Ancient North Eurasians from Siberia

Ancient North Eurasian (ANE) is a genetic component that traveled mainly east into Native Americans, but also west into Europeans. It's represented by pre-historic individuals discovered in Siberia:

Here we sequence the draft genome of an approximately 24,000-year-old individual (MA-1), from Mal'ta in south-central Siberia. [...] Similarly, we find autosomal evidence that MA-1 is basal to modern-day western Eurasians and genetically closely related to modern-day Native Americans, with no close affinity to east Asians. [...] Sequencing of another south-central Siberian, Afontova Gora-2 dating to approximately 17,000 years ago, revealed similar autosomal genetic signatures as MA-1, suggesting that the region was continuously occupied by humans throughout the Last Glacial Maximum.

Though MA-1 is slightly more Western than Eastern Eurasian genetically, it's actually intermediate and way outside the range of Modern Europeans and Caucasoids:

In the first two principal components, MA-1 is intermediate between modern western Eurasians and Native Americans, but distant from east Asians.

When it's included in an admixture analysis, it comes out as a mix of Western Eurasians and the different Eastern Non-Africans that it shares ancestry with:

The ancient samples appear to be mostly West Eurasian in their ancestry, although the hunter gatherers are also inferred to have greater or lesser extents of an eastern non-African (ENA) component lacking in Stuttgart. [...] We note that the ancestry proportions in ancient samples like MA1 are more likely explained by shared ancestry than admixture. This is more likely to explain the nearly three-way distribution of South Asian, West Eurasian and Native American (plus Northeast Siberian) ancestry proportions in MA1, than three-way admixture of established populations.

Admixture from MA1 (and AG2) is highest in Eastern Hunter-Gatherers (EHG), then Scandinavian Hunter-Gatherers (SHG), then Western Hunter-Gatherers (WHG):

Application of this methods highlights the impact of Ancient North Eurasian (ANE) ancestry related to the ~22,000 BCE Mal'ta 1 and ~15,000 BCE Afontova Gora 2 on populations living in Europe, the Americas, and Eastern Eurasia. Eastern Eurasians can be modelled as arrayed along a cline with different proportions of ANE ancestry, ranging from ~40% ANE in Native Americans matching previous findings, to no less than ~5-10% ANE in diverse East Asian groups including Han Chinese. We also document a cline of ANE ancestry across the east-west extent of Eurasia. Eastern Hunter Gatherers (EHG) derive ~3/4 of their ancestry from the ANE; Scandinavian hunter-gatherers (SHG) are a mix of EHG and WHG; and WHG are a mix of EHG and the Upper Paleolithic Bichon from Switzerland.

After Native Americans, then Siberians, it's Northern and Eastern Europeans that have the highest ANE admixture because they have the most EHG (which came mainly with Bronze Age migrations from the Russian Steppe):

We find that genetic affinity to MA-1 is greatest in two regions: first, the Americas; and second, northeast Europe and northwest Siberia, with north-to-south latitudinal clines in shared drift with MA-1 in both Europe and Asia.

(redder circles = more affinity, bluer = less)

Here's the genetic cline formed by Modern Europeans stretching from Caucasoid Neolithic Farmers (Stuttgart) towards ANE:

By using BedouinB instead of Stuttgart, we can also plot Stuttgart in the space of these statistics. Europeans uniformly share more drift with MA1 than with Karitiana, and form a cline in this space with slope >1. Karitiana, because of its Ancient North Eurasian ancestry was crucial in detecting the presence of such ancestry in Europeans but can now be replaced in the study of this ancestry by a better proxy for this ancestry (MA1).

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Raghavan et al. "Upper Palaeolithic Siberian genome reveals dual ancestry of Native Americans". Nature, 2014.

Lazaridis et al. "Ancient human genomes suggest three ancestral populations for present-day Europeans". Nature, 2014.

Lazaridis et al. "Genomic insights into the origin of farming in the ancient Near East". Nature, 2016.

Related: Phenotypes of Hunters and Farmers

Phoenicians and Moors Were Caucasoid

Afrocentrists claim that Phoenicians and Moors were black, but most of these ancient samples from colonies in Iberia are a mix of North African, Levantine and Southern European (Local Iberian), clustering very close to Europeans because North Africans are Caucasoid. Only two samples from the later Muslim period have Sub-Saharan African admixture (37% and 48%) and they cluster far away from the rest. Modern Iberians have much less of all this admixture (~7% North African and 0% Sub-Saharan African) and cluster with other Europeans.

