Age estimates for hominin fossils and the onset of
the Upper Palaeolithic at Denisova Cave
Katerina Douka1,2*, Viviane Slon3, Zenobia Jacobs4,5, Christopher Bronk Ramsey2, Michael V. Shunkov6,7,
Anatoly P. Derevianko6,8, Fabrizio Mafessoni3, Maxim B. Kozlikin6, Bo Li4,5, Rainer Grün9, Daniel Comeskey2, Thibaut Devièse2,
Samantha Brown1, Bence Viola10, Leslie Kinsley11, Michael Buckley12, Matthias Meyer3, Richard G. Roberts4,5, Svante Pääbo3,
Janet Kelso3 & Tom Higham2*
Denisova Cave in the Siberian Altai (Russia) is a key site for
understanding the complex relationships between hominin
groups that inhabited Eurasia in the Middle and Late Pleistocene
epoch. DNA sequenced from human remains found at this
site has revealed the presence of a hitherto unknown hominin
group, the Denisovans1,2, and high-coverage genomes from
both Neanderthal and Denisovan fossils provide evidence for
admixture between these two populations
. Determining the age
of these fossils is important if we are to understand the nature of
hominin interaction, and aspects of their cultural and subsistence
adaptations. Here we present 50 radiocarbon determinations
from the late Middle and Upper Palaeolithic layers of the site.
We also report three direct dates for hominin fragments and
obtain a mitochondrial DNA sequence for one of them. We apply
a Bayesian age modelling approach that combines chronometric
(radiocarbon, uranium series and optical ages), stratigraphic and
genetic data to calculate probabilistically the age of the human
fossils at the site. Our modelled estimate for the age of theoldest
Denisovan fossil suggests that this group was present at the site as
early as 195,000years ago (at 95.4% probability). All Neanderthal
fossils—as well as Denisova11, the daughter of a Neanderthal
and a Denisovan4—date to between 80,000 and 140,000years ago.
The youngest Denisovan dates to 52,000–76,000years ago. Direct
radiocarbon dating of Upper Palaeolithic tooth pendants and bone
points yielded the earliest evidence for the production of these
artefacts in northern Eurasia, between43,000and49,000calibrated
years before present (taken as 1950). On the basis of current
archaeological evidence, it may be assumed that these artefacts
are associated with the Denisovan population. It is not currently
possible to determine whether anatomically modern humans
were involved in their production, as modern-human fossil and
genetic evidence of such antiquity has not yet been identified in
the Altai region.
Denisova Cave preserves the longest and most notable Palaeolithic
sequence in northern Asia. It consists of three chambers: Main
Chamber, East Chamber and South Chamber (Supplementary
Information, section1). Excavations at the site have so far yielded the
remains of 12 hominins (Extended Data Fig.1 and Supplementary
Information, section3); most of these remains are small and highly
fragmentary. Despite this, the preservation of DNA in some of these
remains is very good and has enabled genome-wide data to be obtained
from both Neanderthal and Denisovan human remains, as well as from
cave sediments, enabling comparisons to be madebetween the two
The chronology of the site and the age of the recovered human
remains are key unresolved issues. Previous attempts at building a
chronology at Denisova Cave have used radiocarbon dating in the
uppermost sections, and thermoluminescence dating in the older
layers9. More recently, radiocarbon dating from the uppermost
Pleistocene layers in East Chamber revealed some age variations, which
were ascribed to taphonomic mixing and carnivore bioturbation2.
A set of optical ages
has been obtained from Pleistocene sedimentary
layers in all three chambers.
Here we report 50 radiocarbon determinations from 40 samples,
collected from the upper parts of the Pleistocene sequence (layers
9–12) in Main Chamber and East Chamber (Fig.1 and Extended Data
Table1). A further 23 samples were processed but did not yield suffi-
cient carbon for dating (Supplementary Information, section2). We
selected samples of charcoal, and humanly modified bone and arte-
facts (Extended Data Fig.2 and Supplementary Information, section2)
from locations that were deemed during excavation to be undisturbed.
Where possible, the samples were prepared using robust decontamina-
tion protocols, including collagen ultrafiltration and single amino acid
extraction of hydroxyproline from bones and teeth, and acid-base-wet
oxidation stepped-combustion (ABOx-SC) or acid-wet oxidation
stepped-combustion (AOx-SC) for charcoal (Supplementary
All samples from layers 11.3, 11.4 and 12 in East Chamber, as well
as the directly dated Denisova11 bone11, pre-date the radiocarbon
age limit. In layer 11.2, we found two age clusters: three samples
have infinite ages, and three samples have finite calibrated ages
(Extended Data Table1). A horse tooth from layer 9.2 gave a result of
45,720–50,000calibrated years before present (cal. years ) (Oxford
radiocarbon laboratory code OxA-29859). This date is statistically
indistinguishable from the group of finite dates (treated with ultrafil-
tration and ABOx) from layer 11.2.
In Main Chamber, our radiocarbon ages reveal a depositional hiatus
between layers 12 and 11.4. Samples from layer 12 (at the end of the
Middle Palaeolithic) all gave infinite radiocarbon ages compared to
samples from layer 11.4, which date to between approximately 35,000
and 40,000cal. years (Fig.1).
Four pendants made from red deer (Cervus elaphus) and elk (Alces
alces) teeth—which are often associated with Upper Palaeolithic
technocomplexes—provided results of ~32,000, ~40,000 and ~45,000cal.
years (Fig.1 and Extended Data Fig.2). The oldest of these dates
(OxA-30963) is corroborated by a charcoal date (OxA-31506)
from the same stratigraphic location and year of excavation, and is
the earliest direct date for an artefact of this type in northern Eurasia.
1Department of Archaeology, Max Planck Institute for the Science of Human History, Jena, Germany. 2Oxford Radiocarbon Accelerator Unit, Research Laboratory for Archaeology and the History
of Art, University of Oxford, Oxford, UK. 3Department of Evolutionary Genetics, Max Planck Institute for Evolutionary Anthropology, Leipzig, Germany. 4Centre for Archaeological Science, School
of Earth, Atmospheric and Life Sciences, University of Wollongong, Wollongong, New South Wales, Australia. 5Australian Research Council (ARC) Centre of Excellence for Australian Biodiversity
and Heritage, University of Wollongong, Wollongong, New South Wales, Australia. 6Institute of Archaeology and Ethnography, Russian Academy of Sciences Siberian Branch, Novosibirsk, Russia.
7Novosibirsk State University, Novosibirsk, Russia. 8Altai State University, Barnaul, Russia. 9Australian Research Centre for Human Evolution, Griffith University, Brisbane, Queensland, Australia.
10Department of Anthropology, University of Toronto, Toronto, Ontario, Canada. 11Research School of Earth Sciences, The Australian National University, Canberra, Australian Capital Territory,
Australia. 12Manchester Institute of Biotechnology, University of Manchester, Manchester, UK. *e-mail: firstname.lastname@example.org; email@example.com
640 | NATURE | VOL 565 | 31 JANUARY 2019
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