''Gigantopithecus'' ( ; ) is an extinct

genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...

of ape from roughly 2 million to 350,000 years ago during the

Early Early may refer to: History * The beginning or oldest part of a defined historical period, as opposed to middle or late periods, e.g.: ** Early Christianity ** Early modern Europe Places in the United States * Early, Iowa * Early, Texas * Early ...

Middle Pleistocene The Chibanian, widely known by its previous designation of Middle Pleistocene, is an age in the international geologic timescale or a stage in chronostratigraphy, being a division of the Pleistocene Epoch within the ongoing Quaternary Period. Th ...

of southern China, represented by one species, ''Gigantopithecus blacki''. Potential identifications have also been made in Thailand, Vietnam, and Indonesia. The first remains of ''Gigantopithecus'', two third

molar teeth The molars or molar teeth are large, flat teeth at the back of the mouth. They are more developed in mammals. They are used primarily to grind food during chewing. The name ''molar'' derives from Latin, ''molaris dens'', meaning "millstone ...

, were identified in a drugstore by anthropologist Ralph von Koenigswald in 1935, who subsequently described the ape. In 1956, the first mandible and more than 1,000 teeth were found in Liucheng, and numerous more remains have since been found in at least 16 sites. Only teeth and four mandibles are known currently, and other skeletal elements were likely consumed by porcupines before they could fossilise. ''Gigantopithecus'' was once argued to be a

hominin The Hominini form a taxonomic tribe of the subfamily Homininae ("hominines"). Hominini includes the extant genera ''Homo'' (humans) and '' Pan'' (chimpanzees and bonobos) and in standard usage excludes the genus ''Gorilla'' (gorillas). The ...

, a member of the

human Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedalism and exceptional cognitive skills due to a large and complex brain. This has enabled the development of advanced tools, cultu ...

line, but it is now thought to be closely allied with

orangutan Orangutans are great apes native to the rainforests of Indonesia and Malaysia. They are now found only in parts of Borneo and Sumatra, but during the Pleistocene they ranged throughout Southeast Asia and South China. Classified in the genu ...

s, classified in the

subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus Genus ( plural genera ) is a taxonomic rank used in the biological classifica ...

Ponginae. ''Gigantopithecus'' has traditionally been restored as a massive,

gorilla Gorillas are herbivorous, predominantly ground-dwelling great apes that inhabit the tropical forests of equatorial Africa. The genus ''Gorilla'' is divided into two species: the eastern gorilla and the western gorilla, and either four ...

-like ape, potentially when alive, but the paucity of remains make total size estimates highly speculative. The species may have been

sexually dimorphic Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most ani ...

, with males much bigger than females. The incisors are reduced and the canines appear to have functioned like

cheek teeth Cheek teeth or post-canines comprise the molar and premolar teeth in mammals. Cheek teeth are multicuspidate (having many folds or tubercles). Mammals have multicuspidate molars (three in placentals, four in marsupials, in each jaw quadrant) and p ...

(

premolar The premolars, also called premolar teeth, or bicuspids, are transitional teeth located between the canine and molar teeth. In humans, there are two premolars per quadrant in the permanent set of teeth, making eight premolars total in the mouth ...

s and molars). The premolars are high- 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 enamel by absolute measure of any ape, up to 6 mm (a quarter of an inch) in some areas, though this is only fairly thick when tooth size is taken into account. ''Gigantopithecus'' appears to have been a generalist

herbivore A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpar ...

of C₃ 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 Tubers are a type of enlarged structure used as storage organs for nutrients in some plants. They are used for the plant's perennation (survival of the winter or dry months), to provide energy and nutrients for regrowth during the next growing ...

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 A number of varieties of '' Homo'' are grouped into the broad category of archaic humans in the period that precedes and is contemporary to the emergence of the earliest early modern humans (''Homo sapiens'') around 300 ka. Omo-Kibish I (Omo I) f ...

activity. ''Gigantopithecus'' has become popular in

cryptozoology Cryptozoology is a pseudoscience and subculture that searches for and studies unknown, legendary, or extinct animals whose present existence is disputed or unsubstantiated, particularly those popular in folklore, such as Bigfoot, the Loch Ness ...

circles as the identity of the Tibetan

yeti The Yeti ()"Yeti"
''

bigfoot Bigfoot, also commonly referred to as Sasquatch, is a purported ape-like creature said to inhabit the forest of North America. Many dubious articles have been offered in attempts to prove the existence of Bigfoot, including anecdotal claims o ...

, humanlike creatures in local folklore.

