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{{Speciesbox
| fossil_range = [[Pleistocene|Early–Middle Pleistocene]]<br/>~{{fossil range|2|0.3}}
| image = Gigantopithecus
| image_caption = Reconstructed ''Gigantopithecus'' [[mandible]] at the [[
| display_parents = 2
| taxon = Gigantopithecus blacki
| authority = [[von Koenigswald]], 1935<ref name=Koenigswald1935>{{cite journal|last1=von Koenigswald|first1=G. H. R.|title=Eine fossile Säugetierfauna mit Simia aus Südchina|journal=Proceedings of the Koninklijke Akademie van Wetenschappen te Amsterdam|date=1935|volume=38|issue=8|pages=874–879|url=http://www.dwc.knaw.nl/DL/publications/PU00016780.pdf|access-date=12 December 2017|archive-date=12 December 2017|archive-url=https://web.archive.org/web/20171212084230/http://www.dwc.knaw.nl/DL/publications/PU00016780.pdf|url-status=live}}</ref>
}}
'''''Gigantopithecus''''' ({{IPAc-en|d|ʒ|aɪ|ˌ|g|æ|n|t|oʊ|p|ɪ|ˈ|θ|i|k|ə|s|,_|ˈ|p|ɪ|θ|ɪ|k|ə|s|,_|d|ʒ|ɪ|-}} {{respell|jy|gan|toh|pih|THEE|kəs|,_|-PITH|ih|kəs|,_|jih-}}<ref>{{Cite web|url=https://www.dictionary.com/browse/gigantopithecus|title=Definition of gigantopithecus {{!}} Dictionary.com|website=dictionary.com|language=en|access-date=2 October 2022|archive-date=3 October 2022|archive-url=https://web.archive.org/web/20221003015831/https://www.dictionary.com/browse/gigantopithecus|url-status=live}}</ref>){{efn|name=title|{{lit|giant ape}}; from [[Ancient Greek]] {{lang|grc|γίγας}} ({{translit|grc|gígas}}), meaning 'giant', and {{lang|grc|πίθηκος}} ({{translit|grc|píthekos}}), meaning 'ape'}} is an extinct [[genus]] of [[ape]] that lived in
''Gigantopithecus'' has traditionally been restored as a massive, [[gorilla]]-like ape, potentially {{convert|200|-|300|kg|abbr=in}} when alive, but the paucity of remains make total size estimates highly speculative. The species may have been [[sexual dimorphism|sexually dimorphic]], with males much bigger than females. The incisors are reduced and the canines appear to have functioned like [[cheek teeth]] ([[premolar]]s and molars). The premolars are high-[[crown (tooth)|crowned]], and the fourth premolar is very molar-like. The molars are the largest of any known ape, and have a relatively flat surface. ''Gigantopithecus'' had the thickest [[tooth enamel|enamel]] by absolute measure of any ape, up to {{convert|6|mm|in|abbr=in|frac=4}} in some areas, though this is only fairly thick when tooth size is taken into account.
