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Post by Admin on Mar 14, 2022 20:49:28 GMT
CONCLUSIONS
There is irrefutable evidence that there were incursions of people from the Pontic-steppe
into Old Europe in the late 5th millennium- 4300-4000 BCE. They likely could have been riding
on horseback. They began to implant their culture on this frontier and foreign landscape with
monumental tumuli, or early forms of kurgans with a low mound of earth or stone cairns over
single or double burials with identical features of the steppe including body position, the use of
ochre, shell tempered pottery, copper spiral bracelets, boar tusk pendants and long flint blades
likely used in javelins. They also contained pottery from the local cultures, the Gumelnita and
Tripolye B1 and used fine copper and gold ornamentation like at Varna. They seemed to have
skirted the coast of the Black Sea and chased the locals away, but interacted with the CuceteniTripolye people.
However, it is difficult to tell whether these people moved to this region permanently
since no clear settlements of their culture exist. They likely were highly mobile and maintained
interactions with their home populations along the Dnieper. It is also difficult to say whether
these people were violent and were the cause of so many people abandoning their longestablished settlements in the lower Danube valley. It appears that the settlements of the Bolgrad culture north of the Danube delta were abandoned and burned soon after these Suvorovo
immigrants arrived. Most of the abandonments apparently were planned, since almost
everything was picked up. But at Vulvanesti, radiocarbon dated 4200-4100 BCE, abandonment
was quick, with many whole pots left to burn (Anthony 2007). A second and seemingly smaller
migration seems to have branched off from the first and moved into Transylvania and eastern
Hungary and left cemeteries at Deccea Muresti in the Mures valley and at Csongrad at the end of
the Tiszapolgar and the beginning of the Bodrogkeresztur periods, about 4000-3900 BCE, but
seemed not to disrupt the local cultural traditions.
The causes of such migrations are not clear but it could be related to climate change seen
in the late 5th millennium related to the Piora Oscillation. Winters began to get colder after about
4200 BCE. The marshlands of the Danube delta would have been attractive to mobile
pastoralists for winter refuge as seen by steppe pastoralists in historic times because they offered
good winter forage and cover for cattle. The Danube delta was the richest area for this in the
entire Black Sea. The first Suvorovo herders who appeared here about 4200-4100 might have
been seeking forage for their cattle during a period of particularly cold winters. These steppe
tribes also may have been seeking the highly prized manufactured copper of the Danubian
cultures. It is possible aggressive raiding from year to year frightened the people of the lower
Danube into fleeing to the west or possibly to the northeast. There they might have sought
refuge in the Tripolye cultural area and thus account for a massive increase in settlements of the
Tripolye B1 period as well in an increase in fortifications. However, it is not possible for me to
say for certain whether this was the primary cause of the collapse. Evidence of a massive
migration is not there and leads many to discount an invasion hypothesis as the cause. However,
the few dozen graves that have been found in Southeast Europe likely represent the chiefs of the
nomadic tribes over generations and offer a possibility of repeated incursions over time or
permanently settling.
The Suvorovo migrants brought with them new symbols of power and prestige
embedded in the horse-head mace heads and also probably in horses themselves. They were
almost certainly larger than the pony-like native marsh and mountain horses of central and
Western Europe (Anthony 2007: 341). Considering these steppe horses being ridden, it would
have been an impressive sight. Their mobility and lack of reliance on a failing cereal field
system appealed to the struggling farmers of Old Europe and gave the appearance of strength,
vitality, resilience, and new economic opportunities. This likely led to the first shifts in language
as these Balkan “Old Europeans” speaking perhaps an Afro-Asiatic language began to adopt the
early Proto-Indo-European dialect of these Pontic-steppe Srendy Stog folk. The old language,
associated with the tightly closed village farmers, might have become stigmatized in favor of the
language of stockbreeding and mobility in a model considering these invaders were the carriers
of a proto-Indo-European language. The Gumelnitsa were likely already switching to an
economy more reliant on animal stockbreeding and as these newcomers arrived with their herds
they knew how to manage in new ways, the Gumelnita people may have looked to them for help.