Discovery

Research history

''Gigantopithecus blacki'' was named by anthropologist Ralph von Koenigswald in 1935 based on two third lower

, which, he noted, were of enormous size (the first was "''Ein gewaltig grosser (...) Molar''", the second was described as "''der enorme Grösse besitzt''"), measuring . 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 The Dutch East Indies, also known as the Netherlands East Indies ( nl, Nederlands(ch)-Indië; ), was a Dutch colony consisting of what is now Indonesia. It was formed from the nationalised trading posts of the Dutch East India Company, whic ...

Mineralogical Survey on Java, had found the teeth in a drugstore in

Hong Kong Hong Kong ( (US) or (UK); , ), officially the Hong Kong Special Administrative Region of the People's Republic of China (abbr. Hong Kong SAR or HKSAR), is a List of cities in China, city and Special administrative regions of China, special ...

where they were being sold as " dragon bones" to be used in

traditional Chinese medicine Traditional Chinese medicine (TCM) is an alternative medical practice drawn from traditional medicine in China. It has been described as "fraught with pseudoscience", with the majority of its treatments having no logical mechanism of acti ...

. By 1939, after purchasing more teeth, he determined they had originated somewhere in

Guangdong Guangdong (, ), alternatively romanized as Canton or Kwangtung, is a coastal province in South China on the north shore of the South China Sea. The capital of the province is Guangzhou. With a population of 126.01 million (as of 2020 ...

Guangxi Guangxi (; ; alternately romanized as Kwanghsi; ; za, Gvangjsih, italics=yes), officially the Guangxi Zhuang Autonomous Region (GZAR), is an autonomous region of the People's Republic of China, located in South China and bordering Vietnam ...

. He could not formally describe the

type specimen In biology, a type is a particular specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally attached. In other words, a type is an example that serves to anchor or centralizes th ...

until 1952 due to his

internment Internment is the imprisonment of people, commonly in large groups, without charges or intent to file charges. The term is especially used for the confinement "of enemy citizens in wartime or of terrorism suspects". Thus, while it can simp ...

by Japanese forces during

World War II World War II or the Second World War, often abbreviated as WWII or WW2, was a world war that lasted from 1939 to 1945. It involved the World War II by country, vast majority of the world's countries—including all of the great power ...

. The originally discovered teeth are part of the collection of the

University of Utrecht Utrecht University (UU; nl, Universiteit Utrecht, formerly ''Rijksuniversiteit Utrecht'') is a public research university in Utrecht, Netherlands. Established , it is one of the oldest universities in the Netherlands. In 2018, it had an enrollme ...

. In 1955, a survey team that was led by Chinese palaeontologist Pei Wenzhong was tasked by the Chinese

Institute of Vertebrate Palaeontology and Palaeoanthropology The Institute of Vertebrate Paleontology and Paleoanthropology (IVPP; ) of China is a research institution and collections repository for fossils, including many dinosaur and pterosaur specimens (many from the Yixian Formation). As its name sugges ...

(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 ''In situ'' (; often not italicized in English) is a Latin phrase that translates literally to "on site" or "in position." It can mean "locally", "on site", "on the premises", or "in place" to describe where an event takes place and is used in ...

'' remains, a third molar and

, in a cave (subsequently named "''Gigantopithecus'' Cave") in Niusui Mountain, Guangxi. Also in 1956, Liucheng farmer Xiuhuai Qin discovered more teeth and the first

mandible In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...

on his field. From 1957 to 1963, the IVPP survey team carried out excavations in this area and recovered two more mandibles and more than 1,000 teeth. Confirmed ''Gigantopithecus'' remains have since been found in 16 different sites across southern China. The northernmost sites are Longgupo and Longgudong, just south of the

Yangtze River The Yangtze or Yangzi ( or ; ) is the longest river in Asia, the third-longest in the world, and the longest in the world to flow entirely within one country. It rises at Jari Hill in the Tanggula Mountains (Tibetan Plateau) and flows ...

, and southernmost on Hainan Island in the

South China Sea The South China Sea is a marginal sea of the Western Pacific Ocean. It is bounded in the north by the shores of South China (hence the name), in the west by the Indochinese Peninsula, in the east by the islands of Taiwan and northwestern Ph ...

. 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''. Two mandibular fragments each preserving the last two molars from Semono in Central

Java Java (; id, Jawa, ; jv, ꦗꦮ; su, ) is one of the Greater Sunda Islands in Indonesia. It is bordered by the Indian Ocean to the south and the Java Sea to the north. With a population of 151.6 million people, Java is the world's mo ...

