Ancient DNA From Siberia As A Source For Modern Populations

Different strains of Siberian ancestry spread east to the New World and west to Scandinavia.

Human populations across a vast area in northern Eurasia, from Fennoscandia to Chukotka, share a distinct genetic component often referred to as the Siberian ancestry. Most enriched in present-day Samoyedic-speaking populations such as Nganasans, its origins and history still remain elusive despite the growing list of ancient and present-day genomes from Siberia. 

Here, we reanalyze published ancient and present-day Siberian genomes focusing on the Baikal and Yakutia, resolving key questions regarding their genetic history. First, we show a long-term presence of a unique genetic profile in southern Siberia, up to 6,000 yr ago, which distinctly shares a deep ancestral connection with Native Americans. Second, we provide plausible historical models tracing genetic changes in West Baikal and Yakutia in fine resolution. Third, the Middle Neolithic individual from Yakutia, belonging to the Belkachi culture, serves as the best source so far available for the spread of the Siberian ancestry into Fennoscandia and Greenland. These findings shed light on the genetic legacy of the Siberian ancestry and provide insights into the complex interplay between different populations in northern Eurasia throughout history.

Haechan Gill, Juhyeon Lee, Choongwon Jeon, "Reconstructing the Genetic Relationship between Ancient and Present-Day Siberian Populations" 16(4) Genome Biology and Evolution evae063 (March 25, 2024) https://doi.org/10.1093/gbe/evae063

The Makings Of Hungary

Hungary is one of the most notable examples of a population that experienced elite driven language shift (to the Uralic language of their Magyar conquerors), without experiencing much population genetic (a.k.a. demic) change in the long run. Razib Khan has posted a stunning short history of it.

True wholesale population replacement has been rare in the post-Bronze Age era, when the broad outlines of the genetic makeup of modern Europe were largely established, although the conquering Turks in Anatolia and the Uralic people in Finland both gave rise to more genetic introgression than the Magyars in Hungary. (The non-conquering Jews and Gypsies in medieval Europe roughly contemporaneously with the Magyars in Europe, had even less of a demic impact on the general European population.) 

We know a lot about how this happened because it happened in the historic era recorded by contemporary scribes, and because ancient DNA for Conqueror graves in Hungary rule out alternative hypotheses. 

Linguistic and genetic data show that their Magyar language and people were more closely connected to that of the modern Mansi and Khanty people of Siberia (who combined number under 50,000 speakers today), than to the Finns, Saami, Karelian and Estonian peoples in the vicinity of the Baltic Sea, which split linguistically around 2000 BCE to 1000 BCE (with genetic data favoring a more recent date, around the time of Bronze Age collapse, than linguistic estimates).

But, the Magyar language also borrowed words from Iranian and Turkish peoples (probably in that order), who were also nomadic pastoralists of the Eurasian steppe, the most western extent of which includes Hungary.

Prior to their entry into Europe just before 900 CE, they were subordinate partners in coalitions with Iranian and then Turkish nomadic pastoralists. The Huns, who harried the late Roman Empire, in contrast, were linguistically and ethnically Turkish. Notably:

The first attestation of Magyars refers to their service as mercenaries under the Bulgar Khan in 831 AD. From this date, historians can trace the migration of the Magyars westward in annals of Byzantine historians, until they enter Europe just before 900 AD. Despite their coexistence with Iranian and Turkic people for thousands of years, and a strong cultural imprint of both of these groups on the Magyars, they somehow managed to preserve their ethnolinguistic identity. The same cultural continuity persists down to the present. While the ancient Bulgarians spoke a Turkic language, rather than the Slavic speech dominant there today, the modern Hungarians speak a language descended directly from pastoralist Magyar forebears.

Like most conqueror peoples over time, and certainly all Uralic conqueror peoples. the Uralic contribution to Hungary, in addition to being small, was male dominated with invading men marrying local wives, as shown by comparing uniparental genetic markers (Y-DNA comes from fathers, mtDNA comes from mothers):

[G]raves of elite Magyars, the Conquerors, dated to the period between 1000 and 1200 AD. About 30% of the Y-chromosomal lineages in these graves were East-Eurasian haplogroups, where we see fewer than 5% among modern Hungarians. The mtDNA from these samples shows a substantial East-Asian origin, nearly 40%, as opposed to the 1% in modern Hungarians. . . . Even the Siberian Mansi and Khanty are only 25%-33% East Eurasian in their mtDNA haplogroups, whereas medieval Magyars were 10-20%.

 

The only notable exception to that rule that comes to mind is Japan, where indigenous fisher-gatherer Jomon source Y-DNA is found in a very substantial proportion of the modern Japanese population, while Jomon source mtDNA is much less frequent (despite the fact that there are almost no traces of the Jomon language in the modern Japanese language).

Also notable is the the survival of pagan beliefs to a late date, and even into the modern era, more closely tracks mtDNA than Y-DNA, among Uralic peoples. This lines up with the evidence from psychology and sociology that women tend to be more religious than men.

The demise of the Magyar elite, and the survival of the Magyar language can probably be attributed to a couple of key factors. 

First, the Magyar elite swiftly converted to Western Christianity and allied itself with Western Europe, after a half century period of raiding across Western Europe all of the way to Spain, the boot of the Italian Peninsula, and Belgium, against weak, disorganized, divided and undisciplined local feudal nobles. 

After they converted, they joined the Crusades, defended Europe from the Mongols (first losing in 1241-1242 CE only to win against a second round invasion in the 1280s), and then defended Europe from the Ottoman Muslims. 

Second, the Magyar elite actively participated in these wars and their own internal civil wars, not infrequently suffering catastrophic defeats.