Anthony (2007) suggests the steppe migrants may have become patrons in a patron-client system
with the natives.
Tribal warfare and raiding was likely a part of this pastoral society on the steppe.
Boundaries for grazing needed to be held and enforced through strength. As the climate became
colder and dryer on the steppe, competition for good grazing land led groups to migrate to the
southwest, into the Lower Danube, river valleys of the Karanovo culture, and along the Black
Sea coast. Pedestrians without access to horses were easy targets for these Suvorovo people
Copper and cattle were likely their main targets, along with what grain supplies the tells might
have had. The fact that many of these sites in the lower Danube, such as Telish-Liga, burned
suddenly with much of the pottery and other artifacts were left in place suggests they were
fleeing. The sites of Hotnitsa and Yunasite in Bulgaria show numerous human skeletons in the
final burnt layer, which suggests a massacre took place. But it may just be that these examples
were from any number of causes such as fire spreading from a lightning strike, a prairie or forest
fire facilitated by drought, or an accidental fire from within the village in which some just didn’t
make it out.
Why the Tripolye culture avoided the calamity their Gumelnita neighbors suffered, is
worth pondering. Perhaps it was because their immediate neighbors were the Srendy Stog along
a long established frontier between agricultural Old Europe and the pastoral steppes. Centuries
of direct peaceful trade, exchange, and alliances often sealed by marriage between the two
societies had left some sort of bond that the Suvorovo groups did not want to break with the
Tripolye. If these Eneolithic steppe cattle herders were mounted, they could pick a distant target
that did not threaten these valued gift partnerships. As Anthony mentioned (2007: 239) cattle
raiding was encouraged by Indo-European beliefs and rituals. They likely needed more animals
for their herds and copper to pay for bride-prices in a society that had seen an expanding
population in the Eneolithic but which the changing environment was straining. Whatever the
exact cause raids over several generations could have eventually caused the people living in the
tells to pack up and finally move. The result would have been dispersed and difficult targets for
such raids. Although these tells were often fortified by walls and palisades, this could have done
little to stop even small bands of mounted raiders firing flaming arrows. This likely would have
led to a cycle of thieving raids and revenge killings. The dispersed clans would have practiced
more transhumant pastoralism and built small, single phase settlements like the Gumelnita B1
hamlet of Jilava. Southwest of Bucharest, with just five to six houses and a single-level cultural
deposit, it was burned and abandoned seemingly suddenly abandoned leaving behind whole pots
and many other artifacts. Eventually they began to settle back down alongside these Suvorovo
groups to form the Cernavoda I culture, after about 4000 BC.
In addition to evidence of migrations and raiding emanating from the steppes, I believe
that climate change played a significant role in the crisis. It likely precipitated the initial decline
in agricultural productivity and economy of the tell communities and affected the migrations
from the steppes. Crop failures exacerbated by warfare would have led to a more mobile
economy. Climate change, especially to colder and or dryer conditions is a powerful force that
can easily disrupt the balance of agriculture dependent communities. This left them weaker and
more vulnerable to raiding. Soil degradation caused by centuries of overuse also could have
been a factor, as Dennell showed. This could very well have led to agricultural decline in the
most heavily occupied areas of the Balkans around the same time. This probably would have led
to an economy less dependent on agriculture and the soil and more on pastoral exploitation of
more marginal environments. Vinca began to decline around 4200 BC. Yet there is almost no
evidence of kurgans or any other steppe elements there from this time suggests it was something
else. The mines of the Balkans were abandoned after 4000 BC. It appears that the
Bodrogkeresztur culture in Hungary beginning around 4000 BC began to expand south into this
territory. This is right around the same time that the copper-using cultures in central Europe and
the Carpathians switched to Transylvanian and Hungarian ores (Anthony 2007: 228). The likely
collapse of these mines probably as a result of the decline of the core cultures of Old Europe.
Vinca and many other related sites were still occupied into the Bronze Age but were smaller
This is a theory that can also be tested by soil and pollen analysis from landscapes of this period.
