Germanic-speaking populations historically form an integral component of the North and Northwest European cultural configuration. According to linguistic consensus, the common ancestor of the Germanic languages, which include German, English, Frisian, Dutch as well as the Nordic languages, was spoken in Northern Europe during the Pre-Roman Iron Age. However, important questions remain concerning the earlier Bronze Age distribution of this Indo-European language branch in Scandinavia as well as the driving factors behind its Late Iron Age diversification and expansion across the European continent. A key difficulty in addressing these questions are the existence of striking differences in the interpretation of the archaeological record, leading to various hypotheses of correlations with linguistic dispersals and changes in material culture. Moreover, these interpretations have been difficult to assess using genomics due to limited ancient genomes and the difficulty in differentiating closely related populations. Here we integrate multidisciplinary evidence from population genomics, historical sources, archaeology and linguistics to offer a fully revised model for the origins and spread of Germanic languages and for the formation of the genomic ancestry of Germanic-speaking northern European populations, while acknowledging that coordinating archaeology, linguistics and genetics is complex and potentially controversial. We sequenced 710 ancient human genomes from western Eurasia and analysed them together with 3,940 published genomes suitable for imputing diploid genotypes. We find evidence of a previously unknown, large-scale Bronze Age migration within Scandinavia, originating in the east and becoming widespread to the west and south, thus providing a new potential driving factor for the expansion of the Germanic speech community. This East Scandinavian genetic cluster is first seen 800 years after the arrival of the Corded Ware Culture, the first Steppe-related population to emerge in Northern Europe, opening a new scenario implying a Late rather than an Middle Neolithic arrival of the Germanic language group in Scandinavia. Moreover, the non-local Hunter-Gatherer ancestry of this East Scandinavian cluster is indicative of a cross-Baltic maritime rather than a southern Scandinavian land-based entry. Later in the Iron Age around 1700 BP, we find a southward push of admixed Eastern and Southern Scandinavians into areas including Germany and the Netherlands, previously associated with Celtic speakers, mixing with local populations from the Eastern North Sea coast. During the Migration Period (1575-1200 BP), we find evidence of this structured, admixed Southern Scandinavian population representing the Western Germanic Anglo-Saxon migrations into Britain and Langobards into southern Europe. During the Migration Period, we detect a previously unknown northward migration back into Southern Scandinavia, partly replacing earlier inhabitants and forming the North Germanic-speaking Viking-Age populations of Denmark and southern Sweden, corresponding with historically attested Danes. However, the origin and character of these major changes in Scandinavia before the Viking Age remain contested. In contrast to these Western and Northern Germanic-speaking populations, we find the Wielbark population from Poland to be primarily of Eastern Scandinavian ancestry, supporting a Swedish origin for East Germanic groups. In contrast, the later cultural descendants, the Ostrogoths and Visigoths are predominantly of Southern European ancestry implying the adoption of Gothic culture. Together, these results highlight the use of archaeology, linguistics and genetics as distinct but complementary lines of evidence.### Competing Interest StatementThe authors have declared no competing interest.

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 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. Population-scale ancient genomics are used to infer ancestry, social structure and pathogen infection in 108 Scandinavian Neolithic individuals from eight megalithic graves and a stone cist, showing that Neolithic plague was widespread.

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.

Strontium isotope ratios in human teeth from the three Swedish prehistoric Stone Age hunter-fisher-gathering societies Norje Sunnansund (Maglemose), Skateholm (Ertebølle) and Västerbjers (Pitted Ware Culture) were analysed with laser ablation to produce data on both individual movement patterns and societal mobility trends. The analyses of teeth from both Skateholm and Västerbjers displayed homogeneous ratios and corresponding mobility patterns, while the data from Norje Sunnansund showed larger variances with heterogenous strontium ratios and varied inter-individual mobility patterns. Correlation with the bioavailable baseline suggests that the size of the geographical areas, where human strontium ratios could have originated, was roughly comparable for all three sites. The teeth measurements were reflected within a 50-km radius of the surrounding landscape and the 25–75% data quartile matched with distances between 3 and 30 km from the sites, suggesting limited mobility ranges among aquatically dependent foragers from southernmost Sweden. By applying ethnographic analogies and site-specific contextual inferences, the results suggest that mobility ranges at Norje Sunnansund were likely not delimited by neighbouring group territories. This changed over time and an increasing territorialisation of the landscape may have influenced movement patterns and caused restrictions to the foraging activities at both Skateholm and Västerbjers.