In 1241 and 1242, led by the general Subetai, the Mongols ravaged Hungary. At the Battle of Mohi, nearly the whole Hungarian army was slaughtered, up to 10,000 men. In the year after this defeat, as much as 25% of the population may have died due to the chaos ushered in by Mongol units having free rein over the Pannonian plain. . . .

Between 1396 and 1526, the Hungarians fought the Ottoman Turks for supremacy, ultimately losing. At the Battle of Mohacs, 14,000 Hungarian soldiers died, 1,000 nobles were killed, and 2,000 captured prisoners were executed. Hungary was partitioned between the Habsburgs of Austria and the Ottomans.

The rulers of Transylvania maintained some semblance of Magyar independence in their mountainous domain. Nevertheless, it is notable that the first two ruling Houses of Transylvania’s elective monarchy, the Zápolya and the Báthory, were not descended paternally from Conqueror lineages. Rather, they were Croat and German respectively.

The end result was that in terms of genetic distance, the Hungarians are closer to their European neighbors, bear few traces of the Conquerors, and few genetic traces of Uralic populations (another surprising face is that the Swedes bear genetic similarities to the Welsh people, despite the Welsh people have Celtic linguistic affinities that persist to today and the Swedes speaking a Germanic language).

The Grave Of An Intersex Warrior In Finland

A single individual's tomb in Finland in the municipality of Hattula, which was radiocarbon dated to between 1040 and 1174, contained remains buried with prestige weapons usually associated with men, but also jewelry and other grave goods usually associated with women (via Bernard's Blog in French).

Investigators considered the possibility that it was the grave of a man and a woman, but there was only one set of remains and it was too small for that. They considered the possibility as well that it could have been a woman who had a leadership position for whom the weapons and male associated grave goods were symbols of authority rather than tools that the decedent used in life.

The bones themselves were not in sufficiently good condition due to decomposition to determine the decedent's gender, but two femurs yielded some ancient DNA evidence. As Bernard relates:

there was very little preserved DNA and the authors were only able to test the gender of the individual. The results showed that this individual has an aneuploid karyotype: XXY (Klinefelter syndrome).

This syndrome is found in one out of 1000 to 2000 births, so while it is rare, it is an exceptional genetic condition that is observed now and then. 

Bernard identifies three other cases in the last three years where there is ancient DNA indicating either chromosomal abnormality (aneuploidy), or a gender identity suggested by grave goods that is contrary to what the person's genes would suggest.

Klinefelter syndrome increases the risk of someone's premature death somewhat, but is hardly a death sentence with people who have the condition usually living to adulthood and not discovering that they have the condition until then. According to Wikipedia (linked above):

The individual is then of a masculine character, but infertile. . . .

Under the name of Klinefelter syndrome we group together all or part of all of the following symptoms, a variability of expression being often observed and all the problems of life that cannot be linked to this syndrome: size on average larger than the siblings, possible delay in puberty, possibility during childhood of learning disabilities of language or reading, size of the testicles smaller from puberty, possibility in adolescence if there is a lack of testosterone of a low hairiness, lack of muscle tone, development of mammary glands or gynecomastia, brittle tooth enamel and osteoporosis adulthood.

The atypical expression of this syndrome therefore explains the frequent delay in its diagnosis, which is often done only as part of a search for sterility.

In other words, people with this genetic type, not infrequently, present as intersex individuals that don't fit nearly as neatly into categorization as male or female as the vast majority of people do. 

It seems clear that this medieval Finnish warrior presented in that way, and nonetheless lived a life ending with recognition as a high status member of this community.

A Principal Component Analysis Of Major Branches Of Eurasian Population Genetic Diversity

Via Razib Khan.

This chart is a two dimensional chart of the relative genetic similarities and relationships of many modern and one group of ancient Eurasian populations from a paper analyzing the genetics of the Saami people of Northern Scandinavia (especially Finland with which they have the strongest linguistic ties), who are now herders and historically were one of the last hunter-gatherer populations of Europe.

Finnish Great Winter Myth Partially Validated

Almost every society has a layer of "legendary history" in which the line between fact and fiction is blurred with stories that are on their face impossible in some respect have kernels of reality that undergird them. The Finnish people are not an exception to this rule. 

Recent archaeological work and chemical analysis reveals a time period following a major volcanic eruption that corresponds to "Fimbulwinter," a legendary period with three successive winters (a legend that was probably also an inspiration for the modern fantasy "Game of Thrones" world) and a location the corresponds to the sacred spring containing lakes of Sami legends. Wikipedia explains the legend as follows:

Fimbulwinter is the harsh winter that precedes the end of the world and puts an end to all life on Earth. Fimbulwinter is three successive winters, when snow comes in from all directions, without any intervening summer. Then, there will be innumerable wars. 

The event is described primarily in the Poetic Edda. In the poem Vafþrúðnismál, Odin poses the question to Vafþrúðnir as to who of mankind will survive the Fimbulwinter. Vafþrúðnir responds that Líf and Lífþrasir will survive and that they will live in the forest of Hoddmímis holt.

The mythology might be related to the extreme weather events of 535–536, which resulted in a notable drop in temperature across northern Europe. There have also been several popular ideas about whether the particular piece of mythology has a connection to the climate change that occurred in the Nordic countries at the end of the Nordic Bronze Age from about 650 BC. Before that climate change, the Nordic countries were considerably warmer.

It also bears noting that plagues and famines and major political upheavals and wars often coincide with climate events, with this case again proving that rule.

On a more personal note, the site discussed is quite close to my mother's family's ancestral homeland in Finland.

