Early Humans, Denisova Cave
Facts, Lingjing bone engravings

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When Did Denisovans Live?

Denisovans were hunter-gatherers who inhabited Asia and the Pacific, during the Stone Age between 500,000 and 30,000 BC.

Denisova Cave in Siberia; the type site for the sub-species of archaic Homo known as Denisovans
Denisova Cave - a karst cave in the Bashelaksky Range of the Altai mountains - is the type site for the Denisovans (H. denisova). Image by Alexander Baidukov. (CC BY-SA 4.0)

Who were the Denisovans?

The Denisovans are a species of Stone Age hominins named after the Denisova Cave in Russia where the first fossils (teeth and small bone fragments) were unearthed.

They are the first species of early humans to be identified by genetic evidence, rather than physical remains.

Thus, although they have been assigned to the Homo genus, they have not yet received a species classification, because we have no physical description for them.

When Did Denisovans Live?

Genetic and sediment analysis indicates the species lived between about 500,000 and 30,000 BC, or possibly later.

For example, analysis of deposits at Denisova cave shows that Denisovans occupied the cave from about 300,000 to 50,000 BC, with some indications that they may have been there as late as 14,000 BC.

Archaeogenetics analysis of Denisovan DNA in Papua New Guinea and Indonesia, shows that modern humans dispersing across SE Asia, interbred with indigenous Denisovans in these countries around 44,000 BC.

A second interbreeding phase occurred about 28,000 BC and possibly as recently as 13,000 BC.

If all this is true, it means Denisovans were the last race of hominins to become extinct.

For information about the earliest prehistoric art, which emerged during the Lower Paleolithic, see: World's Oldest art (from 540,000 BC).

Where Did Denisovans Live?

So far, fossil specimens have been found in only three sites: Denisova Cave, a remote site located in the Altai Mountains in Siberian Russia; the Baishiya Karst Cave, a high-altitude Stone Age shelter on the Tibetan Plateau; and Tam Ngu Hao 2 Cave in the Annamite Mountains, in Laos.

But according to genetic studies, the distribution of Denisovans stretched across much of East Asia, and parts of western Eurasia.

Evidence shows they contributed genes to today's Melanesians and Indigenous Australian populations, whose ancestors must have interbred with Denisovans as they dispersed across southeast Asia.

Icelanders also have an anomalously high level of Denisovan DNA, suggesting a Denisovan presence in the polar north.

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Genetic analysis shows that Denisovans, Neanderthals and modern humans (H. sapiens) all evolved from (or share a common ancestor with) Homo heidelbergensis.

According to nuclear DNA obtained from a finger fragment (Denisova 3) in the Denisova Cave, Denisovans and Neanderthals were genetically closer to each other than they were to modern humans.

Further analysis showed that Denisovans/ Neanderthals diverged from modern humans about 800,000 years ago, and from each other about 650,000 years ago.

A likely explanation, is that around this time, an ancestral cohort of H. heidelbergensis left Africa and then separated.

One group migrated into central and eastern Asia and became Denisovans; the other group moved into Europe and the Middle East, becoming Neanderthals.

Meantime, those H. heidelbergensis who remained in Africa (a group sometimes referred to as Homo rhodesiensis) evolved into H. sapiens.

The Denisovan group duly split into three sub-groups: (1) Those who lived in Siberia and East Asia, some of whom took the polar route to Iceland; (2) Those who migrated to New Guinea and nearby islands; and (3) Those who dispersed into Oceania.

For more about the chronology of paleoart, see: Prehistoric Art Timeline (540,000 BC).

Discovery at Denisova Cave

Denisova Cave (named after an 18th century hermit called Denis) is situated in the Altai Mountains close to the border with Kazakhstan, Mongolia and China.

It was first explored for fossils in the 1970s, but it wasn't until 2008 that Russian archaeologists found the finger bone of a juvenile female hominin dating to 75,000–50,000 BC.

Mitochondrial DNA (mtDNA) extracted from the bone revealed it belonged to a completely new hominin, genetically separate from both from Neanderthals and H. sapiens.

Five other fossils have been recovered from the cave, including three teeth and two fragments of bone, dating from about 217,000 to 38,000 BC.

Human occupation of the cave began around 287,000 BC, and involved Neanderthals as well as Denisovans, although it is unclear whether the two species were living in the cave at the same time.

Note: In 2012, a long bone fragment fossil belonging to a 13-year old girl (nicknamed 'Denny') was recovered from Denisova Cave.

According to a genetic study led by Svante Paabo and Viviane Slon from Max Planck Institute for Evolutionary Anthropology in Germany, Denny was the daughter of a Denisovan father and a Neanderthal mother.

