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Citation: Skoglund, Pontus, et al. "Genomic Diversity and Admixture Differs for Stone-Age Scandinavian Foragers and Farmers." Science 344.6185 (2014): 747-750.

Citation: Skoglund, Pontus, et al. “Genomic Diversity and Admixture Differs for Stone-Age Scandinavian Foragers and Farmers.” Science 344.6185 (2014): 747-750.

We are Whigs, whether we want to be or not. History moves in one direction, and that direction is associated with progress. Progress being something we would recognize as associated with ourselves in some fashion. Ergo, the “mystery” of the evacuation of Greenland by Scandinavians in the 1400s. Today that mystery seems to be solved to the satisfaction of most. With the waning of the Medieval Warm Period the Scandinavian agro-pastoral economic system of production was not a viable form of subsistence at high latitudes. Greenland got less green. In contrast the Greenland Inuit’s ancestors, the Thule culture, were eminently well prepared for the shift in climatic regimes. In a previous more Eurocentric age the curiosity was that a European society which was advanced enough to receive bishops from Rome could be replaced by hunter-gatherers in sealskin canoes.

440px-Saami_Family_1900Of course many peoples would not have been shocked, the ancients were well aware of the concept of societies falling from states of greater complexity or social elaboration to ones of simplicity, as a matter of necessity (see: Dark Age Greece). It was in Europe where the Age of Discovery transformed into the industry & science driven era of European colonialism, that gave us the idea that the world was ascending up a ladder of development evermore, under the aegis of the white race. Cases where Europeans gave ground to non-Europeans, and ones in an earlier mode of production in a historical determinist sense (i.e., societies moving through modes of production in sequence), would certainly raise eyebrows in a culture where the Garden of Eden had been turned into a legend and Greek myths of ages of Gold giving way to Silver and Bronze were seen as anthropological curiosities.

Obviously things have changed a great deal, but the shadow of the Whig, and the vision of eternal progress haunts us. Scratch a Critical Race theorist, and you get James Mill. A Whiggish and Eurocentric perspective colors our own perceptions of the past, even if we live in an age of the critique of all things Western and white. Most especially thees sorts of biases are a problem when it comes to prehistory, when we don’t even have to bother to twist and interpret the past’s words to fit our preconceptions. We can simply impute upon it because it is mute. In this blog I have been talking about the impact ancient DNA has had upon our understanding, and the impact it will have. But the inferences we make are only as good as our interpretative framework. The researchers who are working at the cutting edge of the field understand they aren’t explaining everything. Rather, they are attempting to construct some broad sketches which can serve as a scaffold for more specific detailed understanding of events which transpired before history.

Citation: Cramp, Lucy JE, et al. "Neolithic dairy farming at the extreme of agriculture in northern Europe."

In Proceedings of the Royal Society B there is a paper which explores the dynamics of the transition from hunting & gathering to farming as the dominant way of life in Finland, Neolithic dairy farming at the extreme of agriculture in northern Europe. As you can see in the map to the left Finland spans the same latitudes as southern Greenland. Only its position along the western maritime fringe of Eurasia moderates the conditions so as to make agriculture marginally viable. The paper takes as a starting point what we know in general about the transition to farming in the far northeast of Europe. It came late. Around ~2500 BC. It was associated with the Corded Ware culture. Though such suppositions are fraught with uncertainty, believe that the Corded Ware were the first early Indo-Europeans in Northern Europe. The hunter-gatherers preceding the Corded Ware were of the Comb Cermic culture. The culture relatives of these people in Scandinavia were the Pitted Ware culture.

The basic results of the paper are easy to understand from a non-specialist perspective. Around ~2500 BC there was a very rapid shift to agro-pastoralism utilizing dairy from a predominantly marine diet. This correlates with the switch from Comb Ceramic hunter-gatherers, who were specifically reliant on marine animals in much of Finland, to the Corded Ware people. Later it seems that marine organisms made something of a comeback in the diet of the peoples of Finland, and a culturally more synthetic society emerged, with elements from the Comb Ceramic and the Corded Ware.