To evaluate the possibility of obtaining detailed individual mobility data from archaeological teeth, the strontium isotope ratios on 28 human teeth from three separate Early-Mid Holocene, Swedish, foraging contexts (Norje Sunnansund, Skateholm and Västerbjers) were analysed through laser ablation. The teeth/individuals have previously been analysed using traditional bulk sampled thermal ionisation mass spectrometry. To validate the conclusions regarding the archaeological teeth, a tooth from a modern man with a known background was also analysed. The result shows that all of the teeth display less than 0.4% discrepancy between the mean values of the laser ablation profiles and the previously published bulk data and 25 (89%) of the teeth display less than a 0.2% discrepancy. By calculating linear and polynomial trendlines for each ablated tooth, it was possible to illustrate a strong correlation for the transition pattern between the measurements when following a chronological sequence from the tip to the cervix. Such correlations were not reproduced when the data sequence was randomized. The analyses show that the chronologically sequenced ablation data fit with a transition between local bioavailable strontium regions, that the measurements do not fluctuate between extremes and that their values are not caused by end-member mixing. This indicates an increasing data resolution when reducing strontium isotope ratio averaging time by minimizing the sampling area. The results suggest strontium incorporation in human teeth can be measured on an ordinal scale, with a traceable chronological order to enamel mineralization when sampled from tip to cervix at an equal distance from the surface. Micro-sampling enamel is considered a valid method to assess prehistoric, but not modern, human mobility; laser ablation technology increases the amount of information obtained from a single tooth while rendering minimal damage to the studied specimen.

The nature of the Neolithic Pitted Ware Culture (PWC) has been debated in Scandinavian archaeology since the beginning of the 20th century. This material culture post-dates the inception of an agro-pastoral Neolithic economy in the region (TRB) but demonstrate a semi-foraging lifeway. The PWC is considered elusive in the sense that the economy has been interpreted as either based on maritime foraging (isotope analysis), or a mixedNeolithic economy based on boar (Sus scrofa) (archaeozoology). The mobility of the PWC group on Gotland has not been studied previously. We provide an analysis of strontium data from the site Vasterbjers, Gotland, and engage in the question whether this population was mobile (foragers) or stationary (mixed-Neolithic). We also discuss the strontium baseline of the island in the light of new data. The results presented demonstrate a group that was confined to the island and with no strontium data suggesting a supra-regional mobility. We cannot discriminate between a regional mobility pattern on Gotland (mobility between sites) and a local, stationary group (fully sedentary), but we present data that suggest the former. Thus, our conclusion is that the PWC group on Gotland was stationary, the sedentism disclosed being supported by a mixed-Neolithic economy based on boar (Sus scrofa). In fact, the PWC group is depicting less variance with respect to Sr-87/Sr-86 ratios than the TRB and Late Neolithic groups on the island, being more stationary than the agricultural groups.

At Norje Sunnansund, an Early Holocene settlement in southern Sweden, the world’s earliest evidence of fermentation has been interpreted as a method of managing long-term and large-scale food surplus. While an advanced fishery is suggested by the number of recovered fish bones, until now it has not been possible to identify the origin of the fish, or whether and how their seasonal migration was exploited. We analysed strontium isotope ratios ( 87 Sr/ 86 Sr) in 16 cyprinid and 8 pike teeth, which were recovered at the site, both from within the fermentation pit and from different areas outside of it, by using laser ablation multi-collector inductively coupled plasma mass spectrometry. Our investigation indicates three different regions of origin for the fish at the site. We find that the most commonly fermented fish, cyprinids (roach), were caught in the autumn during their seasonal migration from the Baltic Sea to the sheltered stream and lake next to the site. This is in contrast to the cyprinids from other areas of the site, which were caught when migrating from nearby estuaries and the Baltic Sea coast during late spring. The pikes from the fermentation pit were caught in the autumn as by-catch to the mainly targeted roach while moving from the nearby Baltic Sea coast. Lastly, the pikes from outside the fermentation pit were likely caught as they migrated from nearby waters in sedimentary bedrock areas to the south of the site, to spawn in early spring. Combined, these data suggest an advanced fishery with the ability to combine optimal use of seasonal fish abundance at different times of the year. Our results offer insights into the practice of delayed-return consumption patterns, provide a more complete view of the storage system used, and increase our understanding of Early Holocene sedentism among northern hunter-fisher-gatherers. By applying advanced strontium isotope analyses to archaeological material integrated into an ecological setting, we present a methodology that can be used elsewhere to enhance our understanding of the otherwise elusive indications of storage practices and fish exploitation patterns among ancient foraging societies.

