Why did the largest ever primate disappear?

Chinese apothecary shops sell an assortment of fossils. They include shells of brachiopods that when ground up and dissolved in water allegedly treat rheumatism, skin diseases, and eye disorders. Traditional apothecaries also supply  ‘dragons’ teeth’, said by Dr Subhuti Dharmananda, Director of the Institute for Traditional Medicine in Portland, Oregon to treat epilepsy, madness, manic running about, binding qi (‘vital spirit’) below the heart, inability to catch one’s breath, and various kinds of spasms, as well as making the body light, enabling one to communicate with the spirit light, and lengthening one’s life. Presumably have done a roaring trade in ‘dragons’ teeth’ since they were first mentioned in a Chinese pharmacopoeia (the Shennong Bencao Jing) from the First Century of the Common Era. In 1935 the anthropologist Gustav von Koenigswald came across two ‘dragons’ teeth’ in a Hong Kong shop. They were unusually large molars and he realised they were from a primate, but far bigger (20  × 22 mm) than any from living or fossil monkeys, apes or humans.

Eventually, in 1952 (he had been interned by Japanese forces occupying Java), von Koenigswald formally described the teeth and others that he had found. Their affinities and size prompted him to call the former bearer the ‘Huge Ape’ (Gigantopithecus). By 1956 Chinese palaeontologists had tracked down the cave site in Guangxi province where the teeth had been sourced, and a local farmer soon unearthed a complete lower jawbone (mandible) that was indeed gigantic. More teeth and mandibles have since been found at several sites in Southern and Southeast Asia, with an age range from about 2.0 to 0.3 Ma. Anatomical differences between teeth and mandibles suggest that there may have been 4 different species. Using mandibles as a very rough guide to overall size it has been estimated that Gigantopithecus may have been up to 3 m tall weighing almost 600kg.

Above: Size comparison of G. blacki with a 1.8 m tall human male; NB G.blacki probably walked on all fours, as do living orangutans when they rarely descend from the forest canopy. (Credit: Frido Welker) Below: Mandible of Gigantopithecus blacki from India (Credit: Prof. Wei Wang, Photo retouched by Theis Jensen)

Plaque on some teeth contain evidence for fruit, tubers and roots, but not grasses, which suggest suggest that Gigantopithecus had a vegetarian diet based on forest plants. Mandibles also showed affinities with living and fossil orangutans (pongines). Analysis of proteins preserved in tooth enamel confirm this relationship (Welker, F. and 17 others 2019. Enamel proteome shows that Gigantopithecus was an early diverging pongine. Nature, v.576, p. 262–265; DOI: 10.1038/s41586-019-1728-8). It was one of the few members of the southeast Asian megafauna to go extinct at the genus level during the Pleistocene. Its close relative Pongo the orangutan survives as three species in Borneo and Sumatra. Detailed analysis of material from 22 southern Chinese caves that have yielded Gigantopithecus teeth has helped resolve that enigma (Zhang, Y. and 20 others 2024. The demise of the giant ape Gigantopithecus blacki. Nature, v. 625; DOI: 10.1038/s41586-023-06900-0).

At the time Gigantopithecus first appeared in the geological record of China (~2.2 Ma), it ranged over much of south-western China. The early Pleistocene ecosystem there was one of diverse forests sufficiently productive to support large numbers of this enormous primate and also the much smaller orangutan Pongo weidenreichi.  By 295 to 215 ka, the age of the last known Gigantopithecus fossils, its range had shrunk dramatically. The teeth show marked increases in size and complexity by this time, which suggests adaptation of diet to a changing ecosystem. That is confirmed by pollen analysis of cave sediments which reveal a dramatic decrease in forest cover and increases in fern and non-arboreal flora at the time of extinction. One physical sign of environmental stress suffered by individual late G. blacki is banding in their teeth defined by large fluctuations of barium and strontium concentrations relative to calcium. The bands suggest that each individual had to change its diet repeatedly over its lifetime. Closely related orangutans, on the other hand survived into the later Pleistocene of China, having adapted to the changed ecosystem, as did early humans in the area. It thus seems likely that Gigantopithecus was an extreme specialist as regards diet, and was unable to adapt to changes brought on by the climate becoming more seasonal. Today’s orangutans in Indonesia face a similar plight, but that is because they have become restricted to forest ‘islands’ in the midst of vast areas of oil palm plantations. Their original range seems to have been much the same as that of Gigantopithecus, i.e. across south-eastern Asia, but Pongo seems to have gone extinct outside of Indonesia (by 57 ka in China) during the last global cooling and when forest cover became drastically restricted.

