Karl-Göran Sjögren | University of Gothenburg (original) (raw)

Books by Karl-Göran Sjögren

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Papers by Karl-Göran Sjögren

Nature, 2024

In the period between 5,300 and 4,900 calibrated years before present (cal. bp), populations acro... more In the period between 5,300 and 4,900 calibrated years before present (cal. bp), populations across large parts of Europe underwent a period of demographic decline. However, the cause of this so-called Neolithic decline is still debated. Some argue for an agricultural crisis resulting in the decline, others for the spread of an early form of plague. Here we use population-scale ancient genomics to infer ancestry, social
structure and pathogen infection in 108 Scandinavian Neolithic individuals from eight megalithic graves and a stone cist. We find that the Neolithic plague was widespread, detected in at least 17% of the sampled population and across large geographical distances. We demonstrate that the disease spread within the Neolithic community in three distinct infection events within a period of around 120 years. Variant graph-based pan-genomics shows that the Neolithic plague genomes retained ancestral genomic
variation present in Yersinia pseudotuberculosis, including virulence factors associated with disease outcomes. In addition, we reconstruct four multigeneration pedigrees, the largest of which consists of 38 individuals spanning six generations, showing a patrilineal social organization. Lastly, we document direct genomic evidence for Neolithic female exogamy in a woman buried in a different megalithic tomb than her brothers. Taken together, our findings provide a detailed reconstruction of plague
spread within a large patrilineal kinship group and identify multiple plague infections in a population dated to the beginning of the Neolithic decline.

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Nature, 2024

n the period between 5,300 and 4,900 calibrated years before present (cal. bp), populations acros... more n the period between 5,300 and 4,900 calibrated years before present (cal. bp), populations across large parts of Europe underwent a period of demographic decline1,2. However, the cause of this so-called Neolithic decline is still debated. Some argue
for an agricultural crisis resulting in the decline3, others for the spread of an early form of plague4. Here we use population-scale ancient genomics to infer ancestry, social structure and pathogen infection in 108 Scandinavian Neolithic individuals from eight megalithic graves and a stone cist. We find that the Neolithic plague was widespread, detected in at least 17% of the sampled population and across large geographical distances. We demonstrate that the disease spread within the Neolithic community in three distinct infection events within a period of around 120 years. Variant graph-based pan-genomics shows that the Neolithic plague genomes retained ancestral genomic variation present in Yersinia pseudotuberculosis, including virulence factors associated with disease outcomes. In addition, we reconstruct four multigeneration pedigrees, the largest of which consists of 38 individuals spanning six generations, showing a patrilineal social organization. Lastly, we document direct genomic evidence for Neolithic female exogamy in a woman buried in a different megalithic tomb than her brothers. Taken together, our findings provide a detailed reconstruction of plague spread within a large patrilineal kinship group and identify multiple plague infections in a population dated to the beginning of the Neolithic decline.

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Plos One, 2024

The lethally maltreated body of Vittrup Man was deposited in a Danish bog, probably as part of a ... more The lethally maltreated body of Vittrup Man was deposited in a Danish bog, probably as part of a ritualised sacrifice. It happened between c. 3300 and 3100 cal years BC, i.e., during the period of the local farming-based Funnel Beaker Culture. In terms of skull morphological features, he differs from the majority of the contemporaneous farmers found in Denmark, and associates with hunter-gatherers, who inhabited Scandinavia during the previous millennia.
His skeletal remains were selected for transdisciplinary analysis to reveal his life-history in terms of a population historical perspective. We report the combined results of an integrated set of genetic, isotopic, physical anthropological and archaeological analytical approaches. Strontium signature suggests a foreign birthplace that could be in Norway or Sweden. In addition, enamel oxygen isotope values indicate that as a child he lived in a colder climate, i.e., to the north of the regions inhabited by farmers. Genomic data in fact demonstrates that he is closely related to Mesolithic humans known from Norway and Sweden. Moreover, dietary stable isotope analyses on enamel and bone collagen demonstrate a fisher-hunter way of life in his childhood and a diet typical of farmers later on. Such a variable life-history is also reflected by proteomic analysis of hardened organic deposits on his teeth, indicating the consumption of forager food (seal, whale and marine fish) as well as farmer food (sheep/goat). From a dietary isotopic transect of one of his teeth it is shown that his transfer between societies of foragers and farmers took place near to the end of his teenage years.