''Gigantopithecus'' appears to have been a [[generalist and specialist species|generalist]] herbivore of [[C3 carbon fixation|C<sub>3</sub>]] forest plants, with the jaw adapted to grinding, crushing, and cutting through tough, fibrous plants, and the thick enamel functioning to resist foods with abrasive particles such as stems, roots, and [[tuber]]s with dirt. Some teeth bear traces of [[fig family]] fruits, which may have been important dietary components. It primarily lived in subtropical to tropical forest, and went extinct about 300,000 years ago likely because of the retreat of preferred habitat due to climate change, and potentially [[archaic human]] activity. ''Gigantopithecus'' has become popular in [[cryptozoology]] circles as the identity of the Tibetan [[yeti]] or the American [[bigfoot]], apelike creatures in local folklore
==Discovery==
===Research history===
[[Image:F. Schrenk mit Gigantopithecus-Molar 2005.jpg|thumb|left|{{Interlanguage link multi|Friedemann Schrenk|de}} holding the [[holotype]] ''Gigantopithecus blacki'' [[Molar (tooth)|molar]]]]
''Gigantopithecus blacki'' was named by anthropologist [[Ralph von Koenigswald]] in 1935 based on two third lower [[molar teeth]], which, he noted, were of enormous size (the first was "''Ein gewaltig grosser (...) Molar''", the second was described as "''der enorme Grösse besitzt''"), measuring {{convert|20|x|22|mm|abbr=in|frac=8}}.<ref name=Koenigswald1935/> The specific name ''blacki'' is in honour of Canadian palaeoanthropologist [[Davidson Black]], who had studied human evolution in China and had died the previous year. Von Koenigswald, working for the [[Dutch East Indies]] Mineralogical Survey on Java, had found the teeth in a drugstore in [[Hong Kong]] where they were being sold as "[[dragon bones]]" to be used in [[traditional Chinese medicine]]. By 1939, after purchasing more teeth, he determined they had originated somewhere in [[Guangdong]] or [[Guangxi]]. He could not formally describe the [[type specimen]] until 1952 due to his [[internment]] by Japanese forces during [[World War II]]. The originally discovered teeth are part of the collection of the [[University of Utrecht]].<ref name=ZhangHarrison2017/><ref name=Hartwig2002/> While on [[Java]], with the onset of [[World War II]], von Koenigswald put the ''Gigantopithecus'' teeth in a milk bottle and buried them in a friend's backyard before being interned by Japanese forces. After the war, he recovered the fossils and moved to [[New York City]] and could not continue research on the subject.<ref name=Zhang2024/>
In 1955, a survey team that was led by Chinese palaeontologist [[Pei Wenzhong]] was tasked by the Chinese [[Institute of Vertebrate Paleontology and Paleoanthropology|Institute of Vertebrate Palaeontology and Palaeoanthropology]] (IVPP) with finding the original ''Gigantopithecus'' locality. They collected 47 teeth among shipments of "dragon bones" in Guangdong and Guangxi. In 1956, the team discovered the first ''[[in situ]]'' remains, a third molar and [[premolar]], in a cave (subsequently named "''Gigantopithecus'' Cave") in [[Niusui Mountain]], Guangxi. Also in 1956, [[Liucheng County|Liucheng]] farmer
Confirmed ''Gigantopithecus'' remains have since been found in 16 different sites across southern China. The northernmost sites are {{ill|Longgupo|de|Longgupo-Höhle}} and [[Longgu Cave]], just south of the [[Yangtze River]], and southernmost on Hainan Island in the [[South China Sea]]. An isolated canine from [[Thẩm Khuyên Cave]], Vietnam, and a fourth premolar from [[Pha Bong]], Thailand, could possibly be assigned to ''Gigantopithecus'', though these could also represent the extinct orangutan ''[[Pongo weidenreichi]]''.<ref name=ZhangHarrison2017/>
===Classification===
====''G. blacki''====
[[File:Orangutan -Zoologischer Garten Berlin-8a.jpg|thumb|right|200px|''Gigantopithecus'' is
