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Post by Admin on Apr 15, 2024 6:03:02 GMT
 Extended Data Figure 10. A subset of IBD Mixture Modelling results for Iron Age sources. Row 1 shows variation from Denmark_Jutland to the Islands of Denmark, to Southern Sweden. Row two shows Denmark and Sweden during the Migration Period (1575 – 1200 BP, left) and the Viking Period (1200 - 800 BP, right). Row three shows the surrounding regions to the west (left) and east (right).  Extended Data Figure 11. A subset of IBD Mixture Modelling results for Iron Age sources, when including two Southern Scandinavian Iron Age sources. Row 1 shows variation from Denmark_Jutland to the Islands of Denmark, to Southern Sweden. Row two shows Denmark and Sweden during the Migration Period (1575 - 1200 BP, left) and the Viking Period (1200 - 800 BP, right). Row three shows the surrounding regions to the west (left) and east (right). From 1230 BP until 800 BP, including the Viking Age, we see most individuals modelled primarily with small proportions of ancestries that prior to 1575 BP were only found south of Scandinavia: ENS ancestry of the East North Sea coast, Northern German ancestry from Mecklenburg and Celtic ancestry of the Britain and Ireland and France, and European Farming ancestry found in western Europe (Extended Data Figures 9 - 11). On Zealand and the Baltic Islands we also detect a number of individuals with Baltic (Estonian Bronze Age) ancestry, similar to populations associated with the Slavic-related populations. In addition to these non-local ancestries, many of these individuals are modelled with small proportions of East, West and South Scandinavian ancestry primarily found within Scandinavia during the Iron Age. Although in Northern Jutland, we have evidence of admixture between the local Iron Age population and the incoming Migration Period population, suggesting that admixture at this time occurred within Scandinavia. However, we cannot exclude the possibility of admixture between the more southern sources and the Scandinavian IA sources occuring in the unsampled regions of Southern Jutland or continental Europe. The dense sampling and high resolution demographic inference have allowed us to establish a baseline ancestry for various regions, and subsequently identify outliers (Supplementary Note S6.8.1). |
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Post by Admin on Apr 17, 2024 2:43:33 GMT
Discussion
The Germanic Indo-European language group is frequently assumed to have been introduced by the first major Steppe cultures to arrive in Scandinavia. The Corded Ware culture, appearing around 4800 BP, is generally seen as a likely context 3–6, the local Jutlandic Single Grave culture often taking a central role 16,68,69. A comparable model sees the appearance of the Bell Beaker culture to Jutland and Norway around 4400 BP as the moment when this language group was introduced 7. In contrast with these older hypotheses, an East Scandinavian population, which is not detected for another 400-800 years, is revealed here as an alternative vector for the introduction of Germanic, allowing for the proposition of a revised model. Although all Early Bronze Age populations of Scandinavia derive their Steppe ancestry from people of Corded Ware culture, the earliest Scandinavian individuals carry small proportions of local Western Hunter-Gatherer ancestry, whereas the later Eastern Scandinavians are modelled with Lithuanian/Latvian Hunter-Gatherer ancestry (Extended Data Figure 3, Figure S6.5.1.4, Supplementary Note S6.5.1), indicative of a Late Neolithic cross-Baltic migration into Scandinavia. No such migration has to our knowledge been identified in the archaeological record. However, the timing coincides with the introduction of a new, Late Neolithic sheep breed to Scandinavia 70. It also coincides with the spread of a new burial rite of gallery graves in south Sweden, the Danish islands 71 and Norway 72, a new house type 70,73,74, the first durative bronze networks 75, as well as with the end of an east-west divide in Scandinavia between 4050 and 3650 BP 73.