, Indonesia, described in 2016 could represent ''Gigantopithecus''. The oldest remains date to 2 million years ago from Baikong Cave, and the youngest 380,000–310,000 years ago from Hei Cave. In 2014, a fourth confirmed mandible was discovered in Yanliang, Central China. Indicated by extensive

rodent Rodents (from Latin , 'to gnaw') are mammals of the order Rodentia (), which are characterized by a single pair of continuously growing incisors in each of the upper and lower jaws. About 40% of all mammal species are rodents. They are n ...

gnawing marks, teeth primarily accumulated in caves likely due to

porcupine Porcupines are large rodents with coats of sharp spines, or quills, that protect them against predation. The term covers two families of animals: the Old World porcupines of family Hystricidae, and the New World porcupines of family, Erethiz ...

activity. Porcupines gnaw on bones to obtain nutrients necessary for quill growth, and can haul large bones into their underground dens and consume them entirely, except the hard, enamel-capped crowns of teeth. This may explain why teeth are typically found in great quantity, and why remains other than teeth are so rare.

Classification

''G. blacki''

In 1935, von Koenigswald considered ''Gigantopithecus'' to be closely allied with the

Late Miocene The Late Miocene (also known as Upper Miocene) is a sub-epoch of the Miocene Epoch made up of two stages. The Tortonian and Messinian stages comprise the Late Miocene sub-epoch, which lasted from 11.63 Ma (million years ago) to 5.333 Ma. The ...

Sivapithecus ''Sivapithecus'' () (syn: ''Ramapithecus)'' is a genus of extinct apes. Fossil remains of animals now assigned to this genus, dated from 12.2 million years old in the Miocene, have been found since the 19th century in the Siwalik Hills of the I ...

'' from India. In 1939, South African palaeontologist

Robert Broom Robert Broom FRS FRSE (30 November 1866 6 April 1951) was a British- South African doctor and palaeontologist. He qualified as a medical practitioner in 1895 and received his DSc in 1905 from the University of Glasgow. From 1903 to 1910, he ...

hypothesised that it was closely allied with ''

Australopithecus ''Australopithecus'' (, ; ) is a genus of early hominins that existed in Africa during the Late Pliocene and Early Pleistocene. The genus ''Homo'' (which includes modern humans) emerged within ''Australopithecus'', as sister to e.g. ''Australo ...

'' and the last common ancestor of humans and other apes. 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 ''Homo erectus'' (; meaning "upright man") is an extinct species of archaic human from the Pleistocene, with its earliest occurrence about 2 million years ago. Several human species, such as '' H. heidelbergensis'' and '' H. antecessor ...

'' (at the time "''Pithecanthropus''" for early Javan specimens), and envisioned a lineage from ''Gigantopithecus'', to the Javan ape ''

Meganthropus ''Meganthropus'' is an extinct genus of non-hominin hominid ape, known from the Pleistocene of Indonesia. It is known from a series of large jaw and skull fragments found at the Sangiran site near Surakarta in Central Java, Indonesia, alongside ...

'' (then considered a human ancestor), to "''Pithecanthropus''", to "'' Javanthropus''", and finally

Aboriginal Australian Aboriginal Australians are the various Indigenous peoples of the Australian mainland and many of its islands, such as Tasmania, Fraser Island, Hinchinbrook Island, the Tiwi Islands, and Groote Eylandt, but excluding the Torres Strait I ...

s. This was part of his multiregional hypothesis, that all modern races and ethnicities evolved independently from a local archaic human species, rather than sharing a more recent and fully modern common ancestor. In 1952, von Koenigswald agreed that ''Gigantopithecus'' was a

, but believed it was an offshoot rather than a human ancestor. Much debate followed whether ''Gigantopithecus'' was a hominin or not for the next three decades until the Out of Africa hypothesis overturned the

Out of Asia Out may refer to: Arts, entertainment, and media Films * ''Out'' (1957 film), a documentary short about the Hungarian Revolution of 1956 * ''Out'' (1982 film), an American film directed by Eli Hollander * ''Out'' (2002 film), a Japanese film ba ...

and multiregional hypotheses, firmly placing humanity's origins in Africa. Orangutan -Zoologischer Garten Berlin-8a

''Gigantopithecus'' is now classified in the

Ponginae, closely allied with ''Sivapithecus'' and ''Indopithecus''. This would make its closest living relatives the

s. However, there are few similar traits (

synapomorphies In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to ha ...

) 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 to the Chinese '' Lufengpithecus'', which went extinct 4 million years prior to ''Gigantopithecus''. In 2019,

peptide sequencing Protein sequencing is the practical process of determining the amino acid sequence of all or part of a protein or peptide. This may serve to identify the protein or characterize its post-translational modifications. Typically, partial sequencing o ...

of dentine and 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 genetics, the mutation rate is the frequency of new mutations in a single gene or organism over time. Mutation rates are not constant and are not limited to a single type of mutation; there are many different types of mutations. Mutation rates ...

in orangutans has remained constant, shared a common ancestor about 12–10 million years ago in the Middle to Late Miocene. Their last common ancestor would have been a part of the Miocene

radiation In physics, radiation is the emission or transmission of energy in the form of waves or particles through space or through a material medium. This includes: * ''electromagnetic radiation'', such as radio waves, microwaves, infrared, visi ...

of apes. The same study calculated a divergence time between the Ponginae and African great apes about 26–17.7 million years ago.

Cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to ...

according to Zhang and Harrison, 2017:

"''G. bilaspurensis''"

In 1969, an 8.6 million year old mandible from the

Sivalik Hills The Sivalik Hills, also known as the Shivalik Hills and Churia Hills, are a mountain range of the outer Himalayas that stretches over about from the Indus River eastwards close to the Brahmaputra River, spanning the northern parts of the India ...

in northern India was classified as "''G. bilaspurensis''" by palaeontologists

Elwyn L. Simons Elwyn LaVerne Simons (July 14, 1930 – March 6, 2016) was an American paleontologist, paleozoologist, and a wildlife conservationist for primates. He was known as the father of modern primate paleontology for his discovery of some of humanki ...

and , who believed it was an the ancestor of ''G. blacki''. This bore resemblance to a molar discovered in 1915 in the Pakistani Pothohar Plateau then classified as "'' Dryopithecus giganteus''". Von Koenigswald reclassified "''D. giganteus''" in 1950 into its own genus, ''

Indopithecus ''Indopithecus giganteus'' () is an extinct species of large ape that lived in the late Miocene of the Siwalik Hills in northern India. Although frequently assigned to the more well-known genus ''Gigantopithecus'', recent authors consider it to ...

'', but this was changed again in 1979 to "''G. giganteus''" by American anthropologists Frederick Szalay and Eric Delson until ''Indopithecus'' was resurrected in 2003 by Australian anthropologist David W. Cameron. "''G. bilaspurensis''" is now considered a

synonym A synonym is a word, morpheme, or phrase that means exactly or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are al ...

of ''Indopithecus giganteus'', leaving ''Gigantopithecus''

monotypic In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unispe ...

(with only one species), ''G. blacki''.

Description

Size

Total size estimates are highly speculative because only tooth and jaw elements are known, and molar size and total body weight do not always correlate, such as in the case of post-canine megadontia hominins (small-bodied primate exhibiting massive molars and thick enamel). In 1946, Weidenreich hypothesised that ''Gigantopithecus'' was twice the size of male gorillas. In 1957, Pei estimated a total height of about . In 1970, Simons and American palaeontologist Peter Ettel approximated a height of almost and a weight of up to , which is about 40% heavier than the average male gorilla. In 1979, American anthropologist Alfred E. Johnson Jr. used the dimensions of gorillas to estimate a

femur The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates ...

length of and

humerus The humerus (; ) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius and ulna, and consists of three sections. The humeral upper extremity consists of a r ...

length of for ''Gigantopithecus'', about 20–25% longer than those of gorillas. In 2017, Chinese palaeoanthropologist Yingqi Zhang and American anthropologist Terry Harrison suggested a body mass of , though conceded this was likely an overestimate and it is impossible to obtain a reliable body mass estimate without more complete remains. The average maximum length of the upper canines for presumed males and females are and , respectively, and Mandible III (presumed male) is 40% larger than Mandible I (presumed female). These imply

sexual dimorphism Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most an ...

, with males being larger than females. Such a high degree of dimorphism is only surpassed by gorillas among modern apes in canine size, and is surpassed by none for mandibular disparity.

Teeth and jaws

Like other apes, ''Gigantopithecus'' had a

dental formula Dentition pertains to the development of teeth and their arrangement in the mouth. In particular, it is the characteristic arrangement, kind, and number of teeth in a given species at a given age. That is, the number, type, and morpho-physiolog ...

of , with two

incisor Incisors (from Latin ''incidere'', "to cut") are the front teeth present in most mammals. They are located in the premaxilla above and on the mandible below. Humans have a total of eight (two on each side, top and bottom). Opossums have 18, wher ...

