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Thread: How Diverse Were the Vikings: Population genomics of the Viking world

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    How Diverse Were the Vikings: Population genomics of the Viking world

    The Viking maritime expansion from Scandinavia (Denmark, Norway, and Sweden) marks one of the swiftest and most far-flung cultural transformations in global history. During this time (c. 750 to 1050 CE), the Vikings reached most of western Eurasia, Greenland, and North America, and left a cultural legacy that persists till today. To understand the genetic structure and influence of the Viking expansion, we sequenced the genomes of 442 ancient humans from across Europe and Greenland ranging from the Bronze Age (c. 2400 BC) to the early Modern period (c. 1600 CE), with particular emphasis on the Viking Age. We find that the period preceding the Viking Age was accompanied by foreign gene flow into Scandinavia from the south and east: spreading from Denmark and eastern Sweden to the rest of Scandinavia. Despite the close linguistic similarities of modern Scandinavian languages, we observe genetic structure within Scandinavia, suggesting that regional population differences were already present 1,000 years ago. We find evidence for a majority of Danish Viking presence in England, Swedish Viking presence in the Baltic, and Norwegian Viking presence in Ireland, Iceland, and Greenland. Additionally, we see substantial foreign European ancestry entering Scandinavia during the Viking Age. We also find that several of the members of the only archaeologically well-attested Viking expedition were close family members. By comparing Viking Scandinavian genomes with present-day Scandinavian genomes, we find that pigmentation-associated loci have undergone strong population differentiation during the last millennia. Finally, we are able to trace the allele frequency dynamics of positively selected loci with unprecedented detail, including the lactase persistence allele and various alleles associated with the immune response. We conclude that the Viking diaspora was characterized by substantial foreign engagement: distinct Viking populations influenced the genomic makeup of different regions of Europe, while Scandinavia also experienced increased contact with the rest of the continent.
    The rest of the study: Population genomics of the Viking world



    Survive the Jive's take on the study:





    Some interesting points. Particularly how the Norse already had considerable amounts of non-Anglo-Saxon British DNA before the Viking Age, and modern Norwegians averaging around 12 percent.
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    Haven't seen the vid yet but that screenshot suggests they just love pushing the race mixing agenda.
    I grew up on a belief of honour, courage and the old world values. The world isn't about that anymore, preferring to die a slow death of fast food and cheap thrills.

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    Quote Originally Posted by Meister View Post
    Haven't seen the vid yet but that screenshot suggests they just love pushing the race mixing agenda.
    That's just Survive the Jive making fun and being click-bait-y. The study is remarkably free of any pro-race-mixing agenda and sentiments. Although, StJ does bring up an interesting side-point about a sub-Saharan man making his way to Iceland in the 1700s (I believe), whom settled down there and had children, of which every single Icelander is descended from today, due to the extremely small gene pool on the island.
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    I was actually waiting for you to show up three days ago, when the study first appeared, so I could show it to you but now you beat me to it.

    I will reply to the actual study tomorrow but I didn't want to leave this unchallenged:

    Quote Originally Posted by Žoreišar View Post
    Although, StJ does bring up an interesting side-point about a sub-Saharan man making his way to Iceland in the 1700s (I believe), whom settled down there and had children, of which every single Icelander is descended from today, due to the extremely small gene pool on the island.
    Even if he actually came in the early 1700s, even Iceland had a population of over 50.000 people at this time, living in an area of over 100.000 km². It would be impossible for him to be the ancestor of every living Icelander or even merely the majority of them.

    The guy he actually means is probably Hans Jonatan, a Mulatto from the Carribean, who came to Iceland in 1802, fathered two children and died in 1827. He only has 788 descendants nowadays, even most of whom don't appear to carry African traces by now, according to a study done on his descendants("just" 121 of those were tested, though):

    A genome is a mosaic of chromosome fragments from ancestors who existed some arbitrary number of generations earlier. Here, we reconstruct the genome of Hans Jonatan (HJ), born in the Caribbean in 1784 to an enslaved African mother and European father. HJ migrated to Iceland in 1802, married and had two children. We genotyped 182 of his 788 descendants using single-nucleotide polymorphism (SNP) chips and whole-genome sequenced (WGS) 20 of them. Using these data, we reconstructed 38% of HJ’s maternal genome and inferred that his mother was from the region spanned by Benin, Nigeria and Cameroon.


