The Indigenous People of Europe

Do you know how long I've been waiting to use this?

The settlement in Cambridgeshire, which had been buried for 3,000 years, was discovered when the tops of crude protest signs were spotted above layers of mud.

Archaeologist Helen Archer said: “The signs, which include ‘Any old iron? NO THANKS,’ and ‘IRON? IR NO,’ a primitive attempt at wordplay, show that the residents were up in arms about climate-based migration patterns.

Note: The Daily Mash is a humor/satire site, similar to The Onion.

Anyway, on to the genetics!

From Haak et al, rearranged by me
From Haak et al, rearranged by me

Click for full size

Haak et al. made this graph, but I rearranged it so that the oldest samples are on the left and the newest ones are on the right. When multiple samples were about the same age, I ordered them from west to east (that is, from left to right as you look at a standard map. Unless you are in Australia.) I’ve added the dates (shown as ranges) that were in Haak’s paper. Note the asterisk under Karsdorf–those dates are still uncertain.

The first three genomes are from super old skeletons found out in, like, Russia. I don’t know why they look so crazy–maybe because the DNA is really old and so not very good, or maybe because they actually had a bunch of different DNA in them, or maybe because they’re ancestral to a bunch of different groups. I don’t know! Luckily, it doesn’t really matter for today’s post, so I’ll investigate them later.

Approximately 28,000 years later, we have the Blue People, also known as “Western European Hunter Gatherers,” or WHG. There were people in Europe in intervening 28,000 years; they just aren’t on the table, and I don’t know if anyone has successfully sequenced their genomes yet. (More research required.)

As you might guess, the WHG people hunted and gathered. They had stone tools, and were quite widespread, ranging from Spain (the La Brana1 site,) to Sweden to Samara, Russia (and probably beyond.)

And then some new guys showed up: Farmers.

Known as the Early Eurasian Farmers (EEF,) they first appear on our graph in Starcevo, Serbia, their DNA in orange. They came from the Middle East (the birthplace of agriculture,) bringing their wheat, permanent settlements, and livestock.

Why isn't it in English? Oh, well. We'll manage.
Neolithic cultures of Europe–Starcevo is i the lower right-hand corner.

These farmers quickly overran the hunter-gatherers throughout western Europe (though the northern extremes held out longer, most likely due to crops that originated in the Middle East taking a while to adapt to the harsh Scandinavian climate.)

300px-Neolithic_expansion.svg (source: Wikipedia)

The hunter gatherers disappeared (most likely slaughtered by the farmers, but perhaps merely overwhelmed numerically) but their DNA lives on in the descendants of those first farmers. Some groups may have combined willingly–others, as the spoils of war. Within the Farmers’ range, the only place the hunter-gatherers managed to live on appears to be a small island off the coast of Sweden (the second “Skoglund” sample.)

But to the east, out on the Eurasian steppes, the hunter-gatherers lived on. The steppes are known more for their rampaging hordes than their farmers, and this is exactly what they became.

The Yamnaya, as we now call them, are about half WHG and half some new population (I call them the Teal People.) As far as I know, no “pure” teal people have yet been found, but teal DNA is all over the place, from India to Spain.

Teal and blue DNA in India central Asia, and Siberia:

IndiaandSteppe

The Yamnaya are also known as the Proto-Indo-Europeans–the guys who spoke the language ancestral to all of today’s Indo-European languages. And like all conquering barbarian hordes, they expanded out of their homeland in present-day southern Russia (north of the Caucuses,) and conquered everything in their path.

Just eyeballing the graph, it looks like the resulting peoples are about half Yamnaya, and about half EEF. This tri-part inheritance is still seen in every European population (and some of their neighbors) today:

Europe

If we didn’t have the ancient DNA–or if we had less of it–it would be easy to think that the Blue component in modern Europeans had come directly from the ancient WHG population that lived in their particular area. Instead, much (if not most) of the modern “blue” component hails from the steppes of Russia–a remarkable comeback for the WHGs.