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Journal of Neolithic Archaeology, 2023

The excavation of the simple dolmen at Tiarp, Falbygden, dating to around 3500 BCE, has provided ... more The excavation of the simple dolmen at Tiarp, Falbygden, dating to around 3500 BCE, has provided important information for the understanding of the megalithic and early TRB in southern Scandinavia and northern Central Europe. The absolute chronological dating shows that dolmens were erected at about the same time between Falbygden and Altmark, before the main passage grave architectural phase. Although fragmented and affected by taphonomic processes, the bone assemblage provides insights into the burial practices. At least twelve individuals, from neonates to elderly, had been buried within the chamber. The predominance of hand and foot bones suggests that they were primary inhumations. Their isotope values indicate an already agrarian society, which, however, was based only to a certain extent on agriculture. Insofar, the dolmen at Tiarp signals transformations-not only in respect to the introduction of agriculture but also regarding ritual practices-within the Early and Middle Neolithic.

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The Indo-European Puzzle Revisited

This book examines the impact of ancient DNA research and scientific evidence on our understandin... more This book examines the impact of ancient DNA research and scientific evidence on our understanding of the emergence of Indo-European languages in prehistory. Offering cutting-edge contributions from an international team of scholars, it considers the driving forces behind the Indo-European migrations during the 3rd and 2nd millenia BC. The volume explores the rise of the world's first pastoral nomads the Yamnaya Culture in the Russian Pontic steppe including their social organization, expansions, and the transition from nomadism to semi-sedentism when entering Europe. It also traces the chariot conquest in the late Bronze Age and its impact on the expansion of the Indo-Iranian languages into Central Asia. In the final section, the volumes consider the development of hierarchical societies and the origins of slavery. A landmark synthesis of recent, exciting discoveries, the book also includes an extensive theoretical discussion regarding the integration of linguistics, genetics, ...

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Archaeological and Anthropological Sciences, 2023

In northern Europe, the first indications of a Neolithic lifestyle appear around 4000 cal BC from... more In northern Europe, the first indications of a Neolithic lifestyle appear around 4000 cal BC from northern Germany up to middle Sweden and southeastern Norway in an apparently short period of time, largely carried by immigrant populations bringing new species of plants and animals into the region. However, the nature of this domestication "package" is not everywhere the same, whereby both environmental and cultural filters acted on the particular set of species cultivated and bred in different regions. In Neolithic Scandinavia, cattle, pigs, and caprines (sheep and goat) are all present in varying proportions, with cattle more prominent in Denmark and pigs more prominent in more northerly areas. However, little is known about the ratio of sheep to goat remains within this region, largely due to difficulties in morphologically separating the two species. In this paper, we report the results from ZooMS analysis of 45 sheep/goat bone samples from two recently excavated Funnel Beaker settlements in Karleby, Falbygden, Sweden. The ZooMS analyses gave a clear and somewhat surprising result: 33 of the samples were classed as sheep, one as deer, and none of them as goat. In all likelihood, goats have not been present at all on these sites. A survey of the literature shows that while small numbers of goats are likely present in Denmark from the Early Neolithic, their presence in Sweden at this time is ambiguous and the few claims merit reassessment. Furthermore, the low numbers in Scandinavia compared to central and southern Europe suggests an overall geographic trend, with decreasing proportions of caprines as well as goats in the north.

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Journal of Neolithic Archaeology, 2023

The thousands of dolmens and long barrows spread across the Danish landscape are the earliest lon... more The thousands of dolmens and long barrows spread across the Danish landscape are the earliest long-lasting expressions of architectural monumentality in Scandinavia. A series of new AMS dates on human skeletal material from several of them leads to a clarification of the generations-long debate on the relative chronology and typological evolution of this group of monu ments. Earthen long barrows were raised from ca. 3700 cal BC. That is at least two centuries later than the arrival of such elements of the Neolithic world as funnel beaker pottery and domestic cattle to the region. The practice of using large stones (megaliths) for burial chambers was present by 3600 cal BC. Classical Urdolmen were built alongside various types of more complex dolmen chambers during the period ca. 3600-3400 cal BC, after which passage graves were erected.

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Megaliths of the world, 2022

In southern Scandinavia and the northern central European lowlands, megalithic graves were erecte... more In southern Scandinavia and the northern central European lowlands, megalithic graves were erected mainly between 3600 and 3100 BCE. These collective tombs shape the cultural landscape of the so-called older and middle Funnel Beaker (TRB) societies. At this time, a ‘megalithic boom’ occurred with the introduction of new agricultural techniques such
as ard ploughing, animal traction, manuring, and land clearance. Recent research projects have considerably increased our knowledge of the builders of the megalithic sites. In Falbygden, Sweden, and in Holstein, northern Germany, projects have informed us about the environmental conditions, economic practices, and burial customs of the builders of the
Nordic megalithic tombs. On the Cimbrian Peninsula, excavations have helped to clarify the relationship between causewayed enclosures and megaliths. It now seems that the megalithic boom was connected both to agricultural innovations and to a peak in ceremonial activities in general.

