Finnis, Saami, and Fennoscandian DNA

 

800px-Lenguas_finougrias
Distribution of the Finno-Ugric languages

I’ve long wondered which group arrived first in Europe: the Indo-Europeans or the Finno-Ugrics. Most Europeans speak one of the hundreds of languages in the Indo-European family tree, but a few groups speak languages from the mostly Siberian Finno-Ugric branch of the Uralic family.

(Sorry, guys, I’m out of practice writing and these sentences don’t sound good to me, but the only way to improve is to forge ahead, so let’s go.)

Major countries/ethnic groups that speak Finno-Ugric languages include the Finns (obviously,) Saami/Lapps, Hungarians, and Estonians. The most southerly of this family, Hungarian, arrived in the Carpathian Basin within the span of recorded History (in 894 or 895, followed by a few years of warfare to secure their territory,) but the origins of the other European Finno-Ugric languages remains mysterious.

Who arrived first, the Indo Europeans or the Finns? Did the Saami always live in their current homelands, or did they once range much further south or east? Did they migrate here recently or long ago (since the entire area was under ice sheets during the ice age, no one lived there tens of thousands of years ago.)

With the exception of Hungarian, these languages all hail from the far north (especially if you include the Samoyidic languages, which hail from north of Komi on the map,) a cold and forbidding land where herding, hunting, gathering, and fishing have remained the primary way of life until quite recently–the long winters making agriculture very difficult.

Lamnidis et al have a new paper out on Ancient Fennoscandian DNA that sheds a fascinating light on the subject:

Here we analyse ancient genomic data from 11 individuals from Finland and north-western Russia. We show that the genetic makeup of northern Europe was shaped by migrations from Siberia that began at least 3500 years ago. This Siberian ancestry was subsequently admixed into many modern populations in the region, particularly into populations speaking Uralic languages today. Additionally, we show that ancestors of modern Saami inhabited a larger territory during the Iron Age, which adds to the historical and linguistic information about the population history of Finland.

41467_2018_7483_fig1_htmlLet’s cut to the pictures, because they are worth a thousand words:

Just in case you are unclear on the geography, the Modern Saami come from northern part of the Finnoscandian peninsula. Six of the ancient remains came from Bolshoy Oleni Ostrov in the Murmansk Region on the Kola Peninsula–that’s the topmost dot on the map, now in Russia. These remains are very old–dated to about 1610-1436 BC.

Seven remains came from Levänluhta in Isokyrö, Finland, from a more recent burial dated to around 300-800 AD. (Actually, I think Levanluhta is a lake, so  This is the most southwestern burial on the map, in an area where the modern Finns live.

And the remains of two people came from a much more recent Saami cemetery in the Kola peninsula, Chalmny Varre, dating from the 17 or 1800s.

All of this DNA was compared against a variety of reference populations:

41467_2018_7483_fig2_html

(I would just like to pause for a moment to appreciate both the beauty and hard work that went into these graphs.)

PC2 graphs are a little complicated, but what we’re basically looking at (in color) are two different human population axes. They very roughly correlate to north-south (up and down) and east-west, (left to right), because people tend to be more closely related to their neighbors than people thousands of miles away, but there’s another, more fascinating story going on here.

On the right-hand side, we have a cline that maps very nicely to north and south, from the Yukagir–a people from a part of Russia that’s so far to the northeast it’s almost in Alaska–at the top and the Semende of Indonesia and the Atayal, an indigenous Taiwanese group, at the bottom. (Most Taiwanese you meet are either newly arrived Han Chinese or older Han Chinese; the aboriginal Taiwanese are different, but likely the ancestors of Polynesians.)

AsianDNA1
DNA from various Asian peoples

Most east Asian DNA shows up as a blend of these two groups (which we may call roughly polar and tropical). In the chart to the right, taken from Haak et al, the polar DNA is red and the tropical is yellow. So the up-down cline on the right side of the map represents which particular mix of Polar/Tropical DNA these folks have.