    Genealogical reconstruction of HJ’s maternal chromosome 3
    The genealogy connecting HJ to his 182 genotyped descendants (colored chromosomes) via 61 ungenotyped descendants (uncolored chromosomes). Each individual is represented by a pair of chromosomes, with parent–offspring relationships represented by connecting lines. For the genotyped descendants, yellow- and blue-colored fragments indicate European and African ancestry, respectively. Inferred African fragments in ungenotyped ancestors are shown in dark gray. In total, 126.97 (64%) of 198.29 Mb of HJ’s maternal chromosome 3 was reconstructed.
    They tested 151.000, so almost half of Iceland's ethnic Icelandic population for African ancestry:

    Because of the relative isolation of Icelanders, genuine African fragments indescendants of HJ are likely to originate only from HJ. To verify this assumption, we ran ADMIXTURE and HAPMIX for all our chiptyped Icelanders (151,014 individuals; see Methods). The results reveal that among individuals born between 1880 and 1930, only descendants of HJ have evident ( > 1%) African ancestry. Among those born after 1930 and not descended from HJ, we identified 320 with evident African ancestry, none of whom were ancestors of HJ’s descendants.
    So if we add those 320 to his 788 descendants and double the number to account for the untested 150.000, only about 2.200 Icelanders could have any amount of African ancestry. As we have seen from the above figure, that number, purely genetically speaking, is actually much lower. I can't find any concrete number apart from that above figure of Hans Jonatan's descendants, though.
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    Ancient DNA: List of All Studies analyzing DNA of Ancient Tribes and Ethnicities(post-2010)


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    So, danish people are the most closest population to the ancient Swedish vikings?

    There are some incredible Fig. in the paper.



    Map of the Ā“Viking WorldĀ” from 8th till 11th centuries.
    Different symbols on the map (a) correspond to ancient sites of a specific age/culture. The ancient samples are divided into the following five broad categories: Bronze Age (BA) - c. 2500 BC - 900 BC; Iron Age (IA) - c. 900 BC to 700 CE; Early Viking Age (EVA) - c. 700 to 800 CE; VA - c. 800 to 1100 CE; Medieval - c. 1100 to 1600 CE. b, All ancient individuals from this study (n=442) and published VA samples (n=21) from Sigtuna6 are categorized based on their spatio-temporal origin.



    Viking Age archaeological sites.
    Examples of a few archaeological Viking Age sites and samples used in this study. a, Salme II ship burial site of Early Viking Age excavated in present-day Estonia: schematic representation of skeletons (upper left-hand corner image) and aerial images of skeletons (upper right-hand corner and lower images). b, Ridgeway Hill mass grave dated to the 10th or 11th century, located on the crest of Ridgeway Hill, near Weymouth, on the South coast of England. Around 50 predominantly young adult male individuals were excavated. c, The site of Balladoole: around AD 900, a Viking was buried in an oak ship at Balladoole, Arbory in the south east of the Isle of Man. d, Viking Age archaeological site in Varnhem, Sweden: Schematic map of the church foundation (left) and the excavated graves (red markings) at the early Christian cemetery in Varnhem; foundations of the Viking Age stone church in Varnhem (middle) and the remains of a 182 cm long male individual (no. 17) buried in a lime stone coffin close to the church foundations (right).



    Symmetry tests of genetic affinity of ancient individuals with contemporary populations.
    Panels show D-statistics of the form D(YRI,Y; X,Denmark), which contrast allele sharing of an ancient individual Y with either contemporary population X or Denmark. Plot symbols show point estimates, and density plots distributions across all individuals per analysis group.




