Middle Palaeolithic Neanderthals and Denisovans of East Asia

During the Middle Palaeolithic (250 to 30 ka) anatomically modern humans (AMH) and Neanderthals were engaged in new technological developments in Europe and Africa as well as in migration and social interaction. This is reflected in the tools that they left at occupation sites and the fact that most living non-Africans carry Neanderthal DNA. One of the major cultural developments was a novel means of manufacturing stone implements. It developed from the Levallois technique that involved knapping sharp-edged flakes of hard rock from larger blocks or cores. A type of tool first found at a Neanderthal site near La Quina in France is a thick flake of stone with a broad, sharp edge that shows evidence of having been resharpened many times. Most other flake tools seem to have been ‘one-offs’ that were discarded after brief usage. The Quina version was not only durable but seems to have been multipurpose. Analysis of wear patterns on the sharpened edges suggest that they were deployed in carving wood and bone, removing fat and hair from animal hides, and butchery. Such scrapers have been found over a wide area of Europe, the Middle East and NE Asia mostly at Neanderthal sites, including the famous Denisova Cave of southern Siberia that yielded the first Denisovan DNA as well as that of Neanderthals.

Making a typical Quina scraper and related tools. The toolmaker would flake pieces of stone off the core and then carefully shape the Quina scraper. (Image credit: Pei-Yuan Xiao)

Until now, the early humans of East Asia were thought not to have proceeded beyond more rudimentary tools during the Middle Palaeolithic: in fact that archaeological designation hasn’t been applied there. Recent excavations at Longtan Cave in south-west China have forced a complete revision of that view (Ruan, Q.-J., et al. 2025. Quina lithic technology indicates diverse Late Pleistocene human dynamics in East Asia. Proceedings of the National Academy of Sciences, v. 122, article e2418029122; DOI: 10.1073/pnas.2418029122). The Longtan site has yielded more than fifty scrapers and the cores from which they had been struck that clearly suggest the Quina technology had been used there. They occur in cave sediments dated at between 60 and 50 ka. As yet, no human remains have been found in the same level at Longtan, although deeper levels dated at 412 ka have yielded hominin crania, mandibular fragments, and teeth, that have been suggested to be Homo erectus.

Quina type tools in East Asia may previously have been overlooked at other hominin sites in China: re-examination of archived tool collections may show they are in fact widespread. The technology could have been brought in by migrating Neanderthals, or maybe it was invented independently by local East Asian hominins. Because most living people in China carry Denisovan DNA in the genomes so perhaps that group developed the technique before interbreeding with AMH immigrants from the west. Indeed there is no reason to discard the notion that  early AMH may have imported the Quina style. A lot of work lies ahead to understand this currently unique culture at Longtan Cave. However, interpretation of another discovery published shortly after that from Longtan has spectacularly ‘stolen the thunder’ of the Qina tools, and it was made in Taiwan …

Right (top) and downward (lower) views of the partial Penghu mandible. Credit: Yousuke Kaifu University of Tokyo, Japan and Chun-Hsiang Chang Tunghai University, Taichung, from Tsutaya et al. Fig. 1 (inset)Taiwan.

About 10 years ago, Taiwanese fishers trawling in the Penghu Channel between Taiwan and China were regularly finding bones in their nets. Between 70 to 10 ka and 190 to 130 ka ago much lower sea level due to continental ice cap formation exposed the Penghu seabed. Animals and humans were thus able to move between the East Asian mainland and what is now Taiwan. The bones brought to the surface included those of elephants, water buffaloes and tigers, but one was clearly a human lower jawbone (mandible). Its shape and large molar teeth are very different from modern human mandibles and molars. A multinational team from Japan, Denmark, Taiwan and Ireland has extracted proteins from the mandible to check its genetic affinities (Tsutaya, T. and 14 others 2025. A male Denisovan mandible from Pleistocene Taiwan. Science, v. 388, p. 176-180; DOI: 10.1126/science.ads3888). Where DNA has not been preserved in bones proteomics is a useful tool, especially if results are matched with other bones that have yielded both DNA and protein sequences. In the case of the Penghu mandible, proteins from its teeth matched those of Denisovans from the Denisova Cave in Siberia which famously yielded the genome of this elusive human group. They also matched proteins from a rib found in Tibet associated with Denisovan mitochondrial DNA in cave sediments that enclosed the bones.

