=live }}</ref><ref>{{cite journal |last1=Rossie |first1=J. B. |last2=Hill |first2=A. |year=2018 |title=A new species of ''Simiolus'' from the middle Miocene of the Tugen Hills, Kenya |journal=Journal of Human Evolution |volume=125 |pages=50–58 |doi=10.1016/j.jhevol.2018.09.002 |pmid=30502897|bibcode=2018JHumE.125...50R |s2cid=54625375 }}</ref><ref>{{cite journal |last1=Rasmussen |first1=D. T. |last2=Friscia |first2=A. R. |last3=Gutierrez |first3=M. |display-authors=3 |last4=Kappelman |first4=J. |last5=Miller |first5=E. R. |last6=Muteti |first6=S. |last7=Reynoso |first7=D. |last8=Rossie |first8=J. B. |last9=Spell |first9=T. L. |last10=Tabor |first10=N. J. |last11=Gierlowski-Kordesch |first11=E. |last12=Jacobs |first12=B. F. |last13=Kyongo |first13=B. |last14=Macharwas |first14=M. |last15=Muchemi |first15=F. |year=2019 |title=Primitive Old World monkey from the earliest Miocene of Kenya and the evolution of cercopithecoid bilophodonty |journal=Proceedings of the National Academy of Sciences |volume=116 |issue=13 |pages=6051–6056 |doi=10.1073/pnas.1815423116 |pmid=30858323 |pmc=6442627 |bibcode=2019PNAS..116.6051R |doi-access=free}}</ref> The gibbons split from the rest about 18 mya, and the hominid splits happened 14 mya (''Pongo''),<ref>{{Cite journal |last1=Alba |first1=D. M. |last2=Fortuny |first2=J. |last3=Moyà-Solà |first3=S. |date=2010 |title=Enamel thickness in the Middle Miocene great apes ''Anoiapithecus'', ''Pierolapithecus'' and ''Dryopithecus'' |journal=Proceedings of the Royal Society of London B: Biological Sciences |volume=277 |issue=1691 |pages=2237–2245 |doi=10.1098/rspb.2010.0218|issn=0962-8452 |pmc=2880156 |pmid=20335211}}</ref> 7 mya (''Gorilla''), and 3–5 mya (''Homo'' & ''Pan'').<ref>{{Cite journal |last1=Grabowski |first1=M. |last2=Jungers |first2=W. L. |date=2017 |title=Evidence of a chimpanzee-sized ancestor of humans but a gibbon-sized ancestor of apes |journal=Nature Communications |volume=8 |issue=1 |pages=880 |bibcode=2017NatCo...8..880G |doi=10.1038/s41467-017-00997-4|issn=2041-1723 |pmc=5638852 |pmid=29026075}}</ref><!-- note, this figure of 3-5 mya for the Homo/Pan split differs from the figure in the article [[Great ape]], which says 6.5-7.4 mya or alternatively 5.4-6.3 mya--> In 2015, a new genus and species were described, ''[[Pliobates cataloniae]]'', which lived 11.6 mya, and appears to predate the split between Hominidae and Hylobatidae.<ref>{{cite web |url=http://phys.org/news/2015-10-primate-species-root-tree-extant.html |title=A new primate species at the root of the tree of extant hominoids |date=29 October 2015 |access-date=29 October 2015 |archive-date=29 October 2015 |archive-url=https://web.archive.org/web/20151029203847/http://phys.org/news/2015-10-primate-species-root-tree-extant.html |url-status=live }}</ref><ref name=":0">{{Cite journal |last1=Nengo |first1=I. |last2=Tafforeau |first2=P. |last3=Gilbert |first3=C. C. |display-authors=3 |last4=Fleagle |first4=J. G. |last5=Miller |first5=E. R. |last6=Feibel |first6=C. |last7=Fox |first7=D. L. |last8=Feinberg |first8=J. |last9=Pugh |first9=K. D. |year=2017 |title=New infant cranium from the African Miocene sheds light on ape evolution |url=http://discovery.ucl.ac.uk/1570349/1/Spoor_Nengo-et-al_text.pdf |journal=Nature |volume=548 |issue=7666 |pages=169–174 |bibcode=2017Natur.548..169N |doi=10.1038/nature23456|pmid=28796200|s2cid=4397839 |access-date=15 July 2019 |archive-url=https://web.archive.org/web/20180722103154/http://discovery.ucl.ac.uk/1570349/1/Spoor_Nengo-et-al_text.pdf |archive-date=22 July 2018 |url-status=dead}}</ref>{{Sfn|Dixson|1981|p=16}}<ref name="Benton2005p371" />{{clarify |date=July 2019 |reason=It's not clear which of these sources contains the cladogram shown, and it would be SYNTH to combine cladograms from different sources}}
{{Clade|{{Clade
  |label1='''(29)'''
  |1={{Clade
    |1=[[Saadanioidea]] (†28)
    |label2='''[[Cercopithecoidea]] (24)'''
    |2={{Clade
      |1=[[Victoriapithecus|Victoriapithecinae]] (†19)
      |2='''[[Cercopithecoidea|Crown Cercopithecoidea]]'''
    }}
  }}
|label2='''Hominoidea (30)'''
|2={{Clade
       |1=[[Dendropithecidae]] (†7 Mya)
       |label2=
       |2={{clade
         |1=''[[Ekembo|Ekembo heseloni]]'' (†17 Mya)
|2={{clade
           |1={{clade
             |2=''[[Ekembo|Ekembo nyanzae]]'' (†17 Mya)
  |1=[[Proconsulidae]] (†18 Mya)
}}
  |label2='''(29)'''
  |2={{Clade
    |1=''[[Equatorius]]'' (†16)
|2=''[[Pliobates]]'' (†11.6 Mya)
    |label3='''(29)'''
    |3={{Clade
      |label1=[[Afropithecidae]] (28)
      |1={{Clade
         |1=''[[Morotopithecus]]'' (†20)

