ArticlePDF Available

Genome sequence of a 45,000-year-old modern human from western Siberia


Abstract and Figures

We present the high-quality genome sequence of a 45,000-year-old modern human male from Siberia. This individual derives from a population that lived before—or simultaneously with—the separation of the populations in western and eastern Eurasia and carries a similar amount of Neanderthal ancestry as present-day Eurasians. However, the genomic segments of Neanderthal ancestry are substantially longer than those observed in present-day individuals, indicating that Neanderthal gene flow into the ancestors of this individual occurred 7,000–13,000 years before he lived. We estimate an autosomal mutation rate of 0.4 3 10 29 to 0.6 3 10 29 per site per year, a Y chromosomal mutation rate of 0.7 3 10 29 to 0.9 3 10 29 per site per year based on the additional substitutions that have occurred in present-day non-Africans compared to this genome, and a mitochondrial mutation rate of 1.8 3 10 28 to 3.2 3 10 28 per site per year based on the age of the bone.
Content may be subject to copyright.
ARTICLE doi:10.1038/nature13810
Genome sequence of a 45,000-year-old
modern human from western Siberia
Qiaomei Fu
, Heng Li
, Priya Moorjani
, Flora Jay
, Sergey M. Slepchenko
, Aleksei A. Bondarev
, Philip L. F. Johnson
Ayinuer Aximu-Petri
, Kay Pru
, Cesare de Filippo
, Matthias Meyer
, Nicolas Zwyns
, Domingo C. Salazar-Garcı
Yaroslav V. Kuzmin
, Susan G. Keates
, Pavel A. Kosintsev
, Dmitry I. Razhev
, Michael P. Richards
, Nikolai V. Peristov
Michael Lachmann
, Katerina Douka
, Thomas F. G. Higham
, Montgomery Slatkin
, Jean-Jacques Hublin
David Reich
, Janet Kelso
, T. Bence Viola
& Svante Pa
We present the high-quality genome sequence of a 45,000-year-old modern human male from Siberia. This individual
derives from a population that lived before—or simultaneously with—the separation of the populations in western and
eastern Eurasia and carries a similar amount of Neanderthal ancestry as present-day Eurasians. However, the genomic
segments of Neanderthal ancestry are substantially longer than those observed in present-day individuals, indicating
that Neanderthal gene flow into the ancestors of this individual occurred 7,000–13,000 years before he lived. We
estimate an autosomal mutation rate of 0.4 310
to 0.6 310
per site per year, a Y chromosomal mutation rate of
0.7 310
to 0.9 310
per site per year based on the additional substitutions that have occurred in present-day non-
Africans compared to this genome, and a mitochondrial mutation rate of 1.8 310
to 3.2 310
per site per year based
on the age of the bone.
In 2008, a relatively complete left human femoral diaphysis was discov-
ered on the banks of the river Irtysh (Fig. 1a, c, d), near the settlement
of Ust’-Ishim in western Siberia (Omsk Oblast, Russian Federation).
Although the exact locality is unclear, the femur was eroding out of al-
luvial deposits on the left bank of the river, north of Ust’-Ishim. Here,
Late Pleistocene and probably redeposited Middle Pleistocene fossils
are found in sand and gravel layers that are about 50,000–30,000 years
old (that is, from Marine Oxygen Isotope Stage 3).
Morphology, dating and diet
The proximal end of the bone shows a large gluteal buttress and gluteal
tuberosity, while the midshaft is dominated by a marked linea aspera,
resulting in a teardrop-shaped cross-section (Fig. 1e, f) (for details, see
Supplementary Information section 3). The morphology of the prox-
imal end of the shaft is similar to Upper Paleolithic modern humans and
distinct from Neanderthals (Supplementary Table 3.1, Supplementary
Fig. 3.2.),while the teardrop-shaped cross section of the midshaftis sim-
ilar to most Upper Paleolithic humans and early anatomically modern
. Taken together, this suggests that the Ust’-Ishim femur derives
from a modern human.
Two samples of 890 mg and 450mg of the bone were removed on sep-
arate occasionsfor dating. Collagen preservationsatisfied all criteria for
and after ultrafiltration we obtained ages of 41,400 61,300 years
before present (BP) (OxA-25516) and 41,400 61,400 BP (OxA-30190).
These two dates, when combined and corrected for fluctuations of atmo-
C through time, correspond to an age of about 45,000 calibrated
years BP (46,880–43,210 cal BP at 95.4% probability, Supplementary
Information section 1). The Ust’-Ishim individualis therefore the oldest
directly radiocarbon-datedmodern human outside Africa and the Mid-
dle East (Fig. 1b). Carbon and nitrogen isotope ratios indicate that the
diet of theUst’-Ishim individual (Supplementary Information section 4)
was based on terrestrial C
plants and animals that consumed them, but
also that an important part of his dietary protein may have come from
aquatic foods, probably freshwater fish, something that has been ob-
served in other early Upper Palaeolithic humans from Europe
DNA retrieval and sequencing
Nine samples of between41 and 130 mg of bone material wereremoved
from the distal part of the femur and used to construct DNA libraries
using a protocoldesigned to facilitatethe retrieval of short and damaged
. The percentage of DNA fragments in these libraries that could
be mapped to the human genome varied between 1.8% and 10.0% (Sup-
plementary Table 1.1).From the extract containing the highest propor-
tion of human DNA, eight further libraries were constructed. Each of
these libraries was treated with uracil-DNA glycosylase andendonucle-
ase VIII to remove deaminatedcytosine residues, and library molecules
with inserts shorter than approximately 35 base pairs(bp) were depleted
by preparative acrylamide gel electrophoresis before sequencing on the
Illumina HiSeq platform (Supplementary Information section 6). In total,
42-fold sequence coverage ofthe ,1.86 gigabases (Gb) of the autosomal
genome to which short fragments can be confidently mapped was gen-
erated. The coverage of the X and Y chromosomes was approximately
Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, IVPP, CAS, Beijing 100044, China.
Department of Evolutionary Genetics, Max Planck Institute for Evolutionary
Anthropology, D-04103 Leipzig, Germany.
Broad Institute of MIT and Harvard, Cambridge, Massachusetts 02142, USA.
Department of Genetics, Harvard Medical School, Boston, Massachusetts 02115,
Department of Biological Sciences, Columbia University, New York, New York 10027, USA.
Department of Integrative Biology, University of California, Berkeley, California 94720-3140, USA.
Institute for Problems of the Development of the North, Siberian Branch of the Russian Academy of Sciences, Tyumen 625026, Russia.
Expert Criminalistics Center, Omsk Division of the Ministry of
Internal Affairs, Omsk 644007, Russia.
Department of Biology, Emory University, Atlanta, Georgia 30322, USA.
