The transitions from foraging to farming and later to pastoralism in Stone Age Eurasia (c. 11-3 thousand years before present, BP) represent some of the most dramatic lifestyle changes in human evolution. We sequenced 317 genomes of primarily Mesolithic and Neolithic individuals from across Eurasia combined with radiocarbon dates, stable isotope data, and pollen records. Genome imputation and co-analysis with previously published shotgun sequencing data resulted in >1600 complete ancient genome sequences offering fine-grained resolution into the Stone Age populations. We observe that: 1) Hunter-gatherer groups were more genetically diverse than previously known, and deeply divergent between western and eastern Eurasia. 2) We identify hitherto genetically undescribed hunter-gatherers from the Middle Don region that contributed ancestry to the later Yamnaya steppe pastoralists; 3) The genetic impact of the Neolithic transition was highly distinct, east and west of a boundary zone extending from the Black Sea to the Baltic. Large-scale shifts in genetic ancestry occurred to the west of this "Great Divide", including an almost complete replacement of hunter-gatherers in Denmark, while no substantial ancestry shifts took place during the same period to the east. This difference is also reflected in genetic relatedness within the populations, decreasing substantially in the west but not in the east where it remained high until c. 4,000 BP; 4) The second major genetic transformation around 5,000 BP happened at a much faster pace with Steppe-related ancestry reaching most parts of Europe within 1,000-years. Local Neolithic farmers admixed with incoming pastoralists in eastern, western, and southern Europe whereas Scandinavia experienced another near-complete population replacement. Similar dramatic turnover-patterns are evident in western Siberia; 5) Extensive regional differences in the ancestry components involved in these early events remain visible to this day, even within countries. Neolithic farmer ancestry is highest in southern and eastern England while Steppe-related ancestry is highest in the Celtic populations of Scotland, Wales, and Cornwall (this research has been conducted using the UK Biobank resource); 6) Shifts in diet, lifestyle and environment introduced new selection pressures involving at least 21 genomic regions. Most such variants were not universally selected across populations but were only advantageous in particular ancestral backgrounds. Contrary to previous claims, we find that selection on the FADS regions, associated with fatty acid metabolism, began before the Neolithisation of Europe. Similarly, the lactase persistence allele started increasing in frequency before the expansion of Steppe-related groups into Europe and has continued to increase up to the present. Along the genetic cline separating Mesolithic hunter-gatherers from Neolithic farmers, we find significant correlations with trait associations related to skin disorders, diet and lifestyle and mental health status, suggesting marked phenotypic differences between these groups with very different lifestyles. This work provides new insights into major transformations in recent human evolution, elucidating the complex interplay between selection and admixture that shaped patterns of genetic variation in modern populations.
The inland area of southwestern Sweden is well known for its well-preserved archaeological animal and human remains dating back to the Mesolithic and Neolithic (10000–4000 and 4000–1700 BC). They allow application of multiple bioarchaeological methods, giving insights into various and complementary aspects of prehistoric human life, as well as economic and social structures. One important aspect concerns human mobility and its relation to social networks and to circulation of objects. Here, strontium isotope analysis plays a crucial role. The present study aims to construct a strontium isotope baseline of southwestern Sweden with considerably greater coverage and higher resolution than previously published data. As the region has been affected by glacial events, the relation between bedrock geology and isotope signals of the bioavailable strontium in such areas is given special attention. We determined strontium isotope ratios for 61 water and five archaeological animal samples, and combined the data with previous measurements of two water and 21 non-domestic faunal samples. The results reveal a complex pattern. Several areas with distinct baseline ranges can be distinguished, although with overlaps between some of them. Overall, the bioavailable strontium isotope signals mirror the basement geology of the region. The highest ratios occur in the geologically oldest eastern parts of the Precambrian terrain, while lower ratios are found in the western part, and the lowest ratios occur in the youngest Paleozoic areas. At the same time, there are minor deviations compared to the underlying bedrock, due to glacial transport, overlying sediments, and local intrusions of younger rocks. The background data set now available allows for more nuanced and detailed interpretations of human and animal mobility in the region, in particular by identification of subregions with differing strontium isotope ratios within the Precambrian province. Also, we can now identify long distance mobility with greater confidence.
This study investigates possible variation in post-mortem histories during the Neolithic period in southwestern Sweden based on microscopic studies of human bone. Numerous megalithic graves were built in this region and good preservation conditions have left a rich skeletal record. After more than a hundred years of research, it is still a controversy whether or not these skeletal assemblages were the result of primary burials, or ossuaries where skeletonized remains were deposited. In this study we apply histological analysis to obtain insights into post-mortem histories and taphonomic processes affecting the human remains, potentially including funerary rituals. This type of analysis records the condition and traces of degradation found in skeletal material at a microscopic level. Human skeletal material from four different megalithic tombs in the Falbygden area has been sampled and analysed by thin-section light microscopy, and by scanning electron microscopy. The results of the study provide evidence of variation and changes in burial conditions for skeletal remains from the different graves, also for remains from the same grave. Extent of bioerosion varied, from extensive to moderate/arrested, to none. Bone samples from the same graves also differed in the type of staining and mineral inclusions, showing that the non-bioeroded samples relatively early post-mortem must have experienced an anoxic environment, and later a change to an aerated environment. This could be taken as an indication of primary burial somewhere else, but more likely reflect a special micro-environment occurring temporarily in some graves and parts of graves after the tombs were filled with soil and sealed by roof slabs. The study illustrates the usefulness of bone histological analysis in the reconstruction of post-mortem histories, revealing variations not discernible at macro-level that may aid in the interpretations of funerary rituals. However, the results also highlight the issues of equifinality. Based on current data and knowledge, several scenarios are possible. Further histotaphonomic work is advisable, including archaeological remains from megalithic tombs, and bones from taphonomic experiments.
Megalithic tombs have since long been a focus of debate within the archaeological research field, not least regarding their emergence, use life and the various bursts of building activity in different regions and periods. The aim of this study is to investigate the temporal span of the main burial sequences in the conventional megalithic grave types of southern Sweden, with special focus on the less studied gallery graves. In Scandinavia, megalithic tombs are divided into three main types: dolmens, passage graves and gallery graves. Here, this prevailing typological seriation was tested. The study was based on 374 14C dates from unique individuals selected from 66 tombs. The form, layout and dating of the different types of tombs were studied in order to examine regional and chronological variation in the use of megaliths. By comparing sum plots, KDE models, individual 14C dates and typology of artefacts, the existing chronologies were evaluated. The 14C dates from dolmens and passage graves more or less agreed with the conventional chronology, while the presence of early skeletons in gallery graves was unexpected. The results indicate that megalithic graves appeared more or less simultaneously in southern Sweden and were first used around 3500-3300 cal BC. The dolmens and passage graves were used contemporaneously, although the proportion of early dates supports a slightly earlier start of the dolmens. Some of the gallery graves may also have been introduced at this time, although reburial of old bones cannot be ruled out.
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