Oh, and the “indigenous” people of Europe? They’re all indigenous to the continent.

Some more helpful graphs, maps, and information:

From Haak et al.
From Haak et al.
From Haak et al.
From Haak et al.
From Haak et al.
From Haak et al.

On the Iceman, aka Otzi: found in the Alps on the Italian-Austrian border; Same age as Sweden, between 3359 and 3105 BCE. (Hailed from the vicinity of Feldthurns, Italy.)

Analysis of the mtDNA of Ötzi the Iceman, the frozen mummy from 3,300 BC found on the AustrianItalian border, has shown that Ötzi belongs to the K1 subclade. It cannot be categorized into any of the three modern branches of that subclade (K1a, K1b or K1c). The new subclade has provisionally been named K1ö for Ötzi.[14] Multiplex assay study was able to confirm that the Iceman’s mtDNA belongs to a new European mtDNA clade with a very limited distribution amongst modern data sets.[15]” (source)

Otzi ate grain but was lactose intolerant.

His Y DNA is haplogroup G, which is now rare in Europe:

Haplogrupo_G_(ADN-Y)

Various estimated dates and locations have been proposed for the origin of Haplogroup G. The National Geographic Society places haplogroup G origins in the Middle East 30,000 years ago and presumes that people carrying the haplogroup took part in the spread of the Neolithic.[2] Two scholarly papers have also suggested an origin in the Middle East, while differing on the date. …

Haplogroup G2a(SNP P15+) has been identified in neolithic human remains in Europe dating between 5000-3000BC. Furthermore, the majority of all the male skeletons from the European Neolithic period have so far yielded Y-DNA belonging to this haplogroup. The oldest skeletons confirmed by ancient DNA testing as carrying haplogroup G2a were five found in the Avellaner cave burial site for farmers in northeastern Spain and were dated by radiocarbon dating to about 7000 years ago.[5] At the Neolithic cemetery of Derenburg Meerenstieg II, north central Germany, with burial artifacts belonging to the Linear Pottery culture, known in German as Linearbandkeramik (LBK). This skeleton could not be dated by radiocarbon dating, but other skeletons there were dated to between 5,100 and 6,100 years old. The most detailed SNP mutation identified was S126 (L30), which defines G2a3.[6] G2a was found also in 20 out of 22 samples of ancient Y-DNA from Treilles, the type-site of a Late Neolithic group of farmers in the South of France, dated to about 5000 years ago.[7] The fourth site also from the same period is the Ötztal of the Italian Alps where the mummified remains of Ötzi the Iceman were discovered. Preliminary word is that the Iceman belongs to haplogroup G2a2b [8] (earlier called G2a4).

Haplogroup G2a2b is a rare group today in Europe. (source)

Back on the Otzi page:

By autosomal DNA he is most closely related to southern Europeans, especially to the geographically isolated populations of the two Mediterranean islands of Sardinia and Corsica.[41][42]

… In October 2013, it was reported that 19 modern Tyrolean men were related to Ötzi. Scientists from the Institute of Legal Medicine at Innsbruck Medical University had analysed the DNA of over 3,700 Tyrolean male blood donors and found 19 who shared a particular genetic mutation with the 5,300-year-old man, which led them to identify the link.[46]

Hungary Gamba CA= Copper age, 3,300 BC-2,700 AD

From an analysis of the Gamba site:

The Great Hungarian Plain was a crossroads of cultural transformations that have shaped European prehistory. Here we analyse a 5,000-year transect of human genomes, sampled from petrous bones giving consistently excellent endogenous DNA yields, from 13 Hungarian Neolithic, Copper, Bronze and Iron Age burials including two to high (~22 × ) and seven to ~1 × coverage, to investigate the impact of these on Europe’s genetic landscape. These data suggest genomic shifts with the advent of the Neolithic, Bronze and Iron Ages, with interleaved periods of genome stability. The earliest Neolithic context genome shows a European hunter-gatherer genetic signature and a restricted ancestral population size, suggesting direct contact between cultures after the arrival of the first farmers into Europe. The latest, Iron Age, sample reveals an eastern genomic influence concordant with introduced Steppe burial rites. We observe transition towards lighter pigmentation and surprisingly, no Neolithic presence of lactase persistence.