In 1935, von Koenigswald considered ''Gigantopithecus'' to be closely allied with the [[Late Miocene]] ''[[Sivapithecus]]'' from India.<ref name=Koenigswald1935/> In 1939, South African palaeontologist [[Robert Broom]] hypothesised that it was closely allied with ''[[Australopithecus]]'' and the [[last common ancestor]] of humans and other apes.<ref>{{cite journal|last=Broom|first=R.|author-link=Robert Broom|year=1939|title=The dentition of the Transvaal Pleistocene anthropoids, ''Plesianthropus'' and ''Paranthropus''|journal=Annals of the Transvaal Museum|volume=19|issue=3|pages=303–314|url=https://journals.co.za/docserver/fulltext/nfi_annalstm/19/3/484.pdf?expires=1587575563&id=id&accname=guest&checksum=2AB75E7DD6F91D015A8D2145BD8137F9}}</ref> In 1946, Jewish German anthropologist [[Franz Weidenreich]] described ''Gigantopithecus'' as a human ancestor as "''Gigantanthropus''", believing that the human lineage went through a gigantic phase. He stated that the teeth are more similar to those of modern humans and ''[[Homo erectus]]'' (at the time "''Pithecanthropus''" for [[Java Man|early Javan specimens]]), and envisioned a lineage from ''Gigantopithecus'', to the Javan ape ''[[Meganthropus]]'' (then considered a human ancestor), to "''Pithecanthropus''", to "''[[Solo Man|Javanthropus]]''", and finally [[Aboriginal Australian]]s. This was part of his [[multiregional origin of modern humans|
''Gigantopithecus'' is now classified in the subfamily [[Ponginae]], closely allied with ''Sivapithecus'' and ''Indopithecus''. This would make its closest living relatives the [[orangutan]]s. However, there are few similar traits ([[synapomorphies]]) linking ''Gigantopithecus'' and orangutans due to fragmentary remains, with the main morphological argument being its close affinities to ''Sivapithecus'', which is better established as a pongine based on skull features. In 2017, Chinese palaeoanthropologist Yingqi Zhang and American anthropologist Terry Harrison suggested that ''Gigantopithecus'' is most closely allied
In 2019, [[peptide sequencing]] of [[dentine]] and [[Tooth enamel|enamel]] proteins of a ''Gigantopithecus'' molar from [[Chuifeng Cave]] indicates that ''Gigantopithecus'' was indeed closely allied with orangutans, and, assuming the current [[mutation rate]] in orangutans has remained constant, shared a common ancestor about 12–10 million years ago in the [[Middle Miocene|Middle]] to Late Miocene. Their last common ancestor would have been a part of the Miocene [[Adaptive radiation|radiation]] of apes. The same study calculated a divergence time between the Ponginae and African [[great ape]]s about 26–17.7 million years ago.<ref name= "Welker2019">{{cite journal|last1= Welker|first1= F.|last2= Ramos-Madrigal|first2= J.|last3= Kuhlwilm|first3= M.|last4= Liao|first4= W.|last5= Gutenbrunner|first5= P.|last6=de Manuel|first6= M.|last7= Samodova|first7= D.|last8= Mackie|first8= M.|last9= Allentoft|first9=M. E.|last10= Bacon|first10= A.-M.|last11= Collins|first11=M. J.|last12= Cox|first12= J.|last13= Lalueza-Fox|first13 =C.|last14= Olsen|first14=J. V.|last15= Demeter|first15= F.|last16= Wang|first16= W.|last17= Marques-Bonet|first17= T.|last18= Cappellini|first18= E.|display-authors= 3|title= Enamel proteome shows that ''Gigantopithecus'' was an early diverging pongine|journal= Nature|year= 2019|doi= 10.1038/s41586-019-1728-8|pmc= 6908745|pmid= 31723270|volume= 576|issue= 7786|pages= 262–265|bibcode= 2019Natur.576..262W}}</ref>
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[[File:Giganthopithecus blacki, molar.jpeg|thumb|''Gigantopithecus'' molar]]
The tooth enamel on the molars is in absolute measure the thickest of any known ape, averaging {{convert|2.5|-|2.9|mm|abbr=in|frac=32}} in three different molars, and over {{convert|6|mm|abbr=in|frac=4}} on the tongue-side (lingual) cusps of an upper molar.<ref name=Kono2014/> This has attracted comparisons with the extinct ''[[Paranthropus]]'' hominins, which had extremely large molars and thick enamel for their size.<ref name="Dean2003"/><ref name=Kono2014>{{cite journal|first1=R. T.|last1=Kono|first2=Y.|last2=Zhang|first3=C.|last3=Jin|first4=M.|last4=Takai|first5=G.|last5=Suwa|year=2014|title=A 3-dimensional assessment of molar enamel thickness and distribution pattern in ''Gigantopithecus blacki''|journal=Quaternary International|volume=354|pages=46–51|doi=10.1016/j.quaint.2014.02.012|bibcode=2014QuInt.354...46K}}</ref> However, in relation to the tooth's size, enamel thickness for ''Gigantopithecus'' overlaps with that of several other living and extinct apes. Like orangutans and potentially all pongines (though unlike African apes) the ''Gigantopithecus'' molar