On the left side of the graph, we have a farming/hunter-gatherer cline. The first farmers hailed from Anatolia (now Turkey, but that was before the Turks moved to Turkey,) and subsequently spread/conquered most of Europe and probably a few other places, because agriculture was quite successful. So the orange is Middle Easterners; above them are southern Europeans like Albanians and Basques; then the English, French, Hungarians, Finns, etc; and finally some older burials of people with descriptive names like “Eastern Hunter-Gatherer” [EHG] or “Scandinavian Hunter-gatherers” [SHG].

(I have to constantly remind myself what these little abbreviations mean, but The Genetic Prehistory of the Baltic Region probably clears things up a bit:

Similarly, in the Eastern Baltic, where foraging continued to be the main form of subsistence until at least 4000 calBCE15, ceramics technology was adopted before agriculture, as seen in the Narva Culture and Combed Ceramic Culture (CCC). Recent genome-wide data of Baltic pottery-producing hunter-gatherers revealed genetic continuity with the preceding Mesolithic inhabitants of the same region as well as influence from the more northern EHG21,22, but did not reveal conclusively whether there was a temporal, geographical or cultural correlation with the affinity to either WHG or EHG.

The transition from the Late (Final) Neolithic to the Early Bronze Age (LNBA) is seen as a major transformative period in European prehistory, accompanied by changes in burial customs, technology and mode of subsistence as well as the creation of new cross-continental networks of contact seen in the emergence of the pan-European Corded Ware Complex (CWC, ca. 2900–2300 calBCE) in Central2 and north-eastern Europe21.

If you remember your Guns, Germs, and Steel, Turkish farmers had a really hard time getting their wheat to grow up in really cold places like Northern Russia,  Scandinavia and Narva (near the border between Estonia and Russia on the Baltic Sea,) which is why modern Finland is super poor and Turkey and Mexico, where corn was domesticated, are rich–what it doesn’t quite work like that?

Okay.

Anyway.

So most Europeans today are a mix of Anatolian farmers and various European hunter gatherer groups, with exactly how much you got depending a lot on whether the local environment was hospitable to farming. The pure hunter-gatherer genomes therefore show up as “further north” than the mixed, modern genomes of modern French and British folks.

There were additional events besides the Anatolian conquest that shaped modern European genetics–mostly the aforementioned Indo-European conquest–but the Indo-Europeans were at least part hunter-gatherer by DNA (nomadic pastoralists by profession,) so on this scale, their contributions look a lot like the older hunter-gatherer DNA.

So the interesting part of the graph is the middle, where all of the central Eurasian peoples are plotted. The purple band is various Finno-Ugric/Uralic speakers.

Hungarians are solidly in Europe because the ancient conquering Magyars left behind their language, but not much of their DNA (as we’ve discussed previously.) The Nganasan are one of the most thoroughly Siberian peoples you can imagine; they historically survived by hunting reindeer.

The green swaths (light and dark teal) are mostly Turkic-language speaking peoples; the Turkic peoples originated near Mongolia/Korea and spread out from there, mostly absorbing the DNA of whomever they encountered and passing on their language. The authors have also included Mongolian (which is not in the Turkic language family) in the light green group and some Caucuses groups in the dark teal.

Interestingly, the Yukaghir language (far upper right) is (according to Wikipedia,) potentially in the greater Finno-Ugric/Uralic family:

The relationship of the Yukaghir languages with other language families is uncertain, though it has been suggested that they are distantly related to the Uralic languages, thus forming the putative Uralic–Yukaghir language family.[5]

Based on the genetics, I’d say it looks very likely that the ancestors of Uralic-speaking Nganasan and the Yukagirs were conversing in some sort of mutually intelligible language. Unfortunately, Yukaghir has very few speakers and is likely to die, so there’s not much time to research it.

Finally in the Light Teal we have some groups from Pakistan/Afghanistan, like the Balochi.

(Note that all of the colors used in these studies are arbitrary; DNA doesn’t really have a color.)

So where do our ancient DNA remains fall on this graph?