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Journal of Anthropological archaeology, 2021

The Pitted Ware Culture emerged during the Neolithic around 3400 BCE in east-central Sweden and q... more The Pitted Ware Culture emerged during the Neolithic around 3400 BCE in east-central Sweden and quickly spread south and west to various parts of southern Scandinavia. For the next millennium these groups exhibited less interest in agricultural activity and a general return to the hunting and gathering of wild animals and plants with a focus on seals in some coastal areas. Pitted Ware Culture arrived in northeastern Denmark after 3100 BCE. One of the most important sites of this period was at Kainsbakke, on the peninsula of Djursland. One deep pit at the site contained unusual numbers of bear, red deer, European elk, and boar remains. This study focuses on the isotopic proveniencing of some of these animal remains to evaluate their origins. Confirmation of the import of some of the animals, probably from western Sweden, suggests possible shared ritual activity at the central Kainsbakke site. This evidence also confirms the navigating skills of Neolithic peoples in northern Europe.

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Archaeological and Anthropological Sciences, 2021

In this paper, we investigate population dynamics in the Scandinavian Neolithic and Early Bronze ... more In this paper, we investigate population dynamics in the Scandinavian Neolithic and Early Bronze Age in southwestern Sweden. Human mobility patterns in Falbygden were studied by applying strontium isotope analysis combined with archaeological and bioarchaeological data, including mtDNA and sex assessment on a large dataset encompassing 141 individuals from 21 megalithic graves. In combination with other archaeological and anthropological records, we investigated the temporal and spatial scale of individual movement, mobility patterns of specific categories of people and possible social drivers behind them. Our results of strontium and biomolecular analyses suggest that mobility increased in the Late Neolithic and Early Bronze Age compared to the earlier parts of the Neolithic. The data indicate individuals moving both into and away from Falbygden. Mobility patterns and contact networks also shift over time.

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Population genomics of Stone Age Eurasia, 2022

The transitions from foraging to farming and later to pastoralism in Stone Age Eurasia (c. 113 th... more The transitions from foraging to farming and later to pastoralism in Stone Age Eurasia (c. 113 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 1,000years. 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.

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Animals on the Move

In this contribution, we highlight some of the results from analyses of cattle management at two ... more In this contribution, we highlight some of the results from analyses of cattle management at two Neolithic settlement sites in Falbygden, western Sweden. The two sites are located close to each other in the parish of Karleby in central Falbygden. Both were excavated 2012-2017 and yielded abundant animal bones, due to the calcareous bedrock. 14C datings indicate that they were both inhabited continuously during the period c. 3200-2900 BC, i.e. the Middle Neolithic Funnel Beaker (TRB) culture. Bulk and laser ablation strontium isotope ratios in cattle teeth from these settlements suggest a complex and wide-ranging system of animal exchange in the Scandinavian Middle Neolithic. More than half of the cattle found in the Karleby settlements were born outside the Falbygden region, and laser ablation suggests variable and individual trajectories through the landscape, in some cases involving areas in eastern Sweden. Further, origins of cattle at the two settlements are only partly overlapping, suggesting independent contact networks. We suggest an ongoing system of exchanges between settlement groups/social groups throughout Sweden to be behind this variability, rather than transhumance. This may in turn be motivated by a number of factors including biological herd viability, as well as social and economic factors.

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PlosOne, 2021

The Gjerrild burial provides the largest and best-preserved assemblage of human skeletal material... more The Gjerrild burial provides the largest and best-preserved assemblage of human skeletal material presently known from the Single Grave Culture (SGC) in Denmark. For generations it has been debated among archaeologists if the appearance of this archaeological complex represents a continuation of the previous Neolithic communities, or was facilitated by incoming migrants. We sampled and analysed five skeletons from the Gjerrild cist, buried over a period of c. 300 years, 2600/2500-2200 cal BCE. Despite poor DNA preservation, we managed to sequence the genome (>1X) of one individual and the partial genomes (0.007X and 0.02X) of another two individuals. Our genetic data document a female (Gjerrild 1) and two males (Gjerrild 5 + 8), harbouring typical Neolithic K2a and HV0 mtDNA haplogroups, but also a rare basal variant of the R1b1 Y-chromosomal haplogroup. Genome-wide analyses demonstrate that these people had a significant Yamnaya-derived (i.e. steppe) ancestry component and a close genetic resemblance to the Corded Ware (and related) groups that were present in large parts of Northern and Central Europe at the time. Assuming that the Gjerrild skeletons are genetically representative of the population of the SGC in broader terms, the transition from the local Neolithic Funnel Beaker Culture (TRB) to SGC is not characterized by demographic continuity. Rather, the emergence of SGC in Denmark was part of the Late Neolithic and Early Bronze Age population expansion that swept across the European continent in the 3rd millennium BCE, resulting in various degrees of genetic replacement and admixture processes with previous Neolithic populations. PLOS ONE PLOS ONE | https://doi.org/10.1371/journal.pone.