This shows that Neanderthals and Denisovans must have lived in close proximity to each other in the Altai Mountains.

Baishiya Cave

The Russian fossils remained the only known specimens of Denisovans until 2019, when a partial mandible - known as the Xiahe mandible - was found in 1980 by a Buddhist monk, in the Baishiya Karst Cave on the Tibetan Plateau, in China.

The fossil joined the collection of Lanzhou University, but wasn't analyzed until 2010, when it was found to be the same as the Denisovans from Denisova Cave.

Uranium-Thorium dating technology dated the specimen to at least 160,000 BC. Denisovan mtDNA was also recovered from cave deposits and dated from 100,000 to 60,000 BC.

For a simple explanation of all major relative and absolute dating technologies, please see: Dating Methods in Archaeology.

Tam Ngu Hao 2 Cave

In 2018, a team of anthropologists from Laos, France and the United States recovered a human tooth (164,000 BC) from Tam Ngu Hao 2 ('Cobra Cave'), a Stone Age cave in the Annamite Mountains.

According to the scientists, the tooth's close morphological affinity with the Chinese Xiahe mandible shows it belongs to the same taxon and most likely represents a Denisovan.

That said, the study did not rule out the possibility it might be Neanderthal.

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Facts About Denisovans

Because so few Denisovan fossils have been found, almost all of what we know about these extinct hominins derives from their DNA, which suggests the following physical traits:

Denisovan Tool Culture

Although thousands of stone tools and other artifacts have been excavated from Denisova cave, none have been positively identified as belonging to Denisovans, so their contribution to Stone Age culture is as yet unknown.

This lack of cultural data makes it difficult to form a detailed picture of the Denisovan lifestyle and flint-knapping activities.

However, scientists consider they were relatively advanced in terms of intelligence and had a similar lifestyle to other humans of the time, including Neanderthals.

This means that they created Mode 2 technology tools which were employed mostly for hunting and then butchering the kills, similar to those used by H. heidelbergensis.

These included: large tools with flakes removed from front and back to produce bifacial hand axes, carvers and cleavers.

In addition, Denisovans also fashioned tools out of bone and wood, such as scrapers, hammers and wooden spears tipped with stone spearheads or 'points'.

For more, see: History of Stone Tools.

Art Created By Denisovans

No cave art has been positively associated with Denisovans, but both H. heidelbergensis and H. neanderthalensis are known to have practised three types of art:

  1. Cupules
    Asia was a centre of cupule making, see for instance: the Auditorium Cave Cupules in India.
  2. Body painting
    Ochre pigments have been used for decorative purposes since 300,000 BC,or earlier. At Twin Rivers in Zambia, about 266,000 BC, early humans built up a collection of hematite containing reflective metallic flakes, creating a glittery effect.
  3. Engraving
    Engraving on bones and shells was another early form of decorative art practiced by hominins at this time: see, for instance the Trinil Shell Engravings (540,000 BC), and the Bilzingsleben Engravings (350,000 BC). The only example of bone engraving attributed to Denisovans comes from a site at Lingjing in Henan Province, China. The find consisted of two animal bone fragments decorated with abstract engravings - one overpainted with red ochre - dated to between 125,000 and 105,000 BC.

Other Hominins


(1) Reich, D.; Green, R. E.; Kircher, M.; et al. (2010). "Genetic history of an archaic hominin group from Denisova Cave in Siberia" (PDF). Nature. 468 (7327): 1053–60.
(2) Krause, J.; Fu, Q.; Good, J. M.; Viola, B.; et al. (2010). "The complete mitochondrial DNA genome of an unknown hominin from southern Siberia". Nature. 464 (7290): 894–897.
(3) Skov, L.; Macià, M. C.; Sveinbjörnsson, G.; et al. (2020). "The nature of Neanderthal introgression revealed by 27,566 Icelandic genomes". Nature. 582 (7810): 78–83.
(4) Chen, F., Welker, F., Shen, CC. et al. A late Middle Pleistocene Denisovan mandible from the Tibetan Plateau. Nature 569, 409–412 (2019).
(5) Demeter, F., Zanolli, C., Westaway, K.E. et al. A Middle Pleistocene Denisovan molar from the Annamite Chain of northern Laos. Nat Commun 13, 2557 (2022).
(6) Zhanyang Li, Luc Doyon, Hao Li, Qiang Wang, Zhongqiang Zhang, Qingpo Zhao and Francesco d'Errico. "Engraved bones from the archaic hominin site of Lingjing, Henan Province." Antiquity, Volume 93, Issue 370, August 2019 pp. 886-900.
(7) Gibbons, Ann (August 2011). "Who Were the Denisovans?" (PDF). Science. 333 (6046): 1084–87.

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