Today, the Levanluhta site is in Finland, surrounded solidly by Finns (and maybe some random Scandinavians; who knows;) in 300-800 AD, the population was almost identical to modern Saami. So even though Saami and Finns both speak Finno-Ugric languages, the Finns replaces the Saami in this area sometime in the past 1,500 years or so.

One Levanlughta skeleton is an exception–the one marked Levanlughta_B; it is clearly closer to the Finns and English on this graph, but deeper mathematical analysis disputes this conclusion:

One of the individuals from Levänluhta (JK2065/Levänluhta_B) rejects a cladal position with modern Saami to the exclusion of most modern Eurasian populations. This individual also rejects a cladal position with Finns. We analysed low coverage genomes from four additional individuals of the Levänluhta site using PCA (Supplementary Figure 3), confirming the exclusive position of Levänluhta_B compared to all other six individuals (including the four low-coverage individuals) from that site, as is consistent with the ADMIXTURE and qpAdm results. The outlier position of this individual cannot be explained by modern contamination, since it passed several tests for authentication (see Methods) along with all other ancient individuals. However, no direct dating was available for the Levänluhta material, and we cannot exclude the possibility of a temporal gap between this individual and the other individuals from that site.

In other words, it is a mystery.

The remains from Chalmny Varre, which we know was a Saami cemetery, unsurprisingly cluster with the other Saami.

The Bolshoy remains, though, are quite interesting. They are shifted slightly in the direction of the ancient hunter-gatherers (perhaps their descendants, if still around, have mixed a bit with the agriculturalists.) Their physical location is about as far east as the Red Squares (ethnic Russians,) yet the more closely resemble the Mansi or the Selkups. (The modern Mansi live here; the modern Selkups live nearby.)

Getting down to the bar graphs, we see this data presented in a different way.

There are three groups that we can see contributing to most modern Europeans–Farmers, represented by the Orange LBK DNA; exclusively Indo-European, Green, notably not found in the Basque; and hunter-gatherers in Dark Blue. (Note that the ancestors of the Indo-Europeans hailed from the Central Eurasian steppes and so their DNA could have gotten around there, too.)

The modern Saami also have a Purple component to their DNA, which finds its highest expression in the Nganasan of far eastern polar Russia.

So the oldest burials–the Bolshoy–show no agricultural DNA. They are hunter-gatherers+Siberians, with a touch of Indo-European (probably from a steppe population that might have contributed to the Indos as well) and a bit they share with… the Karitiana of Brazil? Well, the Native Americans did descend from Paleo Siberians, so some genetic relatedness is expected.

The more recent burials, which cluster with the modern Saami, all show agricultural DNA–probably due to marrying a few of the local Finns/Russians who carry some agricultural DNA (who are almost genetically identical on this scale) rather than a pure LBK agriculturalist.

Here we see why the one outlier, Levanlughta_B, doens’t group with the Finns, either–modern Finns and Russians have some of that Nganasan-style Siberian DNA (probably from the same process that gifted Finnish/Russian DNA to the Saami), but Levanlughta_B doesn’t. Levanlughta_B looks more like the Baltic BA sample (Baltic Bronze Age.) Perhaps this individual was just a merchant, traveler, or lost–or represents a stage before the modern Finnish population had been produced.

The Finnish population itself is interesting, because it is genetically very similar to the Russian, but obviously speaks a language far more closely related to Saami (Lapp) than anything in the Indo-European tree. While it is therefore likely that the Finns replaced the Saami in the area around Lake Levanlughta, it seems also probable that in the process, they absorbed a large number of Uralic-speaking people. Who conquered (or married) whom? Did an ancient Balto-Slavic population move into what is now Finland, marry the local Saami girls, and adopt their language? Did an ancient Siberian population speaking a Uralic language conquer some ancient group of Russians, take their women, pass on their Uralic language, and later move into Finland and drive out the locals? Or perhaps something even more complicated occurred.

As for the Bolshoy, are they related (closely) to the modern Saami, or are they a group that simply died out?