In the middle of the Levänluhta era, the most severe climate disaster in 2,000 years took place. In the 540s volcanic eruptions initiated a cold and dark period lasting several years, possibly reflected in folktales across the northern hemisphere. Recently the researchers working in the project headed by Oinonen have found a link between the disaster and a reduction in the quantity of light observed in the carbon isotopes found in the annual growth rings of trees in Lapland between 541 and 544. 

"If you want to date Fimbulwinter, the three successive winters mentioned in Scandinavian sagas, this is the best candidate," Oinonen considers. 

Fimbulwinter has been said to have caused a collapse in farming in the areas surrounding Sweden and Estonia. However, the ratio of food from terrestrial sources consumed by the Levänluhta population does not decrease after this period. Instead, the group relying heavily on marine food starts to fade out. The largest group of people continued to supplement their diet with marine food, actually increasing its presence in the human remains buried in the middle of the 7th century. Protein-rich food indicates produce derived from animals, and it appears that, instead of farming, most of the population probably based their sustenance on animal husbandry and hunting. In fact, fur trade has traditionally been thought as the source of wealth during the Iron Age in these southern roots of the Suomenselkä water divide. 

Prior genetic research and place name data indicate a connection between the Levänluhta population and the Sámi. Signs of the diverse livelihoods of Iron Age Sámi have also been previously observed in Sweden on the same latitudes. Indeed, the researchers are considering whether the lake burial site of Levänluhta could be a manifestation of sáivas, the sacred spring-containing lakes in the Sámi mythology.

From here, based upon the following paper: 

Levänluhta is a unique archaeological site with the remains of nearly a hundred Iron Age individuals found from a water burial in Ostrobothnia, Finland. The strongest climatic downturn of the Common Era, resembling the great Fimbulvinter in Norse mythology, hit these people during the 6th century AD.

This study establishes chronological, dietary, and livelihood synthesis on this population based on stable carbon and nitrogen isotopic and radiocarbon analyses on human remains, supported by multidisciplinary evidence. Extraordinarily broad stable isotopic distribution is observed, indicating three subgroups with distinct dietary habits spanning four centuries. This emphasizes the versatile livelihoods practiced at this boundary of marine, freshwater, and terrestrial ecosystems. 

While the impact of the prolonged cold darkness of the 6th century was devastating for European communities relying on cultivation, the broad range of livelihoods provided resilience for the Levänluhta people to overcome the abrupt climatic decline.

Markku Oinonen, et al., "Buried in water, burdened by nature—Resilience carried the Iron Age people through Fimbulvinter." 15(4) PLOS ONE e0231787 (April 21, 2020) (open access). DOI: 10.1371/journal.pone.0231787

The introduction of the paper (citations omitted) provides helpful context:

Mediterranean historical sources identify a mystery cloud obstructing the Sun at AD 536/537. The year without the Sun is observed in tree rings as a negative growth anomaly throughout the Northern Hemisphere (NH). An even larger tree-growth decline is observed during the AD 540s, and discussion of the anomaly has evolved from a single mystery cloud to a decade-scale climatic catastrophe as a result of multiple volcanic eruptions of AD 536–550. The anomalous years probably triggered a longer climatic disturbance lasting until AD 570, or even beyond, known as the “Late Antique Little Ice Age” (henceforth LALIA). This cold and dark period—further stressed by the breakout of the Justinian Plague in AD 542—coincided with the rise and fall of empires, human migrations, and political upheavals. 

The volcanic winter of the AD 540s, recently linked to the Ilopango eruption in El Salvador, featured a drastic reduction of solar irradiance for several years in AD 541–544. This was observed in Northern Finland, reducing the temperature and photosynthetic rate, and thus primary production. The severe effects of distant volcanic eruptions on agricultural communities across the NH have been recently demonstrated—crop losses and famines in 17th century AD Finland have been shown to have resulted from tropical volcanic eruptions. As the AD 536–550 climatic catastrophe was the most severe in the last 2500 years, one can hypothesize that its consequences for communities relying on cultivation were even more catastrophic. Indeed, the abandonment of Iron Age settlements in Scandinavia and Estonia after AD 550 may have been a result of the climatic downturn and cultivational challenges. 

This work studies the dietary habits and livelihoods of a population that experienced the AD 536–550 climatic downturn in Southern Ostrobothnia, Finland. Early scholars only fragmentarily tell of the people of the North, and these frontiers of ancient civilizations have remained a kind of an unknown otherworld, associated with mystical elements of the seasonal change from long darkness to midnight sun. However, already paralleling the early civilizations of the Mediterranean, the Northerners were involved in cultural connections and trading networks across the Europe, linking them to the world system. In the Southern Ostrobothnian inland, nearly a hundred Iron Age (ca. AD 300–800) individuals were buried in water in Levänluhta. The site, presently consisting of three springs releasing red iron-containing water, has been one of the most intriguing archaeological mysteries within Finland. The lives of these people coincided with the greatest global-scale changes of the Common Era: a land uplift of one meter per century and an abrupt period without the Sun. Using dietary stable isotope studies combined with chronological analyses on human bone collagen samples, it is possible to understand the factors that allowed the Iron Age populations at the northern edge of European civilization to carry themselves through these challenges. A synthesis of this watery burial of worldwide uniqueness is established by tying the existing multidisciplinary knowledge together.

Another analysis of the same event can be found here. 

Admixture Analysis Of Global and Ancient Genomes

What Is Admixture Analysis?