The introduction of wheeled carts and wagons in the mid 4th millennium along with heavier
ploughs and traction with the widespread use of the horse likely facilitated a more mobile
existence and exploitation of less ideal soils. This process accelerated along with further
migrations from the steppe.
It is also just as likely that the collapse of Chalcolithic southeastern Europe was directly
caused by the climate change that ended the Atlantic Period, the mid-Holocene optimum. A long
period of harsh winters and or droughts, or even a couple of severe years could have caused
catastrophic crop failures which led to starvation and abandonment of the tells in favor of an
economy dependent on animal herding and mobility. This seems likely due to the apparent
catastrophic nature of the crisis, as Todorovo noted, in which nearly every tell of the lower
Danube Gumeltnita culture was abandoned after about 4000 BCE. This also affected the
culturally related Karanovo IV settlements of southern Bulgaia as many were abandoned, but
likely it wasn’t as severe as the climate here is noticeably more mild. Nevertheless, Karanovo
seems to have been abandoned for nearly 500 years, while Ezero was continuously inhabited.
This catastrophe might have opened the door to migrating pastoral nomads from the steppes to
move in and mingle with the displaced Danubian people to form hybrid cultures such as the
Cernavoda and Ezero cultures. The end of the Vinca culture seems to have happened at this
same time, 4200 BCE, suggesting something widespread and catastrophic that cannot be
explained by small migrating bands of steppe herders.
Likely causes of a massive restructuring of the Late Chalcolithic of southeast Europe may
very well have been a combination three factors; migrations, climate change, and soil depletion.
In identifying the different systemic variables that led to periods of decline in Ancient Egypt,
Karl Butzer (1980) gives us a model for cultural decline that can be used in other instances,
including our Balkan Chalcolithic chiefdoms. It gives us a model with different causes often in
combination. The first variable is a social pathology that leads to overexploitation of the masses
by a growing unproductive elite, with resulting social disequilibrium and eventual
politicoeconomic collapse. This is similar to what Chapman (2006) and Windler et al. (2012)
have recently hypothesized for what happened at Varna and Durankulak based on evidence of
growing inequality in access to prestige items and luxury goods in graves. This elite likely
resulted from the copper trade. The second variable, the strength of leadership is hard to attest in
the Chalcolithic. There is no evidence of any central authority in the Gumelnita-Karanovo VI or
Vinca cultures. There is no site size hierarchy or evidence of monumental architecture and
palatial structures or elite houses at all, only elite burials. Each tell village probably had a chief
who had some sort of authority among the different clans in the community but there is no
evidence their power reached beyond their own village so a loose grip on power over a large
kingdom and territory such as the cultural area doesn’t seem to be a factor in the decline. The
third critical variable, foreign intervention, is relevant here as I have discussed. His fourth
variable, ecological stress is also a serious factor we have considered, in either climate stress or
declining soil potential.
Other variables to consider that may be more appropriate for this situation are agricultural
production, access to resources, technology, settlement aggregation, exchange networks for food,
raw materials, and finished goods, and demography (Butzer 1980: 522). Several processes with
varying periodicities of magnitude may occasionally coincide, reinforcing one another and
creating an overall tendency that is strongly unfavorable at the low point of trends. I believe that
is what happened in this case. It was a likely coincidence of nearly simultaneous negative inputs
to the system. The changing climate, declining soil productivity, and foreign incursions
triggered a catastrophic process of mutually reinforcing events. The common result is that of
rural depopulation and decreasing productivity. A similar situation appears to have happened to
the Akkadian Empire as climate change brought drought to the northern Mesopotamian plains
and weakened the empire it was finally put down by the invasion of the Guti. It seems,
according to Butzer, foreign intervention and invasions are a regular part of cultural process.
Whatever the cause(s), the consequences indeed appear to have been very dramatic, even
catastrophic as Todorova (1995) noted. Almost all the tells of the lower Danube were abandoned
within the span of about 2 centuries. As is common perhaps in archaeology, too much
interpretation can be applied based on theory. However, everyone is entitled to one. Cultural
collapse and change and their causes are age old debates in archaeology and history. They are
often complex events with different factors working in tandem and cannot be reduced to just one
cause. The point I try to make is that migrations are still a factor worth considering in this debate
even though the overall trend in the last decades has been to find other explanations.
