| Gigantopithecus | |
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| ReconstructedGigantopithecusmandible at theUniversity of Iowa Museum of Natural History | |
Scientific classification | |
| Kingdom: | Animalia |
| Phylum: | Chordata |
| Class: | Mammalia |
| Order: | Primates |
| Suborder: | Haplorhini |
| Family: | Hominidae |
| Subfamily: | Ponginae |
| Genus: | †Gigantopithecus von Koenigswald, 1935[1] |
| Species: | †G. blacki |
| Binomial name | |
| †Gigantopithecus blacki von Koenigswald, 1935 | |
Gigantopithecus (/dʒaɪˌɡæntoʊpɪˈθikəs,ˈpɪθɪkəs,dʒɪ-/jy-gan-toh-pih-THEE-kəs, -PITH-ih-kəs, jih-[2])[a] is an extinctgenus ofape that lived in central to southern China from 2 million to approximately 200,000–300,000 years ago during theEarly toMiddle Pleistocene, represented by one species,Gigantopithecus blacki. Potential identifications have also been made inBangladesh,India, andPakistan, but they could be misidentified remains of the orangutanPongo weidenreichi. The first remains ofGigantopithecus, two third-molar teeth, were identified in a drugstore by anthropologistRalph von Koenigswald in 1935 in England, who subsequentlydescribed the ape. In 1956, the first mandible and more than 1,000 teeth were found inLiucheng, and numerous more remains have since been found in at least 16 sites. Only teeth and four mandibles are known currently. Other skeletal elements were likely consumed byporcupines before they could fossilize.[3][4]Gigantopithecus was once argued to be ahominin, a member of thehuman line, but it is now thought to be closely allied withorangutans, classified in the subfamilyPonginae.
Gigantopithecus has traditionally been restored as a massive,gorilla-like ape, potentially 200–300 kg (440–660 pounds) when alive, but the few remains make total size estimates highly speculative. The species may have beensexually dimorphic, with males much bigger than females. The incisors are reduced and the canines appear to have functioned likecheek 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 thickestenamel by absolute measure of any ape, up to 6 mm (1⁄4 inch) in some areas, though this is only fairly thick when tooth size is taken into account.
Gigantopithecus appears to have been ageneralist herbivore ofC3 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, andtubers with dirt. Some teeth bear traces offig 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 potentiallyarchaic human activity.Gigantopithecus has become popular incryptozoology circles as the identity of the Tibetanyeti or the Americanbigfoot, apelike creatures in local folklore.
Gigantopithecus blacki was named by anthropologistRalph von Koenigswald in 1935 based on two third lowermolar teeth, which, he noted, were of enormous size (the first was "Ein gewaltig grosser (...) Molar", the second was described as "der enorme Grösse besitzt"), measuring20 mm × 22 mm (3⁄4 by7⁄8 inch).[1] The specific nameblacki is in honour of Canadian palaeoanthropologistDavidson Black, who had studied human evolution in China and had died the previous year. Von Koenigswald, working for theDutch East Indies Mineralogical Survey on Java, had found the teeth in a drugstore inHong Kong where they were being sold as "dragon bones" to be used intraditional Chinese medicine. By 1939, after purchasing more teeth, he determined they had originated somewhere inGuangdong orGuangxi. He could not formally describe thetype specimen until 1952 due to hisinternment by Japanese forces duringWorld War II. The originally discovered teeth are part of the collection of theUniversity of Utrecht.[3][5] While onJava, with the onset of World War II, von Koenigswald put theGigantopithecus teeth in a milk bottle and buried them in a friend's backyard before being interned by Japanese forces. After the war, he recovered the fossils and moved toNew York City and could not continue research on the subject.[4]
In 1955, a survey team that was led by Chinese palaeontologistPei Wenzhong was tasked by the ChineseInstitute of Vertebrate Palaeontology and Palaeoanthropology (IVPP) with finding the originalGigantopithecus locality. They collected 47 teeth among shipments of "dragon bones" in Guangdong and Guangxi. In 1956, the team discovered the firstin situ remains, a third molar andpremolar, in a cave (subsequently named "Gigantopithecus Cave") inNiusui Mountain, Guangxi. Also in 1956,Liucheng farmer Tan Xiuhuai discovered more teeth and the firstmandible on his field. From 1957 to 1963, the IVPP survey team carried out excavations in this area including most especially Yanyan Cave and recovered two more mandibles and more than 1,000 teeth.[3][5][6][7] As of 2024[update], Yanyan Cave is the most productiveGigantopithecus site.[4] In 2014, a fourth confirmed mandible was discovered inYanliang, Central China.[8][4] Indicated by extensiverodent gnawing marks, teeth primarily accumulated in caves likely due toporcupine 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.[3][4]