s, one canine, two premolars, and three molars in each half of the jaw for both jaws. The canines, due to a lack of honing facets (which keep them sharp) and their overall stoutness, have been suggested to have functioned like premolars and molars (cheek teeth). Like other apes with enlarged molars, the incisors of ''Gigantopithecus'' are reduced. Wearing on the tongue-side of the incisors (the lingual face), which can extend as far down as the tooth root, suggests an underbite. Overall mandibular anatomy and tooth wearing suggests a side-to-side movement of the jaw while chewing (lateral excursion). The incisors and canines have extremely long tooth roots, at least double the length of the tooth crown (the visible part of the tooth). These teeth were closely packed together. In the upper jaw, the first premolar (P³) averages in surface area, the second premolar (P⁴) , the first and/or second molars (M^1/2, which are difficult to distinguish) , and the third molar (M³) . In the lower jaw, P₃ averages , P₄ , M_1/2 , and M₃ . The molars are the biggest of any known ape. Teeth continually evolved to become larger and larger. The premolars are high-crowned, and the lower have two tooth roots, whereas the upper have three. The lower molars are low-crowned, long and narrow, and waist at the midline—which is more pronounced in the lower molars—with low-lying and bulbous cusps and rounded-off crests. Giganthopithecus blacki, molar

The tooth enamel on the molars is in absolute measure the thickest of any known ape, averaging in three different molars, and over on the tongue-side (lingual) cusps of an upper molar. This has attracted comparisons with the extinct ''

Paranthropus ''Paranthropus'' is a genus of extinct hominin which contains two widely accepted species: '' P. robustus'' and '' P. boisei''. However, the validity of ''Paranthropus'' is contested, and it is sometimes considered to be synonymous with ''Austr ...

'' hominins, which had extremely large molars and thick enamel for their size. 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 had a large and flat (tabular) grinding surface, with an even enamel coating, and short dentine horns (the areas of the dentine layer which project upwards into the top enamel layer). The molars are the most

hypsodont Hypsodont is a pattern of dentition with high-crowned teeth and enamel extending past the gum line, providing extra material for wear and tear. Some examples of animals with hypsodont dentition are cows and horses; all animals that feed on gritt ...

(where the enamel extends beyond the gums) of any ape.

Palaeobiology

Diet

''Gigantopithecus'' is considered to have been a

. Carbon-13 isotope analysis suggests consumption of C₃ plants, such as fruits, leaves, and other forest plants. 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 In zoology, a folivore is a herbivore that specializes in eating leaves. Mature leaves contain a high proportion of hard-to-digest cellulose, less energy than other types of foods, and often toxic compounds.Jones, S., Martin, R., & Pilbeam, D. (1 ...

s) or seeds (

granivore Seed predation, often referred to as granivory, is a type of plant-animal interaction in which granivores (seed predators) feed on the seeds of plants as a main or exclusive food source,Hulme, P.E. and Benkman, C.W. (2002) "Granivory", pp. 132 ...

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 The chimpanzee (''Pan troglodytes''), also known as simply the chimp, is a species of great ape native to the forest and savannah of tropical Africa. It has four confirmed subspecies and a fifth proposed subspecies. When its close relative t ...

s, which could indicate a similarly generalist diet. The molar-like premolars, large molars, and long rooted cheeked teeth could point to chewing, crushing, and grinding of bulky and fibrous materials. Thick enamel would suggest a diet of abrasive items, such as dirt particles on food gathered near or on the ground (like bamboo shoots). 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

s. ''Gigantopithecus'' does not appear to have consumed the commonplace savanna grasses ( C4 plants). Nonetheless, in 1990, a few opal

phytolith Phytoliths (from Greek, "plant stone") are rigid, microscopic structures made of silica, found in some plant tissues and persisting after the decay of the plant. These plants take up silica from the soil, whereupon it is deposited within different ...

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

fig The fig is the edible fruit of ''Ficus carica'', a species of small tree in the flowering plant family Moraceae. Native to the Mediterranean and western Asia, it has been cultivated since ancient times and is now widely grown throughout the world ...

mulberry ''Morus'', a genus of flowering plants in the family Moraceae, consists of diverse species of deciduous trees commonly known as mulberries, growing wild and under cultivation in many temperate world regions. Generally, the genus has 64 ident ...

breadfruit Breadfruit (''Artocarpus altilis'') is a species of flowering tree in the mulberry and jackfruit family ( Moraceae) believed to be a domesticated descendant of '' Artocarpus camansi'' originating in New Guinea, the Maluku Islands, and the Phil ...

and

banyan A banyan, also spelled "banian", is a fig that develops accessory trunks from adventitious prop roots, allowing the tree to spread outwards indefinitely. This distinguishes banyans from other trees with a strangler habit that begin life as a ...