Archaeologically, the Nordic Bronze Age is a period of strong cultural homogenisation in south Scandinavia, starting around 3500 BP, creating the so-called Nordic Cultural Zone that lasted until 2500 BP. It was accompanied by widespread mobility not least in relation to forging new alliances supporting metal distribution 76. Although it is possible additional migratory events occurred, our results based on IBD Mixture Modelling (Supplementary Note admixed source) and DATES analyses (Supplementary Note S6.7) suggest that admixture between Bronze Age Southern and Eastern Scandinavians likely occurred in Jutland and the Danish Isles during the Nordic Bronze Age, between 3700 - 3400 BP, and leading to the formation of the Iron Age Southern Scandinavians (Supplementary Note S6.5.1). The formation of the admixed Late Bronze Age Western Scandinavians as Bronze Age Western and Eastern Scandinavian similarly occurred in the overlapping time period of 4200 - 3600 BP (Figure S6.7.1), however by the Iron Age however, Norwegian individuals carry additional East Scandinavian ancestry. Linguistically, the Late Bronze Age is the period during which Palaeo-Germanic donated vocabulary to Finno-Saamic in the east and adopted vocabulary from Celtic in the south, suggesting that it was spoken widely among East Scandinavians distributed between Sweden and Denmark, and possibly also in the Nordic Bronze Age communities in Finland and Estonia 77,78.
The transition from Palaeo- to Proto-Germanic is traditionally characterised by defining phonological changes known as the Germanic sound shifts and took place around the start of the Iron Age (∼2600 BP) 8,79. This defining event has been speculated to result from the assimilation of a different, unknown language 79. Our results reveal no major admixture events around this period, suggesting that this linguistic phase shift was rather induced by other factors, such as changes in mobility patterns or social hierarchies towards the onset of the Iron Age, or by language-internal developments. At any rate, the persistent genetic border between Southern and Eastern Scandinavians throughout the Iron Age suggested that the Proto-Germanic speech community united these different populations until its dissolution around 2000 BP.
We further find that the IA Southern Scandinavians that arose from admixture between Bronze Age Southern and Eastern Scandinavians are central to understanding the Germanic dispersal. After the Pre-Roman Iron Age, around 2000 BP, Proto-Germanic diverged into North, East and West Germanic. The spread of West Germanic to Germany, the Netherlands and Britain, appears to be closely related to populations migrating from the Jutland Peninsula. In these regions, we see the transition from Bell Beaker-related to the Corded Ware-related Southern Scandinavian ancestry. For Germany and Britain, where Celtic was known to be spoken, this period also saw a linguistic transition to Germanic. In the Netherlands, IA Southern Scandinavians’ ancestry became dominant in the place of a distinct Eastern North Sea population. The linguistic affiliation of this population is unknown. According to the linguistic ‘Nordwestblock’ hypothesis, the Netherlands may have harboured a language distinct from both Celtic and Germanic80. Given that ENS is a Bell Beaker subcluster, which is associated with Celtic languages in Britain and France, our results can alternatively be brought in line with theories of Celtic speakers, perhaps including the Frisii of the Roman Period, inhabiting the Dutch North Sea coast during the Early Iron Age 81. Although no unadmixed ENS populations are found during the migration period, the incoming Southern Scandinavians carry small proportions of ENS ancestry, indicating the migrations were not a complete replacement. Dutch coastal areas see a habitation hiatus around 1600 BP and subsequent appearance of a new material culture that is often referred to as Anglo-Saxon in nature 82, mirroring the genetics and timing of the Late Iron Age, linguistically West-Germanic Frisians in this dataset. In addition, we find that the Southern Scandinavian ancestry of these migrating populations is better modelled by individuals near Southern rather than the Northern Jutland, and that the migrating populations often carry varying but minor proportions of ENS ancestry, inherited from the earlier people who previously lived in the region. In contrast to previous studies, which relied on Scandinavian samples postdating the Migration Period 47, we can now reject the Danish Isles and Sweden as a source area for the Anglo-Saxons in Britain, as these were dominated by Eastern Scandinavian ancestry prior to the Viking Age (Figure 6).