ConfirmedGigantopithecus remains have since been found in 16 different sites across southern China. The northernmost sites areLonggupo [de] andLonggu Cave, just south of theYangtze River, and southernmost on Hainan Island in theSouth China Sea. An isolated canine fromThẩm Khuyên Cave, Vietnam, and a fourth premolar fromPha Bong, Thailand, could possibly be assigned toGigantopithecus, though these could also represent the extinct orangutanPongo weidenreichi.[3] In 2016, twoGigantopithecus mandibular fragments each preserving the last two molars were reported fromSemono [nl] inCentral Java, Indonesia.[9] They were collected by a local named Dakri in 2014 who found them at the surface within 4 km2 (1.5 sq mi) of the Semono site; they may have been imported from China and left there since Chinese "dragon bones" were commonly sold in Javanese drugstores.[4] The oldest remains date to 2.2 million years ago fromBaikong Cave, and the youngest to 295 to 215 thousand years ago fromShuangtan andGongjishan Caves.[10]
In 1935, von Koenigswald consideredGigantopithecus to be closely allied with theLate MioceneSivapithecus from India.[1] In 1939, South African palaeontologistRobert Broom hypothesised that it was closely allied withAustralopithecus and thelast common ancestor of humans and other apes.[11] In 1946, Jewish German anthropologistFranz Weidenreich describedGigantopithecus 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 andHomo erectus (at the time "Pithecanthropus" forearly Javan specimens), and envisioned a lineage fromGigantopithecus, to the Javan apeMeganthropus (then considered a human ancestor), to "Pithecanthropus", to "Javanthropus", and finallyAboriginal Australians. This was part of hispolycentric 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.[12] In 1952, von Koenigswald agreed thatGigantopithecus was ahominin, but believed it was an offshoot rather than a human ancestor.[13] Much debate followed whetherGigantopithecus was a hominin or not for the next three decades until theOut of Africa hypothesis overturned the Out of Asia-theory and multiregional hypotheses, firmly placing humanity's origins in Africa.[3][5]
Gigantopithecus is now classified in the subfamilyPonginae, closely allied withSivapithecus andIndopithecus. This would make its closest living relatives theorangutans. However, there are few similar traits (synapomorphies) linkingGigantopithecus and orangutans due to fragmentary remains, with the main morphological argument being its close affinities toSivapithecus, which is better established as a pongine based on skull features. In 2017, Chinese palaeoanthropologist Yingqi Zhang and American anthropologist Terry Harrison suggested thatGigantopithecus is most closely allied with the ChineseLufengpithecus which went extinct 4 million years prior toGigantopithecus.[3]
In 2019,peptide sequencing ofdentine andenamel proteins of aGigantopithecus molar fromChuifeng Cave indicates thatGigantopithecus was indeed closely allied with orangutans, and, assuming the currentmutation rate in orangutans has remained constant, shared a common ancestor about 12–10 million years ago in theMiddle to Late Miocene. Their last common ancestor would have been a part of the Mioceneradiation of apes. The same study calculated a divergence time between the Ponginae and Africangreat apes about 26–17.7 million years ago.[14]
Cladogram according to Zhang and Harrison, 2017:[3]
| Hominoidea (apes) | |
In 1969, an 8.6 million year old mandible from theSivalik Hills in northern India was classified as "G. bilaspurensis" by palaeontologistsElwyn L. Simons andShiv Raj Kumar Chopra [de], who believed it was the ancestor ofG. blacki.[3][5] This bore resemblance to a molar discovered in 1915 in the PakistaniPothohar Plateau then classified as "Dryopithecus giganteus". Von Koenigswald reclassified "D. giganteus" in 1950 into its own genus,Indopithecus, but this was changed again in 1979 to "G. giganteus" by American anthropologists Frederick Szalay and Eric Delson[15] untilIndopithecus was resurrected in 2003 by Australian anthropologistDavid W. Cameron [wd].[3][16] "G. bilaspurensis" is now considered asynonym ofIndopithecus giganteus, leavingGigantopithecusmonotypic (with only one species),G. blacki.[3][17]
Total size estimates are highly speculative because only tooth and jaw elements are known. Molar size and total body weight do not always correlate, such as in the case ofpostcanine megadontia hominins (small-bodied primates exhibiting massive molars and thick enamel).[18] Still,Gigantopithecus is normally reconstructed as the biggest primate ever recorded.[10]
The average maximum length of the uppercanines for presumed males and females are21.1 mm (3⁄4 inch) and15.4 mm (1⁄2 inch), respectively, and Mandible III (presumed male) is 40% larger than Mandible I (presumed female). These implysexual dimorphism, 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.[3]