. This suggests that fruit was a significant dietary component for at least this population of ''Gigantopithecus''. The 400,000–320,000 year old

teeth from Hejiang Cave in southeastern China (near the time of extinction) show some differences from

Early Pleistocene The Early Pleistocene is an unofficial sub-epoch in the international geologic timescale in chronostratigraphy, being the earliest division of the Pleistocene Epoch within the ongoing Quaternary Period. It is currently estimated to span the time ...

material from other sites, which could potentially indicate that the Hejiang ''Gigantopithecus'' were a specialised form adapting to a changing environment with different food resources. The Hejiang teeth display a less level (more crenulated) outer enamel surface due to the presence of secondary crests emanating from the paracone and protocone on the side of the molar closer to the midline (medially), as well as sharper major crests. That is, the teeth are not as flat. 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. This hypothesis is no longer considered viable because its dental anatomy is consistent with herbivory. In 1975, American palaeoanthropologist

Tim D. White Tim D. White (born August 24, 1950) is an American paleoanthropologist and Professor of Integrative Biology at the University of California, Berkeley. He is best known for leading the team which discovered Ardi, the type specimen of ''Ardipithecu ...

drew similarities between the jaws and dentition of ''Gigantopithecus'' and those of the

giant panda The giant panda (''Ailuropoda melanoleuca''), also known as the panda bear (or simply the panda), is a bear species endemic to China. It is characterised by its bold black-and-white coat and rotund body. The name "giant panda" is sometimes u ...

, and suggested they both occupied the same

niche Niche may refer to: Science *Developmental niche, a concept for understanding the cultural context of child development * Ecological niche, a term describing the relational position of an organism's species *Niche differentiation, in ecology, the ...

bamboo Bamboos are a diverse group of evergreen perennial flowering plants making up the subfamily Bambusoideae of the grass family Poaceae. Giant bamboos are the largest members of the grass family. The origin of the word "bamboo" is uncertain, ...

specialists. This garnered support from some subsequent researchers, but thicker enamel and hypsodonty in ''Gigantopithecus'' could suggest different functionality for these teeth.

Growth

A ''Gigantopithecus'' permanent third molar, based on an approximate 600–800 days required for the enamel on the

cusp A cusp is the most pointed end of a curve. It often refers to cusp (anatomy), a pointed structure on a tooth. Cusp or CUSP may also refer to: Mathematics * Cusp (singularity), a singular point of a curve * Cusp catastrophe, a branch of bifurc ...

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 four μm per day; this is seen in only baby teeth for modern apes. 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'' which facilitated biomineralisation of enamel during prolonged amelogenesis (enamel growth).

Pathology

''Gigantopithecus'' molars have a high cavity rate of 11%, which could mean fruit was commonly included in its diet. The molars from ''Gigantopithecus'' Cave frequently exhibit

pitting enamel hypoplasia Enamel hypoplasia can take a variety of forms, but all types are associated with a reduction of enamel formation due to disruption in ameloblast production. One of the most common types, Pitting Enamel Hypoplasia (PEH), ranges from small circular pi ...

, where the enamel improperly forms with pits and grooves. This can be caused by malnutrition during growth years, which could point to periodic food shortages, though it can also be induced by other factors. Specimen PA1601-1 from Yanliang Cave shows evidence of tooth loss of the right second molar before the eruption of the neighboring third molar (which grew slantedly), which suggests this individual was able to survive for a long time despite impaired chewing abilities.

Society

The high levels of sexual dimorphism could indicate relatively intense male–male competition, though considering the upper canines only projected slightly farther than the cheek teeth, canine display was probably not very important in agonistic behaviour, unlike modern non-human apes.

Palaeoecology

''Gigantopithecus'' remains are generally found in what were subtropical evergreen broadleaf forest in South China, except in

Hainan Hainan (, ; ) is the smallest and southernmost province of the People's Republic of China (PRC), consisting of various islands in the South China Sea. , the largest and most populous island in China,The island of Taiwan, which is slightly l ...

which featured a

tropical rainforest Tropical rainforests are rainforests that occur in areas of tropical rainforest climate in which there is no dry season – all months have an average precipitation of at least 60 mm – and may also be referred to as ''lowland equator ...

. Carbon and oxygen isotope analysis of Early Pleistocene enamel suggests ''Gigantopithecus'' inhabited dense, humid, closed-canopy forest. Queque Cave featured a mixed deciduous and evergreen forest dominated by

birch A birch is a thin-leaved deciduous hardwood tree of the genus ''Betula'' (), in the family Betulaceae, which also includes alders, hazels, and hornbeams. It is closely related to the beech- oak family Fagaceae. The genus ''Betula'' cont ...

, oak, and chinkapin, as well as several low-lying

herb In general use, herbs are a widely distributed and widespread group of plants, excluding vegetables and other plants consumed for macronutrients, with savory or aromatic properties that are used for flavoring and garnishing food, for medicina ...

s and

fern A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes exce ...

s. The "''Gigantopithecus''

fauna Fauna is all of the animal life present in a particular region or time. The corresponding term for plants is '' flora'', and for fungi, it is '' funga''. Flora, fauna, funga and other forms of life are collectively referred to as '' biota''. ...