Sherratt (1983: 188) said, “The Early Bronze Age does indeed have the character of a
fresh start, a restructuring of society on entirely new lines.” Each theory presented is worth
considering. Even though I have not amassed any new data from this context, I have attempted
to gather and present as much prior works dealing with the theory of cultural change at this time.
It seems that published detail specifically examining this ‘transitional period’ and the cultural
changes is lacking. There needs to be more intensive study on sites of this period. Interpretations
of cultural change and collapse are notoriously difficult, especially in prehistoric societies. We
can only assume that the same forces and pressures that contributed to the collapse of more
understood and or documented states and chiefdoms were at work here in the Chalcolithic
Balkans. As intriguing as the Chalcolithic cultures were by themselves, much more needs to be
discovered, interpreted or reinterpreted. In focusing on this cultural collapse, I hope to keep the
debate alive and suggest further research.
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Post by Admin on Mar 27, 2022 17:31:07 GMT
Interdisciplinary analyses of Bronze Age communities from Western Hungary reveal complex population histories Dániel Gerber, Bea Szeifert, Orsolya Székely, Balázs Egyed, Balázs Gyuris, Julia I. Giblin, Anikó Horváth, László Palcsu, Kitti Köhler, Gabriella Kulcsár, Ágnes Kustár, Vajk Szeverényi, Szilvia Fábián, Balázs Gusztáv Mende, Mária Bondár, Eszter Ari, Viktória Kiss, Anna Szécsényi-Nagy doi: doi.org/10.1101/2022.02.03.478968Abstract In this study we report 20 ancient shotgun genomes from present-day Western Hungary (3530 – 1620 cal BCE), mainly from previously understudied Baden, Somogyvár-Vinkovci, Kisapostag, and Encrusted Pottery archaeological cultures. Besides analysing archaeological, anthropological and genetic data, 14C and strontium isotope measurements complemented reconstructing the dynamics of the communities discovered at the site Balatonkeresztúr. Our results indicate the appearance of an outstandingly high Mesolithic hunter-gatherer ancestry in the largest proportion (up to ~46%) among Kisapostag associated individuals, despite this component being thought to be highly diluted by the Early Bronze Age. We show that hunter-gatherer ancestry was likely derived from a previously unrecognised source in Eastern Europe that contributed mostly to prehistoric populations in Central Europe and the Baltic region. We revealed a patrilocal residence system and local female exogamy for this Kisapostag population that was also the genetic basis of the succeeding community of the Encrusted Pottery culture, represented by a mass grave that likely resulted from an epidemic. We also created a bioinformatic pipeline dedicated for archaeogenetic data processing. By developing and applying analytical methods for analysing genetic variants we found carriers of aneuploidy and inheritable genetic diseases. Furthermore, based on genetic and anthropological data, we present here the first female facial reconstruction from the Bronze Age Carpathian Basin. Significance Here we present a genomic time transect study from the Carpathian Basin (3530 – 1620 cal BCE), that sheds light on local and interregional population processes. We not only discovered long-distance mobility to provide detailed analysis of yet understudied Bronze Age communities, but we also recovered a previously hidden remnant hunter-gatherer genetic ancestry and its contribution to various populations in Eastern and Central Europe. We integrated 14C and strontium isotope measurements to the interdisciplinary interpretation of a site with 19 individuals analysed, where patrilocal social organisation and several health-related genetic traits were detected. Furthermore, we developed new methods and method standards for computational analyses of archaic DNA, implemented to our newly developed and freely available bioinformatic pipeline. |
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Post by Admin on Mar 27, 2022 20:18:20 GMT