Like other apes,Gigantopithecus had adental formula of2.1.2.32.1.2.3, with twoincisors, one canine, two premolars, and three molars in each half of the jaw for both jaws.[3] 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 ofGigantopithecus are reduced.[22][23] Wearing on the tongue-side of the incisors (the lingual face), which can extend as far down as thetooth root, suggests anunderbite.[3] Overall mandibular anatomy and tooth wearing suggests a side-to-side movement of the jaw while chewing (lateral excursion).[24] The incisors and canines have extremely long tooth roots, at least double the length of thetooth crown (the visible part of the tooth). These teeth were closely packed together.[3]
In the upper jaw, the average size of the 1st premolar (P3), 2nd premolar (P4), 1st and 2nd molar (which are difficult to distinguish, M1/2), and 3rd molar (M3) are:[3]
In the lower jaw:[3]
The molars are the biggest of any known ape.[3] Teeth continually evolved to become larger and larger.[25] 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.[3]
The tooth enamel on the molars is in absolute measure the thickest of any known ape, averaging2.5–2.9 mm (3⁄32–1⁄8 inch) in three different molars, and over6 mm (1⁄4 inch) on the tongue-side (lingual) cusps of an upper molar.[24] This has attracted comparisons with the extinctParanthropus hominins, which had extremely large molars and thick enamel for their size.[22][24] However, in relation to the tooth's size, enamel thickness forGigantopithecus overlaps with that of several other living and extinct apes. Like orangutans and potentially all pongines (though unlike African apes) theGigantopithecus molar has 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).[18] The molars are the mosthypsodont (where the enamel extends beyond the gums) of any ape.[3]
Gigantopithecus is considered to have been aherbivore.Carbon isotope analysis suggests consumption ofC3 plants, such as fruits, leaves, and other forest plants.[26] The robust mandible ofGigantopithecus 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 (folivores) or seeds (granivores).Gigantopithecus teeth have a markedly lower rate of pitting (caused by eating small, hard objects) than orangutans, more similar to the rate seen inchimpanzees, which could indicate a similarlygeneralist diet.[3]
The molar-like premolars, large molars, and long rooted cheeked teeth could point to chewing, crushing, and grinding of bulky and fibrous materials.[27][28] Thick enamel would suggest a diet of abrasive items, such as dirt particles on food gathered near or on the ground (likebamboo shoots).[24] Similarly, oxygen isotope analysis suggestsGigantopithecus consumed more low-lying plants such as stems, roots, and grasses than orangutans.Dental calculus indicates the consumption oftubers.[29]Gigantopithecus does not appear to have consumed the commonplace savanna grasses (C4 plants).[26] Nonetheless, in 1990, a few opalphytoliths adhering to four teeth fromGigantopithecus Cave were identified to have originated from grasses; though, the majority of phytoliths resemble the hairs offig family fruits, which includefigs,mulberry,breadfruit, andbanyan. This suggests that fruit was a significant dietary component for at least this population ofGigantopithecus.[28] Additionally, the extremely high δ44/42Ca values ofG. blacki are indicative of the species likely making frequent use of mineral licks derived from rocks and soils.[30]
The 320,000–400,000-year-old Middle PleistoceneGigantopithecus teeth fromHejiang 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 HejiangGigantopithecus were locally 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 theparacone andprotocone 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.[8][31][32]
In 1957, based on hoofed animal remains in a cave located in a seemingly inaccessible mountain, Pei had believed thatGigantopithecus was a cave-dwelling predator and carried these animals in.[33] This hypothesis is no longer considered viable because its dental anatomy is consistent with herbivory.[26] In 1975, American palaeoanthropologistTim D. White drew similarities between the jaws and dentition ofGigantopithecus and those of thegiant panda, and suggested they both occupied the sameniche asbamboo specialists.[34] This garnered support from some subsequent researchers, but thicker enamel andhypsodonty inGigantopithecus could suggest different functionality for these teeth.[24]
AGigantopithecus permanent third molar, based on an approximate 600–800 days required for the enamel on thecusps to form (which is quite long), was estimated to have taken four years to form, which is within the range (albeit, far upper range) of what is exhibited in humans and chimpanzees. Like many other fossil apes, the rate of enamel formation near the enamel-dentine junction (dentine is the nerve-filled layer beneath the enamel) was estimated to begin at about 4 μm per day; this is seen in only baby teeth for modern apes.[22]
Protein sequencing ofGigantopithecus enamel identifiedalpha-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 inGigantopithecus teeth which facilitatedbiomineralisation of enamel during prolongedamelogenesis (enamel growth).[14]