While previous studies have identified the presence of some northern European ancestry in Migration Period populations with historically documented ancestral myths about origins in northern Europe 45,48,83, they have not had the resolution to identify a source region with the resolution presented here. Here we show that the Scandinavian ancestry in most of the Langobards is from Southern Scandinavia, consistent with post-classical origin legends 84. However, three outlier Langobards from the Czech Republic and Hungary are of Eastern Scandinavian origin. The earliest individuals from Wielbark, Poland (∼1900 BP) are primarily of Eastern Scandinavian ancestry, supporting a population migration from a region and population distinct from that of the West and North Germanic populations, a scenario potentially consistent with Gothic oral history. Further south, the later Ostrogoth and Visigoth individuals (1600 - 1100 BP) who were cultural descendents of the earlier Goths, appear similar to local Southern Europeans. The two outliers from Spain have around 50% northern European ancestry, but unlike the earlier Wielbark individuals, they fall along the Northeast-Southeast Baltic cline. The genetic distinction of the Ostrogoth and Visigoth populations from the Eastern Scandinavian Wielbark Goths suggests an adoption of the culture and East Germanic language by the more southern groups.
The subsequent period (1600 - 1200 BP) was one of great turbulence, including the collapse of the Western Roman Empire, the Barbarian migrations, the Justinian plague and the Late Antique Little Ice Age resulting from volcanic eruptions (Figure 7). In the archaeological and historical literature this is considered a period of genetic continuity in Scandinavia despite a reduction in population size (Supplementary Note S7.3, S7.4), however the genetic record now negates this assumption of pervasive genetic continuity from the Iron Age on the Danish Isles, Northern Jutland and Southern Sweden. Due to the scarcity of genomes from this period we rely on other lines of evidence to provide information on the homeland and timing of this migration.
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Post by Admin on Apr 21, 2024 2:25:38 GMT
 Figure 7. Timeline showing the climatic, cultural, linguistic and genetic shifts in the Danish Isles and Southern Sweden occurring from the Migration Period to the Viking Age. The population in southern Scandinavia after 1200 BP shows hitherto unknown changes compared with the situation in the same areas before 1600 BP. Our results demonstrate the arrival of a strong component of North German IA ancestry, in combination with a series of ancestries previously associated with Celtic-speaking groups and populations carrying European Farmer (in addition to GAC) ancestry from north-western Europe. In the Danish islands, the shift amounts to a virtually complete population replacement. Subsequently these changes are supplemented by a modest arrival of eastern ancestry associated with Slavic populations, who migrated into areas south of the Baltic Sea formerly settled by East Germanic speakers, and noted as a component in Scandinavian samples after 1200 BP. In the period directly following the volcanic activity (1414 and 1411 BP) and the Justinian Plague (1409 BP), Scandinavia saw a population decline that did not fully recover until around 1300 BP (Supplementary Note S7) 85. Linguistically, this period is one of central importance to Northern Europe. Runic inscriptions from across Scandinavia testify to a North Germanic language that remained relatively similar to Proto-Germanic during 2000 - 1500 BP. However, during the Migration period (1575 - 1200 BP) the language underwent far-reaching changes resulting in the formation of Old Norse 54. The glottogenesis of Old Norse thus coincides with a period of social and demographic instability 86. Following this transition, the originally common Germanic script known as the Elder Futhark was likewise fundamentally remodelled, giving rise to the Younger Futhark that was tailored specifically to Old Norse 87,88, and was taken into use all across Scandinavia. Old Norse, spoken across a vast area, including Norway, Iceland and Sweden, was by its speakers referred to as dǫnsk tunga, i.e. the Danish tongue 89,90. Across Scandinavia, we see variation in how the populations associated with this language were established. In Denmark and Sweden we show strong genetic evidence suggesting that observed archaeological and linguistic changes are linked to the migration of Iron Age Danes. Based on the genetic heterogeneity of the migrating population and the inability to identify a suitable source population, it appears that between 1500 and 1200 BP was likely the outcome of an amalgamation among several migrating and local groups, comparable to the formation processes among Germanic groups on the continent. In contrast, in Norway, the adoption of Old Norse and similar social changes as seen in South Scandinavia occurred with limited genetic impact from Southern Scandinavian and must have been more cultural in nature. With the exception of a single early Viking sample, the majority of Viking Age Norwegians appear either to carry local ancestry, or to reflect back migrations from Celtic regions of Britain and Ireland. Of note, the border between the East and West