The three sites (Denisova, Baishiya Cave in Tibet and Penghu Channel) that have produced plausible Denisovan specimens span a large range of latitudes and altitudes. This suggests that Denisovans were capable of successful subsistence across much of East Asia. The Penghu mandible and teeth are similar to several hominin specimens from elsewhere in China that hitherto have been attributed to H. erectus. Apart from the Denisovan type locality, most of the sites have yet to be accurately dated. Having been immersed in sea water for thousands of years isotopes used in dating have been contaminated in the Panghu specimen. It can only be guessed to have lived when the seabed from which it was recovered was dry land; i.e. between 70 to 10 ka and 190 to 130 ka. China was undoubtedly occupied by Homo erectus during the early Pleistocene, but much younger fossils have been attributed to that species by Chinese palaeoanthropologists. Could it be that they are in fact Denisovans? Maybe such people independently developed the Quina knapping technique

See also: Marwick, B. 2025.  Unknown human species in East Asia used sophisticated tools at the same time Neanderthals did in Europe. Live Science, 31 March 2025; Ashworth. J. 2025. Denisovan jawbone helps to reveal appearance of ancient human species. Natural History Museum News 11 April 2025.

A fully revised edition of Steve Drury’s book Stepping Stones: The Making of Our Home World can now be downloaded as a free eBook

How changes in the Earth System have affected human evolution, migration and culture

Refugees from the Middle East migrating through Slovenia in 2015. Credit: Britannica

During the Pliocene (5.3 to 2.7 Ma) there evolved a network of various hominins, with their remains scattered across both the northern and southern parts of that continent. The earliest, though somewhat disputed hominin fossil Sahelanthropus tchadensis hails from northern Chad and lived  around 7 Ma ago, during the late Miocene, as did a similarly disputed creature from Kenya Orrorin tugenensis (~5.8 Ma). The two were geographically separated by 1500 km, what is now the Sahara desert and the East African Rift System.  The suggestion from mtDNA evidence that humans and chimpanzees had a common ancestor, the uncertainty about when it lived (between 13 to 5 Ma) and what it may have looked like, let alone where it lived, makes the notion debateable. There is even a possibility that the common ancestor of humans and the other anthropoid apes may have been European. Its descendants could well have crossed to North Africa when the Mediterranean Sea had been evaporated away to form the thick salt deposits that now lie beneath it: what could be termed the ‘Into Africa’ hypothesis. The better known Pliocene hominins were also widely distributed in the east and south of the African continent. Wandering around was clearly a hominin predilection from their outset. The same can be said about humans in the general sense (genus Homo) during the Early Pleistocene when some of them left Africa for Eurasia. Artifacts dated at 2.1 Ma have been found on the Loess Plateau of western China, and Georgia hosts the earliest human remains known from Eurasia. Since them H. antecessor, heidelbergensis, Neanderthals and Denisovans roamed Eurasia. Then, after about 130 ka, anatomically modern humans progressively populated all continents, except Antarctica, to their geographic extremities and from sea level to 4 km above it.

There is a popular view that curiosity and exploration are endemic and perhaps unique to the human line: ‘It’s in our genes’. But even plants migrate, as do all animal species. So it is best to be wary of a kind of hominin exceptionalism or superior motive force. Before settled agriculture, simply diffusion of populations in search of sustenance could have achieved the enormous migrations undertaken by all hominins: biological resources move and hunter gatherers follow them. The first migration of Homo erectus from Africa to northern China by way of Georgia seems to taken 200 ka at most and covered about ten thousand kilometres: on average a speed of only 50 m per year! That achievement and many others before and later were interwoven with the evolution of brain size, cognitive ability, means of communication and culture. But what were the ultimate drivers? Two recent papers in the journal Nature Communications make empirically-based cases for natural forces driving the movement of people and changes in demography.

The first considers hominin dispersal in the Palaearctic biogeographic realm: the largest of eight originally proposed by Alfred Russel Wallace in the late 19th century that encompasses the whole of Eurasia and North Africa (Zan, J. et al. 2024. Mid-Pleistocene aridity and landscape shifts promoted Palearctic hominin dispersals. Nature Communications, v. 15, article 10279; DOI: 10.1038/s41467-024-54767-0). The Palearctic comprises a wide range of ecosystems: arid to wet, tropical to arctic. After 2 Ma ago, hominins moved to all its parts several times. The approach followed by Zan et al. is to assess the 3.6 Ma record of the thick deposits of dust carried by the perpetual westerly winds that cross Central Asia. This gave rise to the huge (635,000 km2) Loess Plateau. At least 17 separate soil layers in the loess have yielded artefacts during the last 2.1 Ma. The authors radiocarbon dated the successive layers of loess in Tajikistan (286 samples) and the Tarim Basin (244 samples) as precisely as possible, achieving time resolutions of 5 to 10 ka and 10 to 20 ka respectively. To judge variations in climate in these area they also measured the carbon isotopic proportions in organic materials preserved within the layers. Another climate-linked metric that Zan et al. is a time series showing the development of river terraces across Eurasia derived from the earlier work of many geomorphologists. The results from those studies are linked to variations through time in the numbers of archaeological sites across Eurasia that have yielded hominin fossils, stone tools and signs of tool manufacture, many of which have been dated accurately.

