''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 ...

ape Apes (collectively Hominoidea ) are a clade of Old World simians native to sub-Saharan Africa and Southeast Asia (though they were more widespread in Africa, most of Asia, and as well as Europe in prehistory), which together with its sister g ...

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

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 t ...

, 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 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 ...

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

Ponginae Ponginae , also known as the Asian hominids, is a subfamily in the family Hominidae. Once a diverse lineage of Eurasian apes, the subfamily has only one extant genus, '' Pongo'' (orangutans), which contains three extant species; the Sumatran oran ...

. ''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 or fi ...

-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 ( premolars 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 A generalist is a person with a wide array of knowledge on a variety of subjects, useful or not. It may also refer to: Occupations * a physician who provides general health care, as opposed to a medical specialist; see also: ** General pract ...

herbivore 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 growin ...

s with dirt. Some teeth bear traces of

fig family The Moraceae — often called the mulberry family or fig family — are a family of flowering plants comprising about 38 genera and over 1100 species. Most are widespread in tropical and subtropical regions, less so in temperate climates; however ...

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, 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 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 city and special administrative region of China on the eastern Pearl River Delta i ...

where they were being sold as "

dragon bones Oracle bones () are pieces of ox scapula and turtle plastron, which were used for pyromancy – a form of divination – in ancient China, mainly during the late Shang dynasty. ''Scapulimancy'' is the correct term if ox scapulae were used for the ...

" 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) ...

or Guangxi. 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 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 vast majority of the world's countries—including all of the great powers—forming two opposing ...

. 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 PEI or Pei may refer to: Places *Matecaña International Airport, Pereira, Colombia, IATA code PEI * Pei County (沛县), Jiangsu, China *Pei Commandery (沛郡), a commandery in Chinese history *Prince Edward Island, a province of Canada *Pei, ...

was tasked by the Chinese 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 ''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 premolar, 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 list of rivers of Asia, river in Asia, the list of rivers by length, third-longest in the world, and the longest in the world to flow entirely within one country. It rises at Jari Hill in th ...

, 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 Phi ...

. 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 The Chinese orangutan (''Pongo weidenreichi'') is an extinct species of orangutan from the Early Pleistocene of South China South China () is a geographical and cultural region that covers the southernmost part of China. Its precise meaning ...

''. 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 mos ...

, 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 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 ''

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. ''Austral ...

'' and the

last common ancestor In biology and genetic genealogy, the most recent common ancestor (MRCA), also known as the last common ancestor (LCA) or concestor, of a set of organisms is the most recent individual from which all the organisms of the set are descended. The ...

of humans and other apes. In 1946, Jewish German anthropologist

Franz Weidenreich Franz Weidenreich (7 June 1873 – 11 July 1948) was a Jewish German anatomist and physical anthropologist who studied evolution. Life and career Weidenreich studied at the Kaiser-Wilhelm-Universität in Strasbourg where he earned a medical ...

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 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 Isl ...

s. This was part of his

multiregional hypothesis The multiregional hypothesis, multiregional evolution (MRE), or polycentric hypothesis is a scientific model that provides an alternative explanation to the more widely accepted "Out of Africa" model of monogenesis for the pattern of human evoluti ...

, 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 ''Out of Africa'' is a memoir by the Danish author Karen Blixen. The book, first published in 1937, recounts events of the seventeen years when Blixen made her home in Kenya, then called British East Africa. The book is a lyrical meditation on ...

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 subfamily

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

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 to the Chinese ''

Lufengpithecus ''Lufengpithecus'' () is an extinct genus of ape in the subfamily Ponginae. It is known from thousands of dental remains and a few skulls and probably weighed about . It contains three species: ''L. lufengensis'', ''L. hudienensis'' and ''L. k ...

'', which went extinct 4 million years prior to ''Gigantopithecus''. In 2019, peptide sequencing of

dentine Dentin () (American English) or dentine ( or ) (British English) ( la, substantia eburnea) is a calcified tissue of the body and, along with enamel, cementum, and pulp, is one of the four major components of teeth. It is usually covered by e ...