Department of Human Evolution, Max Planck Institute for Evolutionary Anthropology,
D-04103 Leipzig, Germany.
Department of Anthropology, University of California, Davis, California 95616, USA.
Department of Archaeology, University of Cape Town, Cape Town 7701, South Africa.
Departament de Prehisto
`ria i Arqueologia, Universitat de Vale
`ncia, Valencia 46010, Spain.
Research Group on Plant Foods in Hominin Dietary Ecology, Max-Planck Institute for Evolutionary
Anthropology, D-04103 Leipzig, Germany.
Institute of Geology and Mineralogy, Siberian Branch of the Russian Academy of Sciences, Novosibirsk 630090, Russia.
Institute of Plant and Animal Ecology,
Urals Branch of the Russian Academy of Sciences, Yekaterinburg 620144, Russia.
Laboratory of Archaeology, Department of Anthropology, University of British Columbia, Vancouver, British Columbia
V6T 1Z1, Canada.
Siberian Cultural Center, Omsk 644010, Russia.
Santa Fe Institute, Santa Fe, New Mexico 87501, USA.
Oxford Radiocarbon Accelerator Unit, Research Laboratory for Archaeology
and the History of Art, University of Oxford, Oxford OX1 3QY, UK.
Howard Hughes Medical Institute, Harvard Medical School, Boston, Massachusetts 02115, USA.
23 OCTOBER 2014 | VOL 514 | NATURE | 445
Macmillan Publishers Limited. All rights reserved
half that of the autosomes (,22-fold), indicating that the bone comes
from a male. A likelihood method estimated present-day human mito-
chondrial DNA (mtDNA) contamination
to 0.50% (95% confidence
interval (CI) 0.26–0.94%), whereas a method that uses the frequency of
non-consensus bases in autosomal sequences estimated the contam-
ination to be less than 0.13% (Supplementary Information section 7).
Thus, less than 1% of the hominin DNA fragments sequenced are esti-
mated to be extraneous to the bone. After consensus genotype calling,
such low levels of contamination will tend to be eliminated.
Population relationships
About 7.7 positions per 10,000 are heterozygous in the Ust’-Ishim
genome, whereas between 9.6 and 10.5 positions are heterozygous in
present-day Africans and 5.5 and 7.7 in present-day non-Africans (Sup-
plementary Information section 12). Thus, with respect to genetic di-
versity, the population to which the Ust’-Ishim individual belonged was
more similar to present-day Eurasians than to present-day Africans,
which probably reflects the out-of-Africa bottleneck shared by non-
African populations. The Ust’-Ishim mtDNA sequence falls at the root
of a large group of related mtDNAs (the ‘R haplogroup’), which occurs
today across Eurasia (Supplementary Information section 8). The Y
chromosome sequence of the Ust’-Ishim individualis similarly inferred
to be ancestral to a group of related Y chromosomes (haplogroup K(xLT))
that occurs across Eurasia today
(Supplementary Information section 9).
As expected, the number of mutations inferred to have occurred on the
branch leading to the Ust’-Ishim mtDNA is lower than the numbers
inferred to have occurred on the branches leading to related present-
day mtDNAs (Supplementary Fig.8.1). Using this observation and nine
directly carbon-dated ancient modern human mtDNAs as calibration
in a relaxed molecular clock model, we estimate the age of the
Ust’-Ishim bone to be ,49,000 years BP (95% highest posterior den-
sity: 31,000–66,000 years BP), consistent with the radiocarbon date.
In a principal component analysis of the Ust’-Ishim autosomal ge-
nome along with genotyping data from 922 present-day individuals
from 53 populations
(Fig. 2a), the Ust’-Ishim individual clusters with
non-Africans rather than Africans. When only non-African popula-
tions are analysed (Fig. 2b), the Ust’-Ishim individual falls close to zero
on the two first principal component axes, suggesting that it does not share
much more ancestry with any particular group of present-day humans.
To determine how the Ust’-Ishim genome is related to the genomes of
present-day humans, we tested, using Dstatistics
, whether it shares more
derived alleles with one modern human than with another modern human
using pairs of human genomes from different parts of the world (Fig. 3).
Based on genotyping data for 87 African and 108 non-African indivi-
duals (Supplementary Information section 11), the Ust’-Ishim genome
shares more alleles with non-Africans than with sub-Saharan Africans
(jZj541–89), consistent with the principal component analysis, mtDNA
and Y chromosome results. Thus, the Ust’-Ishim individual represents
a population derived from, or related to, the population involved in the
dispersal of modern humans out of Africa. Among the non-Africans,
110° E
100° E
100° E
90° E
90° E
80° E
80° E
70° E
70° E60° E
60° N
60° N
50° N
50° N
Calibrated date (cal BP)
Kostenki 14 (OxA-X-2395-15)
Kostenki 1 (OxA-15055)
Tianyuan Cave (BA-03222)
Pestera cu Oase 1 (GrA-22810)
Ust’-Ishim (OxA-25516 & 30190)
Cavallo B (Model age)
GI 12H5 H4
50 mm
Kent’s Cavern 4 (
Mo del ag e
Figure 1
Geographic location, morphology and dating. a, Map of Siberia
with major archaeological sites. Red triangles: Neanderthal fossils; white
circle within a red (Neanderthal) triangle: Denisovan fossils; blue square:
Initial Upper Palaeolithic sites; yellow asterisk: Ust’-Ishim. 1: Ust’-Ishim; 2:
Chagyrskaya Cave; 3: Okladnikov Cave; 4: Denisova Cave; 5: Kara-Bom.
b, Radiocarbon ages of early modernhuman fossils in northern Eurasia and the
O palaeotemperature record. Specimens in light grey are indirectly
dated (OxCal v4.2.3(ref. 33); r:5 IntCal13 atmospheric curve
). H5: Heinrich 5
event, H4: Heinrich 4 event, GI 12: Greenland Interstadial 12. For a more
extensive comparison see Supplementary Information Fig. 2.1. cf, The Ust’-
Ishim 1 femur. c, Lateral view. d, Posterior view. e, Cross-section at the 80
percent level. f, Cross-section at the midshaft. For other views see
Supplementary Fig. 3.1.
446 | NATURE | VOL 514 | 23 OCTOBER 2014
Macmillan Publishers Limited. All rights reserved
the Ust’-Ishim genome shares more derived alleles with present-day
people fromEast Asia than with present-dayEuropeans (jZj52.1–6.4).