            |2=''[[Afropithecus]]'' (†16)
}}
      |label2='''Crown Hominoidea (22)'''
      |2={{Clade
        |1='''[[Hominidae]]'''
        |2='''[[Hylobatidae]]'''
      }}
    }}
  }}
}}
}}}}}}|label1='''[[Catharrhini|Crown Catharrhini]] (31)'''|style1=line-height:80%;font-size:80%}}
{{Clade|style=line-height:80%;font-size:80%
|label1=[[Catarrhini]] (31.0 Mya)
|1={{Clade
   |grouplabel1=<br><br><br>Traditional apes
   |label1=Hominoidea/apes (20.4 Mya)
   |1={{Clade
      |label1=[[Hominidae]]/great apes (15.7 Mya)
      |1={{Clade
         |label1=[[Homininae]] (8.8 Mya)
         |1={{Clade
            |label1=[[Hominini]] (6.3 Mya)
            |1={{Clade
               |1=humans&nbsp;(genus&nbsp;''Homo'') [[File:Bechuana of Distinction-1841 (white background).jpg|60 px]]
               |2=chimpanzees&nbsp;(genus&nbsp;''Pan'') [[File:PanTroglodytesSmit (white background).jpg|50 px]]|barbegin2=purple
               }}
            |2=gorillas&nbsp;(genus ''Gorilla'') [[File:Gorila de llanura occidental. Gorilla gorilla - Blanca Martí de Ahumada (white background).jpg|60 px]]|bar2=purple
            }}
         |2=orangutans&nbsp;(genus ''Pongo'') [[File:Simia satyrus - 1837 - Print - Iconographia Zoologica - Special Collections University of Amsterdam - White Background.jpg|50 px]]|bar2=purple
         }}
      |2=gibbons/lesser apes (family&nbsp;Hylobatidae) [[File:Le gibbon (white background).jpg|50 px]]|barend2=purple
      }}
      |2=[[Cercopithecoidea]] [[File:Cynocephalus doguera - 1700-1880 - Print - Iconographia Zoologica - Special Collections University of Amsterdam - (white background).tiff|50 px]]&nbsp;Old World monkeys
   }}
}}

[[File:Ape skeletons.png|thumb|510px|Skeletons of members of the ape superfamily, '''Hominoidea'''. There are two extant families: [[Hominidae]], the "great apes"; and [[Hylobatidae]], the gibbons, or "lesser apes".]]

[[File:Comparison of size of gibbon, human, chimpanzee (Pan troglodytes) western gorilla (Gorilla gorilla) (standing), and orangutan (Pongo abelii).svg|thumb|510px|From left: Comparison of size of gibbon, human, chimpanzee, gorilla and orangutan. Non-human hominoids do not stand upright as their normal posture.]]

The families, and extant genera and species of hominoids are:
* '''Superfamily Hominoidea'''<ref name=MSW3/>
** Family [[Hominidae]]: hominids ("great apes")
*** Genus ''[[Orangutan|Pongo]]'': orangutans
**** [[Bornean orangutan]], ''P. pygmaeus''
**** [[Sumatran orangutan]], ''P. abelii''
**** [[Tapanuli orangutan]], ''P. tapanuliensis''<ref name="NYT-20171102">{{cite news |last=Cochrane |first=J. |title=New Orangutan Species Could Be the Most Endangered Great Ape |url=https://www.nytimes.com/2017/11/02/science/new-orangutan-species.html |date=2 November 2017 |work=[[The New York Times]] |access-date=3 November 2017 |archive-date=17 April 2018 |archive-url=https://web.archive.org/web/20180417014924/https://www.nytimes.com/2017/11/02/science/new-orangutan-species.html |url-status=live }}</ref>
*** Genus ''[[Gorilla]]'': gorillas
**** [[Western gorilla]], ''G. gorilla''
**** [[Eastern gorilla]], ''G. beringei''
*** Genus ''[[Homo]]'': humans
**** [[Human]], ''H. sapiens''
*** Genus ''[[Pan (genus)|Pan]]'': chimpanzees
**** [[Chimpanzee]], ''P. troglodytes''
**** [[Bonobo]], ''P. paniscus''
** Family [[Hylobatidae]]: gibbons ("lesser apes")
*** Genus ''[[Hylobates]]''
**** [[Lar gibbon]] or white-handed gibbon, ''H. lar''
**** [[Bornean white-bearded gibbon]], ''H. albibarbis''
**** [[Agile gibbon]] or black-handed gibbon, ''H. agilis''
****[[Western grey gibbon]] or Abbott's grey gibbon, ''H. abbotti''<ref name=":1">{{Cite book |last1=Wilson |first1=Don E. |last2=Cavallini |first2=Paolo |date=2013 |title=Handbook of the Mammals of the World |location=Barcelona |publisher=Lynx Edicions |isbn=978-84-96553-89-7 |oclc=1222638259}}</ref>
**** [[Eastern grey gibbon]] or northern grey gibbon, ''H. funereus''<ref name=":1" />
**** [[Müller's gibbon]] or southern grey gibbon, ''H. muelleri''
**** [[Silvery gibbon]], ''H. moloch''
**** [[Pileated gibbon]] or capped gibbon, ''H. pileatus''
**** [[Kloss's gibbon]] or Mentawai gibbon or bilou, ''H. klossii''
*** Genus ''[[Hoolock]]''
**** [[Western hoolock gibbon]], ''H. hoolock''
**** [[Eastern hoolock gibbon]], ''H. leuconedys''
**** [[Skywalker hoolock gibbon]], ''H. tianxing''<!--as per wiki article on Gibbons-->
*** Genus ''[[Symphalangus]]''
**** [[Siamang]], ''S. syndactylus''
*** Genus ''[[Nomascus]]''
**** [[Northern buffed-cheeked gibbon]], ''N. annamensis''<!--as per wiki article on Gibbons-->
**** [[Black crested gibbon]], ''N. concolor''
**** [[Eastern black crested gibbon]], ''N. nasutus''
**** [[Hainan black crested gibbon]], ''N. hainanus''
**** [[Southern white-cheeked gibbon]] ''N. siki''
**** [[White-cheeked crested gibbon]], ''N. leucogenys''
**** [[Yellow-cheeked gibbon]], ''N. gabriellae''