Stuttgart EN:

To investigate European population history around the time of the agricultural transition, we sequenced complete genomes from a ~7,500 year old early farmer from the Linearbandkeramik (LBK) culture from Stuttgart in Germany and an ~8,000 year old hunter-gatherer from the Loschbour rock shelter in Luxembourg. We also generated data from seven ~8,000 year old hunter-gatherers from Motala in Sweden. We compared these genomes and published ancient DNA to new data from 2,196 samples from 185 diverse populations to show that at least three ancestral groups contributed to present-day Europeans. The first are Ancient North Eurasians (ANE), who are more closely related to Upper Paleolithic Siberians than to any present-day population. The second are West European Hunter-Gatherers (WHG), related to the Loschbour individual, who contributed to all Europeans but not to Near Easterners. The third are Early European Farmers (EEF), related to the Stuttgart individual, who were mainly of Near Eastern origin but also harbored WHG-related ancestry. We model the deep relationships of these populations and show that about ~44% of the ancestry of EEF derived from a basal Eurasian lineage that split prior to the separation of other non-Africans.(bold mine.)

Ancient_North_Eurasian_admixture European_hunter-gatherer_admixture Neolithic_farmer_admixture

Source for the maps.

Iosif Lazaridis et al's model of modern European genetic origins.
Iosif Lazaridis et al’s model of modern European genetic origins.
Also from Iosif Lazaridis et al.'s paper, " Ancient human genomes suggest three ancestral populations for present-day Europeans"
Also from Iosif Lazaridis et al.’s paper, “Ancient human genomes suggest three ancestral populations for present-day Europeans” h/t Dienekes:

 Analysis of ancient DNA can reveal historical events that are difficult to discern through study of present-day individuals. To investigate European population history around the time of the agricultural transition, we sequenced complete genomes from a ~7,500 year old early farmer from the Linearbandkeramik (LBK) culture from Stuttgart in Germany and an ~8,000 year old hunter-gatherer from the Loschbour rock shelter in Luxembourg. We also generated data from seven ~8,000 year old hunter-gatherers from Motala in Sweden. We compared these genomes and published ancient DNA to new data from 2,196 samples from 185 diverse populations to show that at least three ancestral groups contributed to present-day Europeans. The first are Ancient North Eurasians (ANE), who are more closely related to Upper Paleolithic Siberians than to any present-day population. The second are West European Hunter-Gatherers (WHG), related to the Loschbour individual, who contributed to all Europeans but not to Near Easterners. The third are Early European Farmers (EEF), related to the Stuttgart individual, who were mainly of Near Eastern origin but also harbored WHG-related ancestry. We model the deep relationships of these populations and show that about ~44% of the ancestry of EEF derived from a basal Eurasian lineage that split prior to the separation of other non-Africans.

I'm sorry, I forgot where this came from
I’m sorry, I forgot where this came from.

See also:

Significant genetic differentiation between Poland and Germany follows present-day political borders, as revealed by Y-chromosome analysis, by Kayser M. et al:

To test for human population substructure and to investigate human population history we have analysed Y-chromosome diversity using seven microsatellites (Y-STRs) and ten binary markers (Y-SNPs) in samples from eight regionally distributed populations from Poland (n = 913) and 11 from Germany (n = 1,215). Based on data from both Y-chromosome marker systems, which we found to be highly correlated (r = 0.96), and using spatial analysis of the molecular variance (SAMOVA), we revealed statistically significant support for two groups of populations: (1) all Polish populations and (2) all German populations. … The same population differentiation was detected using Monmonier’s algorithm, with a resulting genetic border between Poland and Germany that closely resembles the course of the political border between both countries. The observed genetic differentiation was mainly, but not exclusively, due to the frequency distribution of two Y-SNP haplogroups and their associated Y-STR haplotypes: R1a1*, most frequent in Poland, and R1*(xR1a1), most frequent in Germany. We suggest here that the pronounced population differentiation between the two geographically neighbouring countries, Poland and Germany, is the consequence of very recent events in human population history, namely the forced human resettlement of many millions of Germans and Poles during and, especially, shortly after World War II.

