advances.sciencemag.org/cgi/content/full/4/10/eaat4457/DC1 Supplementary Materials for Ancient genomes suggest the eastern Pontic-Caspian steppe as the source of western Iron Age nomads Maja Krzewińska*, Gülşah Merve Kılınç*, Anna Juras, Dilek Koptekin, Maciej Chyleński, Alexey G. Nikitin, Nikolai Shcherbakov, Iia Shuteleva, Tatiana Leonova, Liudmila Kraeva, Flarit A. Sungatov, Alfija N. Sultanova, Inna Potekhina, Sylwia Łukasik, Marta Krenz-Niedbała, Love Dalén, Vitaly Sinika, Mattias Jakobsson, Jan Storå, Anders Götherström* *Corresponding author. Email: [email protected] (M.K.); [email protected] (G.M.K.); [email protected] (A.G.) Published 3 October 2018, Sci. Adv. 4, eaat4457 (2018) DOI: 10.1126/sciadv.aat4457 The PDF file includes: Archaeological context information Fig. S1. Radiocarbon dating and diet. Fig. S2. Nucleotide misincorporation patterns at last 30-bp sequences. Fig. S3. PCA with modern populations. Fig. S4. Outgroup f3 statistics. Fig. S5. PCA with Bronze Age individuals. Fig. S6. PCA with Paleolithic and Mesolithic individuals. Fig. S7. PCA with Neolithic individuals. Fig. S8. PCA with Chalcolithic individuals. Fig. S9. PCA with Iron Age individuals. Fig. S10. ADMIXTURE analysis. References (56–60) Other Supplementary Material for this manuscript includes the following: (available at advances.sciencemag.org/cgi/content/full/4/10/eaat4457/DC1) Table S1 (Microsoft Excel format). Archaeological information for individuals used in this study. Table S2 (Microsoft Excel format). Stable isotope and radiocarbon dating results information for individuals used in this study. Table S3 (Microsoft Excel format). Sequencing statistics and mitochondrial variants for individuals sequenced in this study.
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Ancient genomes suggest the eastern Pontic-Caspian steppe as the source of western
Iron Age nomads
Maja Krzewińska*, Gülşah Merve Kılınç*, Anna Juras, Dilek Koptekin, Maciej Chyleński, Alexey G. Nikitin, Nikolai Shcherbakov, Iia Shuteleva, Tatiana Leonova, Liudmila Kraeva, Flarit A. Sungatov, Alfija N. Sultanova,
Inna Potekhina, Sylwia Łukasik, Marta Krenz-Niedbała, Love Dalén, Vitaly Sinika, Mattias Jakobsson, Jan Storå, Anders Götherström*
Published 3 October 2018, Sci. Adv. 4, eaat4457 (2018) DOI: 10.1126/sciadv.aat4457
The PDF file includes:
Archaeological context information Fig. S1. Radiocarbon dating and diet. Fig. S2. Nucleotide misincorporation patterns at last 30-bp sequences. Fig. S3. PCA with modern populations. Fig. S4. Outgroup f3 statistics. Fig. S5. PCA with Bronze Age individuals. Fig. S6. PCA with Paleolithic and Mesolithic individuals. Fig. S7. PCA with Neolithic individuals. Fig. S8. PCA with Chalcolithic individuals. Fig. S9. PCA with Iron Age individuals. Fig. S10. ADMIXTURE analysis. References (56–60)
Other Supplementary Material for this manuscript includes the following: (available at advances.sciencemag.org/cgi/content/full/4/10/eaat4457/DC1)
Table S1 (Microsoft Excel format). Archaeological information for individuals used in this study. Table S2 (Microsoft Excel format). Stable isotope and radiocarbon dating results information for individuals used in this study. Table S3 (Microsoft Excel format). Sequencing statistics and mitochondrial variants for individuals sequenced in this study.
Table S4 (Microsoft Excel format). Mitochondrial contamination estimates for individuals sequenced in this study. Table S5 (Microsoft Excel format). Ancient sample data set details including the number of SNPs overlapping with modern reference. Table S6 (Microsoft Excel format). Outgroup f3 statistics for individuals sequenced in this study. Table S7 (Microsoft Excel format). Summary f4 statistics between Srubnaya and Andronovo, Afanasievo, and Sintashta. Table S8 (Microsoft Excel format). Summary f4 statistics between Srubnaya-Alakulskaya tested as a single population or individuals and Karasuk and other Bronze Age populations. Table S9 (Microsoft Excel format). Summary f4 statistics between Srubnaya-Alakulskaya tested as a single population or individuals and Yamnaya and other Bronze Age populations. Table S10 (Microsoft Excel format). Diversity estimates in Bronze Age populations. Table S11 (Microsoft Excel format). Summary f4 statistics between Cimmerians, Scythians, and other ancient populations. Table S12 (Microsoft Excel format). Summary f4 statistics between Cimmerians, Sarmatians, and other ancient populations. Table S13 (Microsoft Excel format). F3 support for different components changing with time in Cimmerians as observed in ADMIXTURE. Table S14 (Microsoft Excel format). Summary f4 statistics between Srubnaya/Srubnaya-Alakulskaya and Andronovo, Afanasievo, and Sintashta. Table S15 (Microsoft Excel format). Summary f4 statistics between Scythians tested as population, Yamnaya, and other Bronze Age populations. Table S16 (Microsoft Excel format). Summary f4 statistics between Cimmerians tested as a population, Srubnaya/Srubnaya-Alakulskaya, and other Bronze Age populations. Table S17 (Microsoft Excel format). Summary f4 statistics of multiway comparisons of individuals form the study together with Srubnaya. Table S18 (Microsoft Excel format). Summary f4 statistics between Sarmatians and other Bronze Age populations.