In the southeast, we recovered genomic data from 45 individuals dated between the 3rd and 16th centuries CE. All analyzed individuals fell outside the genetic variation of preceding Iberian Iron Age populations (Fig. 1, C and D, and fig. S3) and harbored ancestry from both Southern European and North African populations (Fig. 2D), as well as additional Levantine-related ancestry that could potentially reflect ancestry from Jewish groups (21). These results demonstrate that by the Roman period, southern Iberia had experienced a major influx of North African ancestry, probably related to the well-known mobility patterns during the Roman Empire (22) or to the earlier Phoenician-Punic presence (23); the latter is also supported by the observation of the Phoenician-associated Ychromosome J2 (24). Gene flow from North Africa continued into the Muslim period, as is clear from Muslim burials with elevated North African and sub-Saharan African ancestry (Fig. 2D, fig. S4, and table S22) and from uniparental markers typical of North Africa not present among pre-Islamic individuals (Fig. 2D and fig. S11). Present-day populations from southern Iberia harbor less North African ancestry (25) than the ancient Muslim burials, plausibly reflecting expulsion of moriscos (former Muslims converted to Christianity) and repopulation from the north, as supported by historical sources and genetic analysis of present-day groups (25). The impact of Muslim rule is also evident in northeast Iberia in seven individuals from Sant Julià de Ramis from the 8th to 12th centuries CE who, unlike previous ancient individuals from the same region, show North African–related ancestry (Fig. 2C and table S19) and a complete overlap in PCA with present-day Iberians (Fig. 1D).

Olalde et al. "The genomic history of the Iberian Peninsula over the past 8000 years". Science, 2019.

Related: Moors Expelled from Sicily and Southern Italy

"Dark" and "Swarthy" Europeans Are Still Light

We've seen how olive skin is misunderstood by people to mean "tan" or "non-white", now let's look at the same thing with words like "dark" and "swarthy" used to describe the complexions of Europeans. People treat them as evidence against "whiteness", but they're really just exaggerations of reality.

Benjamin Franklin applied them to a lot of groups that are far from dark, basically lumping all whites who weren't Anglo-Saxon into a "swarthy" group with non-whites, including some who are probably lighter than English people:

All Africa is black or tawny. Asia chiefly tawny. America (exclusive of the new Comers) wholly so. And in Europe, the Spaniards, Italians, French, Russians and Swedes, are generally of what we call a swarthy Complexion; as are the Germans also, the Saxons only excepted, who with the English, make the principal Body of White People on the Face of the Earth.

Another well-known example of this kind of exaggeration is English ideas about the so-called "Black Irish". They're really just white people from the British Isles (not just Ireland) who have dark hair and eyes and a Mediterranean appearance — like Colin Farrell, Catherine Zeta Jones, Sean Connery, Mr. Bean, Russell Brand and many others — but old school Nordicists used to claim that part of the Celtic physiognomy was "black-tinted skin":


But even when applied to "darker" Southern Europeans, they're still an exaggeration, like in this passage from White on Arrival about how Italian gangsters were portrayed in the media:

Al Capone was constantly portrayed in books, magazine articles, pulps, and movies as having a "dark" or "swarthy" complexion. When he appeared in court in 1929 in Philadelphia on charges of having concealed a weapon, the Chicago Daily News noticed that his "face, which is rather dark, assumed a dull reddish hue." No one emphasized Italians' dark features more than popular writer and former newsman Walter Burns. In his book, The One-Way Ride, Johnny Torrio was "a slight, dapper, dark young man"; gunmen John Scalise and Albert Anselmi had "dark faces"; the Genna brothers were "swarthy, black haired, black eyed, looked not unlike Arabs, and probably had in their ancestral strain a strong dash of Saracenic [North African] blood".

From these descriptions you'd probably picture really dark Saudi Arabians or maybe even mixed-race Berbers, but here's what those people actually looked like (the rare mugshot of Capone has been skillfully colorized to show his blue eyes):

Al Capone

Genna Brothers

Johnny Torrio

John Scalise and Albert Anselmi

So the lesson is to not take descriptions like that literally or as meaning something "non-white". Europeans (including Southern Europeans) actually have the lightest untanned skin in the world, so even when they're "dark" or "swarthy", they're still lighter than everyone else.

Related: Al Capone: From "Dark" to "Fair"

Who's Really "More European"?

Until a few years ago, people thought that Northern Europeans were descended from indigenous Paleolithic hunter-gatherers, and Southern Europeans from more recent Neolithic farmers from the Middle East. Nordic supremacists claimed that this made them "more European" because their ancestors had been in Europe much longer and they had almost no Middle Eastern or other foreign ancestry. This went with their belief that "pure Nordics" were responsible for all European achievement.

Thanks to the ancient DNA revolution, we now know that Neolithic farmers spread much deeper into Europe demographically than was previously thought (back in 1993 by Luca Cavalli-Sforza and his colleagues):

Cavalli-Sforza was especially interested in interpreting the genetic clusters among present-day people in terms of population history. He and his colleagues analyzed their blood group data by using a technique that identifies combinations of biological variations that are most efficient at summarizing differences across individuals. Plotting these combinations of blood group types onto a map of West Eurasia, they found that the one summarizing the most variation across individuals reached its extreme value in the Near East, and declined along a southeast-to-northwest gradient into Europe. They interpreted this as a genetic footprint of the migration of farmers into Europe from the Near East, known from archaeology to have occurred after nine thousand years ago. The declining intensity suggested to them that after arriving in Europe, the first farmers mixed with local hunter-gatherers, accumulating more hunter-gatherer ancestry as they expanded, a process they called "demic diffusion." Until recently, many archaeologists viewed the demic diffusion model as an exemplary merging of insights from archaeology and genetics.