    Symmetry tests for genetic affinity with Baltic Bronze Age
    Panels show f4-statistics of the form f4(Mbuti,Baltic_BA;Y, Salme.SG_EVA), which contrast allele sharing of Baltic_BA with either a test individual Y or Salme.SG_EVA. a, point estimates and error bars (Ā± 3 standard errors) for each target individual, aggregated by analysis group. Individuals with significant f4-statistics (|Z| ≥ 3) are indicated without transparency and respective sample IDs. b, as in (a), with density plot for distributions across all individuals per analysis group.




    Ancestry diversity of different population groups
    Diversity of different labels (i.e. sample locations combined with historical age) are shown as a function of their sample size. The Diversity measure is the Kullback-Leibler divergence from the label means, capturing the diversity of a group with respect to the average of that group; see text for details. Larger values are more diverse, though a dependence on sample size is expected. The simulation expectation for the best-fit to the data (0=0.2) is shown.



    Ancestry modelling using qpAdm
    a, Ternary plots of ancestry proportions for a three-way model of Mesolithic hunter-gatherer (Loschbour.SG_M), Neolithic farmer (Barcin.SG_EN) and Bronze Age Steppe herders (Yamnaya.SG_EBA). b, Bar plots with ancestry proportions as in (a), with error bars indicating standard errors and transparency/text colors indicating p-value for model fit (no transparency/black: p ≥ 0.05; light transparency/blue: 0.05 > p ≥ 0.01; strong transparency/red: p ≤ 0.01). c, Ancestry proportions of four-way models including additional putative source groups for target groups for which three-way fit was rejected (p ≤ 0.01); transparency/text colors as in (b)










    Genetic structure of VA samples.
    a, Multidimensional scaling (MDS) plot based on a pairwise identity-by-state (IBS) sharing matrix of the VA and other ancient samples (Supplementary Table 3). b, Uniform manifold approximation and projection (UMAP) analysis of the same dataset as in plot (a).




    Genetic structure and diversity of ancient samples.
    a, Uniform manifold approximation and projection (UMAP) analysis of the ancient and modern Scandinavian individuals based on the first 10 dimensions of MDS using identity-by-descent (IBD) segments of imputed individuals. Large symbols indicate median coordinates for each group. b, Genetic diversity in major Scandinavian VA populations. Plots next to the map show MDS analysis based on a pairwise IBS sharing matrix. Here Ā“NorwayĀ” represents all the sites from Norway. The scale is identical for all the plots.






    Spatiotemporal patterns of Viking and non-Viking ancestry in Europe during the IA, EVA and VA.
    UK = Ā‘British-likeĀ’ / Ā‘North AtlanticĀ’ ancient ancestry component. Sweden = Ā‘Swedish-likeĀ’ ancient ancestry component. Denmark = Ā‘Danish-likeĀ’ ancient ancestry component. Norway = Ā‘Norwegian-likeĀ’ ancient ancestry component. Italy = Ā‘Southern European-likeĀ’ ancestry component. See Table S11.2 for statistical tests. The Ā‘Swedish-likeĀ’ ancestry is the highest in present-day Estonia due to the ancient samples from the Salme ship burial, which originated from the MƤlaren Valley of Sweden, according to archaeological sources.

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    Quote Originally Posted by Volk und Rasse View Post
    So, danish people are the most closest population to the ancient Swedish vikings?
    I haven't read all of the original study in detail, but I'm unsure how you gleaned this from the paper. From what I gathered, the distinct nations of Scandinavia correspond quite neatly together with the modern distinct nations, in a genetic sense (with the exception of Skåne still clustering with Danes rather than Swedes).
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    I’m somewhat torn over this study: While I applaud them for sampling so many remains I’m thoroughly disappointed by their methodology for the single sample comparisons. See the supplementary table S6_Ancestry Estimates.

    They are using modern populations (so their descendants/the outcome of possible mixture) to model their ancestors, which is obviously the wrong way round. Especially since they only have British, Danish, Norwegian, Swedish, Polish, Italian and Finnish. No Germans, Dutch (and Flemings) or French, who, apart from Brits, would be the most likely sources of mixture into a Viking Age Scandinavian gene pool.