=== History of hominoid taxonomy ===
{{see|Human taxonomy#History}}

The history of hominoid taxonomy is complex and somewhat confusing. Recent evidence has changed our understanding of the relationships between the hominoids, especially regarding the human lineage; and the traditionally used terms have become somewhat confused. Competing approaches to methodology and terminology are found among current scientific sources. Over time, authorities have changed the names and the meanings of names of groups and subgroups as new evidence — that is, new discoveries of fossils and tools and of observations in the field, plus continual comparisons of anatomy and [[DNA sequences]] — has changed the understanding of relationships between hominoids. There has been a gradual demotion of humans from being 'special' in the taxonomy to being one branch among many. This recent turmoil (of history) illustrates the growing influence on all taxonomy of [[cladistics]], the science of classifying living things strictly according to their lines of descent.{{citation needed|date=December 2021}}

Today, there are eight extant [[genus|genera]] of hominoids. They are the four genera in the family Hominidae, namely ''[[Homo]]'', ''[[Pan (genus)|Pan]]'', ''[[Gorilla]]'', and ''[[Orangutan|Pongo]]''; plus four genera in the family Hylobatidae (gibbons): ''[[Hylobates]]'', ''[[Hoolock]]'', ''[[Nomascus]]'' and ''[[Symphalangus]]''.<ref name=MSW3>{{MSW3 Groves|pages=178–184|id=12100751}}</ref> (The two subspecies of [[hoolock gibbons]] were recently moved from the genus ''[[Bunopithecus]]'' to the new genus ''Hoolock'' and re-ranked as species; a third species was described in January 2017).<ref name="Mootnick">{{cite journal |last1=Mootnick |first1=A. |last2=Groves |first2=C. P. |year=2005 |title=A new generic name for the hoolock gibbon (Hylobatidae) |journal=International Journal of Primatology |issue=4 |pages=971–976 |doi=10.1007/s10764-005-5332-4 |volume=26 |s2cid=8394136}}</ref>

In 1758, [[Carl Linnaeus]], relying on second- or third-hand accounts, placed a second species in ''Homo'' along with ''H. sapiens'': ''Homo troglodytes'' ("cave-dwelling man"). Although the term "Orang Outang" is listed as a variety – ''Homo sylvestris'' – under this species, it is nevertheless not clear to which animal this name refers, as Linnaeus had no specimen to refer to, hence no precise description. Linnaeus may have based ''Homo troglodytes'' on reports of mythical creatures, then-unidentified [[simian]]s, or Asian natives dressed in animal skins.<ref>{{cite book |last1=Frängsmyr |first1=T. |last2=Lindroth |first2=S. |last3=Eriksson |first3=G. |last4=Broberg |first4=G. |title=Linnaeus, the man and his work |year=1983 |publisher=[[University of California Press]] |location=Berkeley and Los Angeles |isbn=978-0-7112-1841-3}}, p. 166</ref> Linnaeus named the orangutan ''Simia satyrus'' ("satyr monkey"). He placed the three genera ''Homo'', ''[[Simia]]'' and ''Lemur'' in the order of Primates.

The ''troglodytes'' name was used for the chimpanzee by [[Johann Friedrich Blumenbach|Blumenbach]] in 1775, but moved to the genus ''Simia''. The orangutan was moved to the genus ''[[Orangutan|Pongo]]'' in 1799 by [[Bernard Germain Étienne de la Ville, Comte de Lacépède|Lacépède]].