The paper goes on:

 While the Siberian genetic component presented here [Purple] has been previously described in modern-day populations from the region1,3,9,10, we gain further insights into its temporal depth. Our data suggest that this fourth genetic component found in modern-day north-eastern Europeans arrived in the area before 3500 yBP. It was introduced in the population ancestral to Bolshoy Oleni Ostrov individuals 4000 years ago at latest, as illustrated by ALDER dating using the ancient genome-wide data from the Bolshoy samples. The upper bound for the introduction of this component is harder to estimate. The component is absent in the Karelian hunter-gatherers (EHG)3 dated to 8300–7200 yBP as well as Mesolithic and Neolithic populations from the Baltics from 8300 yBP and 7100–5000 yBP respectively8

Karelia is a region that crosses the border between Finland and Russia, so it is significant that this Siberian component isn’t found in ancient Karelian hunter-gatherers. Of course, the Siberians could have just been further north, however, the authors note that we have archaeological evidence of the spread of the Bolshoy people:

The large Nganasan-related component in the Bolshoy individuals from the Kola Peninsula provides the earliest direct genetic evidence for an eastern migration into this region. Such contact is well documented in archaeology, with the introduction of asbestos-mixed Lovozero ceramics during the second millennium BC50, and the spread of even-based arrowheads in Lapland from 1900 BCE51,52. Additionally, the nearest counterparts of Vardøy ceramics, appearing in the area around 1,600-1,300 BCE, can be found on the Taymyr peninsula, much further to the East51,52. Finally, the Imiyakhtakhskaya culture from Yakutia spread to the Kola Peninsula during the same period24,53. Contacts between Siberia and Europe are also recognised in linguistics. The fact that the Nganasan-related genetic component is consistently shared among Uralic-speaking populations, with the exceptions of absence in Hungarians and presence in the non-Uralic speaking Russians, makes it tempting to equate this genetic component with the spread of Uralic languages in the area.

The authors qualify this with a bit of “it’s complicated; people move around a lot,” but basically it’s People: not pots.

That was an enjoyable read; I look forward to the next paper from these folks.

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Genetic History of the Finno-Ugrics

Click for full size
From Haak et al.

I often run across people asserting that the Finno-Ugrics are “Mongols” or “Asian” or Chinese,”so today’s post is dedicated to the genetic history of the Finno-Urgrics.

The Finno-Ugrics (which includes the Udmurts but not the Uyghurs,) are people who speak Finno-Ugric languages such as the Khanty, Mansi, Hungarians, Maris, Mordvins, Sámi, Estonians, Karelians, Finns, Udmurts and Komis.[1]

Here’s a map:

Distribution of the Finno-Ugric languages
Distribution of the Finno-Ugric languages

Here are some pictures:

Charles Simonyi, Hungarian
Charles Simonyi, Hungarian
Presidents of the Norwegian Sami Parliament
3 Presidents of the Norwegian Sami Parliament
Erzaya women
Erzaya (Mordvin) women

Edit: I formerly had here pictures of Lennart Meri, President of Estonia, and Linus Torvalds, of Finland, but it turns out they’re actually ethnically Swedish. So I am substituting instead Finish figure skater Kiira Korpi and Estonian soldier Andres Nuiamae (killed in Iraq.) Hopefully they aren’t secretly Swedish.

Kiira Korpi, Finnish
Kiira Korpi, Finnish
Andres Nuiamae, Estonian
Andres Nuiamae, Estonian
Karelian women
Karelian women (Karelia is next door to Finland)
Janne Seurujärvi, Finnish Sami
Janne Seurujärvi, Finnish Sami
Udmurt people
Udmurt people
Khanty family
Khanty family
Mari man
Mari man
Komi People
Komi People
The two men on the right are from the Mansi.
The two men on the right are from the Mansi.

 

The Finno-Ugric languages are a subset of the Uralic Language family that excludes the Samoyedic languages.

Language is always a problematic base for claiming ethnic identity, because conquered people can easily learn a new language. African Americans today speak English, even though their ancestors weren’t Anglo-Saxons. Even the English aren’t majority Anglo-Saxon.