Norse languages closely corresponds closely to that of the Southern Scandinavians and Western Scandinavians clusters during the Viking Period (Figure 6). Combined with linguistic, historical and archaeological evidence, our findings have implications for the prehistory of the Danes. Antique sources mention the Danes living in South Scandinavia by 1450 BP 91,92. According to oral histories, the South Scandinavian royal lineage of the Danes, as well as those of the Swedes and the Norwegians were initiated between 1550-1500 BP 93 and continued throughout the subsequent periods. The appearance of the Danes appears to coincide with prominent cultural changes. By the late Migration Period (1475 - 1400 BP) a new group of large princely halls was introduced in a number of sites, many of which continued in use until the end of the Viking Age 93–96. 1550 - 1450 BP saw the development and spread of Germanic animal art, an expression form that was closely tied with religious concepts, and continued to develop until the conversion to Christianity around 1000 BP 37,97,98. Finally, we see possible evidence of a political shift in the construction of the Dannevirke in Southern Jutland, a south facing moat and rampart earth stretching more than 5 km across the peninsular near Slesvig, whose second phase dates to around 1500 BP 99. Thus, the period between 1550 - 1400 BP in Scandinavia covers a number of potentially major population dynamics. The migrations and plague might have caused abandonment of marginal subsistence areas 55. During the Little Antique Ice Age, although depopulation in marginal areas occurred 100, there was continuity to some degree in more fertile and southern areas 101 also related to intensified food production 102. This is shown in the pollen data from southernmost Sweden, where woodland regeneration occurs in uplands, with continuity of agricultural production in the most favourable areas (Supplementary Note S7.4). Further north, variation between different climatic zones is noted in southern Norway, with different societal impact from place to place which does not directly correspond with the climatic data 55. For those who survived, the subsequent improving conditions and relative abundance of resources due to a lower population size would have created the opportunity for rapid expansion, as attested to in historical sources in other areas. On the present archaeological and historical evidence, we may thus conclude that the major population shift in South Scandinavia between the Roman and the Viking periods was not solely driven by the climate events or plague of 1450 - 1350 BP but instead likely took hold between 1550 and 1450 BP and was associated with the establishment and subsequent expansion of what became the Danes. The major findings from ancient DNA studies over the last 10 years have primarily concerned large scale transitions of genetically distinct populations detected with a relatively small number of genomes. Here we show how the complexities of demographic events between closely related populations can now be exposed through dense sampling through space and time and the application of improved methodologies. Our findings have important implications for the interpretation of the archaeological record after the Middle Neolithic. They additionally allow us to offer a number of revisions to the formation of West Eurasian ancestry as well as the proposition of a new model for the origin and spread of the Germanic languages. However, the present study also has limitations and raises several new questions. With the resolution now shown to be possible here, additional sampling from a series of regions will allow a series of questions to be addressed that are currently not possible with the current dataset. Of particular interest is 1) confirming the proposed Bronze Age source of the East Scandinavians along the Baltic coast, 2) identifying the Iron Age border between the East and South Scandinavian IA in continental Europe between Mecklenburg and Gdansk representing the border between East and North West Germanic, 3) determining the more localised regions both along the East North Sea coast and within Britain representing each of the Angles, Saxons and Jutes, and 4) the regions in North East Europe related to source of Baltic and Slavic populations. Our results additionally call for a reappraisal of the linguistic evidence concerning the hypothetical migration of Germanic from the Baltic into Scandinavia and its trajectory of this linguistic subgroup from the Indo-European steppe. The formation of East Scandinavians out of Baltic populations finds an evident linguistic analogue in the isoglosses shared between the Germanic and Balto-Slavic branches of the Indo-European language family, which point to prehistoric borrowing, a linguistic subclade, or both 103. On the other hand, the relatively late, Bronze Age arrival of agriculture in the Baltic 104,105 vs the presence in Proto-Germanic of agricultural terms inherited from Indo-European 106 raises a question on the suitability of the archaeological context of this area as a linguistic stepping stone during the Late Neolithic. Finally, this study highlights fundamental methodological difficulties in establishing correlations – or lack thereof – between genetic, archaeological and linguistic evidence 107,108. For instance, the immigration of East Scandinavians, central to our new model, has so far not been recognized in the archaeological record. During the Late Iron Age, Northwest Germanic was spoken by both Southern, Eastern and Northern Scandinavians, as demonstrated by runic inscriptions from across Scandinavia, despite persistent genetic boundaries between these populations. Following the Migration Period, southern European individuals exhibit late Germanic burial identities without showing ancestry from Northern Europe. These findings underline the differences in the mechanisms behind the proliferation of genetic, linguistic and cultural features and call for additional interdisciplinary studies on the integration of these diverse lines of evidence on human prehistory. |