Linnaeus's inclusion of humans in the primates with monkeys and apes was troubling for people who denied a close relationship between humans and the rest of the animal kingdom. Linnaeus's Lutheran archbishop had accused him of "impiety". In a letter to [[Johann Georg Gmelin]] dated 25 February 1747, Linnaeus wrote:

{{Blockquote|text=It is not pleasing to me that I must place humans among the primates, but man is intimately familiar with himself. Let's not quibble over words. It will be the same to me whatever name is applied. But I desperately seek from you and from the whole world a general difference between men and simians from the principles of Natural History. I certainly know of none. If only someone might tell me one! If I called man a simian or vice versa I would bring together all the theologians against me. Perhaps I ought to, in accordance with the law of Natural History.<ref>{{cite web |url=http://linnaeus.c18.net/Letters/display_txt.php?id_letter=L0783 |title=Letter, Carl Linnaeus to Johann Georg Gmelin. Uppsala, Sweden, 25 February 1747 |publisher=Swedish Linnaean Society |access-date=4 February 2009 |archive-date=27 February 2009 |archive-url=https://web.archive.org/web/20090227174240/http://linnaeus.c18.net/Letters/display_txt.php?id_letter=L0783 |url-status=live }}</ref>}}

Accordingly, [[Johann Friedrich Blumenbach]] in the first edition of his ''Manual of Natural History'' (1779), proposed that the primates be divided into the [[Quadrumana]] (four-handed, i.e. apes and monkeys) and [[Bimana]] (two-handed, i.e. humans). This distinction was taken up by other naturalists, most notably [[Georges Cuvier]]. Some elevated the distinction to the level of [[order (biology)|order]].

However, the many affinities between humans and other primates – and especially the "great apes" – made it clear that the distinction made no scientific sense. In his 1871 book ''[[The Descent of Man, and Selection in Relation to Sex]]'', [[Charles Darwin]] wrote:

{{Blockquote|text=The greater number of naturalists who have taken into consideration the whole structure of man, including his mental faculties, have followed Blumenbach and Cuvier, and have placed man in a separate Order, under the title of the Bimana, and therefore on an equality with the orders of the Quadrumana, Carnivora, etc. Recently many of our best naturalists have recurred to the view first propounded by Linnaeus, so remarkable for his sagacity, and have placed man in the same Order with the Quadrumana, under the title of the Primates. The justice of this conclusion will be admitted: for in the first place, we must bear in mind the comparative insignificance for classification of the great development of the brain in man, and that the strongly marked differences between the skulls of man and the Quadrumana (lately insisted upon by [[Theodor Ludwig Wilhelm Bischoff|Bischoff]], [[Christoph Theodor Aeby|Aeby]], and others) apparently follow from their differently developed brains. In the second place, we must remember that nearly all the other and more important differences between man and the Quadrumana are manifestly adaptive in their nature, and relate chiefly to the erect position of man; such as the structure of his hand, foot, and pelvis, the curvature of his spine, and the position of his head.<ref>{{cite book |first=C. |last=Darwin |title=The Descent of Man |year=1871 |isbn=978-0-7607-7814-2| title-link=The Descent of Man |publisher=Barnes & Noble |author-link=Charles Darwin}}</ref>}} <!-- ...history from 1870 to 1960 goes here... -->