This should be somewhat familiar. Genetically it seems that in Northern Europe the arrival of agriculture was heralded by a demographic and culture eruption, which was eventually synthesized with the local substrate. If you read ancient DNA papers Scandinavians today are genetically an admixture of farmers and hunter-gatherers, with perhaps a modest bias toward the latter. The figure at the top of the post illustrates that the Pitted Ware populations seem to be genetically distinct from modern Northern Europeans, and in particular Finns. The same goes for the first farmer populations in the north. They were either emulsified in the still dominant hunter-gatherer demographic substrate, or, they experienced a major die off.

A simple model, implied in this paper, is that the modern Finns are a synthesis of the Corded Ware agro-pastoralists and indigenous hunter-gatherers populations. One can then envisage an admixture shock ~2500 BCE, and the past 5,000 years have been an equilibration. Obviously most people will immediately wonder though about the fact that Finns speak a Uralic language. And more specifically a Finno-Permian language. There have long been arguments about whether the Finns, in a cultural sense, are primal to Northern Europe. This plays out in the context of the fact that non-Indo-European languages in Europe always get special attention. What we do know from high density SNP data, as well as earlier Y chromosomal work, is that Finnic peoples seem to have a connection to populations in Siberia. By this, I do not mean the Ancestral North Eurasians. Rather, a population with affinities to modern Northeast Asians. If the Pitted Ware genetic results can be generalized to the Comb Ceramic people, and I do think they can be, then the Siberian admixture in Finns post dates 2500 BC. Since this element is not found in most populations descended from the Corded Ware (the ones where it is found, the Russians, have historical reasons for likely admixture from Asian populations, or, were Russified Finns), I doubt it is from the Corded Ware. Rather, the most likely scenario involves Finnic peoples moving into the population, and adding themselves as a dominant cultural element. The modern Indo-European language spoken in Finland by natives is Swedish, which arrived during the Common Era. With Swedish cultural hegemony and some colonization broad coastal zones of modern Finland are dominated by ethnic Swedes. But if the model I’m outlining above is corrected then Swedish is not the first dominant Indo-European language in Finland. Rather, an earlier Indo-European speaking population were absorbed by the Finns.

Swedish hegemony over Finland after 1200 was to a large extent a function of the fact that Swedes were a post-tribal population which were in the early states of constructing a nation-state. In Finland they encountered a tribal population which was easy to dominate, and integrated into a Swedish Baltic zone of rule. One ultimate basis of the Swedish superiority in domains of statecraft and social mobilization is probably economic, in that the ecology of Sweden was marginally more favorable to agriculture than that of Finland. The Finnish tribes were operating closer to the margins of subsistence, and at a lower limit of population density enforced by Malthusian strictures. But like the Thule conquest of Greenland I suspect the success of the Finnic tribes from the margins of Siberia is the very fact that they were masters of the cultural adaptations necessary for survival on the sub-arctic littoral of Eurasia. The Corded Ware people were like the Greenlanders of their era, agro-pastoralists who attempted to transfer a southern way of life in totality, but who ultimately were transformed and superseded.

• Category: Science • Tags: Finland 
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Ole Magga, Norwegian politician On this blog I regularly get questions about the Sami (Lapp*). That’s because I often talk about Finnish genetics, have readers such as Clark who are of part-Sami origin, and, the provenance and character of the Sami speak to broader questions about the emergence of the modern European gene pool. More precisely questions about the Sami are relevant to the broader nature of the Finnic presence in Europe, and their relationship to other Baltic and northern populations. Are these people “indigenous” to Europe, or relatively newcomers (prehistoric Magyar or Turks).? These questions are prompted by the peculiarity of their languages (as well as the physical appearance of some of the Sami). With Basque they are the only living non-Indo-European European languages whose origins are prehistoric (Magyar and Turkish were arrivals within the last 1,000 years).**