However, when an ,8,000-year-old genome from western Europe (La
or a 24,000-year-old genome from Siberia (Mal’ta 1)
analysed, there is no evidence that the Ust’-Ishim genome shares more
derived alleles with present-day East Asians than with theseprehistoric
individuals (jZj,2). This suggests that the population to which the Ust’-
Ishim individual belonged diverged from the ancestors of present-day
West Eurasian and East Eurasian populations before—or simultaneously
with—theirdivergence from eachother. The finding thatthe Ust’-Ishim
individual is equally closely related to present-day Asians and to 8,000-
to 24,000-year-old individualsfrom western Eurasia, but not to present-
day Europeans, is compatible with the hypothesis that present-day
Europeans derive some of their ancestry from a population that did
not participate in the initial dispersals of modern humans into Europe
and Asia
Mutation rate estimates
The high-quality Ust’-Ishim genome sequence, in combination withits
radiocarbon date, allows us to gauge the rate of mutations by estimating
the numbers of mutations that are ‘missing’ in theUst’-Ishim individual
relative to present-day humans. This results in a mutation rate estimate
of 0.44 310
to 0.63 310
per site per year using the high-coverage
genomes of 14 present-day humans. A challengefor inferring the muta-
tion rate in this way is that differences in error rates among genome
sequences can confound the inference (see discussion in ref. 12). We
therefore developed an alternative approach that leverages the Pairwise
Sequentially Markovian Coalescent (PSMC), a method which estimates
the distribution of coalescence times between the two chromosomes
across a diploid genome to estimate past changes in population size
and which is less influenced by differences in error rates. When the Ust’-
Ishim genomealong with 25 present-day human genomes are analysed
by PSMC, a recent reduction in population size similar to that seen for
11 present-day non-Africans is inferred for the Ust’-Ishim genome. How-
ever, the apparent age of this size reduction is more recent than in present-
day humans,consistent with theUst’-Ishim genomebeing older (Fig. 4).
We then compute the number of additional substitutions that are needed
to best fit the Ust’-Ishim PSMC curve to those of other non-African
genomes. Assuming that this corresponds to the number of mutations
that have accumulated over around 45,000 years, we estimate a muta-
tion rate o f 0.43 310
per site per year (95% CI 0.38 310
to 0.49 3
) that is consistent across all non-African genomes regardless of
their coverage (Supplementary Information section 14). This overall
rate, as well as the relative rates inferred for differentmutational classes
(transversions, non-CpG transitions, and CpG transitions), is similarto
the rate observed for de novo estimates from human pedigrees (,0.5 3
per site per year
) and to the direct estimate of branch shortening
(Supplementary Information section 10). As discussed elsewhere
these rates are slower than those estimated using calibrations based on
the fossil record and thus suggest older dates for the splits of modern
0.00 0.05 0.10
envector1 6.44 % of variance
Eigenvector2 3.94 % of variance
Central and South Asia
North Asia
East Asia
South East Asia
Middle East
North Africa
Sub-Saharan Africa
–0.025 0.000 0.025 0.050
envector1 5.26 % of variance
Eigenvector2 1.79 % of variance
a b
Figure 2
Principal Components(PC) analysis exploring the relationshipof
Ust’-Ishim to present-day humans. a, PC analysis using 922 present-day
individuals from 53 populations and the Ust’-Ishim individual. b, PC analysis
using Eurasian individuals and the Ust’-Ishim individual. The percentages of
the total variance explained by each eigenvector are given.
D(X Y; Ust’−Ishim Chimpanzee)
−0.10 0.00 0.10 0.20 0.30 0.40
La Braña
La Braña
La Braña
La Braña
La Braña
La Braña
La Braña
La Braña
La Braña
African Non-African
Figure 3
Statistics testing whether the Ust’-Ishim genome shares more
derived alleles with one or the other of two modern human genomes (X, Y).
We computed Dstatistics of the form D(X, Y, Ust’-Ishim, Chimpanzee) using
a subset of the genome-wide SNP array data from the Affymetrix Human
Origins array and restricting the analysis to transversions. Error bars
correspond to three standard errors. Red bars indicate that the Dstatistic is
significantly different from 0 (
.2), such that the Ust’-Ishim genome
shares more derived alleles with the genome on the right (X) than the left (Y).
Ancient genomes are given in italics.
23 OCTOBER 2014 | VOL 514 | NATURE | 447
Macmillan Publishers Limited. All rights reserved
human and archaic populations. We caution, however, that rates may
have changed over time and may differ between human populations.
However, we expect this mutation rate estimate to apply at least to
non-African populations over the past 45,000 years.
We also estimated a phylogeny relating the non-recombining part of
the Ust’-Ishim Y chromosome to those of 23 present-day males. Using
this phylogeny, we measured the number of ‘missing’ mutations in the
Ust’-Ishim Y chromosomal lineage relative to the most closely related
present-day Y chromosome analysed. This results in an estimate of the
Y chromosome mutationrate of 0.76310
per site per year (95% CI
0.67 310
to 0.86 310
) (Supplementary Information section 9),
significantly higher than the autosomal mutation rate, consistent with
mutation rates in males being higher than in females
. Finally, using
the radiocarbon date of the Ust’-Ishim femur together with the mtDNAs
of 311 present-day humans,we estimated the mutation rate of the com-
plete mtDNA to be 2.53 310
substitutions per site per year (95%
highest posterior density: 1.76 310
to 3.23 310
) (Supplementary
Information section 8) for mtDNA, in agreement with a previous
Neanderthal admixture
The time of admixture between modernhumans and Neanderthals has
previously been estimated to 37,000–86,000 years BP based on the size
of the DNA segments contributed by Neanderthals to present-day non-
. Thus, the Ust’-Ishim individual could pre-datethe Neander-
thal admixture. From the extent of sharingof derived alleles betweenthe
Neanderthal and the Ust’-Ishim genomes we estimate the proportion
of Neanderthal admixture in the Ust’-Ishim individualto be 2.3 60.3%
(Supplementary Information section 16), similar to present-day east
Asians(1.7–2.1%) and present-day Europeans(1.6–1.8%).Thus, admix-
ture with Neanderthals had already occurred by 45,000 years ago. In
contrast, we fail to detect any contribution from Denisovans, although
such a contribution exists in present-day people not only in Oceania
but to a lesser extent also in mainland east Asia
Information section 17).
The DNA segments contributed by Neanderthals to the Ust’-Ishim
individual are expected to be longer than such segments in present-
day people as the Ust’-Ishim individual lived closer in time to when the
admixture occurred, sothere was less time for the segments to be frag-
mented by recombination. To test if this is indeed the case, we identified
putative Neanderthal DNA segments in the Ust’-Ishim and present-
day genomes based on derived alleles shared with the Neanderthal ge-
nome at positions where Africans are fixed for ancestralalleles. Figure 5
shows that fragments of putative Neanderthal origin in the Ust’-Ishim
individual are substantially longer than those in present-day humans.