When the famous bishop of Lund, Peder Winstrup, died in 1679 he was laid to rest in a family crypt in Lund Cathedral alongside his wife. Following a restoration of the Cathedral in the late 19th century, Peder Winstrup's coffin was moved to the crypt, as the family grave was dismantled. In 2012, a decision was made to move his remains to a different location presenting scientists with an opportunity to investigate bishop's mummified remains. Unexpectedly, hidden underneath Winstrup's body, a small bundle containing mummified remains of a 5-6-month-old foetus was found. This finding prompted questions regarding the possible relation between both individuals with most hypotheses suggesting no relation and an opportunistic character of the placement of the foetus in bishop's coffin. Here we test the hypotheses using ancient DNA genomics, including mitochondrial DNA and Y chromosome data, as tools for kinship analyses. We identified a second-degree kinship relation, which, in combination with genealogical analysis, suggests a grandparent-grandchild relation, as highly probable affiliation.

Additional file 1: Table S1. Assigned reads from all taxonomic levels represented in the metagenomic LUND1 library prior to in-solution capture for MTBC DNA.

Additional file 6: Table S5. List of genomes included in the L4 dataset, with respective publications, accession numbers, lineages, dates, and percentage of total SNPs called as heterozygous.

Background Although tuberculosis accounts for the highest mortality from a bacterial infection on a global scale, questions persist regarding its origin. One hypothesis based on modern Mycobacterium tuberculosis complex (MTBC) genomes suggests their most recent common ancestor followed human migrations out of Africa approximately 70,000 years before present. However, studies using ancient genomes as calibration points have yielded much younger dates of less than 6000 years. Here, we aim to address this discrepancy through the analysis of the highest-coverage and highest-quality ancient MTBC genome available to date, reconstructed from a calcified lung nodule of Bishop Peder Winstrup of Lund (b. 1605–d. 1679). Results A metagenomic approach for taxonomic classification of whole DNA content permitted the identification of abundant DNA belonging to the human host and the MTBC, with few non-TB bacterial taxa comprising the background. Genomic enrichment enabled the reconstruction of a 141-fold coverage M . tuberculosis genome. In utilizing this high-quality, high-coverage seventeenth-century genome as a calibration point for dating the MTBC, we employed multiple Bayesian tree models, including birth-death models, which allowed us to model pathogen population dynamics and data sampling strategies more realistically than those based on the coalescent. Conclusions The results of our metagenomic analysis demonstrate the unique preservation environment calcified nodules provide for DNA. Importantly, we estimate a most recent common ancestor date for the MTBC of between 2190 and 4501 before present and for Lineage 4 of between 929 and 2084 before present using multiple models, confirming a Neolithic emergence for the MTBC.

This study highlights the importance of different protein sources in the diet of Early and Middle Mesolithic humans in southern Scandinavia, and illustrates variation and change in protein consumption patterns during the Early Holocene. By combining previously published stable isotope data with new analyses of human and animal bone remains, a Bayesian mixing model was used to reveal that fishing was more important than previously anticipated in the foraging economy. Incorporating the zooarchaeological record as a prior to guide the Bayesian model enabled further study of Early Holocene foraging in the region. Although primarily a study of human diet, because the results indicate that aquatic systems were more important than previously acknowledged, it is possible to discuss the implications for understanding Early Holocene subsistence strategies and mobility. Furthermore, by incorporating both zooarchaeological data and human stable isotope analysis, the methodology can advance palaeodietary studies, by generating dietary protein estimations that can be used to investigate subsistence strategies across a diverse set of human societies.