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Post by Admin on Apr 24, 2024 12:46:25 GMT
The Genetic Origin of the Indo-Europeans Abstract The Yamnaya archaeological complex appeared around 3300BCE across the steppes north of the Black and Caspian Seas, and by 3000BCE reached its maximal extent from Hungary in the west to Kazakhstan in the east. To localize the ancestral and geographical origins of the Yamnaya among the diverse Eneolithic people that preceded them, we studied ancient DNA data from 428 individuals of which 299 are reported for the first time, demonstrating three previously unknown Eneolithic genetic clines. First, a “Caucasus-Lower Volga” (CLV) Cline suffused with Caucasus hunter-gatherer (CHG) ancestry extended between a Caucasus Neolithic southern end in Neolithic Armenia, and a steppe northern end in Berezhnovka in the Lower Volga. Bidirectional gene flow across the CLV cline created admixed intermediate populations in both the north Caucasus, such as the Maikop people, and on the steppe, such as those at the site of Remontnoye north of the Manych depression. CLV people also helped form two major riverine clines by admixing with distinct groups of European hunter-gatherers. A “Volga Cline” was formed as Lower Volga people mixed with upriver populations that had more Eastern hunter-gatherer (EHG) ancestry, creating genetically hyper-variable populations as at Khvalynsk in the Middle Volga. A “Dnipro Cline” was formed as CLV people bearing both Caucasus Neolithic and Lower Volga ancestry moved west and acquired Ukraine Neolithic hunter-gatherer (UNHG) ancestry to establish the population of the Serednii Stih culture from which the direct ancestors of the Yamnaya themselves were formed around 4000BCE. This population grew rapidly after 3750-3350BCE, precipitating the expansion of people of the Yamnaya culture who totally displaced previous groups on the Volga and further east, while admixing with more sedentary groups in the west. CLV cline people with Lower Volga ancestry contributed four fifths of the ancestry of the Yamnaya, but also, entering Anatolia from the east, contributed at least a tenth of the ancestry of Bronze Age Central Anatolians, where the Hittite language, related to the Indo-European languages spread by the Yamnaya, was spoken. We thus propose that the final unity of the speakers of the “Proto-Indo-Anatolian” ancestral language of both Anatolian and Indo-European languages can be traced to CLV cline people sometime between 4400-4000 BCE.  Summary Figure:The origin of Indo-Anatolian and Indo-European languages. Genetic reconstruction of the ancestry of Pontic-Caspian steppe and West Asian populations points to the North Caucasus-Lower Volga area as the homeland of Indo-Anatolian languages and to the Serednii Stih archaeological culture of the Dnipro-Don area as the homeland of Indo-European languages. The Caucasus-Lower Volga people had diverse distal roots, estimated using the qpAdm software on the left barplot, as Caucasus hunter-gatherer (purple), Central Asian (red), Eastern hunter-gatherer (pink), and West Asian Neolithic (green). Caucasus-Lower Volga expansions, estimated using qpAdm on the right barplot as disseminated Caucasus Neolithic (blue)-Lower Volga Eneolithic (orange) proximal ancestries, mixing with the inhabitants of the North Pontic region (yellow), Volga region (yellow), and West Asia (green). www.biorxiv.org/content/10.1101/2024.04.17.589597v1.full |
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Post by Admin on Apr 26, 2024 2:55:31 GMT
Introduction
Between 3300-1500 BCE, people of the Yamnaya archaeological complex and their descendants, in subsequent waves of migration, spread over large parts of Eurasia, contributing to the ancestry of people of Europe, Central and South Asia, Siberia, and the Caucasus. The spread of Indo-European language and culture1–7 transformed all these regions. Despite the centrality of the Yamnaya expansion to the human story of Bronze Age Eurasia, their ancestral origins are poorly understood. A first challenge has been the sparse sampling of the Yamnaya themselves across their enormous geographic distribution. The remarkable long-range mobility of the Yamnaya, quickly spreading over a vast region, adds further difficulties to tracing, from radiocarbon dating, the origins of their material culture and associated genetic profile. Nor can these origins be traced to the numerous earlier Eneolithic cultures that preceded the Yamnaya, and among whom their ancestors must be sought, as these have been sampled even more poorly and unsystematically.