We use the covariance in such derived alleles of putative Neanderthal
origin across the Ust’-Ishim genome to infer that mean fragment sizes
in the Ust’-Ishimgenome are in the order of ,1.8–4.2 times longer than
in present-day genomes and that the Neanderthal gene flow occurred
232–430 generations before the Ust’-Ishim individual lived (Supplemen-
tary Information section 18; Fig. 6). Under the simplifying assumption
that the gene flow occurred as a singleevent, and assuming a generation
time of 29 years
, we estimate that the admixture between the ances-
tors of the Ust’-Ishim individual and Neanderthals occurred approxi-
mately 50,000 to 60,000 years BP, which is close to the time of the major
expansion of modern humans out of Africa and the Middle East. How-
ever, we also note that the presence of some longer fragments (Fig. 5) may
indicate that additional admixture occurred even later. Nevertheless,
these results suggest that the bulk of the Neanderthal contribution to
present-day people outside Africa does not go back to mixture between
Neanderthals and the anatomically modern humans who lived in the
Middle East at earlier times; for example, the modern humans whose
remains have been found at Skhul and Qafzeh
An Initial Upper Paleolithic individual?
A common model for the modern human colonization of Asia
that an early coastal migration gave rise to the present-day people of
Oceania, while a later more northern migration gave rise to Europeans
and mainland Asians. The fact that the 45,000-year-old individualfrom
Siberia is not more closely related to the Onge from the Andaman
Figure 5
Regions of Neanderthal ancestry on chromosome 12 in the
Ust’-Ishim individual and fifteen present-day non-Africans. The analysis is
based on SNPs where African genomes carry the ancestral allele and the
Neanderthal genome carries the derived allele. Homozygous ancestral alleles
are black, heterozygous derived alleles yellow, and homozygous derived alleles
10–5 10–4 10–3 10–2
10 kya 100 kya 1 Mya 10 Mya
Population size (scaled in units of 4μNe × 103)
Time (scaled in units of 2μT)
Figure 4
Inferred population size changes over time. ‘Time’ on the xaxis
refers to the pairwise per-site sequence divergence. If we erroneously assume
that Ust’-Ishim lived today, its inferred population size history includes an
out-of-Africa-like population bottleneck that is more recent than that seen in
present-day non-Africans (red bold curve). By shifting the Ust’-Ishim curve
to align with those in present-day non-Africans (blue bold curve), and
assuming that the number of mutations necessary to do this corresponds to
45,000 years, we estimate the autosomal mutation rate to be 0.38 310
0.49 310
per site per year. The times indicated on the top of the figure
are based on this mutation rate.
448 | NATURE | VOL 514 | 23 OCTOBER 2014
Macmillan Publishers Limited. All rights reserved
Islands (putative descendants of an early coastal migration) than he is
to present-day East Asians or Native Americans (putative descendants
of a northern migration) (Fig. 3) shows that at least one other group to
which the ancestors of the Ust’-Ishim individual belonged colonized
Asia before 45,000 years ago. Interestingly, the Ust’-Ishim individual
probably lived during a warm period (Greenland Interstadial 12) that
has been proposed to be a time of expansion of modern humans into
. However, the latter hypothesis is based only on the appear-
ance of the so-called ‘Initial Upper Paleolithic’ industries (Supplemen-
tary Information section 5), and not on the identification of modern
human remains
. It is possible that the Ust’-Ishim individual was asso-
ciated with the Asian variant of Initial Upper Paleolithic industry, doc-
umented at sites such as Kara-Bom in the Altai Mountains at about
47,000 years BP. This individual would then represent an early modern
human radiation into Europe and Central Asia that may have failed to
leave descendants among present-day populations
Online Content Methods, along with any additional Extended Data display items
and SourceData, are available in theonline version of the paper;references unique
to these sections appear only in the online paper.
Received 15 May; accepted 29 August 2014.
1. Trinkaus, E. & Ruff, C. B. Diaphyseal cross-sectional geometry of Near Eastern
Middle Paleolithic humans: the femur. J. Archaeol. Sci. 26, 409–424 (1999).
2. Brock, F. et al. Reliability of nitrogen content (%N) and carbon:nitrogen atomic
ratios (C:N) as indicators of collagen preservation suitable for radiocarbondating.
Radiocarbon 54, 879–886 (2012).
3. Richards,M. P. & Trinkaus, E. Out of Africa:modern human origins specialfeature:
isotopic evidence for the diets of European Neanderthals and early modern
humans. Proc. Natl Acad. Sci. USA 106, 16034–16039 (2009).
4. Meyer, M. et al. A high-coverage genome sequence from an archaic Denisovan
individual. Science 338, 222–226 (2012).
5. Fu, Q. et al. A revised timescale for human evolution based on ancient
mitochondrial genomes. Curr. Biol. 23, 553–559 (2013).
6. The Y Chromosome Consortium A nomenclature system for the tree of human
Y-chromosomal binary haplogroups. Genome Res. 12, 339–348 (2002).
7. Shapiro, B. et al. A Bayesian phylogenetic method to estimate unknown sequence
ages. Mol. Biol. Evol. 28, 879–887 (2011).
8. Patterson,N. et al. Ancient admixture in human history.Genetics 192, 1065–1093
9. Olalde, I. et al. Derivedimmune and ancestral pigmentation alleles in a 7,000-year-
old Mesolithic European. Nature 507, 225–228 (2014).
10. Raghavan, M. et al. Upper Palaeolithic Siberian genome reveals dual ancestry of
Native Americans. Nature 505, 87–91 (2014).
11. Lazaridis,I. et al. Ancient human genomes suggest three ancestral populations for
present-day Europeans. Nature 513, 409–413 (2014).
12. Pru
¨fer, K. et al. The complete genome sequence of a Neanderthal from the Altai
Mountains. Nature 505, 43–49 (2014).
13. Li, H. & Durbin, R. Inference of human population history from individual
whole-genome sequences. Nature 475, 493–496 (2011).
14. Scally, A. & Durbin, R. Revising the human mutation rate: implications for
understanding human evolution. Nature Rev. Genet. 13, 745–753 (2012).
15. Kong, A. et al. Rate of de novo mutations and the importance of father’s age to
disease risk. Nature 488, 471–475 (2012).
16. Langergraber, K. E. et al. Generation times in wild chimpanzees and gorillas
suggest earlier divergence times in great ape and human evolution. Proc. Natl
Acad. Sci. USA 109, 15716–15721 (2012).
17. Pru
¨fer, K. et al. The bonobo genome compared with the chimpanzee and human
genomes. Nature 486, 527–531 (2012).
18. Xue, Y. et al. Human Y chromosomebase-substitution mutation rate measured by
direct sequencing in a deep-rooting pedigree. Curr. Biol. 19, 1453–1457 (2009).
19. Kuroki, Y. et al. Comparative analysis of chimpanzee and human Y chromosomes
unveils complex evolutionary pathway. Nature Genet. 38, 158–167 (2006).