The authors use sophisticated statistics to find correlations between times of climatic change and the signs of hominin occupation. Episodes of desertification in Palaearctic Eurasia clearly hindered hominins’ spreading across the continent either from west to east of vice versa. But there were distinct, periodic windows of climatic opportunity for that to happen that coincide with interglacial episodes, whose frequency changed at the Mid Pleistocene Transition (MPT) from about 41 ka to roughly every 100 ka. That was suggested in 2021 to have arisen from an increased roughness of the rock surface over which the great ice sheets of the Northern Hemisphere moved. This suppressed the pace of ice movement so that the 41 ka changes in the tilt of the Earth’s rotational axis could no longer drive climate change during the later Pleistocene, despite the fact that the same astronomical influence continued. The succeeding ~100 ka pulsation may or may not have been paced by the very much weaker influence of Earth changing orbital eccentricity. Whichever, after the MPT climate changes became much more extreme, making human dispersal in the Palearctic realm more problematic. Rather than hominin’s evolution driving them to a ‘Manifest Destiny’ of dominating the world vastly larger and wider inorganic forces corralled and released them so that, eventually, they did.

Much the same conclusion, it seems to me, emerges from a second study that covers the period since ~ 9 ka ago when anatomically modern humans transitioned from a globally dominant hunter-gatherer culture to one of ‘managing’ and dominating ecosystems, physical resources and ultimately the planet itself. (Wirtz, K.W et al. 2024. Multicentennial cycles in continental demography synchronous with solar activity and climate stability. Nature Communications, v. 15, article 10248; DOI: 10.1038/s41467-024-54474-w). Like Zan et al., Kai Wirtz and colleagues from Germany, Ukraine and Ireland base their findings on a vast accumulated number (~180,000) of radiocarbon dates from Holocene archaeological sites from all inhabited continents. The greatest number (>90,000) are from Europe. The authors applied statistical methods to judge human population variations since 11.7 ka in each continental area. Known sites are probably significantly outweighed by signs of human presence that remain hidden, and the diligence of surveys varies from country to country and continent to continent: Britain, the Netherlands and Southern Scandinavia are by far the best surveyed. Given those caveats, clearly this approach gives only a blurred estimate of population dynamics during the Holocene. Nonetheless the data are very interesting.

The changes in population growth rates show distinct cyclicity during the Holocene, which Wirtz et al. suggest are signs of booms and busts in population on all six continents. Matching these records against a large number of climatic time series reveals a correlation. Their chosen metric is variation in solar irradiance: the power per unit area received from the Sun. That has been directly monitored only over a couple of centuries. But ice cores and tree rings contain proxies for solar irradiance in the proportions of the radioactive isotopes 10Be and 14C contained in them respectively. Both are produced by the solar wind of high-energy charged particles (electrons, protons and helium nuclei or alpha particles) penetrating the upper atmosphere. The two isotopes have half-lives long enough for them to remain undecayed and thus detectable for tens of thousand years. Both ice cores and tree rings have decadal to annual time resolutions. Wirtz et al. find that their crude estimates of booms and busts in human populations during the Holocene seem closely to match variations in solar activity measured in this way. Climate stability favours successful subsistence and thus growth in populations. Variable climatic conditions seem to induce subsistence failures and increase mortality, probably through malnutrition.

A nice dialectic clearly emerges from these studies. ‘Boom and bust’ as regards populations in millennial and centennial to decadal terms stem from climate variations. Such cyclical change thus repeatedly hones natural selection among the survivors, both genetically and culturally, increasing their general fitness to their surroundings. Karl Marx and Friedrich Engels would have devoured these data avidly had they emerged in the 19th century. I’m sure they would have suggested from the evidence that something could go badly wrong – negation of negation, if readers care to explore that dialectical law further . . . And indeed that is happening. Humans made ecologically very fit indeed in surviving natural pressures are now stoking up a major climatic hiccup, or rather the culture and institutions that humans have evolved are doing that.

How did African humans survive the 74 ka Toba volcanic supereruption?