And Iron Age and Anglo-Saxon genomes from East England reveal British migration history by Schiffels et al., h/t Steve Sailer

British population history has been shaped by a series of immigrations, including the early Anglo-Saxon migrations after 400 CE. … Here, we present whole-genome sequences from 10 individuals excavated close to Cambridge in the East of England, ranging from the late Iron Age to the middle Anglo-Saxon period. … we estimate that on average the contemporary East English population derives 38% of its ancestry from Anglo-Saxon migrations. … Using rarecoal we find that the Anglo-Saxon samples are closely related to modern Dutch and Danish populations, while the Iron Age samples share ancestors with multiple Northern European populations including Britain.

Judeo Ethnogenesis

Disclaimer: I had 3 hours of sleep last night. Weigh that carefully before testing my patience.

Ethnogenesis, as the name implies, is the process whereby a new ethnic group is created. An ethnicity is more or less a group with a shared culture, belief in common ancestry, and that preferentially marries within itself rather than outside of itself. Over time, this creates a group that is ethnically distinct from its neighbors, even under conditions of close proximity.

The Amish, for example, after splitting off from the Swiss in the 1600s over religious differences (remember, religion is ethnicity,) arrived in Pennsylvania in the early 1700s, so we may mark Amish ethnogenesis around the mid 1600s or early 1700s People today make fun of Ben Franklin for complaining that the German-speaking immigrants to Pennsylvania were problematic and not integrating with the rest of the population, but you know, the Amish still haven’t integrated. They still speak German, follow their own religion and traditions, and don’t inter-marry with the rest of the Pennsylvania population, such that they are quite ethnically distinct, at least on a genetic level.

The Hui of China are another example; they were not really considered an ethnic group before the establishment of the People’s Republic of China circa 1949. The Chinese decided to just lump all of their Muslim minorities–some of them quite distinct–under one term. (Historically, the term “Hui” also referred to Christians and Jews and was just a general catch-all.) Hui now marry other Hui preferentially enough that the Wikipedia page goes into detail on known cases of inter-marriage with the Han, but a fellow Hui from across the country may be regarded as just another Hui, and so a preferred partner.

Anyway, so that got me thinking about the establishment of Israel. Normally when I think of Jews, I am actually thinking of Askenzim, and you probably are, too. But Israel is actually 61% Mizrahi Jews–Jews from predominantly Muslim countries.

You know the general story: Once upon a time, all of the Jews lived in Israel. These people were probably pretty similar, ethnically, to the Palestinians, assuming the Palestinians are anything like the region’s residents 2000 years ago, and don’t have a massive influx of Turkish DNA or something like that.

Then the Jews got conquered and scattered to the winds. Most famously after the destruction of the Second Temple by the Romans, but also during the Babylonian  and Assyrian eras, etc. Anyway, for the past 2000+ years or so, there have been significant Jewish communities in a lot of places that aren’t Israel, eg:

Uzbeki Jews
Uzbeki Jews
Chinese Jews
Chinese Jews
Ethiopian Jews
Ethiopian Jews
Cochin Jews
Cochin Jews
Lemba Jews?
Lemba Jews?

Then in the late 1800s, the Jews–mostly Ashkenazim, I think–got sick of this state of affairs and decided to exit Europe and go back to Israel. Unfortunately, they didn’t really succeed until 1948, at which point Jews from all over the world started pouring in.

Since most people are genetically similar to their neighbors, eg the Palestinians and Syrians, or Han and She, I began wondering how similar Jews were to their neighbors of millenia verses their similarity to each other.