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PlosOne, 2020

We present a high-resolution cross-disciplinary analysis of kinship structure and social institut... more We present a high-resolution cross-disciplinary analysis of kinship structure and social institutions in two Late Copper Age Bell Beaker culture cemeteries of South Germany containing 24 and 18 burials, of which 34 provided genetic information. By combining archaeological, anthropological, genetic and isotopic evidence we are able to document the internal kinship and residency structure of the cemeteries and the socially organizing principles of these local communities. The buried individuals represent four to six generations of two family groups, one nuclear family at the Alburg cemetery, and one seemingly more extended at Irl-bach. While likely monogamous, they practiced exogamy, as six out of eight non-locals are women. Maternal genetic diversity is high with 23 different mitochondrial haplotypes from 34 individuals, whereas all males belong to one single Y-chromosome haplogroup without any detectable contribution from Y-chromosomes typical of the farmers who had been the sole inhabitants of the region hundreds of years before. This provides evidence for the society being patrilocal, perhaps as a way of protecting property among the male line, while in-marriage from many different places secured social and political networks and prevented inbreeding. We also find evidence that the communities practiced selection for which of their PLOS ONE PLOS ONE | https://doi.org/10.1371/journal.pone.

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Science, 2020

Dogs were the first domestic animal, but little is known about their population history and to wh... more Dogs were the first domestic animal, but little is known about their population history and to what extent it was linked to humans. We sequenced 27 ancient dog genomes and found that all dogs share a common ancestry distinct from present-day wolves, with limited gene flow from wolves since domestication but substantial dog-to-wolf gene flow. By 11,000 years ago, at least five major ancestry lineages had diversified, demonstrating a deep genetic history of dogs during the Paleolithic. Coanalysis with human genomes reveals aspects of dog population history that mirror humans, including Levant-related ancestry in Africa and early agricultural Europe. Other aspects differ, including the impacts of steppe pastoralist expansions in West and East Eurasia and a near-complete turnover of Neolithic European dog ancestry.

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Rui Boaventura, Rui Mataloto and André Pereira (eds): Megaliths and Geology: Megálitos e Geologia. MEGA-TALKS 2, 19-20 November 2015 (Redondo, Portugal), 2020

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Nature, 2024

In the period between 5,300 and 4,900 calibrated years before present (cal. bp), populations acro... more In the period between 5,300 and 4,900 calibrated years before present (cal. bp), populations across large parts of Europe underwent a period of demographic decline. However, the cause of this so-called Neolithic decline is still debated. Some argue for an agricultural crisis resulting in the decline, others for the spread of an early form of plague. Here we use population-scale ancient genomics to infer ancestry, social
structure and pathogen infection in 108 Scandinavian Neolithic individuals from eight megalithic graves and a stone cist. We find that the Neolithic plague was widespread, detected in at least 17% of the sampled population and across large geographical distances. We demonstrate that the disease spread within the Neolithic community in three distinct infection events within a period of around 120 years. Variant graph-based pan-genomics shows that the Neolithic plague genomes retained ancestral genomic
variation present in Yersinia pseudotuberculosis, including virulence factors associated with disease outcomes. In addition, we reconstruct four multigeneration pedigrees, the largest of which consists of 38 individuals spanning six generations, showing a patrilineal social organization. Lastly, we document direct genomic evidence for Neolithic female exogamy in a woman buried in a different megalithic tomb than her brothers. Taken together, our findings provide a detailed reconstruction of plague
spread within a large patrilineal kinship group and identify multiple plague infections in a population dated to the beginning of the Neolithic decline.

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Nature, 2024

n the period between 5,300 and 4,900 calibrated years before present (cal. bp), populations acros... more n the period between 5,300 and 4,900 calibrated years before present (cal. bp), populations across large parts of Europe underwent a period of demographic decline1,2. However, the cause of this so-called Neolithic decline is still debated. Some argue
for an agricultural crisis resulting in the decline3, others for the spread of an early form of plague4. Here we use population-scale ancient genomics to infer ancestry, social structure and pathogen infection in 108 Scandinavian Neolithic individuals from eight megalithic graves and a stone cist. We find that the Neolithic plague was widespread, detected in at least 17% of the sampled population and across large geographical distances. We demonstrate that the disease spread within the Neolithic community in three distinct infection events within a period of around 120 years. Variant graph-based pan-genomics shows that the Neolithic plague genomes retained ancestral genomic variation present in Yersinia pseudotuberculosis, including virulence factors associated with disease outcomes. In addition, we reconstruct four multigeneration pedigrees, the largest of which consists of 38 individuals spanning six generations, showing a patrilineal social organization. Lastly, we document direct genomic evidence for Neolithic female exogamy in a woman buried in a different megalithic tomb than her brothers. Taken together, our findings provide a detailed reconstruction of plague spread within a large patrilineal kinship group and identify multiple plague infections in a population dated to the beginning of the Neolithic decline.