20. Wang, J., Fan, H. C., Behr, B. & Quake, S. R. Genome-wide single-cell analysis of
recombination activity and de novo mutation rates in human sperm. Cell 150,
402–412 (2012).
21. Sankararaman, S., Patterson, N., Li, H., Pa
¨bo, S. & Reich, D. The date of
interbreeding between Neandertals and modern humans. PLoS Genet. 8,
e1002947 (2012).
22. Reich, D. et al. Genetic history of an archaic hominin group from Denisova Cave in
Siberia. Nature 468, 1053–1060 (2010).
23. Reich, D. et al. Denisova admixture and the first modern human dispersals into
Southeast Asia and Oceania. Am. J. Hum. Genet. 89, 516–528 (2011).
24. Skoglund,P. & Jakobsson, M. Archaichuman ancestry in EastAsia. Proc. Natl Acad.
Sci. USA 108, 18301–18306 (2011).
25. Fenner, J. N. Cross-cultural estimation of the human generation intervalfor use in
genetics-based population divergence studies. Am. J. Phys. Anthropol. 128,
415–423 (2005).
26. McCown, T. D. & Keith, A. The StoneAge of Mount Carmel Vol. 2 (Clarendon,Oxford,
27. Vandermeersch, B. Les Hommes Fossiles de Qafzeh (Israel) 319 (E
´ditions du CNRS,
28. Rasmussen, M. et al. An Aboriginal Australian genome reveals separate human
dispersals into Asia. Science 334, 94–98 (2011).
29. Hublin,J. J. The earliest modern humancolonization of Europe.Proc. Natl Acad. Sci.
USA 109, 13471–13472 (2012).
30. Mu
¨ller, U. C. et al. The role of climatein the spread of modern humans into Europe.
Quat. Sci. Rev. 30, 273–279 (2011).
31. Goebel, T. A.,Derevianko, A. P. & Petrin, V. T.Dating the Middle to Upper Paleolithic
transition at Kara-Bom. Curr. Anthropol. 34, 452–458 (1993).
32. Kuhn, S. L. & Zwyns, N. Rethinking the initial Upper Paleolithic. Quat. Int. http:// (2014).
33. Bronk Ramsey, C., Scott, M. & van der Plicht, H. Calibration for archaeological and
environmental terrestrial samples inthe time range 26–50 ka cal
55, 2021–2027 (2013).
34. Reimer, P. J. et al. IntCal13 and Marine13 radiocarbon age calibration curves
0–50,000 Years cal
.Radiocarbon 55, 1869–1887 (2009).
Supplementary Information is available in the online version of the paper.
Acknowledgements We are grateful to P. Gunz, M. Kircher, A. I. Krivoshapkin, P. Nigst,
M. Ongyerth, N. Patterson, G. Renaud, U. Stenzel, M. Stoneking and S. Talamo for
valuable input, comments and help; T. Pfisterer and H. Temming for technical
assistance.Q.F. is funded in part by the ChineseAcademy of Sciences (XDA05130202)
and the Ministryof Science and Technologyof China (2007FY110200); P.A.K. by Urals
Branch, Russian Academy of Sciences (12-C-4-1014) and Y.V.K. by the Russian
Foundation for Basic Sciences (12-06-00045); F.J. and M.S. by the National Institutes
of Health of the USA (R01-GM40282); P.J. by the NIH (K99-GM104158); and
T.F.G.H. by ERC advanced grant 324139. D.R. is a Howard Hughes Medical Institute
Investigator and supported by the National Science Foundation (1032255) and the
NIH (GM100233). Major funding for this work was provided by the Presidential
Innovation Fund of the Max Planck Society.
Author Contributions Q.F., S.M.S., A.A.B., Y.V.K., J.K., T.B.V. and S.P. designed the
research. A.A.P. and Q.F. performed the experiments; Q.F., H.L., P.M., F.J., P.L.F.J., K.P.,
C.d.F., M.M., M.L., M.S., D.R., J.K. and S.P. analysed genetic data; K.D. and T.F.G.H.
C dating; D.C.S.-G. and M.P.R. analysed stable isotope data; N.V.P., P.A.K.
and D.I.R. contributed samples and data; S.M.S., A.A.B., N.Z., Y.V.K., S.G.K., J.-J.H. and
T.B.V. analysed archaeological andanthropological data;Q.F., J.K., T.B.V. and S.P. wrote
and edited the manuscript with input from all authors.
Author Information All sequence data have been submitted to the European
Nucleotide Archive (ENA) and are available under the following Ust’-Ishim
accession number: PRJEB6622. The data from the 25 present-day human
genomes are available from (
simons-genome-diversity-project/) and from (
altai/). Reprints and permissions information is available
The authors declare no competing financial interests. Readers are welcome to
comment on the online version of the paper. Correspondence and requests for
materials should be addressed to Q.F. (, D.R.
(, J.K. ( or T.B.V.
0.0 0.2 0.4 0.6 0.8 1.0
Genetic distance (cM)
Weighted covariance coefcient
Figure 6
Dating the Neandertal admixture in Ust’-Ishim and present-day
non-Africans. Exponentially fitted curves showing the decay of pairwise
covariance for variable positions where Africans carry ancestral alleles and the
Neanderthal genome carries derived alleles.
23 OCTOBER 2014 | VOL 514 | NATURE | 449
Macmillan Publishers Limited. All rights reserved
All sequencing was performed on the Illumina HiSeq 2000 and base-calling was
carried out using Ibis 1.1.6 9 (ref. 35). Reads were merged and remaining adaptor
sequences trimmed before being aligned to the Human reference genome (GRCh37/
1000 Genomes) using BWA (version0.5.10)
. GATK version1.3 (v1.3-14-g348f2b)
was used to produce genotype calls for each site. We excluded from analysis tan-
dem repeats and regions of the genome that are not unique. We considered only
genomic regions that fall withinthe 95% coverage distribution(Supplementary In-
formation section 7) and whereat least 99% of overlapping 35mers covering a pos-
ition map uniquely, allowing one mismatch.
35. Kircher, M., Stenzel,U. & Kelso, J. Improved base callingfor the Illumina Genome
Analyzer using machine learning strategies. Genome Biol. 10, R83 (2009).
36. Li, H. & Durbin, R. Fast and accurate short readalignment with Burrows–Wheeler
transform. Bioinformatics 25, 1754–1760 (2009).
Macmillan Publishers Limited. All rights reserved
... lithics). Given the assumption of Late Pleistocene population expansion into east through SCC, this area might have been highly populated at the warm and moist stages of MIS 5 [50] and later as refugium during MIS 4 [37,38,115] when between 50 and 45 ka the Ust'-Ishim man lived in western Siberia [116]. The genomic history of the Ust'-Ishim man shows that the admixture between the ancestors of the Ust'-Ishim and Neanderthals occurred PLOS ONE between ca. ...