Archaeological context information
Nomadic world of the Eurasian steppe zone
The Eurasian steppe zone is a vast region united by shared ecology. The extensive grasslands
of the steppe are able to support large herds of grazing animals such as cattle, horses, sheep
and goats, which in the past facilitated the development of nomadic herding. The horse
became an essential element of the nomadic herders’ life allowing people to follow their herds
moving between seasonal pastures. Horse domestication, invention of horse-back riding and
using steppe resources for the maintenance of large herds of hoses was of key importance in
the emergence of the Western Nomads providing an unprecedented mobility and military
advantage over the settled populations in the area (1). Economically, they were nomadic
pastoralists that often depended on sedentary populations for acquisition of pottery and iron
tools. Metal trading and iron production at the time occurred in settlements of various types.
The nomadic groups relied on trade with the sedentary groups as there was no place for
specialized workshops within the mobile societies (2). Prior to emergence of the Nomadic
peoples the steppe zone was inhabited by various Bronze Age populations (7). They had
different genetic origin but shared similar subsistence strategies where keeping domestic
animals and small scale primitive farming was practiced alongside hunting and gathering. The
Bronze Age populations of the Eurasian steppe are divided into two main social and cultural
complexes; the Srubnaya (Timber Grave) and the Andronovo (Alakulskaya) cultures. The
Srubnaya complex was dispersed the Western steppe region while the Andronovo
(Alakulskaya) complex was associated with the Eastern Eurasian steppe, beyond the Ural
Mountains. The Volga-Kama region, south of the Urals was a shared contact zone between
the two groups. In the developed classical stage of Late Bronze Age (LBA) people had begun
using horses for transportation. The skill was crucial for increased mobility of the steppe zone
inhabitants in the 10th
century BCE which was further induced by a changing climate and a
search for more hospitable conditions. This increased mobility gradually resulted in the
adaptation of nomadic cattle breeding as the main form of subsistence among some of the
LBA groups. Between the 10th
and the 8th
century BCE the mobile herders dominated the
steppe and started using iron. Also, a development of a more stratified social structure is seen.
The Iron Age Nomads first appear in the eastern fringes of Europe in the early first
millennium BC. The steppe nomads consisted of various ethnically and culturally distinct
groups united by certain common cultural elements, including differently shaped and sized
barrow or kurgan burials, similar weaponry (including bows, arrows and swords), presence of
bronze cauldrons and mirrors, shared animalistic art motifs, similar horse bridles and olenniye
kamni (tree-legged carved stone stelae) (2). The Iron Age nomadic groups were part of what
has been referred to as the ‘Scytho-Siberian world’ suggesting a certain degree of ‘unity’, but
despite a number of commonly shared characteristics there were distinct local variation
between those groups. The contact zones between the nomadic world and both the eastern and
the western states are sources of available historical information about the nomadic peoples,
including Persian, Chinese or Byzantine sources. However, little historical information is
available from the heart of the nomadic world such as southern Urals.
Srubnaya (Timber Grave culture)
The Srubnaya culture is known from the eastern part of the Pontic-Caspian Steppe region. It
was an LBA complex dated between 1800-1200BCE, characterized by a burial tradition in
which the dead were interred in individual, often timber-framed burials and were
accompanied by animal remains. Despite its name the timber construction is not as common
in Srubnaya burials as previously thought (2). The Srubnaya complex succeeds the earlier
Catacomb, Poltavaka, Sintashta and Potapovka cultures. The latter two may have been direct
predecessors of both Srubnaya and Andronovo (Alakulskaya), which became the dominating
cultural complexes over both the eastern and the western parts of the steppe region in LBA.
The two LBA cultures coexisted for centuries and in the southern Urals there is evidence of
extensive contacts with many sites with an archeological record exhibiting traits and
characteristics of both traditions (1, 56, 57). According to many scholars representatives of
the Srubanya cultural complex were the predecessors of the Cimmerians (6).
Cimmerians
The Cimmerians managed to create the first extensive union of the nomadic tribes in Southern
Russia (7). According to the written sources the Cimmerians were occupants of the North
Black Sea region and directly preceded the Scythians. Little is known about their origins and
the definition of ‘Cimmerian’ is to some extent controversial due to little archaeological
uniformity allowing for a confident identification of Cimmerian sites. For that reason, few
human remains have been identified as Cimmerian thus far. The Cimmerians appear in the
North Pontic Region (NPR) in the developed classical stage of late Bronze Age and beginning
of the Iron Age (2). They probably mastered the art of iron coasting sometime in the 10th
century BCE. According to Herodotus the Cimmerians were driven from NPR sometime
between the 8th
and 7th
centuries BCE by incoming Scythians that followed the retreating
Cimmerians to Asia Minor (7).