=== Changes in taxonomy and terminology ===
{{see also|Hominidae|Human taxonomy}}
{| class="wikitable"
| '''Humans the non-apes''': Until about 1960, taxonomists typically divided the superfamily Hominoidea into two families. The science community treated humans and their extinct relatives as the [[outgroup (cladistics)|outgroup]] within the superfamily; that is, humans were considered as quite distant from kinship with the "apes". Humans were classified as the family [[Hominidae]] and were known as the "hominids". All other hominoids were known as "apes" and were referred to the family [[Pongidae]].<ref>{{cite journal |last=G. G. |first=Simpson |year=1945 |title=The principles of classification and a classification of mammals |journal=Bulletin of the American Museum of Natural History |volume=85 |pages=1–350 |author-link=George Gaylord Simpson}}</ref>
| [[File:Hominoid taxonomy 1.svg|400px]]
|-
| '''The "great apes" in Pongidae''': The 1960s saw the methodologies of [[molecular biology]] applied to primate taxonomy. Goodman's 1964 immunological study of serum proteins led to re-classifying the hominoids into three families: the humans in Hominidae; the great apes in Pongidae; and the "lesser apes" (gibbons) in [[Gibbon|Hylobatidae]].<ref name=Goodman>{{cite book |first=M. |last=Goodman |year=1964 |chapter=Man's place in the phylogeny of the primates as reflected in serum proteins |editor-first=S. L. |editor-last=Washburn |title=Classification and Human Evolution |publisher=Aldine |location=Chicago |pages=204–234}}</ref> However, this arrangement had two [[Speciation#Genetics|trichotomies]]: ''Pan'', ''Gorilla'', and ''Pongo'' of the "great apes" in Pongidae, and Hominidae, Pongidae, and Hylobatidae in Hominoidea. These presented a puzzle; scientists wanted to know which genus [[speciation|speciated]] first from the common hominoid ancestor.
| [[File:Hominoid taxonomy 2.svg|400px]]
|-
| '''Gibbons the outgroup''': New studies indicated that [[gibbon]]s, not humans, are the [[outgroup (cladistics)|outgroup]] within the superfamily Hominoidea, meaning: the rest of the hominoids are more closely related to each other than (any of them) are to the gibbons. With this splitting, the gibbons (''Hylobates'', ''et al''.) were isolated after moving the great apes into the same family as humans. Now the term "hominid" encompassed a larger collective taxa within the family Hominidae. With the family trichotomy settled, scientists could now work to learn which genus is 'least' related to the others in the subfamily Ponginae.
| [[File:Hominoid taxonomy 3.svg|400px]]
|-
| '''Orangutans the outgroup''': Investigations comparing humans and the three other hominid genera disclosed that the African apes (chimpanzees and gorillas) and humans are more closely related to each other than any of them are to the Asian orangutans (''Pongo''); that is, the [[orangutan]]s, not humans, are the [[Outgroup (cladistics)|outgroup]] within the family Hominidae. This led to reassigning the African apes to the subfamily Homininae with humans—which presented a new three-way split: ''Homo'', ''Pan'', and ''Gorilla''.<ref name=Goodman1>{{cite journal |title=Biochemical evidence on hominid phylogeny |first=M. |last=Goodman |journal=Annual Review of Anthropology |volume=3 |issue=1 |pages=203–228 |year=1974 |doi=10.1146/annurev.an.03.100174.001223}}</ref>
| [[File:Hominoid taxonomy 4.svg|400px]]
|-
| '''Hominins''': In an effort to resolve the trichotomy, while preserving the nostalgic "outgroup" status of humans, the subfamily Homininae was divided into two tribes: Gorillini, comprising genus ''Pan'' and genus ''Gorilla''; and Hominini, comprising genus ''Homo'' (the humans). Humans and close relatives now began to be known as "hominins", that is, of the tribe Hominini. Thus, the term "hominin" succeeded to the previous use of "hominid", which meaning had changed with changes in Hominidae (see above: 3rd graphic, "Gibbons the outgroup").
| [[File:Hominoid taxonomy 5.svg|400px]]
|-
| '''Gorillas the outgroup''': New DNA comparisons now provided evidence that [[gorillas]], not humans, are the outgroup in the subfamily Homininae; this suggested that chimpanzees should be grouped with humans in the tribe Hominini, but in separate subtribes.<ref name=Goodman2>{{cite journal  |title=Primate evolution at the DNA level and a classification of hominoids |first1=M. |last1=Goodman |first2=D. A. |last2=Tagle |first3=D. H. |last3=Fitch |first4=W. |display-authors=3 |last4=Bailey |first5=J. |last5=Czelusniak |first6=B. F. |last6=Koop |first7=P. |last7=Benson |first8=J. L. |last8=Slightom |journal=Journal of Molecular Evolution |year=1990 |volume=30 |pages=260–266 |doi = 10.1007/BF02099995 |pmid=2109087 |issue=3 |bibcode=1990JMolE..30..260G |s2cid=2112935}}</ref> Now the name "hominin" delineated ''Homo'' plus those earliest ''Homo'' relatives and ancestors that arose ''after'' the divergence from the chimpanzees. (Humans are no longer recognized as an outgroup, but are a branch, [[human evolutionary genetics|deep in the tree]] of the pre-1960s ape group).
| [[File:Hominoid taxonomy 6.svg|400px]]
|-
| '''Speciation of gibbons''': Later DNA comparisons disclosed previously unknown speciation of genus ''Hylobates'' (gibbons) into four genera: ''[[Hylobates]]'', ''[[Hoolock]]'', ''[[Nomascus]]'', and ''[[Symphalangus]]''.<ref name=MSW3/><ref name=Mootnick/> The ordering of speciation of these four genera are being [[Gibbon#Evolutionary_history|investigated]] {{As of|2022|lc=yes}}.
| [[File:Hominoid taxonomy 7.svg|400px]]
|}

== Characteristics ==
{{Unreferenced section|date=July 2011}}
[[File:SD Zoo Orangs.jpg|thumb|left|Like those of the orangutan, the shoulder joints of hominoids are adapted to [[brachiation]], or movement by swinging in tree branches.]]

The lesser apes are the gibbon family, Hylobatidae, of sixteen species; all are native to Asia. Their major differentiating characteristic is their long arms, which they use to [[brachiation|brachiate]] through trees. Their wrists are [[ball and socket joint]]s as an evolutionary adaptation to their [[arboreal]] lifestyle. Generally smaller than the African apes, the largest gibbon, the [[siamang]], weighs up to {{convert|14|kg|lb|0|abbr=on}}; in comparison, the smallest "great ape", the [[bonobo]], is {{convert|34|to|60|kg|lb|0|abbr=on}}.

The superfamily Hominoidea falls within the [[parvorder]] [[Catarrhini]], which also includes the Old World monkeys of Africa and Eurasia. Within this grouping, the two families Hylobatidae and Hominidae can be distinguished from Old World monkeys by the number of [[Cusp (dentistry)|cusps]] on their [[molar (tooth)|molars]]; hominoids have five in the "Y-5" molar pattern, whereas Old World monkeys have only four in a [[bilophodont]] pattern.

Further, in comparison with Old World monkeys, hominoids are noted for: more mobile shoulder joints and arms due to the dorsal position of the [[scapula]]; broader ribcages that are flatter front-to-back; and a shorter, less mobile spine, with greatly reduced caudal (tail) vertebrae—resulting in complete loss of the tail in extant hominoid species. These are anatomical adaptations, first, to vertical hanging and swinging locomotion ([[brachiation]]) and, later, to developing balance in a [[bipedal]] pose. Note there are primates in other families that also lack tails, and at least one, the [[pig-tailed langur]], is known to walk significant distances bipedally. The front of the ape skull is characterised by its sinuses, fusion of the frontal bone, and by [[post-orbital constriction]].

=== Distinction from monkeys ===
{{See also|Monkey#Historical and modern terminology}}

[[Cladistics|Cladistically]], apes, catarrhines, and extinct species such as [[Aegyptopithecus]] and [[Parapithecidae]]a, are monkeys,{{citation needed|date=October 2016}} so one can only specify ape features not present in other monkeys.