Isotopic clues to diet of early hominins

‘We are what we eat’ is certainly a truism, but it is neither a trope nor a cliché. The phrase is especially appropriate when scientists examine isotopes of a variety of elements in bones or teeth. For instance the relative proportions of two stable isotopes of the metal strontium – 87Sr and 86Sr – differ from place to place in soil because 87Sr is the daughter isotope of radioactive 87Rb. The older the rock from which a soil has formed the more of the radioactive rubidium isotope will have decayed. Not only does this increase the 87Sr/ 86Sr ratio in the rock and the soil derived from it, but vegetation inherits it too. So it gets into an animal’s diet and ultimately its teeth. A human who has migrated will carry the ratio of the geology of her early home geology in her adult teeth – fully developed by about 13 years-old – to wherever she dies. Likewise, the different oxygen isotopes in rainwater, which result from climate variation, end up in teeth thanks to what a person ate before adulthood. The two ‘signatures’ together allowed archaeologists to backtrack the famous ‘Amesbury Archer’, who may have brought Bronze Age culture to Britain, back to the Alps of Central Europe. Just what a human diet comprised can be roughly assessed from the carbon and nitrogen isotopes in collagen that fossil bone sometimes preserves: the proportion of seafood relative to the meat of land herbivores and the amount of terrestrial grains, nuts and fruits. The trouble is, collagen degrades with the age of human remains and another approach is needed to assess the diets of our distant forebears.

Calcium isotope data from early hominins and some modern primates. Increasingly negative values of δ44/42Ca signify lower values of the ratio compared with a standard. (Credit: Martin et al. 2020; Fig. 1)

It turns out that calcium isotopes in teeth, which do not degrade over extremely long time spans, offer clues to diet. In particular the dental 44Ca/42Ca ratio decreases as its hosts rise in the food chain; effectively as the meat content in their diet increases. This approach has been applied to the hominin and non-human primate fauna of the Turkana Basin in Kenya (Martin, J.E. et al. 2020. Calcium isotopic ecology of Turkana Basin hominins. Nature Communications, v. 11, article 3587; DOI: 10.1038/s41467-020-17427-7). The shores of a large lake in the vicinity of modern Lake Turkana were occupied from 3.5 to about 2 Ma ago by early Homo, australopithecines, paranthropoids and baboons. Using dental Ca isotopes fails to distinguish Australopithecus anamensis and Kenyanthropus platyops, whereas carbon isotopes suggest that the first had a purely C3 plant diet – fruiting plants that thrive under cool, wet conditions, as beneath woodland canopies – whereas Kenyanthropus foraged on both these and the C4 plants – many grasses and sedges – that favour open, well-lit grassland. The 44Ca/42Ca ratios in Homo teeth span a wide range of values that point to omnivory and even a high dietary meat content: a similar isotopic pattern to those of fossil baboons and geladas. Paranthropus boisei is definitely the odd-one-out, among both ancient and modern primates, and even among paranthropoids as a whole. It most likely had a specialised diet. Its teeth show wear patterns that suggest soft plant material, which seems to rule out grasses which are abrasive. Perhaps it fed on succulent semi-aquatic plants of the lake shore. When Mary Leakey first discovered P. boisei in 1959, she and husband Louis considered that its huge molars with thick enamel indicated that it ate hard vegetable matter, hence its original nickname ‘Nutcracker Man’. It also had hands capable of precise manipulation, indeed the association of the first specimen with Oldowan-type stone tools led to speculation that it had made them. Some specimens are associated with long bones with worn ends, suggesting that they may have used them for digging.