Introduction A number of studies addressed population historical questions in Prehistoric Europe by recovering major events connected to the pre-Neolithic hunter-gatherers (HG)1–3, their assimilation to early European farmers during the Neolithic era2,4–6, and the appearance, expansion and admixture of steppe ancestry during the Eneolithic / Late Copper Age to the dawn of Early Bronze Age4,7,8. While some of these studies are essential for understanding the foundation of the European gene pool, studies are sparse in the literature that uncover regional interactions or social stratification via kinship9–11. Additionally, except for a few well-known markers in most archaic studies – e.g. basic pigmentation markers or lactose intolerance analysed large-scale in Mathieson et al. 201512 – no deeper analyses have been made e.g. on clinical variants. Our study aimed to make a transect analysis on a single site concerning understudied archaeological assemblies, as well as introducing the PAPline (Performing Archaeogenetic Pipeline, Supplementary Information section 6), a new bioinformatic pipeline for archaic DNA analysis. We analysed the archaeological finds from Balatonkeresztúr-Réti-dűlő site in Western Hungary, where - among others - Bronze Age assemblies were found during roadwork in 2003. Three Bronze Age archaeological horizons can be distinguished, from the Somogyvár-Vinkovci culture (~2500-2200 BCE), Kisapostag culture (~2200–1900 BCE) and to the Encrusted pottery culture (~1900–1450 BCE) that were named into Bk-I, II and III phases in this study, respectively (Table 1, Supplementary Information section 1, and Fig. S.1.2.1). In order to provide additional proxy to population ancestry of the region one further Late Copper Age individual from a multiple grave of the Baden culture (3600-2800 BCE) excavated at site Balatonlelle-Rádpuszta, ~30 km away from Balatonkeresztúr was added to our dataset.  Table 1 Summary of the investigated samples. MtDNA and ChrY denote mitochondrial haplogroup and Y chromosome haplogroup. In column “Kinship” 1st and 2nd mean the degree relations. For the feature, grave IDs and details on newly reported radiocarbon dates see the Supplementary Table S1. Results We shotgun sequenced genomes of 20 individuals with 0.008 to 0.17x coverage. We also sequenced reads of a capture set consisting 3000 nuclear SNPs (single nucleotide polymorphisms), and whole mitochondrial DNAs (mtDNAs) of all individuals. The shotgun and the capture sequenced samples ultimately resulted in an average ~104k SNPs/individuals using the 1240k SNP panel for genotype calling12, see Materials and Methods and Supplementary Tables S4 and S7. We utilised STR analysis of the Y chromosome to recover paternal kinship patterns. Furthermore, we reconstructed the face of individual S13 (Bk-II), where all known biological and archaeological details were considered, see Supplementary Information section 4. The bioarchaeological analyses were completed with radiocarbon and strontium isotope analyses, the latter can be used to trace individual mobility. Archaeological and anthropological evaluation of samples Bk-I contained the remains of a single male individual having a very long (ultradolichocran) skull type which differentiates it from most individuals found at Bk-II and Bk-III that have a very short (brachycranic) skull type13 (Table 1). In Bk-II and Bk-III male dominance (~78%) suggest distinctive funeral treatment for males and females. Bk-II is represented by 3 juveniles (16-19 years olds) and 7 adults (30+ years olds) distributed into two grave groups of A and B (Table 1, Supplementary Information Fig. S.1.2.1), and one child grave (individual S10) far from the others. Most of the burials contained no remaining grave goods except for small copper jewellery in S10 and S13. Radiocarbon dates place these inhumations to ca. 2200-1770 cal BCE, however, with Bayesian analysis using the OxCal software the timespan of the Bk-II burials can be reduced to ca. 2120-1900 cal BCE (95.4% CI), with two graves (individuals S10 and S11) possibly being slightly earlier (Supplementary Information section 1.8). The absence of children from the site is a common phenomenon that can be traced back to different preservation dynamics or burial practises to adults14, while the reason for the absence of young adults (~20-30 year olds) is unknown. Bk-III is represented by a single mass grave of 8 skeletal remains of all ages that turned out to be an unusual find in a period when the cremation practises and single inhumations were common, from ca. 1870-1620 cal BCE (95.4% CI). For details, see Supplementary Information section 1. |
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Post by Admin on Mar 27, 2022 22:16:18 GMT