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 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 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 The Pothohar Plateau ( ur, ) is a plateau in north-eastern Pakistan, located between Indus River and the Jhelum River, forming the northern part of Punjab. Geography Potohar Plateau is bounded on the east by the Jhelum River, on the west by the ...

then classified as "''

Dryopithecus ''Dryopithecus'' is a genus of extinct great apes from the middle– late Miocene boundary of Europe 12.5 to 11.1 million years ago (mya). Since its discovery in 1856, the genus has been subject to taxonomic turmoil, with numerous new specie ...

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 ...

'', 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 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 Post-canine megadontia is a relative enlargement of the molars and premolars compared to the size of the incisors and canines. This phenomenon is seen in some early hominid ancestors such as ''Paranthropus aethiopicus.'' Archaeological evidence ...

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 wit ...

length of and humerus 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 Canine may refer to: Zoology and anatomy * a dog-like Canid animal in the subfamily Caninae ** ''Canis'', a genus including dogs, wolves, coyotes, and jackals ** Dog, the domestic dog * Canine tooth, in mammalian oral anatomy People with the surn ...

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 Dental anatomy is a field of anatomy dedicated to the study of human tooth structures. The development, appearance, and classification of teeth fall within its purview. (The function of teeth as they contact one another falls elsewhere, under de ...

, suggests an

underbite In orthodontics, a malocclusion is a misalignment or incorrect relation between the teeth of the upper and lower dental arches when they approach each other as the jaws close. The English-language term dates from 1864; Edward Angle (1855-1930) ...

. 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 In dentistry, crown refers to the anatomical area of teeth, usually covered by enamel. The crown is usually visible in the mouth after developing below the gingiva The gums or gingiva (plural: ''gingivae'') consist of the mucosal tiss ...

(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'' 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 herbivore. 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. 13 ...

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 chimpanzees, 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 shoot Bamboo shoots or bamboo sprouts are the edible shoots (new bamboo culms that come out of the ground) of many bamboo species including ''Bambusa vulgaris'' and ''Phyllostachys edulis''. They are used as vegetables in numerous Asian dishes and ...

s). Similarly, oxygen isotope analysis suggests ''Gigantopithecus'' consumed more low-lying plants such as stems, roots, and grasses than orangutans.

Dental calculus In dentistry, calculus or tartar is a form of hardened dental plaque. It is caused by precipitation of minerals from saliva and gingival crevicular fluid (GCF) in plaque on the teeth. This process of precipitation kills the bacterial cells wit ...

indicates the consumption of

s. ''Gigantopithecus'' does not appear to have consumed the commonplace savanna grasses (

C4 plants carbon fixation or the Hatch–Slack pathway is one of three known photosynthetic processes of carbon fixation in plants. It owes the names to the 1960's discovery by Marshall Davidson Hatch and Charles Roger Slack that some plants, when suppl ...

). 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

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 ...

s, mulberry, breadfruit 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 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 A cusp is a pointed, projecting, or elevated feature. In animals, it is usually used to refer to raised points on the crowns of teeth. The concept is also used with regard to the leaflets of the four heart valves. The mitral valve, which has two ...

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 drew similarities between the jaws and dentition of ''Gigantopithecus'' and those of the giant panda, 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 alpha-2-HS-glycoprotein (AHSG, Alpha-2-Heremans-Schmid Glycoprotein) also known as fetuin-A is a protein that in humans is encoded by the ''AHSG'' gene. Fetuin-A belongs to the fetuin class of plasma binding proteins and is more abundant in feta ...

(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 Biomineralization, also written biomineralisation, is the process by which living organisms produce minerals, often to harden or stiffen existing tissues. Such tissues are called mineralized tissues. It is an extremely widespread phenomenon ...

of enamel during prolonged

amelogenesis Amelogenesis is the formation of enamel on teeth and begins when the crown is forming during the advanced bell stage of tooth development after dentinogenesis forms a first layer of dentin. Dentin must be present for enamel to be formed. Amelob ...