Gigantopithecus molars have a highcavity rate of 11%, which could mean fruit was commonly included in its diet.[3][28] The molars fromGigantopithecus Cave frequently exhibitpitting 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.[28]
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.[8]
Gigantopithecus as well as orangutan fossil teeth from Queque, Shuangtan, and Chuifeng cave systems have high levels oflead, some upwards of 50parts per million. Concentrations this high may have causedlead poisoning, which can affect general health, development, and social behaviour.[35]
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.[3]
Gigantopithecus remains are generally found in what were subtropicalevergreen broadleaf forest in South China, except inHainan which featured atropical rainforest. Carbon and oxygen isotope analysis of Early Pleistocene enamel suggestsGigantopithecus inhabited dense, humid, closed-canopy forest.Queque Cave featured a mixed deciduous and evergreen forest dominated bybirch,oak, andchinkapin, as well as several low-lyingherbs andferns.[3]
The "Gigantopithecusfauna", 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 ancientNeogene animals such as thegomphotheriid proboscidean (relative of elephants)Sinomastodon, thechalicothereHesperotherium, the suidHippopotamodon, thetragulidDorcabune [it], and the deerCervavitus. The middle stage is indicated by the appearance of the pandaAiluropoda wulingshanensis, thedholeCuon antiquus, and thetapirTapirus sinensis. The late stage features more typical Middle Pleistocene animals such as the pandaAiluropoda baconi and thestegodontid proboscideanStegodon.[36] Other classic animals typically include orangutans,macaques,rhinos, the extinct pigsSus xiaozhu andSus peii,muntjac,Cervus (a deer),gaur (a cow), thebovidMegalovis, and more rarely the largesaber-toothed catMegantereon.[37] In 2009, American palaeoanthropologistRussell Ciochon hypothesised an undescribed, chimp-sized ape he identified from a few teeth coexisted withGigantopithecus,[38] which in 2019 was identified as the closely relatedMeganthropus.[39] Longgudong Cave may have represented a transitional zone between thePalaearctic andOriental realms, featuring, alongside the typicalGigantopithecus fauna, moreboreal animals such ashedgehogs,hyenas,horses, the bovidLeptobos, andpikas.[36]
Gigantopithecus fossil sites range across Guangxi, Guizhou, Hainan and Hubei Provinces, but those post-dating about 400,000 years ago are only known from Guangxi. Its youngest definitive remains in China are roughly 295,000 to 215,000 years old.[10] Two possible teeth (PIN 5792/439 and PIN 5792/490) have been reported from theLate Pleistocene deposit in Vietnam,[40] but these could actually belong toP. weidenreichi.[4] The extinction ofGigantopithecus correlates with a cooling trend marked by intensifying seasonality andmonsoon strength in the region, which led to the encroachment of open grasslands on rainforest habitats.[41][25] BecauseGigantopithecus teeth dating to this time show evidence of dietary shifts and chronic nutritional stress,Gigantopothecus may have been less successful at adapting to these environmental stressors compared to contemporary great apes — namelyP. weidenreichi andHomo — which could have led to its extinction.[10] Similarly,Gigantopithecus seems to only have been consuming C3 forest plants, instead of the C4 savannah plants which were becoming more common during this time.[42] Savannas remained the dominant habitat of Southeast Asia until the Late Pleistocene.[41][43]
Human activity in southern China is known as early as 800,000 years ago but does not become prevalent until after the extinction ofGigantopithecus, so it is unclear if pressures such as competition over resources or overhunting were factors.[38] In 2024, Zhang and colleagues found no evidence of archaic hominin involvement in the extinctions of any southern Chinese animal.[10]
Gigantopithecus has been used incryptozoology circles as the identity of the Tibetanyeti or Americanbigfoot, apelike monsters in local folklore. This began in 1960 with zoologist Wladimir Tschernezky, briefly describing in the journalNature a 1951 photograph of alleged yeti tracks taken by Himalayan mountaineersMichael Ward andEric Shipton. Tschernezky concluded that the yeti walked like a human and was similar toGigantopithecus. Subsequently, the yeti attracted short-lived scientific attention, with several more authors publishing inNature andScience, 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 anthropologistGrover Krantz, who continued pushing for a connection betweenGigantopithecus and bigfoot from 1970 to his death in 2002. Among thebinomial names he came up with for bigfoot included "Gigantopithecus canadensis". Scientists and amateur monster hunters both dismissed Krantz's arguments, saying he readily accepted clearly false evidence.[44]
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