However, combining language, genetics, archaeology, and whatever historical records we have may result in a pretty trustworthy picture.

In this case, all of the Finno-Ugric people from within “Europe”–Finns, Estonians, Sami, Hungarians, etc.–all look very much like their neighbors. If you just randomly asked me to guess Torvalds or Meri’s ethnicity, the one thing I would not say is “Mongol.”

The groups that hail from Russia’s Siberia look more like other folks from Siberia.

Here are some genetic profiles (these are closeups of the graph at the top of the page):

DNA from various European peoples
DNA from various European peoples

With a few isolated exceptions (eg, the Basque,) almost all Europeans have a fairly similar genetic profile reflecting three main ancestral groups. The original “orange” and “blue” tribes have been identified via DNA sequencing of ancient European skeletons; at some point they seem to have merged. The “teal” component looks like it came in when a “blue” tribe migrated east and merged with a “teal” tribe, then came back and conquered the “orange-blue” tribes, resulting in blue-orange-teal tribes. (You can see the ancient skeleton sequences at the far left on the graph at the top of the page.)

A few groups don’t show this pattern–the Basques, for example, who don’t speak an Indo-European language, have very little teal. Based on this and other evidence, “Blue-Teal” tribe is therefore believed to be the original Indo-Europeans.

The Finns, Estonians, Mordovans, and Sami all have the blue, teal, and orange of other European groups and they also share a bit of red that is also found in the Russians. This group (including Russians) also seems to have a bit more blue than the other Europeans. The Sami in particular seem to have a fair amount of this red; they look rather similar to the Chuvash, a Russian ethnic group:

World's most famous Chumash
World’s most famous Chuvash

The Hungarians have a tiny bit of red if you look very closely, but this is not much at all; several other groups have similarly tiny smidgeons of red and no claims of Finno-Ugric ancestry. The Wikipedia page on Hungarians also states that, despite the well-documented Magyar invasion around 1100, modern Hungarians appear to be genetically continuous with pre-Magyar Hungarians. Perhaps there were never enough Magyars to have much of an impact besides imparting their language; or they just failed to reproduce and so gradually died out in their new land, leaving their language behind; or the red-DNA contained specific adaptations that help people survive in the arctic, and so have been selected against in warmer Hungary; or perhaps the Magyars themselves never had much of the red-DNA for whatever reasons.

By contrast, various tribes from central Eurasia (the Chuvash may perhaps be included) show quite mixed ancestries:

DNA from various steppe peoples
DNA from various Eurasian peoples

The Hazara are from Pakistan/Afghanistan; the Uygurs are primarily from the far western end of China; Turkmen and Uzbeks you’re probably familiar with; and the Evens are a Siberian people who live in far eastern Russia.

The Mansi are one of our Finno-Ugric people, with large sections of blue, red, and even a little teal. Based on the photos, I’m not surprised to see essentially a mix of Siberian and typically European DNA. The Wikipedia has this to say about their origins:

“The ancestors of Mansi people populated the areas west of the Urals.[3] Mansi findings have been unearthed in the vicinity of Perm.[3]

In the first millennium BC, they migrated to Western Siberia where they assimilated with the native inhabitants.[3] According to others they are originated from the south Ural steppe and moved into their current location about 500 AD.” (wikiepdia)

The Selkups are a Samoyed people–the Samoyed languages are cousins to the Finno-Ugric languages under the larger family of Uralic Languages.

It looks like the original Finno-Ugric speakers who settled in Finland, Lapland, Estonia, etc., looked like the Mansi or Selkups, this might explain the slightly higher quantities of blue in these groups.

The red DNA reaches its greatest dominance in the Nganasan, a Samoyedic people living in north central Siberia:

DNA from various Siberian Peoples.
DNA from various Siberian Peoples.

An old picture of the Nganasan:

Ngasani People
It’s cold there.

