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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.
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
present study; K103B1, sample: scy192; K75B1, sample: scy193; K50B1S1, sample: scy197;
K89B3S2, sample: scy300; K89B1, sample: scy301; K75B2, sample: 303; K81B1S1, sample:
scy304; K87B1, sample: scy305; K43B1, sample: scy311; K65B1, sample: scy332.
Glinoe Sad
The Glinoe SAD archaeological site is located in the south-eastern part of present-day
Moldova, about 2 km from the Glinoe site (46.4107°N 29.4901°E). The cemetery consists of
10 barrows dated from Early Bronze Age to Early Iron Age (from the 2nd half of the 4th
millennium BCE to the 4th century BCE). In 2015 the barrow no 4 containing a total of 18
burials was excavated. Three of them were Cimmerian secondary burials: K4B13, K4B16,
K4B17. The assessment of the cultural affiliation of the recovered individuals was made on
the basis of grave goods. In the burial K4B16, a hand-made round-shaped pot with a narrow
neck was found. It is a typical kitchen vessel of the burials of the Chernogorovka culture
(which is the historical name of the Cimmerians) of the second half of the 8th century BCE to
the first half of the 7th century BCE (58). In the present study 2 out of 3 Cimmerian
individuals were included: K4B16 (sample: cim357) and K4B17 (sample: cim358). As the
skeletal remains of the third individual were poorly preserved, aDNA analyses could not be
performed. The skeleton of the first individual belonged to a male aged 20-50 years, whereas
the skeleton of the second one represented a male aged 20-35 years. The burial mound of this
barrow was totally destroyed by plowing.
Mokra
The Mokra archaeological site is located on the left bank of the Lower Dniester in the
Rybnitsa district, in the north-eastern part of Moldova (47.3753°N 29.0907°E). This site is
dated from the Early Bronze Age to the Early Iron Age (from the end of 4th millennium BCE
to 8th century BCE). In 1994 excavations of barrow no 1 were conducted. This barrow
embraced in total 18 burials. One of them was a secondary Cimmerian (K1B16, sample:
cim359) burial. The individual was a female aged 20-35 years. The archaeological affiliation
of this individual was assessed according to two vessels (large pots) found during the
excavations. One of them is polished and has ledges in the lower part of the body. At the
second vessel’s shoulder there is a geometric ornament, stamped with a spiked stamp. The
vessels can be dated within a wide range from the 9th century BCE to the 8th century BCE
(58).
Starosillya (790-410 BCE) and Nesterivka (300-100 BCE), Ukraine
The Scythian individuals SCY006 and SCY009-SCY011 from mainland Ukraine included
kurgan groups from Starosillya, Cherkasy Region (SCY006, SCY009, SCY010, 7th
century
BCE) and Nesterivka, Cherkasy Region (dated to 4th
century BCE).
Starosillya kurgan group
The Kurgan burial ground of the Scythian period near the village of Starosillya (Gorodishche
District, Cherkasy Region) was excavated by the Forest-Steppe Right-Bank Expedition of the
Institute of Archaeology of the National Academy of Sciences of Ukraine from 1984
onwards. Two complexes with barrow groups were identified, in total containing about 400
mounds.
The individuals selected for DNA analysis come from mounds 1 and 3 from the first burial
group, investigated in 1997 by the expedition of the Cherkasy Regional Archaeological
Inspection (59). The burial goods are few, and are partially or completely destroyed by
plowing. The burials were carried out in pits of rectangular shape, oriented along the line
NW-SE, covered with wooden partly burnt logs, with a gently sloping entrance from the
south-eastern side. Most often the burial mounds contained one burial, but sometimes the
skeletal remains were found in concentration, which suggests the dismemberment of bodies
before the burial. Anthropological research has not yet been published. In mound 1, a deep
scoop, a miniature pot, a round spindle and a bronze nail pin were found. The burial mound of
burial no 3 consisted of a ladle, a cup, two bronze pins, glass beads, paste, ceramics and lead,
a ceramic spindle and a bronze pendant in the form of a ring. In addition, remains of dyes of
bright red color were found in this burial. Based on the inventory and burial rites, the
Scythian burials at Starosillya can be dated to the 7th
century BC.
Nesterivka Burial
During the construction of the Kyiv-Odessa Highway in 2003-2004, on the outskirts of the
village of Nesterivka, in the Mankivsky District of Cherkasy Region, a late-Scythian burial
containing a golden treasure was discovered during rescue excavations by the Cherkasy
Regional Archaeological Inspection. The burial was of a woman belonging to the Scythian
military nomadic nobility, topologically dated to the IV century BC. The burial was of the so-
called catacomb type. At first, a mine was excavated to a depth of 7.4 m, from which the
burial chamber was extended. A total of 181 gold objects were discovered within the burial,
including two pendants to the headgear, as well as embroidery bands for clothing. Six of the
bands had images of human face, one contained a plant-geometric ornament. Together with
the woman, a horse was also buried. Additional material culture finds included 16 silver
jewelry harnesses, as well as a bronze lantern, 10 arrows, ceramic loom weight and spindle.
The burial appeared to have been looted in ancient times.