... The genomic history of the Ust'-Ishim man shows that the admixture between the ancestors of the Ust'-Ishim and Neanderthals occurred PLOS ONE between ca. 50 to 60 ka [116]. Recent findings from Eskouldar Rockshelter at southern piedmonts of Alborz with the Initial Upper Palaeolithic industry changes our view and show the complex story of human evolution [117,118]. ...
Full-text available
The study of the cultural materials associated with the Neanderthal physical remains from the sites in the Caucasus, Central Asia and Siberian Altai and adjacent areas documents two distinct techno-complexes of Micoquian and Mousterian. These findings potentially outline two dispersal routes for the Neanderthals out of Europe. Using data on topography and Palaeoclimate, we generated computer-based least-cost-path modelling for the Neanderthal dispersal routes from Caucasus towards the east. In this regard, two dispersal routes have been identified: A northern route from Greater Caucasus associated with Micoquian techno-complex towards Siberian Altai and a southern route from Lesser Caucasus associated with Mousterian towards Siberian Altai via the Southern Caspian Corridor. Based on archaeological, bio- and physio-geographical data, our model hypothesises that during climatic deterioration phases (e.g. MIS 4) the connection between Greater and Lesser Caucasus was limited. This issue perhaps resulted in the separate development and spread of two cultural groups of Micoquian and Mousterian with an input from two different population sources of Neanderthal influxes: eastern and southern Europe refugia for these two northern and southern dispersal routes respectively. Of these two, we focus on the southern dispersal route, for it comprises a 'rapid dispersal route' towards east. The significant location of the Southern Caspian corridor between high mountains of Alborz and the Caspian Sea, provided a special biogeographical zone and a refugium. This exceptional physio-geographic condition brings forward the Southern Caspian corridor as a potential place of admixture of different hominin species including Neanderthals and homo sapiens.
... We identified default parameters that optimize performance when using a set of ca. 1.1 million autosomal SNPs that have been targeted for in-solution enrichment experiments that have produced more than 70% of genome-wide human ancient DNA data sets to date [Fu et al., 2014, 2015. Our tests show that ancIBD robustly identifies IBD longer than 8 centimorgan in aDNA data for SNP capture aDNA with at least 1x average coverage depth (on target), and for whole-genome sequencing (WGS) even as low as 0.25x average genomic coverage. ...
Full-text available
Long DNA sequences shared between two individuals, known as Identical by descent (IBD) segments, are a powerful signal for identifying close and distant biological relatives because they only arise when the pair shares a recent common ancestor. Existing methods to call IBD segments between present-day genomes cannot be straightforwardly applied to ancient DNA data (aDNA) due to typically low coverage and high genotyping error rates. We present ancIBD, a method to identify IBD segments for human aDNA data implemented as a Python package. Our approach is based on a Hidden Markov Model, using as input genotype probabilities imputed based on a modern reference panel of genomic variation. Through simulation and downsampling experiments, we demonstrate that ancIBD robustly identifies IBD segments longer than 8 centimorgan for aDNA data with at least either 0.25x average whole-genome sequencing (WGS) coverage depth or at least 1x average depth for in-solution enrichment experiments targeting a widely used aDNA SNP set (‘1240k’). This application range allows us to screen a substantial fraction of the aDNA record for IBD segments and we showcase two downstream applications. First, leveraging the fact that biological relatives up to the sixth degree are expected to share multiple long IBD segments, we identify relatives between 10,156 ancient Eurasian individuals and document evidence of long-distance migration, for example by identifying a pair of two approximately fifth-degree relatives who were buried 1410km apart in Central Asia 5000 years ago. Second, by applying ancIBD, we reveal new details regarding the spread of ancestry related to Steppe pastoralists into Europe starting 5000 years ago. We find that the first individuals in Central and Northern Europe carrying high amounts of Steppe-ancestry, associated with the Corded Ware culture, share high rates of long IBD (12-25 cM) with Yamnaya herders of the Pontic-Caspian steppe, signaling a strong bottleneck and a recent biological connection on the order of only few hundred years, providing evidence that the Yamnaya themselves are a main source of Steppe ancestry in Corded Ware people. We also detect elevated sharing of long IBD segments between Corded Ware individuals and people associated with the Globular Amphora culture (GAC) from Poland and Ukraine, who were Copper Age farmers not yet carrying Steppe-like ancestry. These IBD links appear for all Corded Ware groups in our analysis, indicating that individuals related to GAC contexts must have had a major demographic impact early on in the genetic admixtures giving rise to various Corded Ware groups across Europe. These results show that detecting IBD segments in aDNA can generate new insights both on a small scale, relevant to understanding the life stories of people, and on the macroscale, relevant to large-scale cultural-historical events.
... Our species' origins lie firmly in the African continent (Scerri et al. 2019), but Homo sapiens has expanded its range several times throughout the millennia (Hershkovitz et al. 2018). The number of Homo sapiens human remains located outside of the African continent increases during the Marine Isotope Stage 3 (MIS 3 -60-30 ka) and are associated, directly or indirectly, to the Initial Upper Palaeolithic (IUP), the Uluzzian and finally the eUP technological traditions (Benazzi et al. 2011(Benazzi et al. , 2015Fu et al. 2014Fu et al. , 2015Hublin 2015;Hublin et al. 2020;Prüfer et al. 2021;Boaretto et al. 2021). While the IUP coincides with an expansion including the Near East, Southeastern Europe and Central-Northern Asia (Zwyns 2021), Uluzzian sites are distributed along the Italian and Peloponnese peninsulas (Koumouzelis et al. 2001;. ...
Full-text available
The early Upper Palaeolithic marks a technological turning point in Western Eurasia, evidenced by the increased spread of bladelet production. The two main technocomplexes, the Aurignacian and the Ahmarian, have long histories of research and have always formed part of the debate on the Homo sapiens dispersal into Europe, with changing interpretations. A large aspect of the debate surrounding the recognition of different technocomplexes revolves around the question of whether or not bladelet production is independent of blade production. Here we present a first-hand analysis of three early Upper Palaeolithic assemblages in Europe and the Levant, conventionally attributed to different technocomplexes: Al-Ansab 1, Românești-Dumbrăvița I GH3, Grotta di Fumane A1-A2. Results show that the lithic technologies at the three sites display almost identical knapping concepts, geared around bladelet production. These results and other recent reassessments support a revision of the early Upper Palaeolithic technological and taxonomical models. Supplementary Material see here:
... version 42.4) for downstream analysis 1,4,7,14,16,23,[36][37][38][39][40]42,43,45,[79][80][81][82][83][84][85][86][87][88][89][90][91][92][93][94] . ...