    Accordingly, those comparisons seem next to useless to me, at least for those samples which get very variable percentages.
    The only samples for which this kind of modeling seems meaningful, is something like the sample VK528 for example, which is from 8-9th century Troms in northern Norway but genetically 97% British and has the paternal haplogroup R1b-L21 (which is very British).

    For example, their “Swedish-like” can’t be a real historic component. There’s a single sample and granted it’s quite early, from 4th-5th century CE Oland, that has 99% Swedish and another from 9th-11th century CE Oland has 96% but the average of 70 samples is only 19% Swedish and the Median is actually 1%!
    If we drop the above mentioned two samples and three others with 78%, 73% and 69%, the average actually drops to 12%!
    So, until someone believes there was a total population replacement in Sweden (of all places, a land whose people didn’t participate much at all in the Viking raids in the west) by the 10th century, this makes no sense. In all likelihood those samples with 99% and 96% Swedish were actually outliers.

    As for the “Italian-like” component, that’s in all likelihood just an increased connection to the Neolithic farmers. That’s because it appears consistently and fairly early at relatively low levels (although there are outliers in both directions), when no actual Italians could have reached Scandinavia in any numbers.
    In fact, I suspect that many samples which get a tripartite split of “British”, “Danish” and “Italian” are just Germans/continental Germanics.

    For example:


    Sample Country Division Division Age Finer_age Y-DNA mtDNA British Danish Swedish Norwegian Polish Italian Finnish
    VK369 Denmark Sealand Bakkendrup Viking 850-900 CE P312 H1a 4% 41% 0% 0% 1% 53% 1%
    VK385 Denmark Sealand Lejre Viking 850-900 CE Frau U4c1 9% 31% 0% 9% 2% 49% 1%
    VK301 Denmark Funen Ladby Viking 640-890 CE I1-Z58 R0a2b 9% 43% 0% 1% 1% 46% 2%
    VK319 Denmark Funen Ladby Viking 800 CE Frau H1bq 15% 32% 0% 2% 3% 44% 3%
    VK84 Denmark Jutland Hesselbjerg Viking 850-900 CE Frau J1c2p 26% 29% 1% 1% 5% 36% 4%
    VK65 Denmark Sealand Tollemosegård Early Viking Late Germanic Iron Age/
    early Viking
    Frau V25 28% 39% 0% 0% 2% 30% 0%
    VK87 Denmark Jutland Hesselbjerg Viking 850-900 CE DF27 K1c2 41% 29% 9% 3% 0% 19% 0%
    VK300 Denmark Jutland Hesselbjerg Viking 850-900 CE Frau H3 19% 62% 0% 0% 0% 18% 0%
    VK315 Denmark Sealand Bakkendrup Viking 850-900 CE I1-L22 T1a1b 11% 68% 0% 3% 0% 18% 0%
    VK69 Denmark Sealand Tollemosegård Early Viking Late Germanic Iron Age/
    early Viking
    Frau H2a2a1 17% 71% 0% 0% 0% 12% 0%


    Quote Originally Posted by Žoreišar View Post
    Some interesting points. Particularly how the Norse already had considerable amounts of non-Anglo-Saxon British DNA before the Viking Age, and modern Norwegians averaging around 12 percent.
    But the thing is, that this “British” DNA actually incorporates Anglo-Saxon DNA, it’s not some sort of pure Island Celtic component because it’s based on modern DNA.

    In one of the pictures VuR already linked and in supplementary table S7_qpAdm they use ancient populations like Yamnaya (the probable original Indo-Europeans or rather a population closely related to them), Barcin (the Anatolian Neolithic farmers) and Loschbour (the original hunter Gatherers of Europe) for sample groups, that’s a better approach. But actually those are too old because they are from the Neolithic or even earlier and it doesn’t tell us much about more recent population mixtures.