(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, 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 botany, an evergreen is a plant which has foliage that remains green and functional through more than one growing season. This also pertains to plants that retain their foliage only in warm climates, and contrasts with deciduous plants, which ...

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 equa ...

. 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'' contains ...

oak An oak is a tree or shrub in the genus ''Quercus'' (; Latin "oak tree") of the beech family, Fagaceae. There are approximately 500 extant species of oaks. The common name "oak" also appears in the names of species in related genera, notably ''L ...

, and chinkapin, as well as several low-lying herbs 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 (ecology ...

", 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 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 confus ...

'', the chalicothere '' Hesperotherium'', the pig '' Hippopotamodon'', the

mouse-deer Chevrotains, or mouse-deer, are small even-toed ungulates that make up the family Tragulidae, the only extant members of the infraorder Tragulina. The 10 extant species are placed in three genera, but several species also are known only f ...

'' Dorcabune'', and the deer '' Cervavitus''. The middle stage is indicated by the appearance of the panda '' Ailuropoda wulingshanensis'', the

dhole The dhole (''Cuon alpinus''; ) is a canid native to Central, South, East and Southeast Asia. Other English names for the species include Asian wild dog, Asiatic wild dog, Indian wild dog, whistling dog, red dog, red wolf, and mountain wolf. It ...

'' 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 ''Ailuropoda baconi'' is an extinct panda known from cave deposits in south China, Laos, Vietnam, Myanmar and Thailand from the Late Pleistocene, 750 thousand years ago, and was preceded by ''A. wulingshanensis'' and '' A. microta'' as an ancesto ...

'' and the stegodontid proboscidean ''

Stegodon ''Stegodon'' ("roofed tooth" from the Ancient Greek words , , 'to cover', + , , 'tooth' because of the distinctive ridges on the animal's molars) is an extinct genus of proboscidean, related to elephants. It was originally assigned to the fami ...

''. Other classic animals typically include orangutans, macaques,

rhino A rhinoceros (; ; ), commonly abbreviated to rhino, is a member of any of the five extant species (or numerous extinct species) of odd-toed ungulates in the family Rhinocerotidae. (It can also refer to a member of any of the extinct species o ...

s, 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 ago, ...

, ''

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 (a ...

'' Megalovis'', 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 ''Megantereon'' was a genus of prehistoric machairodontine saber-toothed cat that lived in North America, Eurasia, and Africa. It may have been the ancestor of ''Smilodon''. Taxonomy Fossil fragments have been found in Africa, Eurasia, and No ...

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

s, hyenas,

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'', and

pika A pika ( or ; archaically spelled pica) is a small, mountain-dwelling mammal found in Asia and North America. With short limbs, very round body, an even coat of fur, and no external tail, they resemble their close relative, the rabbit, but wi ...

s. ''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 osci ...

strength, and regional climatic cooling and/or wettening. Savanna would remain the dominant habitat of Southeast Asia until the Late Pleistocene. 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 The Qinling () or Qin Mountains, formerly known as the Nanshan ("Southern Mountains"), are a major east–west mountain range in southern Shaanxi Province, China. The mountains mark the divide between the drainage basins of the Yangtze and Yellow ...

, 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"
''

bigfoot, 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. 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 Scientific method, builds and organizes knowledge in the form of Testability, testable explanations and predictions about the universe. Science may be as old as the human species, and some of the earli ...

'', 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 Grover Sanders Krantz (November 5, 1931 – February 14, 2002) was an American anthropologist and cryptozoologist; he was one of few scientists not only to research Bigfoot, but also to express his belief in the animal's existence. Throughout his ...

, who continued pushing for a connection between ''Gigantopithecus'' and bigfoot from 1970 to his death in 2002. Among the binomial names 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