Full-text available
Modern humans have populated Europe for more than 45,000 years1,2. Our knowledge of the genetic relatedness and structure of ancient hunter-gatherers is however limited, owing to the scarceness and poor molecular preservation of human remains from that period³. Here we analyse 356 ancient hunter-gatherer genomes, including new genomic data for 116 individuals from 14 countries in western and central Eurasia, spanning between 35,000 and 5,000 years ago. We identify a genetic ancestry profile in individuals associated with Upper Palaeolithic Gravettian assemblages from western Europe that is distinct from contemporaneous groups related to this archaeological culture in central and southern Europe⁴, but resembles that of preceding individuals associated with the Aurignacian culture. This ancestry profile survived during the Last Glacial Maximum (25,000 to 19,000 years ago) in human populations from southwestern Europe associated with the Solutrean culture, and with the following Magdalenian culture that re-expanded northeastward after the Last Glacial Maximum. Conversely, we reveal a genetic turnover in southern Europe suggesting a local replacement of human groups around the time of the Last Glacial Maximum, accompanied by a north-to-south dispersal of populations associated with the Epigravettian culture. From at least 14,000 years ago, an ancestry related to this culture spread from the south across the rest of Europe, largely replacing the Magdalenian-associated gene pool. After a period of limited admixture that spanned the beginning of the Mesolithic, we find genetic interactions between western and eastern European hunter-gatherers, who were also characterized by marked differences in phenotypically relevant variants.
Full-text available
The increase of brain dimensions and complexity has characterized the evolution of the genus Homo. According to the available fossil and genetic evidence, a crucial stage came before the divergence of Neanderthals, Denisovans and Homo sapiens, during the Middle Pleistocene. We consider a specimen of about 400 ka, whose phenotype is at the roots of this divergence: Ceprano calvarium (Italy). Here, we show a derived cerebrovascular organization with a mosaic of modern human and primitive features characteristics. Computed microtomography shows vascular variation and ontogenetic defects associated with ventricular and lymphatic involvement while phylogenetic analyzes highlight a dysregulation of the Tet1 gene that shows an accelerated mutation rate between 1.2 Ma and 466 ka, in contrast with the expected neutral evolution of the human genome. These results shed light on the dynamics of cranio-cerebral growth during the encephalization process and on the cerebral vascular and lymphatic system involved in this process. The results of this study could have implications for the research of many of the diseases of the central nervous system that have become predominant in an increasingly structured and long-lived brain system such as that of modern Homo sapiens. One-Sentence Summary: Genetic mutation, anatomical variations and glymphatic system in the process of encephalization towards Homo sapiens.
Full-text available
The increase of brain dimensions and complexity has characterized the evolution of the genus Homo. According to the available fossil and genetic evidence, a crucial stage came before the divergence of Neanderthals, Denisovans and Homo sapiens , during the Middle Pleistocene. We consider a specimen of about 400 ka, whose phenotype is at the roots of this divergence: Ceprano calvarium (Italy). Here, we show a derived cerebrovascular organization with a mosaic of modern human and primitive features characteristics. Computed microtomography shows vascular variation and ontogenetic defects associated with ventricular and lymphatic involvement while phylogenetic analyzes highlight a dysregulation of the Tet1 gene that shows an accelerated mutation rate between 1.2 Ma and 466 ka, in contrast with the expected neutral evolution of the human genome. These results shed light on the dynamics of cranio-cerebral growth during the encephalization process and on the cerebral vascular and lymphatic system involved in this process. The results of this study could have implications for the research of many of the diseases of the central nervous system that have become predominant in an increasingly structured and long-lived brain system such as that of modern Homo sapiens .
Full-text available
The X chromosome in non-African humans shows less diversity and less Neanderthal introgression than expected from neutral evolution. Analyzing 162 human male X chromosomes worldwide, we identified fourteen chromosomal regions where nearly identical haplotypes spanning several hundred kilobases are found at high frequencies in non-Africans. Genetic drift alone cannot explain the existence of these haplotypes, which must have been associated with strong positive selection in partial selective sweeps. Moreover, the swept haplotypes are entirely devoid of archaic ancestry as opposed to the non-swept haplotypes in the same genomic regions. The ancient Ust'-Ishim male dated at 45,000 before the present (BP) also carries the swept haplotypes, implying that selection on the haplotypes must have occurred between 45,000 and 55,000 years ago. Finally, we find that the chromosomal positions of sweeps overlap previously reported hotspots of selective sweeps in great ape evolution, suggesting a mechanism of selection unique to X chromosomes.
Full-text available
The term Initial Upper Paleolithic (IUP) was originally proposed to describe a specific assemblage from the site of Boker Tachtit (level 4). The use of the term was subsequently extended to cover the earliest Upper Paleolithic assemblages in the Levant, characterized by forms of blade production that combines elements of Levallois method (faceted platforms, hard hammer percussion, flat-faced cores) with features more typical of Upper Paleolithic blade technologies. More recently, the term IUP has been broadened again to include any early Upper Paleolithic assemblage with Levallois-like features in methods of blade production, irrespective of location. Artifact assemblages conforming to this broadest definition of the IUP have been reported from a vast area, stretching from the Levant through Central and Eastern Europe to the Siberian Altai and Northwest China. Whereas it is indisputable that similar lithic technologies can be found in all of these areas, it is not self-evident that they represent a unified cultural phenomenon. An alternative possibility is convergence, common responses to adapting Mousterian/MSA Levallois technology to the production of blade blanks, or some combination of multiple local origins with subsequent dispersal. In this paper, we suggest that the current definition of IUP has become too broad to address such issues, and that understanding the origins of this phenomenon requires a more explicit differentiation between analogies and homologies in lithic assemblages.
Full-text available
Ancient genomic sequences have started to reveal the origin and the demographic impact of farmers from the Neolithic period spreading into Europe. The adoption of farming, stock breeding and sedentary societies during the Neolithic may have resulted in adaptive changes in genes associated with immunity and diet. However, the limited data available from earlier hunter-gatherers preclude an understanding of the selective processes associated with this crucial transition to agriculture in recent human evolution. Here we sequence an approximately 7,000-year-old Mesolithic skeleton discovered at the La Braña-Arintero site in León, Spain, to retrieve a complete pre-agricultural European human genome. Analysis of this genome in the context of other ancient samples suggests the existence of a common ancient genomic signature across western and central Eurasia from the Upper Paleolithic to the Mesolithic. The La Braña individual carries ancestral alleles in several skin pigmentation genes, suggesting that the light skin of modern Europeans was not yet ubiquitous in Mesolithic times. Moreover, we provide evidence that a significant number of derived, putatively adaptive variants associated with pathogen resistance in modern Europeans were already present in this hunter-gatherer.