Uniparental genetics and kinship analyses Both Bk-II and Bk-III show phylogeographic signals for their maternal and paternal lineages. Accordingly, Bk-II is mostly defined by mtDNA connections to the region of present-day Poland and its surroundings, whereas Bk-III has more diverse maternal composition, see Supplementary Information section 2.1. Male lineages in both Bk-II and Bk-III are mostly defined by Y chromosome haplogroup I2a-L1229 (Table 1), for which network analysis (Supplementary Information section 2.2) narrowed down regional affinities to the North European plain and shows continuity between these two horizons. Uniparental diversity makeup points to a patriarchal social structure similar to previously reported Bronze Age findings9,11,15. Results are highly similar to previous observations on Encrusted Pottery culture at the Jagodnjak site (Croatia)10. The kinship network of Bk-II follows the distribution of grave groups (Fig. 1) which were likely established along family relationships and chronology. Individuals buried in the Bk-III mass grave only show a few blood relations, like a half-brother and father-son and a dizygotic twin, the latter is the most archaic detection to date to our knowledge. However, Bk-III as an extended family group can not be excluded. For further details, see Supplementary Information section 2 and Supplementary Tables S1-S3.  Fig. 1 Kinship network at Balatonkeresztúr site based on the biological age of the individuals, and the results of the uniparental and and the READ/MPMR analyses. Blue colour represents the Bk-II grave group “A” that consists of descendant individuals, while the green coloured individuals – found in Bk-II grave group “B” – are mostly the ancestors, which suggests a kinship or chronological based geographical distribution of graves. In the mass grave Bk-III (orange, east to Bk-II graves, for full site map see Supplementary Information Fig. S.1.2.1) only a partial kinship network can be observed. Specimens buried in Bk-II S10 (purple) and S13 (green), and Bk-III S14, S16 and S18 (orange) do not have 1st and 2nd degree relatives in the uncovered graves. Genetic disorders and pigmentation Investigating genetic disorders in archaic datasets is potentially valuable for history of health and medicine, and also highlights the overall genetic health of past populations. Inherited genetic disorders, if accompanied with severe phenotypic anomalies, could also explain unusual burial practises, as it was described in cases of dwarfism16. For detailed results of this topic, see Supplementary Information section 3. Aneuploidies The abnormal number of chromosomes result in a few well known diseases which we tested thoroughly. We found one individual, S10, the only child burial in Bk-II, with XYY gonosomal genotype, described as Jacob’s syndrome. Although this syndrome remains in most cases silent as it is relatively frequent (~0.1%) in today’s populations, it occasionally comes with a wide scale of symptoms17, which may be linked to its separate burial, but due to poor bone preservation for S10 no further assessments could be made. Mitochondrial DNA diseases We examined the clinical significances of the polymorphisms that can be found in the mtDNA by using the mitopathotool software on the AmtDB database18, and found that individual S1 (40+ years old) from Bk-II had one of the defining mutations (T14484C) of Leber’s hereditary optic neuropathy (LHON) causing vision loss in ~10% of females and in ~50% of males between 20-40 years of age, rarely accompanying with other neuropathies19. Pigmentation Pigmentation patterns highly differ between horizons, as Bk-I mostly possess variants for light pigmentation, blue eyes and blonde hair, while Bk-II is more similar to populations of Neolithic Europe (Fig. 2), although some variants for lighter pigmentation exist within this group too. Members of Bk-III on the other hand show a wide range from dark to light tones and even the presence of variants for red hair (Supplementary Table S5, Supplementary Information section 3.2.1).  Fig. 2 Reconstruction of individual S13. Her mouth was partly open due to maxillary prognathia and her burial position differs from the others by her unusual arm position. She likely had higher social status for the rare copper beads she had around her head. Her genomic makeup and pigmentation pattern blends well to other Bk-II individuals, and while she did not have any blood relatives at the site up to second degree, according to strontium isotope data she lived in the region, suggesting her origin from a nearby community of the same population. For more information, see Supplementary Information sections 1 and 4. |
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Post by Admin on Mar 28, 2022 18:13:33 GMT