Scythians
The Scythians are probably the most famous of the nomadic peoples, nowadays defined by
characteristic animal art forms and the kurgan burial tradition. For the Black Sea Scythians
most evidence for an origin point to Central Asia based on the absence of animalistic art
forms in the local Black Sea Late Bronze Age populations (known from East Asian Bronze
Age populations), the records of Herodotus reports, and the finds of horse harnesses that are
earlier in the Asiatic burials, etc. However, Grakov suggested that the Scythians rather were
genetically linked to the Black Sea Late Bronze Age populations and that the culture was
autochthonous to North Pontic-Caspian Steppe region. Finally, a third hypothesis, supporting
a polycentric origin, suggests an independent local development with stable contacts between
neighboring groups. Based on similarities in material culture there seems the have been well
developed connections in Early Iron Age between eastern and west steppe populations.
However, to the contemporaries, Scythians could have been any of the nomadic tribes from
the East (1). The area of Scythian influence stretched from Altai to the Carpathian Mountains,
and from Caucasus in the South to the North European Plain to the North. Scythians were
horseback warriors, settlers, craftsmen and tradesmen who formed Great Scythia in the 6th
century BC. Known for their military skills Scythians sometimes also served as mercenaries:
for instance in ancient Greece. The 3rd
century BCE was characterized by intensifying hostile
relations with Macedonians in the west and the invasion of the Sarmatians from the east,
which eventually led to the Scythian downfall. The Sarmatians and the Scythians coexisted
for a few centuries but eventually the Sarmatians succeeded the Scythians in the region (1).
Sauromatians and Sarmatians
The Sarmatians, became the politically most influential nomadic group in the eastern fringes
of the Roman Empire. Herodotus reported that Sarmatians were related to the Scythians,
though they had a different belief system. Those new tribes probably arose in response to
Alexander the Great’s expansive campaigns. The Sauromatians (Sarmatians) first appear in
6th
century BCE and the term is usually used to refer to numerous nomadic tribes that
inhabited the territories to the east of Scythia. The cultural and social relationships between
the Sauromatians and Sarmatians are unclear. Chronologically, the entire period of the Sauro-
Sarmatian culture is roughly divided into four stages: the Sauromatian period (6th
-4th
century
BCE), the Early Sarmatian period (4th
-2nd
century BCE), the Middle Sarmatian (2nd
century
BCE to 2nd
century CE) period, and the Late Sarmatian period (2nd
-4th
century CE).
According to K. F. Smirnov the origin of Sauro-Sarmatian tribes should be sought in the
Andronovo and Srubnaya (Timber Grave) cultures. It has been suggested that the Early
Sarmatians emerged from nomadic peoples of southern Urals under the influence from trans-
Uralian and Kazakh steppe peoples (7).
Description of the sites and individuals
Kazburun 1/Muradym 8 (1890-1750 BCE)
The Muradym 8 settlement is situated between the Urshak River and Bely Kluch in
Bashkorstan territory (57). The steppe plane between Dejma and Urshak Rivers is marked by
extensive settlement remains of the developed classical stage of Late Bronze Age, and formed
the contact zone between the Srubnaya and the Alakulskaya cultural complexes. The
Kazburun 1 site is a barrow burial ground associated with a Muradymovo settlement site (ca
1,500 BCE), located close to the Usmanovo village. The burial ground consists of 33 barrows
(six barrows were destroyed during the construction of the Usmanovo-Turumbet highway).
The barrows are located in four clusters; in the south-west (nine mounds), in the north-west
(nine barrows), in the central part (tree barrows) and in the south-east (six mounds) (56). Both
the settlement and the burial ground are associated with the same population, however, the
area is, as mentioned above, a mixing zone between Srubnaya and Andronovo (Alakulskaya).
The observed mixed burial customs and relatively short site occupation (see section: 2.
Radiocarbon dating) suggested the population may have consisted of people with various
backgrounds. We analyzed four individuals from the Muradym 8 settlement and nine
individuals from three barrows format the Kazburun 1 burial ground.
Glinoe (400-170 BCE)
The Glinoe site is located in the Slobodzeya district, in the southeastern part of Moldova
(46.6684°N, 29.8001°E). It consists of 114 Scythian barrows dated from the end of the 4th
century BCE to the 2nd century BCE. The assessment of the chronology of the cemetery and
cultural affiliation of the buried individuals are based on burial inventory found during the
excavations (mainly amphorae and epigraphic data, as well as ceramics and lamps). The
majority of the Scythian barrows contained single graves, rarely double graves, while
multiple graves occurred even less frequently. Human skeletal remains were deposited at the
depth of 1-6 m below the ground level. Ten of the excavated individuals were included in