(The Yukagir are from further east in Siberia than the Nganasan (the olive-brown shade is shared with the Eskimo;) the Daur and Oroqen live in inner Mongolia, China; the Henzhen live in northern Manchuria/the region north of there along the Sea of Okhotsk; the Ulchis live just north of them. The Tubalar and Altaian people hail from the meeting point of Russia, China, Mongolia, and Kazakhstan; the Dolgans from north central Siberia; the Yakuts live to their east.)

The red/yellow combination is found throughout most of the “Asian” countries–Japan, China, Korea, Mongolia, etc., but not in Cambodia or Thailand. You can see them on the big chart at the top. The two pure yellow groups, the Ami and Atayal, are indigenous people of Taiwan.

The Red, therefore, is found in large quantities in Siberia/polar peoples. In Asia it mixes with the yellow, with the ration of yellow/red increasing as you go south. Red finds its maximum in far northern Siberia, and yellow in Taiwan. I therefore speculate that the red started in Siberia and worked its way south, while the yellow started somewhere around southern China and moved outwards from there.

The Blue is found in all Europeans but is rare in the Middle East; it appears in small quantities in Central Asia, India, and Siberia. Small quantities could just be the result of thousands of years of people moving around ancient trade routes, but the relatively larger quantities in Siberia seem less likely to be the result of trade.

Teal appears to be found in all Indo-European and Middle Eastern regions; it is even more wide-spread than orange, which never made it to India.

Therefore I suspect that a band of blue and a band of red people merged to form the original Uralic people from which the Finno-Ugrics later split off. (The lack of red in Hungary could be due to the branch which eventually became the Magyars having split off before the red-blue merger, but they lack the extra blue found in Finns, so this seems unlikely. Plus, their language would be quite different from the other Finno-Ugric languages if they had, perhaps similar to the relationship between Anatolian and the other Indo European languages.) More likely, as the original Red/Blue people spread out across Siberia, mostly toward Europe, they were spread thinner and thinner, or mixed with and taught their languages to more and more new until they were only a small percent of the total population, leaving behind only a smidgen of their DNA in Finnland, Estonia, and Hungary.

Here is a map of the distribution of Haplogroup N, which appears to have emerged about 20,000 years ago:

Distribution haplogroup N
Distribution Haplogroup N

According to Wikipedia, Subtype N-P43 is estimated at 4,000 to 6,000 years old, frequently among the Samoyedic peoples, with a sub-clade common in Finno-Ugric and other Uralic speakers in Europe. Additionally,

“The subclade N-M178 … has higher average frequency in Northern Europe than in Siberia, reaching frequencies of approximately 60% among Finns and approximately 40% among Latvians, Lithuanians & 35% among Estonians (Derenko 2007 and Lappalainen 2008).

“Miroslava Derenko and her colleagues noted that there are two subclusters within this haplogroup, both present in Siberia and Northern Europe, with different histories. The one that they labelled N3a1 first expanded in south Siberia (approximately 10,000 years ago on their calculated by the Zhivotovsky method) and spread into Northern Europe where its age they calculated as around 8,000 years ago.”

Here’s a beautiful map showing the spread of Y Chromosome Haplogroups all over the world:

World map of Y-DNA Haplotypes
Isn’t it beautiful?

Since Haplogroup N is found on the Y chromosome, this probably implies armed invasion that resulted in many of the local men dying and the invaders marrying (or raping) the remaining women.

Note that this scenario does not depend on whether the Indo-Europeans or Finno-Ugrics arrived first; it merely describes their relative ratios in the population. We know they arrived after the Indo Europeans in Hungary, for example, but the Sami are considered the indigenous people of Finno-Scandia. Genetically, the Sami have some teal and orange, which the Red-Blue people basically lacked, so they have at least some Indo-European; just eyeballing the graph, it looks like the Sami are a little more than half Indo-European and a little less than half Red-Blue people.

Overall: the Finno-Ugrics living in Europe proper are genetically closest to other Europeans; their Siberian component is quite small. The Sami are the one exception, with a larger chunk of Siberian DNA, but they are still mostly European.

The Finno-Ugrics who live within the heart of Russian Siberia, however, appear to have quite a bit more Siberian DNA, some European, but not Indo-European DNA.