", one of the most important mammalian faunal groups of the Early Pleistocene of southern China, includes tropical or subtropical forest species. This group has been subdivided into three stages spanning 2.6–1.8 million years ago, 1.8–1.2 million years ago, and 1.2–0.8 million years ago. The early stage is characterised by more ancient

Neogene The Neogene ( ), informally Upper Tertiary or Late Tertiary, is a geologic period and system that spans 20.45 million years from the end of the Paleogene Period million years ago ( Mya) to the beginning of the present Quaternary Period Mya. ...

animals such as the gomphotheriid proboscidean (relative of elephants) ''

Sinomastodon ''Sinomastodon'' ("Chinese mastodont") is an extinct gomphothere genus (of order Proboscidea), from the Late Miocene to the Early Pleistocene deposits of Asia ( China, Japan, Thailand, Myanmar, Kashmir, and Indonesia). It is not to be conf ...

'', the

chalicothere Chalicotheres (from Greek '' chalix'', "gravel" and '' therion'', "beast") are an extinct clade of herbivorous, odd-toed ungulate (perissodactyl) mammals that lived in North America, Eurasia, and Africa from the Middle Eocene until the Early Ple ...

'' Hesperotherium'', the pig ''

Hippopotamodon ''Hippopotamodon'' is a genus of extinct suid even-toed ungulates that existed during the Miocene and Pliocene in Europe and Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a contine ...

'', the mouse-deer '' Dorcabune'', and the deer '' Cervavitus''. The middle stage is indicated by the appearance of the panda '' Ailuropoda wulingshanensis'', the dhole '' Cuon antiquus'', and the

tapir Tapirs ( ) are large, herbivorous mammals belonging to the family Tapiridae. They are similar in shape to a pig, with a short, prehensile nose trunk. Tapirs inhabit jungle and forest regions of South and Central America, with one species inh ...

'' Tapirus sinensis''. The late stage features more typical Middle Pleistocene animals such as the panda '' Ailuropoda baconi'' and the stegodontid proboscidean '' Stegodon''. Other classic animals typically include orangutans,

macaque The macaques () constitute a genus (''Macaca'') of gregarious Old World monkeys of the subfamily Cercopithecinae. The 23 species of macaques inhabit ranges throughout Asia, North Africa, and (in one instance) Gibraltar. Macaques are principall ...

s, rhinos, the extinct pigs '' Sus xiaozhu'' and '' S. peii'',

muntjac Muntjacs ( ), also known as the barking deer or rib-faced deer, (URL is Google Books) are small deer of the genus ''Muntiacus'' native to South Asia and Southeast Asia. Muntjacs are thought to have begun appearing 15–35 million years a ...

, ''

Cervus ''Cervus'' is a genus of deer that primarily are native to Eurasia, although one species occurs in northern Africa and another in North America. In addition to the species presently placed in this genus, it has included a whole range of othe ...

'' (a deer),

gaur The gaur (''Bos gaurus''; ), also known as the Indian bison, is a bovine native to South Asia and Southeast Asia, and has been listed as Vulnerable on the IUCN Red List since 1986. The global population was estimated at a maximum of 21,000 m ...

(a cow), the

goat-antelope The subfamily Caprinae, also sometimes referred to as the tribe Caprini, is part of the ruminant family Bovidae, and consists of mostly medium-sized bovids. A member of this subfamily is called a caprine, or, more informally, a goat-antelope ...

Megalovis ''Megalovis'' is an extinct genus of bovid that lived in Eurasia during the Plio-Pleistocene. Taxonomy ''Megalovis'' is typically considered a member of the subtribe Ovibovina of the Caprini tribe within the Antilopinae subfamily, which includes ...

'', and more rarely the large

saber-toothed cat Machairodontinae is an extinct subfamily of carnivoran mammals of the family Felidae (true cats). They were found in Asia, Africa, North America, South America, and Europe from the Miocene to the Pleistocene, living from about 16 million ...

'' Megantereon''. In 2009, American palaeoanthropologist Russel Ciochon hypothesised an undescribed, chimp-sized ape he identified from a few teeth coexisted with ''Gigantopithecus'', which in 2019 was identified as the closely related ''

''. Longgudong Cave may have represented a transitional zone between the

Palaearctic The Palearctic or Palaearctic is the largest of the eight biogeographic realms of the Earth. It stretches across all of Eurasia north of the foothills of the Himalayas, and North Africa. The realm consists of several bioregions: the Euro-Sibe ...

and

Oriental The Orient is a term for the East in relation to Europe, traditionally comprising anything belonging to the Eastern world. It is the antonym of '' Occident'', the Western World. In English, it is largely a metonym for, and coterminous with, the ...