Because of affinities to other Uralic languages which are found in Central Siberia it has often been conjectured that the Finns, Sami, and Estonians are relative newcomers to Norden from that region. This has some equivocal support from Y chromosomal lineages. On the other hand, there are those who argue that the Finnic peoples were present in the north of Europe before the arrival of Indo-European speakers (often these are Finnish nationalists). This has some support from maternal lineages. Naturally, some have been tempted to synthesize these two genetic lines of evidence, and the linguistic affinities, to argue that Finns are a hybrid population of Asiatic men and Paleolithic European women! But we need to go further than uniparental markers, the direct male and female ancestral lines. We need to look across the broader swath of the genome. It just happens that a new paper was published in The European Journal of Human Genetics on autosomal Sami affinities to other populations, A genome-wide analysis of population structure in the Finnish Saami with implications for genetic association studies:

The understanding of patterns of genetic variation within and among human populations is a prerequisite for successful genetic association mapping studies of complex diseases and traits. Some populations are more favorable for association mapping studies than others. The Saami from northern Scandinavia and the Kola Peninsula represent a population isolate that, among European populations, has been less extensively sampled, despite some early interest for association mapping studies. In this paper, we report the results of a first genome-wide SNP-based study of genetic population structure in the Finnish Saami. Using data from the HapMap and the human genome diversity project (HGDP-CEPH) and recently developed statistical methods, we studied individual genetic ancestry. We quantified genetic differentiation between the Saami population and the HGDP-CEPH populations by calculating pair-wise FST statistics and by characterizing identity-by-state sharing for pair-wise population comparisons. This study affirms an east Asian contribution to the predominantly European-derived Saami gene pool. Using model-based individual ancestry analysis, the median estimated percentage of the genome with east Asian ancestry was 6% (first and third quartiles: 5 and 8%, respectively). We found that genetic similarity between population pairs roughly correlated with geographic distance. Among the European HGDP-CEPH populations, FST was smallest for the comparison with the Russians (FST=0.0098), and estimates for the other population comparisons ranged from 0.0129 to 0.0263. Our analysis also revealed fine-scale substructure within the Finnish Saami and warns against the confounding effects of both hidden population structure and undocumented relatedness in genetic association studies of isolated populations.

They had 352 Sami samples, and looked at ~38,000 SNPs. For the questions they’re focusing on 38 K SNPs seems fine. That’s enough to smoke out inter-population variation. In their paper they compared the Sami to the HGDP populations using standard techniques. Assuming 7 ancestral populations in the data set, this what ADMIXTURE popped out:


There is a definite “eastern” affinity among the Sami. Interestingly, it is broken down into a major and minor component. The major one is what is found among the Han, while the minor one resembles Native Americans. The natural interpretation for this is that what one is seeing is the shadow of the circumpolar northern Eurasian populations which spanned eastern Europe to Siberia. In comparison with other European populations the Sami affinity with Russians is clear, though interestingly they lack the “blue” component which peaks in northwest South Asian populations, which the Russians have, and Sardinians and French Basque lack.

samieigenTo the left you see a PCA which breaks out the top two components of genetic variation for the data set. The two axes seem to be roughly west-east, north-south. Whatever ancient affinities the Sami may have with Southern Europeans via mtDNA haplogroup U5, it is not evident in the total genome content. The position of the Sami between Russians and Orcadians (from north of Scotland) is probably attributable to the fact that the Sami share much genetically with other Scandinavians, who are closer to British populations than the Russians are.

I’m not sure these analyses really shed any light on the on the questions I mentioned earlier. The authors themselves note that the “eastern” component of the ancestry in the Sami is probably very old, so they may be an ancient stabilized hybrid population, mostly indigenous with a non-trivial exogenous element. That does not tell us whether Finnic languages are indigenous to Europe, or whether they are indigenous to Central Siberia (indigenous here is in reference to the Indo-European languages). Additionally, there is the matter that for such fine-grained questions the HGDP sample is suboptimal as reference populations. Dienekes Pontikos points this out:

It is unfortunate that they included Native American HGDP populations, but did not include the most relevant published data on Siberians that I first used to study population structure across north Eurasia here and here and here.