Unlike most [[monkey]]s, apes do not possess a tail. Monkeys are more likely to be in trees and use their tails for balance. While the great apes are considerably larger than monkeys, [[gibbon]]s (lesser apes) are smaller than some monkeys. Apes are considered to be more intelligent than monkeys, which are considered to have more primitive brains.<ref>{{cite book |title=The Gestural Communication of Apes and Monkeys |first1=J. |last1=Call |first2=M. |last2=Tomasello |year=2007 |publisher=Taylor & Francis Group/Lawrence Erlbaum Associates}}</ref>

The enzyme [[urate oxidase]] has become inactive in all apes, its function having been lost in two [[primate]] lineages during the middle Miocene; first in the common ancestors of Hominidae, and later in the common ancestor of Hylobatidae. It has been hypothesized that in both incidents it was a mutation that occurred in apes living in Europe when the climate was getting colder, leading to starvation during winter. The mutation changed the biochemistry of the apes and made it easier to accumulate fat, which allowed the animals to survive longer periods of starvation. When they migrated to Asia and Africa, this genetic trait remained.<ref>{{cite journal | pmc=3203212 | year=2011 | last1=Johnson | first1=R. J. | last2=Lanaspa | first2=M. A. | last3=Gaucher | first3=E. A. | title=Uric acid: A Danger Signal from the RNA World that may have a role in the Epidemic of Obesity, Metabolic Syndrome and CardioRenal Disease: Evolutionary Considerations | journal=Seminars in Nephrology | volume=31 | issue=5 | pages=394–399 | doi=10.1016/j.semnephrol.2011.08.002 | pmid=22000645 }}</ref><ref>{{cite journal | url=https://www.scientificamerican.com/article/ancient-mutation-in-apes-may-explain-human-obesity-and-diabetes/ | doi=10.1038/scientificamerican1015-64 | title=The Fat Gene | year=2015 | last1=Johnson | first1=Richard J. | last2=Andrews | first2=Peter | journal=Scientific American | volume=313 | issue=4 | pages=64–69 | bibcode=2015SciAm.313d..64J }}</ref>

== Behaviour ==

Major studies of behaviour in the field were completed on the three better-known "great apes", for example by [[Jane Goodall]], [[Dian Fossey]] and [[Birutė Galdikas]]. These studies have shown that in their natural environments, the non-human hominoids show sharply varying social structure: gibbons are monogamous, territorial pair-bonders, orangutans are solitary, gorillas live in small troops with a single adult male leader, while chimpanzees live in larger troops with bonobos exhibiting promiscuous sexual behaviour. Their diets also vary; gorillas are [[foliovore]]s, while the others are all primarily [[frugivore]]s, although the common chimpanzee hunts for meat. Foraging behaviour is correspondingly variable.

In November 2023, scientists reported, for the first time, evidence that groups of [[primate]]s, including apes, and, particularly [[bonobo]]s, are capable of cooperating with each other.<ref name="NYT-20231116">{{cite news |last=Zimmer |first=Carl |authorlink=Carl Zimmer |title=Scientists Find First Evidence That Groups of Apes Cooperate - Some bonobos are challenging the notion that humans are the only primates capable of group-to-group alliances. |url=https://www.nytimes.com/2023/11/16/science/bonobos-cooperation-study.html |date=16 November 2023 |work=[[The New York Times]] |url-status=live |archiveurl=https://archive.today/20231116194259/https://www.nytimes.com/2023/11/16/science/bonobos-cooperation-study.html |archivedate=16 November 2023 |accessdate=17 November 2023 }}</ref><ref name="SCI-20231116">{{cite journal |author=Samuni, Liran |display-authors=et al. |title=Cooperation across social borders in bonobos
|url=https://www.science.org/doi/10.1126/science.adg0844 |date=16 November 2023 |journal=[[Science (journal)|Science]] |volume=382 |issue=6672 |pages=805–809 |doi=10.1126/science.adg0844 |pmid=37972165 |bibcode=2023Sci...382..805S |url-status=live |archiveurl=https://archive.today/20231117125744/https://www.science.org/doi/10.1126/science.adg0844 |archivedate=17 November 2023 |accessdate=17 November 2023 }}</ref>