A computer program called Admixture uses a mathematical algorithm whose application to genome data includes a little art as well as a lot of science, to fit data from large numbers of genomes as well as possible into a model.  This model assumes that each person's genome is mix of different proportions of a preset number of hypothetical ancestral populations determined in a manner that maximizes the quality of the fit using linear algebra.  It is also possible to tweak the program, for example, by designating some real individual as the exemplar of an ancestry component, rather than having the computer derive its clusters entirely without outside input.

I don't know precisely which choices were used to generate the latest and greatest result from Eurogenes analyzing a sample of more than 2059 modern and ancient genomes that maximally capture all varieties of modern human genetic variation in an Admixture run at K=10 (i.e. requiring the program to fit the individuals in the sample into percentage contributions from ten ancestral populations generated by the computer).  This sample includes almost every available complete ancient genome (which number in the hundreds) and some global databases of genomes that are widely used in the professional literature (such as the thousand genomes database) to represent the rest of the world.

An Example Of Admixture Analysis

For example, the first ten samples in his analysis are African-Americans from Denver (because AA for African-American comes first in the alphabetical listing).  For each individual a percentage of ancestry from each of ten groups that have been labeled for convenience after the fact to give a sense of where that component is most often found.  These categories are (with abbreviations spelled out):

1. Middle Eastern
2. San Bushman
3. American Indian
4. Northern Siberian
5. East Asian
6. Hindu Kush
7. Sub-Saharan African
8. European Hunter-Gatherer
9. Oceanian.
10. East Siberian.

For example, individual number 15 from the African-Americans from Denver sample included in his Admixture run is determined to be:

88.6% Sub-Saharan African
8.5% European Hunter-Gatherer
1.6% Middle Eastern
0.5% San Bushman
0.4% American Indian
0.4% Hindu Kush

The proportion of the other four ancestral components is negligible.

In terms an average person would understand, this individual is 89% black, 10.5% white, and 0.4% American Indian, these proportions reflect the American reality that African-Americans typically have higher proportions of African ancestry, and lower proportions of non-African ancestry, than is typical of people with some African ancestry in Latin America and the Caribbean.

This individual's African ancestry overwhelmingly from populations more like typical Niger-Congo language speaking West Africans and much less like "Paleo-African" populations like the Khoi-San bushmen of the Kalahari desert and the Pygmies of the Congo jungle.  This reflects the typical sources of individuals in the American slave trade.

The mix of "European hunter-gatherer" ancestry, "Middle Eastern" ancestry, and "Hindu Kush" ancestry in the "white" component of his ancestry is roughly in line with what you would expect in someone of Scottish origins (which is typical of Southern whites in the U.S., many of whom were Scotch-Irish).

Small but measurable amounts of Native American ancestry are common in African Americans.

All of this is exactly what one would expect from other data in a typical African-American from Denver.  One of the African-Americans from Denver in the sample, however, who is an exception, is almost half-white and not quite half-black, and is probably light skinned relative to a typical African-American in Denver.

Similar break downs are available for all 2059 people, modern and ancient alike, in the database, although it takes a certain amount of familiarity with how the individuals are identified to know which are modern, which are ancient, and what modern ethnic groups or his archaeological cultures are represented by the label given to an individual in the spreadsheet.

Insights: Genetic Variation Is Highly Structured And Far From Maximal

The fact that a reasonably accurate description of someone's ancestry, relative to seven billion or so living people and untold numbers of deceased individuals who preceded us, can be summed up with a fair degree of specificity with percentages of ten ancestral components, is itself remarkable.

The reality of human genetic variation observed in the real world is dramatically narrower than the default assumption that each SNP is random relative to the entire human population, in which each individual would be their own "special snowflake".  Each individual is unique, but the differences within ethnic communities often colloquially described in terms of race, linguistic affiliation and ancestral religious identification, are often quite subtle.

Indeed, vast areas of the human genome are totally ignored by people interesting in genealogy, forensic applications, or ancient DNA research, because all modern humans are identical in that part of the genome.  Indeed, a significant component of the part of the genome that has reached fixation in modern humans has also reached fixation in archaic hominins (like Neanderthals and Denisovans for which we have ancient DNA to compare to), for primates, for mammals, and for vertebrates.  Indeed, all multi-celled animals, no matter how primitive, share more than 40% of their DNA at locations that are so functionally important that they have reached fixation.  The more that parts of a person's genome are ancestry informative and variable within modern humans, the more likely it is that those parts of a person's genome are not important to evolutionary fitness.

Every Ethnicity Has At Least One Distinctive Genetic Profiles

Still, a person's genomes are ancestry informative and can often pin down a person's likely self-identified ethnicity, race, ancestral religious affiliation and familial place of origin with great specificity, in Europe, for example, pinning down the likely place of origin of someone with ancestors all from the same region, to a location within a hundred miles or so.

For example, when I compared the mix of "white" ancestral components of the African-American individual from Denver described above, it was possibly to obviously rule out a white ancestor from the Near East, Southern Europe or Iceland (because the Middle Eastern component was proportionately too small compared to the other ancestral components), or from Russians (who generally have a significant Northern Siberian component).

Similarly, a recent African immigrant from Somolia would have about ten times or more as much of the San Bushman ancestry component as someone descended from slaves from the American Southeast, as is typically the case with African-Americans, even though the former would not be unheard of in Denver's African-American population.

There are some cases where a population that culturally is just one ethnicity, such as "African-Americans" in the United States, can actually have several distinct genetic profiles (e.g. Ethiopian-Americans and other Africa-Americans in Denver might be classified socially as both being African-American, but would have different genetic profiles).