Cherniy Yar
The burial site Cherniy Yar is located in the south of Orenburg region in the Southern Urals
(N51°04,4090' E55°04,0210'). It consists of about 70 barrows dated from the Bronze Age to
the Middle Ages. In 2007 a unique burial belonging to the Late Sarmatian culture, was
discovered in barrow no 42 (60). The barrow was 1 m high and 30 m in diameter. The burial
represents a deep square grave surrounded by a rampart and ditch and covered by a wooden
funerary building. It is a double burial of a man and a woman. The woman's age is estimated
between 55 and 60, the man’s - 40-45 years. The skulls were artificially deformed during their
lifetime. In ancient times the barrow had been robbed, the bones were found scattered all over
the grave at a depth of 2-3.5 m below the ground level. The items found in the burial are: an
imported glass vessel, a Chinese mirror of the Han period in a wooden case, a sharpening
stone, a lot of gold and bronze sewn plaques, a pendant, a bronze cauldron, ceramic vessels
imported from the Caucasus and Central Asia, ceramic spindle whorls, a sword-belt silver
plaque with a bronze buckle.
Temyasovo
The analyzed individual were recovered from barrow no 9 of the Temyasovo-1 burial ground,
located in the Baymak District, east of the Temyasovo village, in the Republic of
Bashkortostan, Russia. The burial ground contained 13 burial mounds. The remains of three
graves lying parallel to each other were found under one mound excavated from the burial
chamber. Anthropological research has shown that they belong to an adolescent aged 10-12
years, a woman and a man of mature age. The skulls have apparent traces of artificial
deformation of circular type. Anthropological analyses of the skeletal remains and dental
traits indicate a mixture of morphological traits. The distribution of various genetic traits
suggested possibility of a kinship burial. Analysis of activity related osteological markers,
suggests that the man and woman were horsemen.
The discovered inventory comprised of a variety of products: a bronze fibula with articulated-
curved back with a curl at the end, a biconical whorl, a brooch, a navicular earring, a long
knife with a bone handle, and a composite bronze cauldron with loop-handles. Analyses of
available analogs and prevalence of chronologically distinct artefacts allows to date the
complex to the III century and attribute it to the Hun-Sarmatian time.
Archaeological context details and radiocarbon dates for investigated individuals are given in
table S1.
Supplementary Figures
Fig. S1. Radiocarbon dating and diet. (A) Calibrated radiocarbon dates. (B) Diet.
Fig. S2. Nucleotide misincorporation patterns at last 30-bp sequences. The reduced
pattern as seen in kzb002 is a result of UGD treatment.
Fig. S3. PCA with modern populations. Ancient individuals sequenced in this study are
projected on present day genetic variation. Iron Age individuals including Cimmerians,
Scythians and Sarmatians are plotted as triangles. Bronze Age individuals, Srubnaya-
Alakulskaya, are plotted as pentagons.
Fig. S4. Outgroup f3 statistics. (A) Outgroup f3-statistics calculated in the form of
f3(Yoruba, SrubnayaAlakulskaya_individual, Modern_population) projected on a
geographical map. Warmer colors show more genetic drift shared between Srubnaya-
Alakulskaya individual and modern population. (B) Outgroup f3-statistics calculated in the
form of f3(Yoruba, Scythian_individual, Modern_population) projected on a geographical
map. Warmer colors show more genetic drift shared between Scythian individual and modern
population. (C) Outgroup f3-statistics calculated in the form of f3(Yoruba,
Cimmerian_individual, Modern_population) projected on a geographical map. Warmer colors
show more genetic drift shared between Cimmerian individual and modern population. (D)
Outgroup f3-statistics calculated in the form of f3(Yoruba, Scythian_individual,
Modern_population) projected on a geographical map. Warmer colors show more genetic
drift shared between Scythian individual and modern population.
Fig. S5. PCA with Bronze Age individuals. Ancient individuals sequenced in this study and
published Bronze Age individuals are projected on present day genetic variation. Iron age
individuals including Cimmerians, Scythians and Sarmatians are plotted as triangles. Bronze
Age individuals including Srubnaya-Alakulskaya and published samples are plotted as
pentagons. Present-day individuals are plotted as gray points.
Fig. S6. PCA with Paleolithic and Mesolithic individuals. Ancient individuals sequenced in
this study and published Paleolithic and Mesolithic individuals are projected on present day
genetic variation. Iron Age individuals including Cimmerians, Scythians are plotted as
triangles. Bronze Age individuals, Srubnaya-Alakulskaya, are plotted as pentagons.
Paleolithic and Mesolithic individuals are plotted as circles. Present-day individuals are
plotted as gray points.
Fig. S7. PCA with Neolithic individuals. Ancient individuals sequenced in this study and
published Neolithic individuals are projected on present day genetic variation. Iron Age
individuals including Cimmerians, Scythians are plotted as triangles. Bronze Age individuals,
Srubnaya-Alakulskaya, are plotted as pentagons. Neolithic individuals are plotted as squares.
Present-day individuals are plotted as gray points.
Fig. S8. PCA with Chalcolithic individuals. Ancient individuals sequenced in this study and
published Chalcolithic individuals are projected on present day genetic variation. Iron Age
individuals including Cimmerians, Scythians are plotted as triangles. Bronze Age individuals,
Srubnaya-Alakulskaya, are plotted as pentagons. Chalcolithic individuals are plotted as
circles. Present-day individuals are plotted as gray points.
Fig. S9. PCA with Iron Age individuals. Ancient individuals sequenced in this study and
published Iron Age individuals are projected on present day genetic variation. Iron Age
individuals including Cimmerians, Scythians and Sarmatians and published Iron Age samples
are plotted as triangles. Bronze Age individuals, Srubnaya-Alakulskaya, are plotted as
pentagons. Present-day individuals are plotted as gray points.
Fig. S10. ADMIXTURE analysis. (A) ADMIXTURE analysis for K=2 to K=15. (B)
ADMIXTURE results for modern populations projected on a world map for K=15.