    They should have simply used Bronze Age and Iron Age samples from Scandinavia, Germany or even those Longobards from Hungary and Italy who were obviously unmixed or their own early pre-Viking samples which are likely to be yet unmixed with whatever the Vikings brought to Scandinavia.

    The study also doesn’t include a PCA of Western Eurasia and/or Europe, which is pretty much standard in such studies nowadays and from which you can easily get an idea to which population is close.

    I’m afraid we’ll have to wait for the raw data, which will be available once the study has actually been published (so far it’s just a pre-print). Then “amateur” geneticists can analyse the individual samples properly.
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    What do they know of Europe, Who only Europe know?



    Ancient DNA: List of All Studies analyzing DNA of Ancient Tribes and Ethnicities(post-2010)


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    In connection to the recent publishing of the complete study in question, and the following media wave concerning it, I thought this response by one of the co-authors of the study was due.

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    Who cares ... all Germanic people were bright-haired and bright-eyed with bright skin, that's all that matters.

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    Mad Population genomics of the Viking world

    World's largest DNA sequencing of Viking skeletons reveals they weren't all Scandinavian

    by University of Cambridge
    An artistic reconstruction of 'Southern European' Vikings emphasising the foreign gene flow into Viking Age Scandinavia. Credit: Jim Lyngvild

    Invaders, pirates, warriors—the history books taught us that Vikings were brutal predators who travelled by sea from Scandinavia to pillage and raid their way across Europe and beyond.
    Now cutting-edge DNA sequencing of more than 400 Viking skeletons from archaeological sites scattered across Europe and Greenland will rewrite the history books as it has shown:

    • Skeletons from famous Viking burial sites in Scotland were actually local people who could have taken on Viking identities and were buried as Vikings.
    • Many Vikings actually had brown hair not blonde hair.
    • Viking identity was not limited to people with Scandinavian genetic ancestry. The study shows the genetic history of Scandinavia was influenced by foreign genes from Asia and Southern Europe before the Viking Age.
    • Early Viking Age raiding parties were an activity for locals and included close family members.
    • The genetic legacy in the UK has left the population with up to six percent Viking DNA.

    The six-year research project, published in Nature today, debunks the modern image of Vikings and was led by Professor Eske Willerslev, a Fellow of St John's College, University of Cambridge, and director of The Lundbeck Foundation GeoGenetics Centre, University of Copenhagen.
    He said: "We have this image of well-connected Vikings mixing with each other, trading and going on raiding parties to fight Kings across Europe because this is what we see on television and read in books—but genetically we have shown for the first time that it wasn't that kind of world. This study changes the perception of who a Viking actually was—no one could have predicted these significant gene flows into Scandinavia from Southern Europe and Asia happened before and during the Viking Age."
    The word Viking comes from the Scandinavian term 'vikingr' meaning 'pirate'. The Viking Age generally refers to the period from A.D. 800, a few years after the earliest recorded raid, until the 1050s, a few years before the Norman Conquest of England in 1066. The Vikings changed the political and genetic course of Europe and beyond: Cnut the Great became the King of England, Leif Eriksson is believed to have been the first European to reach North America—500 years before Christopher Columbus—and Olaf Tryggvason is credited with taking Christianity to Norway. Many expeditions involved raiding monasteries and cities along the coastal settlements of Europe but the goal of trading goods like fur, tusks and seal fat were often the more pragmatic aim.