Full-text available
We sequenced the genomes of a ~7,000 year old farmer from Germany and eight ~8,000 year old hunter-gatherers from Luxembourg and Sweden. We analyzed these and other ancient genomes1–4 with 2,345 contemporary humans to show that most present Europeans derive from at least three highly differentiated populations: West European Hunter-Gatherers (WHG), who contributed ancestry to all Europeans but not to Near Easterners; Ancient North Eurasians (ANE) related to Upper Paleolithic Siberians3, who contributed to both Europeans and Near Easterners; and Early European Farmers (EEF), who were mainly of Near Eastern origin but also harbored WHG-related ancestry. We model these populations’ deep relationships and show that EEF had ~44% ancestry from a “Basal Eurasian” population that split prior to the diversification of other non-African lineages.
Full-text available
We present a high-quality genome sequence of a Neanderthal woman from Siberia. We show that her parents were related at the level of half-siblings and that mating among close relatives was common among her recent ancestors. We also sequenced the genome of a Neanderthal from the Caucasus to low coverage. An analysis of the relationships and population history of available archaic genomes and 25 present-day human genomes shows that several gene flow events occurred among Neanderthals, Denisovans and early modern humans, possibly including gene flow into Denisovans from an unknown archaic group. Thus, interbreeding, albeit of low magnitude, occurred among many hominin groups in the Late Pleistocene. In addition, the high-quality Neanderthal genome allows us to establish a definitive list of substitutions that became fixed in modern humans after their separation from the ancestors of Neanderthals and Denisovans.
Full-text available
The origins of the First Americans remain contentious. Although Native Americans seem to be genetically most closely related to east Asians, there is no consensus with regard to which specific Old World populations they are closest to. Here we sequence the draft genome of an approximately 24,000-year-old individual (MA-1), from Mal'ta in south-central Siberia, to an average depth of 1×. To our knowledge this is the oldest anatomically modern human genome reported to date. The MA-1 mitochondrial genome belongs to haplogroup U, which has also been found at high frequency among Upper Palaeolithic and Mesolithic European hunter-gatherers, and the Y chromosome of MA-1 is basal to modern-day western Eurasians and near the root of most Native American lineages. Similarly, we find autosomal evidence that MA-1 is basal to modern-day western Eurasians and genetically closely related to modern-day Native Americans, with no close affinity to east Asians. This suggests that populations related to contemporary western Eurasians had a more north-easterly distribution 24,000 years ago than commonly thought. Furthermore, we estimate that 14 to 38% of Native American ancestry may originate through gene flow from this ancient population. This is likely to have occurred after the divergence of Native American ancestors from east Asian ancestors, but before the diversification of Native American populations in the New World. Gene flow from the MA-1 lineage into Native American ancestors could explain why several crania from the First Americans have been reported as bearing morphological characteristics that do not resemble those of east Asians. Sequencing of another south-central Siberian, Afontova Gora-2 dating to approximately 17,000 years ago, revealed similar autosomal genetic signatures as MA-1, suggesting that the region was continuously occupied by humans throughout the Last Glacial Maximum. Our findings reveal that western Eurasian genetic signatures in modern-day Native Americans derive not only from post-Columbian admixture, as commonly thought, but also from a mixed ancestry of the First Americans.
Full-text available
The IntCal09 and Marine09 radiocarbon calibration curves have been revised utilizing newly available and updated data sets from 14C measurements on tree rings, plant macrofossils, speleothems, corals, and foraminifera. The calibration curves were derived from the data using the random walk model (RWM) used to generate IntCal09 and Marine09, which has been revised to account for additional uncertainties and error structures. The new curves were ratified at the 21st International Radiocarbon conference in July 2012 and are available as Supplemental Material at The database can be accessed at © 2013 by the Arizona Board of Regents on behalf of the University of Arizona.
Full-text available
Kara-Bom est un site de plein-air de Mongolie Russe, au pied des montagnes de Sayan, a 4 km au sud du village de Elo. Les fouilles ont mis en evidence 7 niveaux d'occupation du pleistocene superieur, 2 sont caracteristiques du Mousterien, 4 du Paleolithique superieur ancien, 1 du Paleolithique superieur recent. Les datation C14 s'echelonnent entre 30.990 ± 460 BP et 43.300 ± 1.600 BP
For the older part of the radiocarbon dating range, the IntCal13 curve provides the "state of the art" for terrestrial calibration based on all available data. It is constructed from different records, each of which by themselves could be used as a "comparison tool," depending on the research objectives. This paper discusses the pros and cons of different approaches that can be taken when using 14C dates from this time range where the agreement amongst the underlying data sets is poorer than in other time periods. The discussion is illustrated with example calibrations against IntCa09, IntCal13, and comparisons to the Suigetsu record. The examples and discussion are aimed at users of terrestrial 14C dates, in particular Upper Paleolithic archaeologists and those working with environmental terrestrial materials in the same time range. © 2013 by the Arizona Board of Regents on behalf of the University of Arizona.
Previous assessments of Near Eastern Middle Palaeolithic late archaic and early modern human femoral diaphyses have noted differences in shaft shape and robusticity, which have been used to support arguments of differential locomotor activity levels or patterns between the groups. Cross-sectional biomechanical analysis of these femoral remains, however, documents that the perceived differences in overall strength are largely the product of contrasting ecogeographically related body proportions; once these proportions are taken into account and diaphyseal strength measures are appropriately scaled, any differences in overall shaft hypertrophy disappear. At the same time, clear differences in midshaft shape remain, with slightly more antero-posterior strength in the early modern human sample and more medio-lateral reinforcement in the late archaic human sample. It remains unclear to what extent these differences, as well as structural similarities in the subtrochanteric region, might be due to differences in body shape and especially pelvic and hip proportions. In combination with contrasts in femoral neck-shaft angles, it remains possible that the similarities in overall robusticity may hide a subtle mosaic of differences in factors contributing to that uniformity.
Background: Recent analyses of de novo DNA mutations in modern humans have suggested a nuclear substitution rate that is approximately half that of previous estimates based on fossil calibration. This result has led to suggestions that major events in human evolution occurred far earlier than previously thought. Results: Here, we use mitochondrial genome sequences from ten securely dated ancient modern humans spanning 40,000 years as calibration points for the mitochondrial clock, thus yielding a direct estimate of the mitochondrial substitution rate. Our clock yields mitochondrial divergence times that are in agreement with earlier estimates based on calibration points derived from either fossils or archaeological material. In particular, our results imply a separation of non-Africans from the most closely related sub-Saharan African mitochondrial DNAs (haplogroup L3) that occurred less than 62-95 kya. Conclusions: Though single loci like mitochondrial DNA (mtDNA) can only provide biased estimates of population divergence times, they can provide valid upper bounds. Our results exclude most of the older dates for African and non-African population divergences recently suggested by de novo mutation rate estimates in the nuclear genome.