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Plos One, 2024

The lethally maltreated body of Vittrup Man was deposited in a Danish bog, probably as part of a ... more The lethally maltreated body of Vittrup Man was deposited in a Danish bog, probably as part of a ritualised sacrifice. It happened between c. 3300 and 3100 cal years BC, i.e., during the period of the local farming-based Funnel Beaker Culture. In terms of skull morphological features, he differs from the majority of the contemporaneous farmers found in Denmark, and associates with hunter-gatherers, who inhabited Scandinavia during the previous millennia.
His skeletal remains were selected for transdisciplinary analysis to reveal his life-history in terms of a population historical perspective. We report the combined results of an integrated set of genetic, isotopic, physical anthropological and archaeological analytical approaches. Strontium signature suggests a foreign birthplace that could be in Norway or Sweden. In addition, enamel oxygen isotope values indicate that as a child he lived in a colder climate, i.e., to the north of the regions inhabited by farmers. Genomic data in fact demonstrates that he is closely related to Mesolithic humans known from Norway and Sweden. Moreover, dietary stable isotope analyses on enamel and bone collagen demonstrate a fisher-hunter way of life in his childhood and a diet typical of farmers later on. Such a variable life-history is also reflected by proteomic analysis of hardened organic deposits on his teeth, indicating the consumption of forager food (seal, whale and marine fish) as well as farmer food (sheep/goat). From a dietary isotopic transect of one of his teeth it is shown that his transfer between societies of foragers and farmers took place near to the end of his teenage years.

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Journal of Neolithic Archaeology, 2023

The excavation of the simple dolmen at Tiarp, Falbygden, dating to around 3500 BCE, has provided ... more The excavation of the simple dolmen at Tiarp, Falbygden, dating to around 3500 BCE, has provided important information for the understanding of the megalithic and early TRB in southern Scandinavia and northern Central Europe. The absolute chronological dating shows that dolmens were erected at about the same time between Falbygden and Altmark, before the main passage grave architectural phase. Although fragmented and affected by taphonomic processes, the bone assemblage provides insights into the burial practices. At least twelve individuals, from neonates to elderly, had been buried within the chamber. The predominance of hand and foot bones suggests that they were primary inhumations. Their isotope values indicate an already agrarian society, which, however, was based only to a certain extent on agriculture. Insofar, the dolmen at Tiarp signals transformations-not only in respect to the introduction of agriculture but also regarding ritual practices-within the Early and Middle Neolithic.

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The Indo-European Puzzle Revisited

This book examines the impact of ancient DNA research and scientific evidence on our understandin... more This book examines the impact of ancient DNA research and scientific evidence on our understanding of the emergence of Indo-European languages in prehistory. Offering cutting-edge contributions from an international team of scholars, it considers the driving forces behind the Indo-European migrations during the 3rd and 2nd millenia BC. The volume explores the rise of the world's first pastoral nomads the Yamnaya Culture in the Russian Pontic steppe including their social organization, expansions, and the transition from nomadism to semi-sedentism when entering Europe. It also traces the chariot conquest in the late Bronze Age and its impact on the expansion of the Indo-Iranian languages into Central Asia. In the final section, the volumes consider the development of hierarchical societies and the origins of slavery. A landmark synthesis of recent, exciting discoveries, the book also includes an extensive theoretical discussion regarding the integration of linguistics, genetics, ...

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Archaeological and Anthropological Sciences, 2023

In northern Europe, the first indications of a Neolithic lifestyle appear around 4000 cal BC from... more In northern Europe, the first indications of a Neolithic lifestyle appear around 4000 cal BC from northern Germany up to middle Sweden and southeastern Norway in an apparently short period of time, largely carried by immigrant populations bringing new species of plants and animals into the region. However, the nature of this domestication "package" is not everywhere the same, whereby both environmental and cultural filters acted on the particular set of species cultivated and bred in different regions. In Neolithic Scandinavia, cattle, pigs, and caprines (sheep and goat) are all present in varying proportions, with cattle more prominent in Denmark and pigs more prominent in more northerly areas. However, little is known about the ratio of sheep to goat remains within this region, largely due to difficulties in morphologically separating the two species. In this paper, we report the results from ZooMS analysis of 45 sheep/goat bone samples from two recently excavated Funnel Beaker settlements in Karleby, Falbygden, Sweden. The ZooMS analyses gave a clear and somewhat surprising result: 33 of the samples were classed as sheep, one as deer, and none of them as goat. In all likelihood, goats have not been present at all on these sites. A survey of the literature shows that while small numbers of goats are likely present in Denmark from the Early Neolithic, their presence in Sweden at this time is ambiguous and the few claims merit reassessment. Furthermore, the low numbers in Scandinavia compared to central and southern Europe suggests an overall geographic trend, with decreasing proportions of caprines as well as goats in the north.