The largest volcanic eruption during the 2.5 million year evolution of the genius Homo, about 74 thousand years (ka) ago, formed a huge caldera in Sumatra, now filled by Lake Toba. A series of explosions lasting just 9 to 14 days was forceful enough to blast between 2,800 to 6,000 km3 of rocky debris from the crust. An estimated 800 km3 was in the form of fine volcanic ash that blanketed South Asia to a depth of 15 cm. Thin ash layers containing shards of glass from Toba occur in marine sediments beneath the Indian Ocean, the Arabian and South China Seas. Some occur as far off as sediments on the floor of Lake Malawi in southern Africa. A ‘spike’ of sulfates is present at around 74 ka in a Greenland ice core too. Stratospheric fine dust and sulfate aerosols from Toba probably caused global cooling of up to 3.5 °C over a modelled 5 years following the eruption. To make matters worse, this severe ‘volcanic winter’ occurred during a climatic transition from warm to cold caused by changes in ocean circulation and falling atmospheric CO2 concentration, known as a Dansgaard-Oeschger event.

There had been short-lived migrations of modern humans out of Africa into the Levant since about 185 ka. However, studies of the mitochondrial DNA (mtDNA) of living humans in Eurasia and Australasia suggest that permanent migration began about 60 ka ago. Another outcome of the mtDNA analysis is that the genetic diversity of living humans is surprisingly low. This suggests that human genetic diversity may have been sharply reduced globally roughly around the time of the  Toba eruption. This implies a population bottleneck with the number of humans alive at the time to the order of a few tens of thousands (see also: Toba ash and calibrating the Pleistocene record; December 2012). Could such a major genetic ‘pruning’ have happened in Africa? Over six field seasons, a large team of geoscientists and archaeologists drawn from the USA, Ethiopia, China, France and South Africa have excavated a rich Palaeolithic site in the valley of the Shinfa River, a tributary of the Blue Nile in western Ethiopia. Microscopic studies of the sediments enclosing the site yielded glass shards whose chemistry closely matches those in Toba ash, thereby providing an extremely precise date for the human occupation of the site: during the Toba eruption itself (Kappelman, Y. and 63 others 2024. Adaptive foraging behaviours in the Horn of Africa during Toba supereruption. Nature, v. 627; DOI: 10.1038/s41586-024-07208-3).

Selection of possible arrowheads from the Shinfa River site (Credit: Kappelman et al.; Blue Nile Survey Project)

The artifacts and bones of what these modern humans ate suggest a remarkable scenario for how they lived. Stone tools are finely worked from local basalt lava, quartz and flint-like chalcedony found in cavities in lava flows. Many of them are small, sharp triangular points, some of which show features consistent with their use as projectile tips that fractured on impact; they may be arrowheads, indeed the earliest known. Bones found at the site are key pointers to their diet. They are from a wide variety of animal, roughly similar to those living in the area at present: from monkeys to giraffe, guinea fowl to ostrich, and even frogs. There are remains of many fish and freshwater molluscs. Although there are no traces of plant foods, clearly those people who loved through the distant effects of Toba were well fed. Although a period of global cooling may have increased aridity at tropical latitudes in Africa, the campers were able to devise efficient strategies to obtain victuals. During wet seasons they lived off terrestrial prey animals, and during the driest times ate fish from pools in the river valley. These are hardly conditions likely to devastate their numbers, and the people seem to have been technologically flexible. Similar observations were made at the Pinnacle Point site in far-off South Africa in 2018, where Toba ash is also present. Both sites refute any retardation of human cultural progress 74 ka ago. Rather the opposite: people may have been spurred to innovation, and the new strategies may have allowed them to migrate more efficiently, perhaps along seasonal drainages. In this case that would have led them or their descendants to the Nile and a direct route to Eurasia; along ‘blue highway’ corridors as Kappelman et al. suggest.

Yet the population bottleneck implied by mtDNA analyses is only vaguely dated: it may have been well before or well after Toba. Moreover, there is a 10 ka gap between Toba and the earliest accurately dated migrants who left Africa – the first Australians at about 65 ka. However, note that there is inconclusive evidence that modern humans may have occupied Sumatra by the time of the eruption.  Much closer to the site of the eruption in southeast India, stone artifacts have been found below and above the 74 ka datum marked by the thick Toba Ash. Whether these were discarded by anatomically modern humans or earlier migrants such as Homo erectus remains unresolved. Either way, at that site there is no evidence for any mass die-off, even though conditions must have been pretty dreadful while the ash fell. But that probably only lasted for little more than a month. If the migrants did suffer very high losses to decrease the genetic diversity of the survivors, it seems just as likely to have been due to attrition on an extremely lengthy trek, with little likelihood of tangible evidence surviving. Alternatively, the out-of-Africa migrants may have been small in number and not fully representative of the genetic richness of the Africans who stayed put: a few tens of thousand migrants may not have been very diverse from the outset.