=== Diet ===

Apart from humans and gorillas, apes eat a predominantly [[frugivorous]] diet, mostly fruit, but supplemented with a variety of other foods. Gorillas are predominantly [[folivorous]], eating mostly stalks, shoots, roots and leaves with some fruit and other foods. Non-human apes usually eat a small amount of raw animal foods such as insects or eggs. In the case of humans, migration and the invention of hunting tools and cooking has led to an even wider variety of foods and diets, with many human diets including large amounts of cooked [[tuber]]s ([[root]]s) or [[legume]]s.<ref>{{cite web |url=https://www.newscientist.com/article/mg23230980-600-every-human-culture-includes-cooking-this-is-how-it-began |title=Every human culture includes cooking – this is how it began |first=G. |last=Lawton |date=2 November 2016 |website=[[New Scientist]] |access-date=27 August 2021 |archive-date=29 July 2021 |archive-url=https://web.archive.org/web/20210729081037/https://www.newscientist.com/article/mg23230980-600-every-human-culture-includes-cooking-this-is-how-it-began/ |url-status=live }}</ref> Other food production and processing methods including animal husbandry and industrial refining and processing have further changed human diets.<ref>{{cite journal |url=http://www.nature.com/news/humans-are-becoming-more-carnivorous-1.14282 |title=Humans are becoming more carnivorous |last=Hoag |first=Hannah |date=2 December 2013 |journal=[[Nature (journal)|Nature]] |doi=10.1038/nature.2013.14282 |s2cid=183143537 |access-date=26 November 2014 |archive-date=29 November 2014 |archive-url=https://web.archive.org/web/20141129232443/http://www.nature.com/news/humans-are-becoming-more-carnivorous-1.14282 |url-status=live }}</ref> Humans and other apes occasionally eat other primates.<ref>{{cite web |url=https://www.newscientist.com/article/dn14926-loving-bonobos-have-a-carnivorous-dark-side.html#.VHXsaclWqUU |title=Loving bonobos have a carnivorous dark side |last=Callaway |first=E. |date=13 October 2006 |work=[[New Scientist]] |access-date=26 November 2014 |archive-date=29 October 2014 |archive-url=https://web.archive.org/web/20141029130152/http://www.newscientist.com/article/dn14926-loving-bonobos-have-a-carnivorous-dark-side.html#.VHXsaclWqUU |url-status=live }}</ref> Some of these primates are now close to extinction with habitat loss being the underlying cause.<ref>{{cite web |url=http://www.bbc.com/earth/story/20150728-chimps-nearly-wiped-out-monkeys |title=Chimpanzees over-hunt monkey prey almost to extinction |first=Michael |last=M. |publisher=[[BBC Earth]] |access-date=28 May 2018 |archive-date=6 June 2018 |archive-url=https://web.archive.org/web/20180606155344/http://www.bbc.com/earth/story/20150728-chimps-nearly-wiped-out-monkeys |url-status=live }}</ref><ref>{{cite news |url=https://www.sciencedaily.com/releases/2008/08/080804222912.htm |title=Extinction threat to monkeys and other primates due to habitat loss, hunting |website=[[Science Daily]] |access-date=28 May 2018 |archive-date=28 May 2018 |archive-url=https://web.archive.org/web/20180528134736/https://www.sciencedaily.com/releases/2008/08/080804222912.htm |url-status=live }}</ref>

=== Cognition ===
[[File:Gorrila tool use-Efi.jpg|thumb|This series of images shows a gorilla using a small tree trunk as a tool. The trunk is used to maintain balance while fishing for aquatic plants]]

All the non-human hominoids are generally thought of as highly intelligent, and scientific study has broadly confirmed that they perform very well on a wide range of cognitive tests—though there is relatively little data on gibbon cognition. The early studies by [[Wolfgang Köhler]] demonstrated exceptional [[problem-solving]] abilities in chimpanzees, which Köhler attributed to [[insight]]. The [[Tool use by animals|use of tools]] has been repeatedly demonstrated; more recently, the manufacture of tools has been documented, both in the wild and in laboratory tests. [[Imitation]] is much more easily demonstrated in "great apes" than in other primate species. Almost all the studies in [[animal language acquisition]] have been done with "great apes", and though there is continuing dispute as to whether they demonstrate real language abilities, there is no doubt that they involve significant feats of learning. Chimpanzees in different parts of Africa have developed tools that are used in food acquisition, demonstrating a form of [[animal culture]].<ref>{{cite book |first=W. |last=McGrew |title=Chimpanzee Material Culture: Implications for Human Evolution |year=1992}}</ref>

== Threats and conservation ==
All non-human hominoids are rare and threatened with [[extinct]]ion. The [[eastern hoolock gibbon]] is the least threatened, only being [[vulnerable species|vulnerable]] to extinction. Five gibbon species are [[critically endangered]], as are all species of orangutan and gorilla. The remaining species of gibbon, the bonobo, and all four subspecies of chimpanzees are [[endangered species|endangered]]. The chief threat to most of the endangered species is loss of tropical [[rainforest]] habitat, though some populations are further imperiled by hunting for [[bushmeat]]. The great apes of Africa are also facing threat from the [[Ebola virus]].<ref>{{cite news |last=Rush |first=J. |date=23 January 2015 |title=Ebola virus 'has killed a third of world's gorillas and chimpanzees' – and could pose greatest threat to their survival, conservationists warn |url=https://www.independent.co.uk/news/world/africa/ebola-virus-has-killed-a-third-of-the-worlds-gorillas-and-chimpanzees--and-could-pose-greatest-threat-to-their-survival-conservationists-warn-9998386.html |url-status=dead |archive-url=https://web.archive.org/web/20150330035716/http://www.independent.co.uk/news/world/africa/ebola-virus-has-killed-a-third-of-the-worlds-gorillas-and-chimpanzees--and-could-pose-greatest-threat-to-their-survival-conservationists-warn-9998386.html |archive-date=30 March 2015 |access-date=26 March 2015 |work=The Independent}}</ref> Currently considered to be the greatest threat to survival of African apes, [[Ebola]] infection is responsible for the death of at least one third of all gorillas and chimpanzees since 1990.<ref>{{cite news |url=https://www.independent.co.uk/news/world/africa/ebola-virus-has-killed-a-third-of-the-worlds-gorillas-and-chimpanzees--and-could-pose-greatest-threat-to-their-survival-conservationists-warn-9998386.html |title=Ebola virus 'has killed a third of world's gorillas and chimpanzees' – and could pose greatest threat to their survival, conservationists warn |last=Rush |first=J. |date=23 January 2015 |work=The Independent |access-date=26 March 2015 |archive-url=https://web.archive.org/web/20150330035716/http://www.independent.co.uk/news/world/africa/ebola-virus-has-killed-a-third-of-the-worlds-gorillas-and-chimpanzees--and-could-pose-greatest-threat-to-their-survival-conservationists-warn-9998386.html |archive-date=30 March 2015 |url-status=dead}}</ref>