Hence, they discover a “Native American”-like component in Saami, which in all likelihood can be further resolved into Siberian-specific components utilizing the Rasmussen et al. dataset.

The “closest approximation” to the East Eurasian component in Saami in the HGDP panel are the Yakuts, but finer-scale analysis (see my previous posts) reveals that the Yakuts are made up almost entirely of an Altaic-specific component tying them to Turkic, Mongol, and Tungusic populations, while the eastern component in European Finns, Vologda Russians and Chuvashs has relationships with Central Siberians such as Kets, Selkups, and Nganasans, all of which are missing in this paper.

Below is a re-edited ADMIXTURE plot from Dienekes:


Note: There are many ways to spell Sami. They used two a’s, but I find that confusing, so I just used one in my text.

Citation: Maki-Torkko, Elina, Aikio, Pekka, Sorri, Martti, Huentelman, Matthew J, & Camp, Guy Van (2010). A genome-wide analysis of population structure in the Finnish Saami with implications for genetic association studies European Journal of Human Genetics : 10.1038/ejhg.2010.179

* Apparently “Lapp” is considered derogatory among Norwegians, though Finnish Sami refer to themselves as lappalainen. I will use Sami to avoid irritating Norwegian terminology police.

** I am implicitly excluding much of European Russia west of the Urals, but so be it.

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If you haven’t, you should keep an eye on DienekesDodecad Ancestry Project (RSS). The pilot phase of data collection is over, and the first population level statistics are now coming out. Of particular interest to me is a new analysis of various northern European ethnicities just published.

The samples used in this analysis are:

– 25 HapMap-3 White Americans. These are the Mormons of predominantly Northwest European heritage

– 5 Dodecad Project Finns

– 25 HGDP-CEPH Russians from Vologda, in north-central European Russia

– 12 Dodecad Project continental Germanics (Scandinavians and Germans)

– 10 Behar et al. (2010) Lithuanians

– 9 Behar et al. (2010) Belorussians

– 3 Dodecad Project Northern Slavs

Below are two visualizations of the genetic structure. First, an MDS. And second, a bar plot of ancestral quanta derived from ADMIXTURE. I’ve added some clarifying labels.



Remember that the data you input into these analyses shape the nature of the outcomes to some extent. All these populations are very genetically close when scaled to average worldwide inter-population genetic variation. So what Dienekes is smoking out here are subtle differences between relatively close groups.

The first clear result supports previous research using uniparental markers: the ethnogenesis of the “Great Russians” involved both demographic expansion, and, cultural assimilation. The process on the southern and eastern frontiers is well documented, because it continued into the early modern period via a series of private wars of expansion. Turkic and Ugric groups were defeated by “Cossacks”, and often themselves integrated into the Cossack population as it expanded further into Siberia and the Steppe. Lenin’s paternal grandmother for example is often claimed to have been a Kalmyk, a branch of the Dzungar Mongol Confederacy which had settled in the lower Volga region. Whatever the truth, Lenin’s father clearly had an Asiatic cast to his features. The ancestral quanta estimates always seem to show that Russians, though not other Slavs further to the west, seem to average around ~5% or so “eastern” ancestry (by analogy, this is about the amount of African ancestry in the typical Levantine Arab).

But the expansion into the Finnic north is less well documented. To some extent the process of Russification began far earlier, as even Kievan Rus at the turn of the first millennium has been claimed to have had Finnic elements (the Rus were Swedes, but they probably picked up Finns in their warbands as they swept south, in addition to the numerous indigenous Finnic groups in northeast Europe). Additionally, unlikely the Muslim Turks these Finnic groups were often small-scale societies without international connections or affiliation with any “higher civilization” which could serve as an oppositional ideology to Orthodox Russian culture. The wide geographic expanse of the Russian ethnos means that one must be exceedingly sensitive to sample representativeness. Readers of Russian or Finnish origin are often aware of which localities in northern Russia were only recently Slavicized, and so express caution in comments as to utilization of those samples as representatives of Slavs more generally.