Departing from two recently dated finds of human bones in wetlands from the area of Falbygden in western Sweden, this paper describes the finds in detail and sets them in a wider context of depositional practices in the south Scandinavian Early Neolithic. The two finds are both of girls in the age of 15-20 years, and in one of the cases it is probable that she was bound and possibly intentionally drowned in shallow water. The Falbygden finds have clear parallels in Denmark and in southern Sweden in the same period. The Danish material indicates that a particular segment of the population was treated in this way, primarily young individuals around 15-20 years. Some of them show signs of disease or deformations, many have signs of trauma, and a couple have been found with cords around the neck, suggesting violent deaths. In a number of cases, they were also found in pairs. One of the Falbygden finds, the so-called Hallonflickan, was also unusual in that her Sr isotope ratio indicates that the was born far away, probably in Scania in southern Sweden. Whether other finds of this kind were also long distance movers is not known. Direct bone datings show that this practice is established at the same time as the introduction of agriculture and of the Funnel Beaker culture in Scandinavia, around 4000 BC, and continues until ca 3000 BC after which it disappears for ca 1000 years. Deposition of humans in wetlands is paralleled by deposition of animals, pottery and stone tools, and we suggest that these form parts of a complex of ritual practices established over a large area at the time of the earliest Scandinavian agriculture.

Departing from two recently dated finds of human bones in wetlands from the area of Falbygden in western Sweden, this paper describes the finds in detail and sets them in a wider context of depositional practices in the south Scandinavian Early Neolithic. The two finds are both of girls in the age of 15-20 years, and in one of the cases it is probable that she was bound and possibly intentionally drowned in shallow water.The Falbygden finds have clear parallels in Denmark and in southern Sweden in the same period. The Danish material indicates that a particular segment of the population was treated in this way, primarily young individuals around 15-20 years. Some of them show signs of disease or deformations, many have signs of trauma, and a couple have been found with cords around the neck, suggesting violent deaths. In a number of cases, they were also found in pairs.One of the Falbygden finds, the so-called Hallonflickan, was also unusual in that her Sr isotope ratio indicates that the was born far away, probably in Scania in southern Sweden. Whether other finds of this kind were also long distance movers is not known.Direct bone datings show that this practice is established at the same time as the introduction of agriculture and of the Funnel Beaker culture in Scandinavia, around 4000 BC, and continues until ca 3000 BC after which it disappears for ca 1000 years. Deposition of humans in wetlands is paralleled by deposition of animals, pottery and stone tools, and we suggest that these form parts of a complex of ritual practices established over a large area at the time of the earliest Scandinavian agriculture.

Migration is a complex subject to approach in archeology and the new materials and methods available, such as isotope analysis and DNA, make it possible, and necessary, to ask new questions. The objective of this paper is to highlight the possibilities with using a new approach to migration on a population level by applying Bayesian mixing analysis of strontium isotopes. The selected case, the island of Oland in the Baltic, was based on 109 human samples dated to the Early (500 BC-AD 400, n = 71) and Late (AD 400-1050, n = 38) periods. The results from both periods demonstrate that the distribution of Strontium (Sr) is multimodal with several peaks not associated with the local variation. Our results show a large immigration to Oland from other geological areas, with 32% of the population in the Early period and 47% in the Late period being nonlocal. In order to unravel these distributions, we use a Bayesian mixing analysis. The Bayesian mixing analysis provides us with a mean to disentangle the distribution of Sr that is not uninformed. The gravity model, however simplistic, is relevant for explaining the strontium variation in the population in Oland both in the Early and Late period. Our results indicate a significant internal migration in Scandinavia that is increasing in the Late Iron Age at the same time as the Viking expansions (the more well studied external migration), which is usually the only migration discussed. We argue that the method proposed and tested on the case of Oland adds new perspectives for approaching migration patterns in general on a population level, a perspective that is hitherto lacking in archaeology. (C) 2015 Elsevier Ltd. All rights reserved.