    Professor Willerslev added: "We didn't know genetically what they actually looked like until now. We found genetic differences between different Viking populations within Scandinavia which shows Viking groups in the region were far more isolated than previously believed. Our research even debunks the modern image of Vikings with blonde hair as many had brown hair and were influenced by genetic influx from the outside of Scandinavia."
    DNA from a female skeleton named Kata found at a Viking burial site in Varnhem, Sweden, was sequenced as part of the study. Credit: Västergötlands Museum

    The team of international academics sequenced the whole genomes of 442 mostly Viking Age men, women, children and babies from their teeth and petrous bones found in Viking cemeteries. They analysed the DNA from the remains from a boat burial in Estonia and discovered four Viking brothers died the same day. The scientists have also revealed male skeletons from a Viking burial site in Orkney, Scotland, were not actually genetically Vikings despite being buried with swords and other Viking memorabilia.
    There wasn't a word for Scandinavia during the Viking Age—that came later. But the research study shows that the Vikings from what is now Norway travelled to Ireland, Scotland, Iceland and Greenland. The Vikings from what is now Denmark travelled to England. And Vikings from what is now Sweden went to the Baltic countries on their all male 'raiding parties'.
    Dr. Ashot Margaryan, Assistant Professor at the Section for Evolutionary Genomics, Globe Institute, University of Copenhagen and first author of the paper, said: "We carried out the largest ever DNA analysis of Viking remains to explore how they fit into the genetic picture of Ancient Europeans before the Viking Age. The results were startling and some answer long-standing historical questions and confirm previous assumptions that lacked evidence.
    "We discovered that a Viking raiding party expedition included close family members as we discovered four brothers in one boat burial in Estonia who died the same day. The rest of the occupants of the boat were genetically similar suggesting that they all likely came from a small town or village somewhere in Sweden."
    DNA from the Viking remains were shotgun sequenced from sites in Greenland, Ukraine, The United Kingdom, Scandinavia, Poland and Russia.
    Professor Martin Sikora, a lead author of the paper and an Associate Professor at the Centre for GeoGenetics, University of Copenhagen, said: "We found that Vikings weren't just Scandinavians in their genetic ancestry, as we analysed genetic influences in their DNA from Southern Europe and Asia which has never been contemplated before. Many Vikings have high levels of non-Scandinavian ancestry, both within and outside Scandinavia, which suggest ongoing gene flow across Europe."
    The team's analysis also found that genetically Pictish people 'became' Vikings without genetically mixing with Scandinavians. The Picts were Celtic-speaking people who lived in what is today eastern and northern Scotland during the Late British Iron Age and Early Medieval periods.
    A mass grave of around 50 headless Vikings from a site in Dorset, UK. Some of these remains were used for DNA analysis. Credit: Dorset County Council/Oxford Archaeology

    Dr. Daniel Lawson, lead author from The University of Bristol, explained: "Individuals with two genetically British parents who had Viking burials were found in Orkney and Norway. This is a different side of the cultural relationship from Viking raiding and pillaging."
    The Viking Age altered the political, cultural and demographic map of Europe in ways that are still evident today in place names, surnames and modern genetics.
    Professor Sųren Sindbęk, an archaeologist from Moesgaard Museum in Denmark who collaborated on the ground-breaking paper, explained: "Scandinavian diasporas established trade and settlement stretching from the American continent to the Asian steppe. They exported ideas, technologies, language, beliefs and practices and developed new socio-political structures. Importantly our results show that 'Viking' identity was not limited to people with Scandinavian genetic ancestry. Two Orkney skeletons who were buried with Viking swords in Viking style graves are genetically similar to present-day Irish and Scottish people and could be the earliest Pictish genomes ever studied."
    Assistant Professor Fernando Racimo, also a lead author based at the GeoGenetics Centre in the University of Copenhagen, stressed how valuable the dataset is for the study of the complex traits and natural selection in the past. He explained: This is the first time we can take a detailed look at the evolution of variants under natural selection in the last 2,000 years of European history. The Viking genomes allow us to disentangle how selection unfolded before, during and after the Viking movements across Europe, affecting genes associated with important traits like immunity, pigmentation and metabolism. We can also begin to infer the physical appearance of ancient Vikings and compare them to Scandinavians today."
    The genetic legacy of the Viking Age lives on today with six percent of people of the UK population predicted to have Viking DNA in their genes compared to 10 percent in Sweden.
    Professor Willeslev concluded: "The results change the perception of who a Viking actually was. The history books will need to be updated."

    https://phys.org/news/2020-09-world-...ng-viking.html


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