The first formal study of the origins of the Yamnaya identified two disparate sources of ancestry: a northern, “Eastern Hunter-Gatherer” (EHG) source from far eastern Europe, and a southern, West Asian source related to present-day Armenians.2 The latter source was revealed, by ancient DNA, to be related to some of the region’s earliest inhabitants: Paleolithic-Mesolithic “Caucasus Hunter-Gatherers” (CHG) of Georgia,8 and Neolithic people of the Zagros9 and South Caucasus.6,10,11 Additional discoveries further complicated the stories of both the northern and southern ancestors of the Yamnaya. First, it was noted that both CHG and EHG were part of an interaction sphere across the boundary between West Asia and eastern Europe,9 suggesting the existence of intermediate populations and raising the question of when and where these came together to form the Eneolithic antecessors of the Yamnaya. Second, it was recognized that the steppe itself was an admixture zone of EHG with “Western Hunter-Gatherers” (WHG12).
Mesolithic hunter-gatherers from Ukraine were succeeded by more WHG-admixed Neolithic hunter-gatherers in the Dnipro valley,13 representing a local reshuffling within the European portion of a ∼7,000km-long trans-Eurasian cline of boreal hunter-gatherers.14 What was the relative contribution of the EHG (who were present in the Volga River at Lebyazhinka2 ca. 5660-5535 BCE) and these more western Ukraine Neolithic hunter-gatherers (UNHG) of the Dnipro to later populations? Third, it was discovered that the Yamnaya had not only CHG-related, but also Anatolian Neolithic ancestry, absent in the early known steppe inhabitants, and derived from European farmer neighbors west of the steppe5. This ancestry was later shown to be of rather Anatolian-Levantine-Mesopotamian origin, and to be mediated not from Europe but from the Caucasus neighbors south of the steppe.6 Such ancestry must have been added following the expansion of Neolithic farmers into the Caucasus, introduced thence into the steppe as a later exogenous element, distinct from the earlier CHG-related one. Finally, it was recognized that European steppe populations were formed not only by northern-southern admixture, but included, in at least some Eneolithic and Bronze Age people of the North Caucasus, contributions related to Siberians from further east.5 What was the extent of the spread of this eastern ancestry and did the Yamnaya themselves possess it?
Here we present a unified population genetic analysis of 372 newly reported individuals dating from 6400-2000 BCE, as well as increased quality data for 61 individuals. The present study serves as the formal technical report for 299 of the newly reported individuals and 55 of the individuals with increased quality data; more than 80% of the individuals are from Russia, but the dataset is also significant in including dozens of individuals from westward expansion of Steppe cultures along the Danube (Supplementary Information, section 1, Online Table 1). Technical details of the 803 ancient DNA libraries that are the basis for the newly reported data (and an additional 195 libraries that failed our screening) are presented in Online Table 2, while details of 198 newly generated radiocarbon dates on these individuals are presented in Online Table 3. A parallel study15 presents a combined archaeological and genetic analysis of population transformations in the North Pontic Region (Ukraine and Moldova) and serves as the formal report for the data from the other 73 of the newly analyzed individuals and the other 5 individuals with increased quality data, with both studies co-analyzing the full dataset. We grouped individuals into analysis labels based on geographical and temporal information, archaeological context, and genetic clustering (Online Table 4 lists all individuals used for analyses, with their labels). The potential of the combined dataset for shedding light on this period can be appreciated from the fact that it adds 79 analyzed Eneolithic people from the steppe and its environs (from Russia or Ukraine, west of 60E longitude and south of 60N latitude, between 5000-3500BCE) to 82 published5,7,13,15–20 and a total of 286 Yamnaya/Afanasievo individuals compared to 75 in the literature.2,4–6,13,21–29
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