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Journal of Neolithic Archaeology, 2023

The thousands of dolmens and long barrows spread across the Danish landscape are the earliest lon... more The thousands of dolmens and long barrows spread across the Danish landscape are the earliest long-lasting expressions of architectural monumentality in Scandinavia. A series of new AMS dates on human skeletal material from several of them leads to a clarification of the generations-long debate on the relative chronology and typological evolution of this group of monu ments. Earthen long barrows were raised from ca. 3700 cal BC. That is at least two centuries later than the arrival of such elements of the Neolithic world as funnel beaker pottery and domestic cattle to the region. The practice of using large stones (megaliths) for burial chambers was present by 3600 cal BC. Classical Urdolmen were built alongside various types of more complex dolmen chambers during the period ca. 3600-3400 cal BC, after which passage graves were erected.

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Megaliths of the world, 2022

In southern Scandinavia and the northern central European lowlands, megalithic graves were erecte... more In southern Scandinavia and the northern central European lowlands, megalithic graves were erected mainly between 3600 and 3100 BCE. These collective tombs shape the cultural landscape of the so-called older and middle Funnel Beaker (TRB) societies. At this time, a ‘megalithic boom’ occurred with the introduction of new agricultural techniques such
as ard ploughing, animal traction, manuring, and land clearance. Recent research projects have considerably increased our knowledge of the builders of the megalithic sites. In Falbygden, Sweden, and in Holstein, northern Germany, projects have informed us about the environmental conditions, economic practices, and burial customs of the builders of the
Nordic megalithic tombs. On the Cimbrian Peninsula, excavations have helped to clarify the relationship between causewayed enclosures and megaliths. It now seems that the megalithic boom was connected both to agricultural innovations and to a peak in ceremonial activities in general.

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Journal of Anthropological archaeology, 2021

The Pitted Ware Culture emerged during the Neolithic around 3400 BCE in east-central Sweden and q... more The Pitted Ware Culture emerged during the Neolithic around 3400 BCE in east-central Sweden and quickly spread south and west to various parts of southern Scandinavia. For the next millennium these groups exhibited less interest in agricultural activity and a general return to the hunting and gathering of wild animals and plants with a focus on seals in some coastal areas. Pitted Ware Culture arrived in northeastern Denmark after 3100 BCE. One of the most important sites of this period was at Kainsbakke, on the peninsula of Djursland. One deep pit at the site contained unusual numbers of bear, red deer, European elk, and boar remains. This study focuses on the isotopic proveniencing of some of these animal remains to evaluate their origins. Confirmation of the import of some of the animals, probably from western Sweden, suggests possible shared ritual activity at the central Kainsbakke site. This evidence also confirms the navigating skills of Neolithic peoples in northern Europe.

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Archaeological and Anthropological Sciences, 2021

In this paper, we investigate population dynamics in the Scandinavian Neolithic and Early Bronze ... more In this paper, we investigate population dynamics in the Scandinavian Neolithic and Early Bronze Age in southwestern Sweden. Human mobility patterns in Falbygden were studied by applying strontium isotope analysis combined with archaeological and bioarchaeological data, including mtDNA and sex assessment on a large dataset encompassing 141 individuals from 21 megalithic graves. In combination with other archaeological and anthropological records, we investigated the temporal and spatial scale of individual movement, mobility patterns of specific categories of people and possible social drivers behind them. Our results of strontium and biomolecular analyses suggest that mobility increased in the Late Neolithic and Early Bronze Age compared to the earlier parts of the Neolithic. The data indicate individuals moving both into and away from Falbygden. Mobility patterns and contact networks also shift over time.

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Population genomics of Stone Age Eurasia, 2022

The transitions from foraging to farming and later to pastoralism in Stone Age Eurasia (c. 113 th... more The transitions from foraging to farming and later to pastoralism in Stone Age Eurasia (c. 113 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 1,000years. 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.