==Palaeobiology==
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''Gigantopithecus'' is considered to have been a [[herbivore]]. [[Isotope analysis#Carbon-13|Carbon-13 isotope analysis]] suggests consumption of [[C3 carbon fixation|C<sub>3</sub>]] plants, such as fruits, leaves, and other forest plants.<ref name= "Bocherens2017"/> The robust mandible of ''Gigantopithecus'' indicates it was capable of resisting high strains while chewing through tough or hard foods. However, the same mandibular anatomy is typically seen in modern apes which primarily eat soft leaves ([[folivore]]s) or seeds ([[granivore]]s). ''Gigantopithecus'' teeth have a markedly lower rate of pitting (caused by eating small, hard objects) than orangutans, more similar to the rate seen in [[chimpanzee]]s, which could indicate a similarly [[generalist and specialist species|generalist diet]].<ref name=ZhangHarrison2017/>
The molar-like premolars, large molars, and long rooted cheeked teeth could point to chewing, crushing, and grinding of bulky and fibrous materials.<ref>{{cite journal|last1=Kupczik|first1=K.|last2=Dean|first2=M. C.|year=2008|title=Comparative observations on the tooth root morphology of ''Gigantopithecus blacki''|journal=Journal of Human Evolution|volume=54|issue=2|pages=196–204|doi=10.1016/j.jhevol.2007.09.013|pmid=18045651|bibcode=2008JHumE..54..196K }}</ref><ref name="Ciochon1991">{{cite journal|last1=Ciochon |first1=R. |author-link=Russell Ciochon |last2= Piperno|first2=D. R.|last3=Thompson|first3=R. G.|year=1990|title=Opal phytoliths found on the teeth of the extinct ape ''Gigantopithecus blacki'': implications for paleodietary studies|journal=Proceedings of the National Academy of Sciences|volume=87|issue=20|pages=8120–8124|doi=10.1073/pnas.87.20.8120|pmid=2236026 |pmc=54904 |bibcode=1990PNAS...87.8120C |doi-access=free}}</ref> Thick enamel would suggest a diet of abrasive items, such as dirt particles on food gathered near or on the ground (like [[bamboo shoot]]s).<ref name=Kono2014/> Similarly, oxygen isotope analysis suggests ''Gigantopithecus'' consumed more low-lying plants such as stems, roots, and grasses than orangutans. [[Dental calculus]] indicates the consumption of [[tuber]]s.<ref>{{cite journal|first1=Y.|last1=Qu|first2=C.|last2=Jin|first3=Y.|last3=Zhang|year=2014|display-authors=et al.|title=Preservation assessments and carbon and oxygen isotopes analysis of tooth enamel of ''Gigantopithecus blacki'' and contemporary animals from Sanhe Cave, Chongzuo, South China during the Early Pleistocene|journal=Quaternary International|volume=354|pages=52–58|doi=10.1016/j.quaint.2013.10.053|bibcode=2014QuInt.354...52Q}}</ref> ''Gigantopithecus'' does not appear to have consumed the commonplace savanna grasses ([[C4 carbon fixation|C<sub>4</sub>]] plants).<ref name= "Bocherens2017">{{cite journal |last1=Bocherens |first1=H. |last2=Schrenk |first2=F. |last3=Chaimanee |first3=Y. |last4=Kullmer |first4=O. |last5=Mörike |first5=D. |last6=Pushkina |first6=D. |last7=Jaeger |first7= J.-J. |title=Flexibility of diet and habitat in Pleistocene South Asian mammals: Implications for the fate of the giant fossil ape ''Gigantopithecus'' |journal=Quaternary International |volume=434 |year=2017 |pages=148–155 |doi=10.1016/j.quaint.2015.11.059|bibcode=2017QuInt.434..148B }}</ref> Nonetheless, in 1990, a few opal [[phytolith]]s adhering to four teeth from ''Gigantopithecus'' Cave were identified to have originated from grasses; though, the majority of phytoliths resemble the hairs of [[fig family]] fruits, which include [[Common fig|fig]]s, [[mulberry]], [[breadfruit]], and [[banyan]]. This suggests that fruit was a significant dietary component for at least this population of ''Gigantopithecus''.<ref name="Ciochon1991"/>
The