The second peculiarity are the “Germans” who affiliate with the Finns in the MDS, and contribute to the Finnish element among the Germans. Dienekes says: “without revealing any information, I’ll just say that this is contributed primarily by 3 Dodecad Project members who deviate towards Finns and whose ADMIXTURE analysis shows a higher than expected Northeast Asian component. Their outlier status is also visible in the MDS plot.” By “Northeast Asian” he presumably means one of the 10 ancestral components he’d found in earlier analyses. Without any more information I assume there’s a high probability that these are simply Germanized part-Sami. Much of northern Scandinavia was inhabited by Sami down to the early modern period. For example, the Sami were ethnically cleansed and assimilated across the north half of what is today Sweden as late as the 1600s and 1700s. Though I haven’t done the requisite reading, I wouldn’t be surprised if this was just a function of more advanced farming techniques as well as hardy New World crops such as potatoes which pushed the possible limits of Swedish settlement north.

Finally, there’s a clear Finnic component in the results. As Dienekes noted this Finnic component itself may be a composite of East and West Eurasian elements, just as the South Asian component in Eurasia may be a composite of “Ancient North Indians” and “Ancient South Indians.” One thing to remember about the Finnic component is there’s evidence for a fair amount of genetic variation within Finland. Representativeness is probably key here, just as it is for Russians. Ethnic Finnish individuals with ancestry along the southern and western coasts probably have more affinity with Germanic populations than Karelians.

For many decades there have been arguments as to the provenance of the Finns. Specifically, are they outsiders to Norden who arrived from the east, bringing with them their language? Or are they are indigenous vis-a-vis Germanic speakers? The past is complex, so a simple model is going to shave off a lot of the detail, but I suspect that the truth is closer to the second. It seems that the Finnic groups, or at least their languages, have an ultimate origin in Central Eurasia after the last Ice Age. But they are possibly a circumpolar population which expanded north and practiced hunter-gatherer lifestyles following the ice sheets. Over time agriculturalists expanded north and squeezed them on the margin, but I believe there were natural ecological limits to the practice of techniques derived from Middle Eastern crops. Though northern Finns adopted some agricultural techniques, there was enough of a slowdown of the spread agriculture by Indo-European speakers and their precursors that they managed to hold their own in the north. In much of European Russia, and later in pre-19th century Finland, we see plenty of evidence of language-switching from Finnic to Indo-European (in Finland nationalism resulted in a back-switch over the past 150 years). If the Malthusian pre-modern age had persisted for another two or three centuries I would not be surprised if Finnic languages were totally absorbed by Russian and Scandinavian Indo-European dialects. As it is, 19th century language based nationalism stopped the process of elite culture assimilation, and in some cases reversed it (many elite Finland Swedes abandoned Swedish language and identity in the 19th and early 20th centuries).

Addendum: The picture I present above is simple, and I don’t believe it captures a lot of what happened. For example, from my reading there was a pause of about 1,000 years in the expansion of agriculture once it reached the Kattegat between Denmark and Sweden. I suspect that these long pauses were a function of ecology and geography, as they’re often just too long to be determined by social-political inertia. Additionally, it seems unlikely to me that the first agriculturalists in Europe were Indo-European speakers. Rather, that is possibly a subsequent linguistic overlay, especially in the western regions of Europe.

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Dienekes points to a new paper which highlights genetic variation in Fenno-Scandinavia (or in this case, Finland, Sweden and Denmark). A two-dimensional plot with the variation is pretty illustrative of what you’d expect:


Finns are genetic outliers in Europe, to some extent even in comparison to Estonians, who speak a very similar language. But, I wonder if the situation will change a bit when we have more samples from Finnic populations of northern Russia. Remember that the nature of these representations is sensitive to the variation which we throw into the equation in the first place.

• Category: Science • Tags: Finland, Genetics 
Razib Khan
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