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Animals on the Move

In this contribution, we highlight some of the results from analyses of cattle management at two ... more In this contribution, we highlight some of the results from analyses of cattle management at two Neolithic settlement sites in Falbygden, western Sweden. The two sites are located close to each other in the parish of Karleby in central Falbygden. Both were excavated 2012-2017 and yielded abundant animal bones, due to the calcareous bedrock. 14C datings indicate that they were both inhabited continuously during the period c. 3200-2900 BC, i.e. the Middle Neolithic Funnel Beaker (TRB) culture. Bulk and laser ablation strontium isotope ratios in cattle teeth from these settlements suggest a complex and wide-ranging system of animal exchange in the Scandinavian Middle Neolithic. More than half of the cattle found in the Karleby settlements were born outside the Falbygden region, and laser ablation suggests variable and individual trajectories through the landscape, in some cases involving areas in eastern Sweden. Further, origins of cattle at the two settlements are only partly overlapping, suggesting independent contact networks. We suggest an ongoing system of exchanges between settlement groups/social groups throughout Sweden to be behind this variability, rather than transhumance. This may in turn be motivated by a number of factors including biological herd viability, as well as social and economic factors.

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PlosOne, 2021

The Gjerrild burial provides the largest and best-preserved assemblage of human skeletal material... more The Gjerrild burial provides the largest and best-preserved assemblage of human skeletal material presently known from the Single Grave Culture (SGC) in Denmark. For generations it has been debated among archaeologists if the appearance of this archaeological complex represents a continuation of the previous Neolithic communities, or was facilitated by incoming migrants. We sampled and analysed five skeletons from the Gjerrild cist, buried over a period of c. 300 years, 2600/2500-2200 cal BCE. Despite poor DNA preservation, we managed to sequence the genome (>1X) of one individual and the partial genomes (0.007X and 0.02X) of another two individuals. Our genetic data document a female (Gjerrild 1) and two males (Gjerrild 5 + 8), harbouring typical Neolithic K2a and HV0 mtDNA haplogroups, but also a rare basal variant of the R1b1 Y-chromosomal haplogroup. Genome-wide analyses demonstrate that these people had a significant Yamnaya-derived (i.e. steppe) ancestry component and a close genetic resemblance to the Corded Ware (and related) groups that were present in large parts of Northern and Central Europe at the time. Assuming that the Gjerrild skeletons are genetically representative of the population of the SGC in broader terms, the transition from the local Neolithic Funnel Beaker Culture (TRB) to SGC is not characterized by demographic continuity. Rather, the emergence of SGC in Denmark was part of the Late Neolithic and Early Bronze Age population expansion that swept across the European continent in the 3rd millennium BCE, resulting in various degrees of genetic replacement and admixture processes with previous Neolithic populations. PLOS ONE PLOS ONE | https://doi.org/10.1371/journal.pone.

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PlosOne, 2020

We present a high-resolution cross-disciplinary analysis of kinship structure and social institut... more We present a high-resolution cross-disciplinary analysis of kinship structure and social institutions in two Late Copper Age Bell Beaker culture cemeteries of South Germany containing 24 and 18 burials, of which 34 provided genetic information. By combining archaeological, anthropological, genetic and isotopic evidence we are able to document the internal kinship and residency structure of the cemeteries and the socially organizing principles of these local communities. The buried individuals represent four to six generations of two family groups, one nuclear family at the Alburg cemetery, and one seemingly more extended at Irl-bach. While likely monogamous, they practiced exogamy, as six out of eight non-locals are women. Maternal genetic diversity is high with 23 different mitochondrial haplotypes from 34 individuals, whereas all males belong to one single Y-chromosome haplogroup without any detectable contribution from Y-chromosomes typical of the farmers who had been the sole inhabitants of the region hundreds of years before. This provides evidence for the society being patrilocal, perhaps as a way of protecting property among the male line, while in-marriage from many different places secured social and political networks and prevented inbreeding. We also find evidence that the communities practiced selection for which of their PLOS ONE PLOS ONE | https://doi.org/10.1371/journal.pone.

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Science, 2020

Dogs were the first domestic animal, but little is known about their population history and to wh... more Dogs were the first domestic animal, but little is known about their population history and to what extent it was linked to humans. We sequenced 27 ancient dog genomes and found that all dogs share a common ancestry distinct from present-day wolves, with limited gene flow from wolves since domestication but substantial dog-to-wolf gene flow. By 11,000 years ago, at least five major ancestry lineages had diversified, demonstrating a deep genetic history of dogs during the Paleolithic. Coanalysis with human genomes reveals aspects of dog population history that mirror humans, including Levant-related ancestry in Africa and early agricultural Europe. Other aspects differ, including the impacts of steppe pastoralist expansions in West and East Eurasia and a near-complete turnover of Neolithic European dog ancestry.