In 1957, based on hoofed animal remains in a cave located in a seemingly inaccessible mountain, Pei had believed that ''Gigantopithecus'' was a cave-dwelling predator and carried these animals in.<ref>{{cite journal |last=Pei |first=Wen-chung |author-link=Pei Wenzhong |year=1957 |title=Giant ape's jaw bone discovered in China |journal=American Anthropologist |volume=59 |issue=5 |pages=834–838 |doi=10.1525/aa.1957.59.5.02a00080 |doi-access=free}}</ref> This hypothesis is no longer considered viable because its dental anatomy is consistent with herbivory.<ref name= "Bocherens2017"/> In 1975, American palaeoanthropologist [[Tim D. White]] drew similarities between the jaws and dentition of ''Gigantopithecus'' and those of the [[giant panda]], and suggested they both occupied the same [[Ecological niche|niche]] as [[bamboo]] specialists.<ref>{{cite journal|first=T. D.|last=White|author-link=Tim D. White|year=1975|title=Geomorphology to paleoecology: ''Gigantopithecus'' reappraised|journal=Journal of Human Evolution|volume=4|issue=3|pages=219–233|doi=10.1016/0047-2484(75)90009-3|bibcode=1975JHumE...4..219W }}</ref> This garnered support from some subsequent researchers, but thicker enamel and [[hypsodont]]y in ''Gigantopithecus'' could suggest different functionality for these teeth.<ref name=Kono2014/>
===Growth===
A ''Gigantopithecus'' permanent third molar, based on an approximate 600–800 days required for the enamel on the [[Cusp (anatomy)|cusp]]s to form (which is quite long), was estimated to have taken four years to form, which is within the range (albeit, far upper range) of what is exhibited in humans and chimpanzees. Like many other fossil apes, the rate of enamel formation near the enamel-dentine junction (dentine is the nerve-filled layer beneath the enamel) was estimated to begin at about 4 μm per day; this is seen in only baby teeth for modern apes.<ref name=Dean2003>{{cite journal|first1=M. C.|last1=Dean|first2=F.|last2=Schrenk|year=2003|title=Enamel thickness and development in a third permanent molar of ''Gigantopithecus blacki''|journal=Journal of Human Evolution|volume=45|issue=5|pages=381–388|doi=10.1016/j.jhevol.2003.08.009|pmid=14624748|bibcode=2003JHumE..45..381D }}</ref>
Protein sequencing of ''Gigantopithecus'' enamel identified [[alpha-2-HS-glycoprotein]] (AHSG), which, in modern apes, is important in bone and dentine mineralisation. Because it was found in enamel, and not dentine, AHSG may have been an additional component in ''Gigantopithecus'' teeth which facilitated [[biomineralisation]] of enamel during prolonged [[amelogenesis]] (enamel growth).<ref name= "Welker2019"/>
===Pathology===
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== Extinction ==
''Gigantopithecus'' fossil sites range across Guangxi, Guizhou, Hainan and Hubei Provinces, but those post-dating about 400,000 years ago are only known from Guangxi. Its youngest definitive remains in China are roughly 295,000 to 215,000 years old.<ref name="NAT-20240110"/> Two possible teeth (PIN 5792/439 and PIN 5792/490) have been reported from the [[Late Pleistocene]] deposit in Vietnam,<ref>{{Cite journal |last1=Lopatin |first1=A. V. |last2=Maschenko |first2=E. N. |last3=Dac |first3=Le Xuan |year=2022 |title=''Gigantopithecus blacki'' (Primates, Ponginae) from the Lang Trang Cave (Northern Vietnam): The Latest ''Gigantopithecus'' in the Late Pleistocene? |url=https://link.springer.com/10.1134/S0012496622010069 |journal=Doklady Biological Sciences |language=en |volume=502 |issue=1 |pages=6–10 |doi=10.1134/S0012496622010069 |pmid=35298746 |s2cid=254413457 |issn=0012-4966|url-access=subscription }}</ref> but
Human activity in southern China is known as early as 800,000 years ago but does not become prevalent until after the extinction of ''Gigantopithecus'', so it is unclear if pressures such as competition over resources or overhunting were factors.<ref name="Ciochon2009">{{cite journal |last=Ciochon |first=Russell L. |author-link=Russell Ciochon |date=17 June 2009 |title=The mystery ape of Pleistocene Asia |journal=Nature |volume=459 |issue=7249 |pages=910–911 |bibcode=2009Natur.459..910C |doi=10.1038/459910a |pmid=19536242 |doi-access=free}}</ref>
==Cryptozoology==
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