Nuclear variants with clinical significance We also examined the nuclear genomes to find regions with clinical significance. Since a complete panel for determining disease susceptibility only exists in commercial DNA kits, and detailed description for the 1240k SNP set is not available, we created our own SNP calling panel (included in PAPline) focusing on various conditions including amyotrophic lateral sclerosis, Alzheimer disease, autism, Crohn’s disease, diabetes, lactose intolerance, tumor markers, mental disorders, Parkinson disease, schizophrenia and ulcerative colitis. For this study we used 3,874 clinically significant SNPs, which were marked as “pathogenic” or “likely pathogenic” in the ClinVar database20, by ignoring deletion, duplication and copy number variants, as well as SNPs with questionable (“reported”, “conflicting reports”, etc.) contribution to diseases. From this set we found ~2,200 SNPs which covered at least one individual, out of these 27 positions showed clinically relevant substitutions. However, test runs on database data resulted in numerous positive hits for pathogenic variants most likely related to DNA damage, which highlights the unreliability of low coverage SNP data for variant identification (Supplementary Information section 3.2.2). To overcome this issue, we considered positions with more than 0.99 genotype likelihood (GL) values calculated using ANGSD v0.93121 (Supplementary Table S6) or when skeletal features supported results. A variant for Lig4 syndrome (rs10489442122, GL=0.999) in individual S15 was detected, and some of his skeletal features (e.g. congenital hip dysplasia) show possible onset of symptoms (Supplementary Information section 3.2.2.1). In another case the physical manifestation of hereditary spastic paraplegia is likely for S11 and S6, father and son but the genotype likelihood is lower (0.67; see Supplementary Information section 3.2.2.3). Interestingly, a tumor marker on the BRCA2 gene (rs80358920, GL=0.999) in individual S9 is nowadays only prevalent in Asian populations22. For further discussions, see Supplementary Information section 3.2. Whole genome composition and genetic ancestry Balatonkeresztúr site samples According to the principal component analysis (PCA) based on 590k nuclear SNPs (Fig. 3 a.) Bk-I is clearly separated from Bk-II and Bk-III, where Bk-II has a strong shift towards HG samples23 overlapping with only a fraction of known archaic samples23 and Bk-III. Admixture and qpAdm analyses for assessing genetic components (Supplementary Tables S9, S12-16; Supplementary Information sections 5.2 and 5.5.2) revealed ~17% HG, ~40% European farmer, and ~43% steppe ancestry for Bk-I, similar to average Bronze Age Europeans. Bk-I is most likely derived from a single source that is genetically related to a Poland Southeast Bell Beaker culture (BBC) associated population (p = 0.784) in line with archaeological observations24. Bk-II comprises a unique makeup of ~42% HG, ~41% European farmer, and ~17% steppe ancestries. qpAdm analysis revealed most plausible sources of Neolithic Sweden Funnel Beaker culture (~32±8%), Poland Southeast BBC (~41±6%) and an extra HG (~29±3%) ancestry of yet unknown origin. Despite the lower supported models Bk-I can not be excluded as an ancestry component for Bk-II, while the affinity of Sweden Funnel Beaker culture associated population likely reflects a more closely related group, such as population related to the Poland Globular Amphora culture, see Supplementary Information section 5.5.2.2 and Supplementary Tables S12 and S15. Bk-III shows a shift in ancestry composition (~29% HG, ~46% European farmer, ~25% steppe). qpAdm analyses revealed that the main ancestry component for Bk-III is Bk-II (~53±5%), while “dilution” of Bk-II to Bk-III is mostly driven by population events that are yet to be uncovered.  Fig. 3 a) Principal Component Analysis based on 590k SNPs calculated by the smartpca software. Bk-II clearly separated from any known archaic Central-Eastern European populations. b) Highlighted genetic outliers from previous studies (Supplementary Information section 5) that show a similar hunter-gatherer (HG) ancestry component to Bk-II among ancient datasets. The origin of the HG component likely lies between the Carpathian mountains and the Dnieper River or the Black Sea, from where it expanded further North and from where the Bk-II population most likely originated from. Outliers signalise a westward migration route of this component detected in Bk-II. |
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