This reflects that fact that the history of human migration and diversification through mutations, isolation into separate reproductive populations, adaptation to new environments, and admixture, has been highly structured and has involved a finite number of populations, that these populations had enough time to homogenize while isolated from other populations, and that there have been a modest and finite number of significant admixture events in modern human history.

Few People Are Pure Types

Another descriptive observation of the data set is that at the K=10 level, few individuals are "pure types" with 99%+ ancestry from a single ancestry category.

There is no one in the sample with more than 83.1% Middle Eastern ancestry. There is no one with more than 86.6% Hindu Kush ancestry (a component that would be more accurately described as Kalish).

There are 2 of 2059 individuals with more than 99% European hunter-gatherer ancestry from many thousands of years ago (both of whom are ancient DNA samples with sequences released within the last couple of years).

There are 8 of 2059 individuals with more than 99% San Bushman ancestry.  There are 35 of 2059 individuals with more than 99% American Indian ancestry.  There are 6 of 2059 individuals with more than 99% North Siberian ancestry.  There are 19 of 2059 individuals with more than 99% East Asian ancestry.  There are 79 of 2059 individuals with more than 99% Sub-Saharan ancestry.    There are 14 of 2059 individuals with more than 99% Oceanian component (a component that would be more accurately described as Papuan).  There are 2 of 2059 individuals who are more than 99% East Siberian.

Thus, in a sample of 2059 modern and ancient individuals, only 163 are "pure type" individuals (less than 8% of the sample), while the rest have at least two measurable ancestral components in their genomes.  Two of the ten ancestral populations have no "pure type" representative (Middle Eastern and Hindu Kush), and a third has no modern "pure type" representatives.

Also, it is worth recognizing that representation in the sample is not proportionate to modern population size, and indeed, is deliberately chosen to over represent genetically distinct populations.  This is a maximally diverse sample, rather than a representative sample of human genetic diversity.

The populations that are pure types for the San Bushman, for Northern Siberians, and East Siberian (three of the seven ancestral types with modern representatives) are tiny relict populations that subsist in large part on hunting and gathering.

Pure type Papuans are present only on an island between Australia and China that has little contact with the outside world and uses traditional indigeneous non-mechanized agriculture.  Only a very small percentage of Native Americans a "pure blooded" and those who are generally live in economically marginal reservations or remote jungles or mountain villages.  The "pure type" individuals in all of these populations combined in the entire world alive today make up considerably less than 1% of the world's entire population.

Only the Sub-Saharan component and East Asian components have pure type individuals who are present in modern populations that are not tiny and marginalized.

No One Has Measurable Amounts Of All Components:

While few individuals are "pure types" almost no one has measurable contributions to their genome (defined as more than one part per 100,000) from all ten of the globally determined ancestral components.

I was able to identify only six individuals who had measurable amounts of nine of the ten ancestral components in the sample: Turkish4BA57, SaudiA7, HGDP00148 (Makrani, a South Asian Muslim ethnicity), Jordan646, usb25 (an Uzbek) and Yemenese1529.  Given that all of these individuals are from predominantly Muslim areas, it is plausible to infer that global, religiously mandated pilgrimages to Mecca have led to trace admixture in many Muslim populations from all over the Muslim world, and indirectly from almost everywhere around the globe.

Everyone in the sample of 2059 individuals (modern and ancient), except the 163 pure types and 6 nine type individuals, had two to eight ancestral components, and the lion's share have fewer than eight.

With a cutoff that excludes negligible contribution from an ancestral component (say, e.g., less than 0.1%), there would be no individuals with nine ancestral components, and the average number of ancestral components per person would be much smaller.

In any given region or ethnicity, individuals typically have slightly varying percentages of just a few components.

For example, of the seven Finnish people in the sample, all have generally similar percentages of four of the ten ancestral components (Middle Eastern 9.1%-14.8%, Northern Siberian (4.6%-9.7%), Hindu Kush (4.8%-13.0%), European Hunter-Gatherer (66.6%-76.2%).  Six of the seven had small amounts of Eastern Siberian 0.7%-2.5% ancestry, and four also having trace amounts of American Indian ancestry (0.2%-1.2%) including the one with non East Siberian ancestry.  None of the Finnish individuals had any San Bushman, East Asian, Sub-Saharan African, or Oceanian ancestry.

Happy May 12

Finnish flag image via Wikipedia

Today is the Day of the Finnish Identity, which is celebrated on the birthday of an early Finnish statesman.

How curiously humble to have a day in which you acknowledge that your national identity is something that came into being, or at least, into currency, at a time in historical memory, as opposed to trying to plant the false history that it was eternally in existence since time immemorial.

Genealogy v Genetics

Genealogy still towers over genetics in establishing family relationships.  23andMe located just under a thousand second and more distant cousins and relatives.  But, a genealogy website used in Finland has located approximately 350,000 of my closest relatives on my mother's side!  The Finns love their family histories, apparently.

Seriously, more than 6% of the population of Finland has a publicly documented family relationship to me.

I've met about half a dozen of my Finnish relatives and one stayed with my family for a summer after I had left for college.  The most illustrious of them was awarded a knighthood for his service to Finland by revolutionizing their waste treatment system, banishing diseases that would have otherwise afflicted Finland's people.

Finland From Its Prehistory To The Modern Era (UPDATED August 8, 2014)

Razib Khan has a nice little post summing up, among other things, the broad outlines of the prehistory of Finland and the means by which it arrived at its present population genetic and linguistic composition.