realms, featuring, alongside the typical ''Gigantopithecus'' fauna, more

boreal Boreal may refer to: Climatology and geography *Boreal (age), the first climatic phase of the Blytt-Sernander sequence of northern Europe, during the Holocene epoch *Boreal climate, a climate characterized by long winters and short, cool to mild ...

animals such as

hedgehog A hedgehog is a spiny mammal of the subfamily Erinaceinae, in the eulipotyphlan family Erinaceidae. There are seventeen species of hedgehog in five genera found throughout parts of Europe, Asia, and Africa, and in New Zealand by introductio ...

hyena Hyenas, or hyaenas (from Ancient Greek , ), are feliform carnivoran mammals of the family Hyaenidae . With only four extant species (each in its own genus), it is the fifth-smallest family in the Carnivora and one of the smallest in the cl ...

horse The horse (''Equus ferus caballus'') is a domesticated, one-toed, hoofed mammal. It belongs to the taxonomic family Equidae and is one of two extant subspecies of ''Equus ferus''. The horse has evolved over the past 45 to 55 million yea ...

s, the cow ''

Leptobos ''Leptobos'' is an extinct genus of large bovine, known from the Late Pliocene and Early Pleistocene of Eurasia, extending from the Iberian Peninsula to northern China. Species of ''Leptobos'' reached a weight up to 320 kg. ''Leptobos'' is con ...

'', and pikas. ''Gigantopithecus'' appears to have gone extinct about 300,000 years ago, possibly due to the southward retreat of forest and prime habitat throughout the Middle Pleistocene caused by increasing seasonality, intensifying winter and summer

monsoon A monsoon () is traditionally a seasonal reversing wind accompanied by corresponding changes in precipitation but is now used to describe seasonal changes in atmospheric circulation and precipitation associated with annual latitudinal oscil ...

strength, and regional climatic cooling and/or wettening. Savanna would remain the dominant habitat of Southeast Asia until the

Late Pleistocene The Late Pleistocene is an unofficial age in the international geologic timescale in chronostratigraphy, also known as Upper Pleistocene from a stratigraphic perspective. It is intended to be the fourth division of the Pleistocene Epoch withi ...

. It has been speculated that immigrating ''H. erectus'' also contributed to its extinction. 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. In 2009, Ciochon suggested ''Gigantopithecus'' was the dominant primate inhabiting the tropical forests south of the Qinling Mountains, whereas ''H. erectus'' remained in the savannas straddling it.

Cryptozoology

''Gigantopithecus'' has been used in

circles as the identity of the Tibetan

yeti The Yeti ()"Yeti"
''

, humanlike monsters in local folklore. This began in 1960 with zoologist Wladimir Tschernezky, briefly describing in the journal ''

Nature Nature, in the broadest sense, is the physical world or universe. "Nature" can refer to the phenomena of the physical world, and also to life in general. The study of nature is a large, if not the only, part of science. Although humans are ...

'' a 1951 photograph of alleged yeti tracks taken by Himalayan mountaineers Michael Ward and

Eric Shipton Eric Earle Shipton, CBE (1 August 1907 – 28 March 1977), was an English Himalayan mountaineer. Early years Shipton was born in Ceylon (now Sri Lanka) in 1907 where his father, a tea planter, died before he was three years old. When he was eig ...

. Tschernezky concluded that the yeti walked like a human and was similar to ''Gigantopithecus''. Subsequently, the yeti attracted short-lived scientific attention, with several more authors publishing in ''Nature'' and ''

Science Science is a systematic endeavor that builds and organizes knowledge in the form of testable explanations and predictions about the universe. Science may be as old as the human species, and some of the earliest archeological evidence ...

'', but this also incited a popular monster hunting following for both the yeti and the similar American bigfoot which has persisted into the present day. The only scientist who continued trying to prove such monsters exist was anthropologist Grover Krantz, who continued pushing for a connection between ''Gigantopithecus'' and bigfoot from 1970 to his death in 2002. Among the

binomial name In taxonomy, binomial nomenclature ("two-term naming system"), also called nomenclature ("two-name naming system") or binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, bo ...

s he came up with for bigfoot included "Gigantopithecus canadensis". Krantz did not meet support from either mainstream science or amateurs who said he readily accepted clearly false evidence.

References

{{Authority control Prehistoric apes Prehistoric primate genera Pleistocene primates Pleistocene mammals of Asia Fossils of China Extinct animals of China Fossils of India Extinct animals of India Fossils of Vietnam Extinct animals of Vietnam Fossils of Indonesia Extinct animals of Indonesia Tortonian first appearances Pleistocene genus extinctions Fossil taxa described in 1935 Taxa named by Gustav Heinrich Ralph von Koenigswald Mammals described in 1935