Here’s a graph showing major genetic lineages of a bunch of different ethnic groups, including several Jewish ones:

nature09103-f3.2

Broad cultural zones are easily distinguished, like East Asians in yellow, South Asia in greens, Europeans with their large dark blue chunk, Middle Easterners with their big patches of light green and light blue, and the rust-tones in sub-Saharan Africa. This data set is great, because it lets us compare various Jewish groups to their immediate neighbors, eg:

J1

I made a condensed version of the graph that highlights the measured Jewish groups and their neighbors, (sadly, some of the samples are pretty small, making them hard to read):

J1

And an even more condensed version that just compares the Jews to each other:

j2

(Note that the pure green section on the right-hand side is not a Jewish group, but just a chunk of the graph that happened to overlap the text due to the Cochin Jewish section being so small.)

Observations: Most Jewish groups are significantly more similar to their immediate neighbors than they are to other Jewish groups, especially when we look at the furthest-flung folks. Cochin Jews and Ethiopian Jews, for example, show almost no DNA in common (in this graph.)

Given what all Middle Eastern groups look like in the sample, we may speculate that the original Jewish group primarily had a large section of light blue and a slightly smaller section of light green, with probably a smidge of sub-Saharan. Several of the Middle Eastern Jewish groups still have this genetic makeup.

Three Jewish groups show a more European makeup, with a large dark blue chunk characteristic of Europeans and North Africans: the Ashkenazim, Sephardim, and Moroccan Jews. They look closest to Cypriots, though I compared them to Spaniards and Tuscans as their nearest neighbors in the graph.

Since the Ashkenazim are estimated to be about half Italian, it’s not surprising that they have about half as much dark blue as the Italians. Even within European groups, while they look fairly similar at this level of resolution, some groups are quite distinct from each other–Italians and Germans, for example, or Brits and Greeks. Geneticists can determine whether your ancestors were Italians or Germans or Greeks just by looking at your DNA, but those kinds of small details don’t really show up all that well on a graph that is trying to show the differences between Sub-Saharan Africanss and Asians. So while Moroccans, Sephardim, and Ashkenazim all look rather similar here, there may be finer grained differences that just don’t show up at this scale.

What’s up with the Moroccan Jews? They do not look like Moroccans; I therefore speculate a more recent migration of Moroccan Jews from somewhere else that’s not Morocco, like Spain.

The Jews who migrated to the East, however, lost a significant portion–almost all–of their light blue component, replacing it with dark green more typical of Indians and other SE Asian populations.

I don’t think this dataset contains Uzbeki Jews (or the Lemba, who are not Jewish enough to be considered Jewish, but still have a few Jewish traditions and folktales and a bit of Jewish ancestry,) which is sad, but I’d wager the Uzbeki Jews look a lot like other Uzbeks.

One of the things I’ve heard often from Jews is that all Jews are Jews, part of one great big Jewish family descended from Abraham (even the atheist ones!) and thus Jews should always try to be kind to each other, all Jews are welcome in Israel, etc. This is a perfectly sensible philosophy when you’re a peasant in Poland and the only foreign Jews you’ve ever met were from Lithuania. But 2000+ years of diaspora have resulted in far flung groups becoming quite ethnically distinct from each other. Like the Amish, isolated groups in Cochin or Ethiopia have become their own ethnies distinct from their ancestors, but unlike the Amish, they have inter-married significantly with the locals. (The Amish do not marry non-Amish.)

The Roman Exile, therefore, should be regarded as a major ethnogenesis event–the beginning of the creation of most current Jewish ethnic groups.

The creation of the state of Israel constitutes a second major ethnogenesis event, a bringing together of these multiple ethnic groups into one population that views itself as one population. I expect a great deal of mixing between these historically distinct groups into a more homogenous whole, (though some groups don’t mix terribly well, like certain Ultra Orthodox communities that haven’t been accepting of Ethiopian Jews.)