For full disclosure, I note than I am 50% Swede-Finn (my Lutheran Swede-Finn ancestors, who are on my maternal side, migrated to the Upper Peninsula of Michigan in the late 1800s), and it is possible that my account may be influenced by this fact. Some key points (with some of my own additions):

Mesolithic Finland

Finland was repopulated after the Last Glacial Maximum (which covered all of Northern Eurasia in an ice sheet ca. 20,000 BCE and completely ended modern human and any other hominin population of this region) only around 9,000 BCE (some sources indicate a date closer to 7300 BCE which would be about 9300 BP which could be caused by misunderstandings regarding the units in which dates are quoted), in an era sometimes called the Mesolithic or Epipaleolithic era.

Mesolithic Finnish Genetics

Uniparental genetics suggest that the source population for the original resettlement, which has left a strong serial founder effect impact on the Saami population (albeit with strong Uralic contributions) made its way up the Atlantic Coast (e.g. the Dutch and Belgian coast), probably from a Franco-Cantabrian refugium which also contributed to the proto-Berber populations with which it shares two of its three predominant mtDNA haplogroups, in what is now Algeria and Morocco to the South as it expanded in that direction as well.

The rare mtDNA haplogroup V is found all along the maritime path of this group of Mesolithic people's expansion (although later studies have shown mtDNA V in Eastern and Central Europe as well).  For example, a new study documents a Kurgan burial of an individual from the Novosvobodnaya culture in a part of what is now Southern Russian in the Northern Caucasus mountains around 3,000 BCE with mtDNA haplogroup V7.  The author of this new paper argues from this one ancient DNA sample that the mtDNA V7 points to a link with the contemporaneous Funnelbeaker culture (often abbreviated TRB from its German spelling) further North which was characterized by herding, fishing, marginal grain farming, and the use of imported copper but not Bronze, on the grounds that there are some indications of cultural linkage between the two cultures and on the grounds that mtDNA V is found in some remains from the Linear Pottery culture (often abbreviated LBK) who were the first farmers of the region from which the author argues that the TRB is derived.  (The lack of references to ancient TRB DNA to back up this claim, and reliance on a very outdated 1999 paper from Brian Sykes for the largely disproven hypothesis that most European mtDNA haplogroups have been widespread in Europe since the Upper Paleolithic era as a foundation for his analysis, undermines confidence in his conclusion, however.)  Also, who is to say that mtDNA V didn't introgress from pre-existing Mesolithic European hunter-gathers of whom the early Finns were a part, into farmers on the frontier of the Neolithic revolution, rather than the other way around?

The other leading mtDNA haplogrop in the Saami is U5b, which is common in all European hunter-gatherer populations. But, the U5b is predominantly U5b1b1 (284 out of 292 individuals with U5b with all 8 outliers located in Southern Sweden at the fringe of the area where Saami people are found), compared to about 20% of Continental Europeans with mtDNA U5b.

Disregarding the editorial white lines of distribution provided by the author (source here), I would say that the U5b1b1 distribution is quite a good fit for the coastal migration route.

On the Y-DNA side, one of the three most common Y-DNA haplogroups of the Finns, I1, is associated with the pre-Neolithic population of Europe (or more precisely, with sister Y-DNA haplogroup I2) much like mtDNA U5b and to a lesser extent mtDNA V (which is also found in ancient DNA in some early Neolithic populations in Europe, although possibly as a result of the first farmers of Europe taking wives from prior hunter-gatherer populations).

These people may have been the source population designed Western European Hunter-Gatherer based in ancient autosomal DNA, one of the three populations, together with "Early European Farmers" and "Ancestral Northern Eurasians" who were the main contributors to modern European population genetics in most of Europe (although as explained below, Finland had another Uralic population that contributed to its population genetics).

Mesolithic Archaeological Cultures In Finland

From 9000 BCE (or 7300 BCE if the reference is inaccurate) until about 2500 BCE, Finland was inhabited by the descendants of these people, who fed themselves mostly by fishing supplemented by hunting and gathering, without farming or herding.

Ulitmately, the Comb Ceramic culture, which was closely akin to the Pitted Ware people of mainland Europe nearby on the Baltic Sea, emerged from these people, around 5300 BCE, as they engaged in trade with Neolithic cultures to the South.

For example, flint from Scandinavia and the Valdai Hills, amber from Scandinavia and the Baltic region and slate from Scandinavia and Lake Onega found their way into Finnish archeological sites, while asbestos and soap stone from e.g. the area of Saimaa spread outside of Finland. Rock paintings—apparently related to shamanistic and totemistic belief systems—have been found, especially in Eastern Finland, e.g. Astuvansalmi.

Farming and Herding Arrive In Finland

Around 2500 BCE (or perhaps as late as 2300 BCE), Finland's Comb Ceramic maritime culture merged with the linguistically Indo-European Corded Ware culture derived Battle Axe culture from the south (that made its first inroads into Southwest Finland ca. 3200 BCE) to give rise to the Kiukainen culture. The Battle Axe culture brought dairy farming and greatly disrupted the local culture, although less the ideal climate conditions impaired the desirability of the area for herding and farming, ultimately led to a resurgence in maritime fishing people's to the local culture, so that both cultures made significant contributions.

It hasn't been clear until this week how much of a contribution this archaeological culture made to the further Northern Saami people, but the sudden appearance of dairy farming as evidenced by residues on pottery suddenly appearing around 2500 BCE even above the Arctic Circle, puts to rest the possibility that dairying didn't reach the far northern reaches of Finland at that time.  It is now clear that almost everyone in the region was affected as the Neolithic revolution finally reached Finland, even if some populations subsequently reverted to a different subsistence mode.

Indo-European Farmer Genetics

The Battle Axe culture is the likely source of Y-DNA haplogroup R1a in Finns and possibly also mtDNA H mostly arrive at this time (although the antiquity of mtDNA H in the region in ancient suggests that this haplogroup could have arrived in the Mesolithic, rather than a Neolithic era.