All the species of [[great apes]] in Africa, are considered endangered. Hunting, logging, agricultural expansion and mining are among the main threats. Recently mining has expanded due to the [[energy transition]]. According to researchers "This means that current climate solutions could lead to more industrialization in these places, which could worsen the climate crisis". The [[ Sustainable Critical Minerals Alliance]] was created for solve problems like this.<ref>{{cite journal |title=Threat of mining to African great apes |journal=Science Advances |date=3 April 2024 |volume=10 |issue=14 |doi=10.1126/sciadv.adl0335 |url=https://www.science.org/doi/10.1126/sciadv.adl0335 |access-date=19 September 2024 |last1=Junker |first1=Jessica |last2=Quoss |first2=Luise |last3=Valdez |first3=Jose |last4=Arandjelovic |first4=Mimi |last5=Barrie |first5=Abdulai |last6=Campbell |first6=Geneviève |last7=Heinicke |first7=Stefanie |last8=Humle |first8=Tatyana |last9=Kouakou |first9=Célestin Y. |last10=Kühl |first10=Hjalmar S. |last11=Ordaz-Németh |first11=Isabel |last12=Pereira |first12=Henrique M. |last13=Rainer |first13=Helga |last14=Refisch |first14=Johannes |last15=Sonter |first15=Laura |last16=Sop |first16=Tenekwetche |pmid=38569032 |bibcode=2024SciA...10L.335J }}</ref>

== See also ==
* {{Portal inline|Mammals}}
* ''[[Dawn of Humanity]]'' (2015 PBS film)
* [[Great Ape Project]]
* [[Great Apes Survival Partnership]]
* [[International Primate Day]]
* [[Kinshasa Declaration on Great Apes]]
* [[List of individual apes]] (for notable non-fictional non-human apes)
* [[List of fictional primates]]
* [[List of primates by population]]
* [[Great Ape Project#World Declaration on Great Apes|World Declaration on Great Apes]] from the [[Great Ape Project]]
{{Clear}}

==Notes==
{{noteslist}}

==References==
{{Reflist}}

===Literature cited===
{{refbegin}}
* {{Cite book |last=Dawkins |first=R. |year=2005 |title=The Ancestor's Tale |edition=p/b |location=London |publisher=Phoenix (Orion Books) |isbn=978-0-7538-1996-8}}
* {{Cite book |last=Dixson |first=A. F. |year=1981 |title=The Natural History of the Gorilla |location=London |publisher=Weidenfeld & Nicolson |isbn=978-0-297-77895-0}}
* {{Cite book |last=Mishler |first=Brent D |year=2009 |editor-last=Ayala |editor-first=F. J. |editor2-last=Arp |editor2-first=R. |contribution=Species are not uniquely real biological entities |title=Contemporary Debates in Philosophy of Biology |pages=110–122 |isbn=978-1-4443-1492-2 |doi=10.1002/9781444314922.ch6 |name-list-style=amp}} 
* {{Cite journal |last=Stace |first=C. A. |author-link=Clive A. Stace |year=2010 |title=Classification by molecules: what's in it for field botanists? |journal=Watsonia |volume=28 |pages=103–122 |url=http://www.watsonia.org.uk/Wats28p103.pdf |access-date=7 February 2010 |url-status=dead |archive-url=https://web.archive.org/web/20110726104027/https://www.watsonia.org.uk/Wats28p103.pdf |archive-date=26 July 2011}}
* {{cite book |last=Terry |first=M. W. |chapter=Use of common and scientific nomenclature to designate laboratory primates |editor-first=A. M. |editor-last=Schrier |title=Behavioral Primatology: Advances in Research and Theory |volume=1 |location=Hillsdale, N.J., US |publisher=Lawrence Erlbaum |year=1977}}
{{refend}}

==External links==
{{Sister project links}}
* {{Wikispecies-inline|Hominoidea}}
* {{Wikibooks-inline|Dichotomous Key|Hominoidea}}
* {{cite journal |author=Pilbeam D. |author-link=David Pilbeam |title=Hominoid systematics: The soft evidence |journal=Proc. Natl. Acad. Sci. U.S.A. |volume=97 |issue=20 |pages=10684–6 |date=September 2000 |pmid=10995486 |pmc=34045 |doi=10.1073/pnas.210390497 |bibcode=2000PNAS...9710684P |doi-access=free }} Agreement between cladograms based on molecular and anatomical data.
* [http://humanorigins.si.edu/evidence/human-evolution-timeline-interactive Human Timeline (Interactive)] – [[Smithsonian Institution|Smithsonian]], [[National Museum of Natural History]] (August 2016).

{{Apes}}
{{Notable apes}}
{{Primates}}
{{Haplorhini|Ho.}}
{{Taxonbar|from=Q102470}}
{{Authority control}}

[[Category:Apes| ]]
[[Category:Extant Chattian first appearances]]
[[Category:Taxa named by John Edward Gray]]