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Rui Boaventura, Rui Mataloto and André Pereira (eds): Megaliths and Geology: Megálitos e Geologia. MEGA-TALKS 2, 19-20 November 2015 (Redondo, Portugal), 2020

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Proceedings of the National Academy of Sciences

The European continent was subject to two major migrations of peoples during the Holocene: the no... more The European continent was subject to two major migrations of peoples during the Holocene: the northwestward movement of Anatolian farmer populations during the Neolithic and the westward movement of Yamnaya steppe peoples during the Bronze Age. These movements changed the genetic composition of the continent's inhabitants. The Holocene was also characterized by major changes in vegetation composition, which altered the environment occupied by the original hunter-gatherer populations. We aim to test to what extent vegetation change through time is associated with changes in population composition as a consequence of these migrations, or with changes in climate. Using ancient DNA in combination with geostatisti-cal techniques, we produce detailed maps of ancient population movements, which allow us to visualize how these migrations unfolded through time and space. We find that the spread of Neolithic farmer ancestry had a two-pronged wavefront, in agreement with similar findings on the cultural spread of farming from radiocarbon-dated archaeological sites. This movement, however, did not have a strong association with changes in the vegetational landscape. In contrast, the Yamnaya migration speed was at least twice as fast and coincided with a reduction in the amount of broad-leaf forest and an increase in the amount of pasture and natural grasslands in the continent. We demonstrate the utility of integrating ancient genomes with archaeometric datasets in a spatiotemporal statistical framework, which we foresee will enable future studies of ancient popu-lations' movements, and their putative effects on local fauna and flora. migrations | ancient DNA | Neolithic | Bronze Age | land cover U p until about 8,500 y before present (BP), Europe was largely populated by groups of hunter-gatherers living at relatively low densities. This scenario changed when a wave of populations from the Middle East entered Europe via Ana-tolia, as evinced by recent ancient DNA studies (1-3). Studies based on radiocarbon-dated domestic plants, animals, and finds from associated contexts suggest that this migration wave spread farming practices into the region, initiating the Neolithic revolution in Europe (4-9). A second massive wave of movement occurred later, at the beginning of the Bronze Age, when populations associated with the Yamnaya culture in the Pontic steppe entered the continent from the east (10-12). These groups may have introduced horse herding and proto-Indo-European languages as they moved westward and are associated with the Corded Ware culture in central and northern Europe and, later on, the Bell Beaker phenomenon in northwestern Europe (13-16). Over the last 10,000 y, the continent also underwent major changes in its land-cover composition, but it is unclear how much the Neolithic and Yamnaya migrations contributed to these changes. Recent pollen-based studies suggest that a dramatic reduction of broad-leaf forests occurred from about 6,000 BP until the present (17). This deforestation intensified from around 2,200 BP, resulting in a replacement of these forests by grassland and arable land throughout the continent (18, 19). These processes , however, did not occur at the same rate throughout all regions. For example, while considerable decreases in broad-leaf forests occurred in central Europe starting around 4,000 BP, the Atlantic seaboard was predominantly occupied by semiopen vegetation since well before this time, while southern Scandinavia experienced less significant reductions in forest cover, at least until the Middle Ages (19-21). Presumably, these phenomena were partly effected by new human land-use activities involving forest clearance and the establishment of farming and herding practices, as earlier hunter-gatherer groups likely had limited effects on their surrounding flora and fauna (although see refs. 22 and 23). Changes in climate patterns may have also played a role in vegetation changes. Additionally, changes in vegetation may have opened up new areas for populations to expand. Until now, however, few efforts have been carried out to explicitly link changes in paleovegetation to particular human population movements, or to distinguish between climatic and human-based factors, assuming these had causal roles in these changes (but see refs. 18 and 24). In this study, we aim to trace how the major Holocene migrations unfolded across the European continent over time and to understand how they were associated with changes in the Significance We present a study to model the spread of ancestry in ancient genomes through time and space and a geostatistical framework for comparing human migrations and land-cover changes, while accounting for changes in climate. We show that the two major migrations during the European Holocene had different spatiotemporal structures and expansion rates. In addition, we find that the Yamnaya expansion had a stronger association with vegetational landscape changes than the earlier Neolithic farmer expansion. Our approach paves the way for future work linking paleogenomics with other archaeometric datasets in the study of the past.

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Archaeological and Anthropological Sciences, 2020

Megalithic tombs have since long been a focus of debate within the archaeological research field,... more 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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THE PITTED WARE CULTURE ON DJURSLAND Supra-regional significance and contacts in the Middle Neolithic of southern Scandinavia, 2020

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GOTARC Serie D nr 74

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Germania, 2020

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Cambridge Archaeological Journal, 2017

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