Notably, in Japan, whose Jomon culture was also maritime fishing based before the arrival of the Yaoyi rice farmers with horse riding warriors, there the indigeneous maritime culture also had a considerable population genetic contribution, rather than the predominant replacement of terrestrial hunter-gatherers seen elsewhere.  More sedentary fishing cultures may have had more staying power vis-a-vis early farmers than terrestrial hunters and gatherers.

The Case For An Indo-European Language Shift

At this point in time, the population of Finland probably came to speak an Indo-European pre-Slavic Baltic language. (A Finnish linguist argues that this Corded Ware language was proto-Germanic, but in my view a Baltic language is more likely.)

The ancestral language of the Comb Ceramic and Pitted Ware peoples was probably lost at this time in what was probably one of its last outposts in Europe (after earlier episodes of Neolithic replacement or conquest in Continental Europe starting ca. 5500-4600 BCE), except for some place names and perhaps some twice removed substrate influences.

This first farmer Neolithic era probably persisted until the arrival of the that persisted until the Finish Bronze Age (ca. 1500 BCE-500 BCE).

The Finnish Bronze Age

Another Language Shift

Sometime after 2500 BCE, Finland received an influx of a Uralic language speaking population from Siberia, not closely related to either of its source populations.

This influx gave rise to a language shift during which the Finnish language (or at least the Finno-Saami language family) probably emerged. The Finnish Bronze Age exactly coincides with a statistical estimate of the time of origin of the Finno-Saami branch of the Uralic languages.

Realistically, the Uralic language probably arrived as part of the advent of the Finnish Bronze Age ca. 1500 BCE which arrived with Bronze using cultures from Northern and Eastern Russia according to Wikipedia with is in accord with this source regarding the earliest appearance of Bronze artifacts being located inland. But, an unsourced Internet resource claims a Western rather than Eastern source for the Finnish Bronze Age and the appearance of the Western practice of cremation around this time in Finland argues for a Western source in coastal areas.  These differences of opinion regarding the origin of the Finnish Bronze Age had nationalist implications for Finland.

Like Razib, I disagree with the Wikipedia analysis that puts the arrival of the Uralic languages according to older scholarship, in Finland ca. 4000 BCE.  The Wikipedia analysis acknowledges the possibility that this is inaccurate, but suggests an Iron Age arrival for Uralic, which I suspect is too late, although a differentiation of Finnish from neighboring Uralic languages may date to the Iron Age ca. 500 BCE.

The Genetic Impact of Bronze Age Uralic Migration Into Finland

Razib notes that they left an east Asian autosomal genetic contribution (about 5%-8% in Saami populations today), mtDNA (mtDNA haplogroups D5 and Z which make up a similar percentage of Saami mtDNA with the mtDNA Z1 subclade contribution dated to 2,000-3,000 years ago by mtDNA mutation rate dating, i.e. the Iron Age) and Y-DNA legacy (e.g. Y-DNA haplogroup N1b and N1c1), which is distinct from the "ancestral North Asian" autosomal genetic legacy in Europe (that may have arrived with first and/or subsequent waves of Indo-Europeans together with, for example, Y-DNA haplogroup R1a).

About the Uralic Language Family

The Uralic language family is shared by the people of Estonia, Latvia, the Karelian region of Russia, and the Saami people of Finland. A more distant branch of this language family is Hungarian, and another more distant branch of this language family is shared by many Siberian ethnic populations including the Mari who are arguably the last population to have continuously practiced pagan religions of Northern Asia into the present.

Some linguists such as Michael Fortescue writing in 1998, have argued that the Eskimo-Aleut languages, including Inuit, are a sister language family to the Uralic languages that together form a circumpolar mega-language family.  Morris Swadesh in 1962, Holst in 2005, and others dating back to the 1746 CE, have made similar proposals.

From a historical timing perspective, however, the Saqqaq (Arctic Paleo-Eskimos) which was present 2000-2500 BCE (the best ancient DNA example of which had Y-DNA Q1a, mtDNA D2a1, and autosomal DNA similar to the modern Koryak people of the Northeast Asia coast) , or the Dorset (second wave Arctic Paleo-Eskimos) would be a better match, and perhaps left a substrate influence on later Eskimo-Aleut languages (although there are genetic indications that these earlier populations were almost totally replaced in the Americas) which arrived with the Thule around 500 CE. Inuits lack Y-DNA N1b and N1c found in modern Finns and contributed mostly mtDNA haplogroups A (which is absent in the Finns) to the existing indigenous American mtDNA pool, while the Dorset contributed mostly mtDNA D3 (found in modern Paleo-Siberian populations and which is a sister clade to Finnish and Asian mtDNA D5), according to the ancient DNA samples.

The Bronze Age Transition Was Demographically More Important Than The Iron Age Transition

The fact that the Finnish Bronze Age arrived from the Uralic heartland with a probable demic component, while the Finnish Iron Age appears to have include more cultural exchange than demic migration, and the likely powerful capacity of a metal age culture to overwhelm a pre-metal age culture militarily, argues for an arrival ca. 1500 BCE, rather than earlier or later.  Certainly, this Siberian centered linguistic family would not have appeared at a time when the cultural influences in Finland were from Rome or Western Europe as it was from 0 CE onward.

One archaeological site from the period is this one and a number of Finnish Bronze Age papers from 2009 can be found here).

The Finnish Iron Age and the Middle Ages

The Finnish Iron Age began around 500 BCE (several hundred years later than in the Mediterranean) and around 0 CE, began to show the influence of trade with the Roman Empire that persisted until about 400 CE. The Migration Period during which there were mass folk migrations of "barbarian" Germanic tribes like the Goths and the Visigoths through Europe extended to Finland's Iron Age from ca. 400 CE to 575 CE and showed increasing Germanic influence in cultural artifacts.

The Migration period and the Merovingian period that followed also coincide historically with Slavic expansion into what is now Orthodox Christian Eastern Europe.

From 575 CE to 800 CE, "The Merovingian period in Finland gave birth to distinctive fine crafts culture of its own, visible in the original decorations of domestically produced weapons and jewelry. Finest luxury weapons were, however imported from Western Europe. The very first Christian burials are found from the latter part of this era as well. The Leväluhta burial findings suggest that the average height of a man was 158 cm [i.e. 5'2"] and that of a woman was 147 cm [i.e. 4'10"]."

Trade with the linguistically Germanic (Indo-European) Vikings (many from Sweden and some of whom started to colonize Finland) followed from 800 CE to 1025 CE during which hill forts started to be erected in Southern Finland in the earliest signs of urbanization.

The Christianization of Finland began in earnest around 1150 CE which was also around the time that Finland begins to appear in the written historic record. Swedish colonization efforts directed at Finland were stepped up in Northern Crusades in the early 1200s CE, bringing with them the other of Finland's two major languages and adding a significant Swedish population genetic component to the overall mix. According to Wikipedia the story then continued as follows:

In the early 13th century, Bishop Thomas became the first bishop of Finland. There were several secular powers who aimed to bring the Finns under their rule. These were Sweden, Denmark, the Republic of Novgorod in Northwestern Russia and probably the German crusading orders as well. Finns had their own chiefs, but most probably no central authority. Russian chronicles indicate there were conflict between Novgorod and the Finnic tribes from the 11th or 12th century to the early 13th century.

The name "Finland" originally signified only the southwestern province that has been known as "Finland Proper" since the 18th century. Österland (lit. Eastern Land) was the original name for the Swedish realm's eastern part, but already in the 15th century Finland began to be used synonymously with Österland. The concept of a Finnish "country" in the modern sense developed only slowly during the period of the 15th–18th centuries.

It was the Swedish regent, Birger Jarl, who established Swedish rule in Finland through the Second Swedish Crusade, most often dated to 1249, which was aimed at Tavastians who had stopped being Christian again. Novgorod gained control in Karelia, the region inhabited by speakers of Eastern Finnish dialects. Sweden however gained the control of Western Karelia with the Third Finnish Crusade in 1293. Western Karelians were from then on viewed as part of the western cultural sphere, while eastern Karelians turned culturally to Russia and Orthodoxy. While eastern Karelians remain linguistically and ethnically closely related to the Finns, they are considered a people of their own by most. Thus, the northern border between Catholic and Orthodox Christendom came to lie at the eastern border of what would become Finland with the Treaty of Nöteborg in 1323.

During the 13th century, Finland was integrated into medieval European civilization. The Dominican order arrived in Finland around 1249 and came to exercise huge influence there. In the early 14th century, the first documents of Finnish students at Sorbonne appear. In the south-western part of the country, an urban settlement evolved in Turku. Turku was one of the biggest towns in the Kingdom of Sweden, and its population included German merchants and craftsmen. Otherwise the degree of urbanization was very low in medieval Finland. Southern Finland and the long coastal zone of the Bothnian Gulf had a sparse farming settlement, organized as parishes and castellanies. In the other parts of the country a small population of Sami hunters, fishermen and small-scale farmers lived. These were exploited by the Finnish and Karelian tax collectors.

Trade exchanges during the early Iron Age probably did not lead to language shift, and even the influx of Swedish and other Scandinavian colonists in starting in the Viking era around 1200 CE only led to a bilingual situation with coastal colony towns speaking Germanic Swedish languages with a Finnish substrate and Finnish and Saami remaining living languages elsewhere.

Finland In The Early Modern Era

Swedish domination would continue for centuries, and in the Reformation, the Swedish sided with the Protestant Lutherans against the Catholics and had their own round of witch hunting in the 1600s, including well as a short lived effort to establish a Swedish colony in America near the Delaware-Pennsylvania area from 1638-1655 CE, with at least half of the colonists coming from Finland.

Very hard times followed for the next quarter century resulting in the death of a third of the population in a four year long famine, followed by the death of half of the population in a twenty-one year long war.  The population of Finland fell by about two-thirds in a single generation.

In 1696–1699, a famine caused by climate decimated Finland. A combination of an early frost, the freezing temperatures preventing grain from reaching Finnish ports, and a lackluster response from the Swedish government saw about one-third of the population die. Soon afterwards, another war determining Finland's fate began (the Great Northern War of 1700–21). The Great Northern War (1700–1721) was devastating, as Sweden and Russia fought for control of the Baltic. Harsh conditions—worsening poverty and repeated crop failures—among peasants undermined support for the war, leading to Sweden's defeat. Finland was a battleground as both armies ravaged the countryside, leading to famine, epidemics, social disruption and the loss of nearly half the population. By 1721 only 250,000 remained.

Constitutional monarchy with a powerful parliament followed in Sweden, but "Finland by this time was [still] depopulated, with a population in 1749 of 427,000." Potato farming (which was a dietary staple of my ancestors) arrived after the 1750s.

In 1809, Finland was annexed to Russia with the assent of a popular assembly.

Mass migration to the United States in the late 19th century was also accompanied by hard times at home in Finland.

Finland secured independence in 1917, followed by a brief civil war in 1918, as a result of the Russian revolution.