The model that Cavalli-Sforza and colleagues proposed to describe the data was intellectually attractive, but it was wrong. Its flaws became apparent beginning in 2008, when John Novembre and colleagues demonstrated that gradients like those observed in Europe can arise without migration. They then showed that a Near Eastern farming expansion into Europe might counter-intuitively cause the mathematical technique that Cavalli-Sforza used to produce a gradient perpendicular to the direction of migration, not parallel to it as had been seen in the real data.

It took the revolution wrought by the ability to extract DNA from ancient bones — the "ancient DNA revolution" — to drive a nail into the coffin of the demic diffusion model. The ancient DNA revolution documented that the first farmers even in the most remote reaches of Europe — Britain, Scandinavia, and Iberia — had very little hunter-gatherer-related ancestry. In fact, they had less hunter-gatherer ancestry than is present in diverse European populations today. The highest proportion of early farmer ancestry in Europe is today not in Southeast Europe, the place where Cavalli-Sforza thought it was most common based on the blood group data, but instead is in the Mediterranean island of Sardinia to the west of Italy.

Compare the early estimates of Anatolian farmer ancestry in Europe to the more accurate estimates of today:


It's still a bit lower in Northern Europe, but that's because it was displaced by a group of pastoralists from the Russian steppe called Yamnaya who arrived even later (less than 5000 years ago during the Bronze Age) and had a lot of another kind of Middle Eastern ancestry (from the Caucasus/Iran) plus a component from way out in Siberia called Ancient North Eurasian that's related to Native American Indians:

The extraordinary fact that emerges from ancient DNA is that just five thousand years ago, the people who are now the primary ancestors of all extant northern Europeans had not yet arrived.

The Tide from the East

[...]

The Yamnaya emerged from previous cultures of the steppe and its periphery and exploited the steppe resources far more effectively than their predecessors. They spread over a vast region, from Hungary in Europe to the foothills of the Altai Mountains in central Asia, and in many places replaced the disparate cultures that had preceded them with a more homogeneous way of life.

[...]

Our analysis of DNA from the Yamnaya — led by Iosif Lazaridis in my laboratory — showed that they harbored a combination of ancestries that did not previously exist in central Europe. The Yamnaya were the missing ingredient, carrying exactly the type of ancestry that needed to be added to early European farmers and hunter-gatherers to produce populations with the mixture of ancestries observed in Europe today. Our ancient DNA data also allowed us to learn how the Yamnaya themselves had formed from earlier populations. From seven thousand until five thousand years ago, we observed a steady influx into the steppe of a population whose ancestors traced their origin to the south — as it bore genetic affinity to ancient and present-day people of Armenia and Iran — eventually crystallizing in the Yamnaya, who were about a one-to-one ratio of ancestry from these two sources.

[...]

...the frequencies of mutations in northern Europeans today tend to be intermediate between those of southern Europeans and Native Americans. He hypothesized that these findings could be explained by the existence of a "ghost population" — the Ancient North Eurasians — who were distributed across northern Eurasia more than fifteen thousand years ago and who contributed both to the population that migrated across the Bering land bridge to people the Americas and to northern Europeans. A year later, Eske Willerslev and colleagues obtained a sample of ancient DNA from Siberia that matched the predicted Ancient North Eurasians—the Mal'ta individual whose skeleton dated to around twenty-four thousand years ago.

How could the finding of an Ancient North Eurasian contribution to present-day northern Europeans be reconciled with the two-way mixture of indigenous European hunter-gatherers and incoming farmers from Anatolia that had been directly demonstrated through ancient DNA studies? The plot became even thicker as we and others obtained additional ancient DNA data from hunter-gatherers and farmers between eight thousand and five thousand years ago and found that they fit the two-way mixture model without any evidence of Ancient North Eurasian ancestry. Something profound must have happened later — a new stream of migrants must have arrived, introducing Ancient North Eurasian ancestry and transforming Europe.

What's most ironic is that it's actually all the later "non-European" Middle Eastern ancestry, including a component called Basal Eurasian, that made Europeans become "more European" (i.e. more Caucasoid, i.e. "whiter") and more homogeneous than they would have been, because the original Paleolithic hunter-gatherers, like Ancient North Eurasians, actually had affinities to East Asians:

Lazaridis was trying to understand a peculiar Four Population Test result showing that East Asians, present-day Europeans, and pre-farming European hunter-gatherers from around eight thousand years ago are not related to one another according to the tree model. Instead, his analysis showed that East Asians today are genetically more closely related on average to the ancestors of ancient European hunter-gatherers than they are to the ancestors of present Europeans. Ancient DNA studies prior to his work had already shown that present-day Europeans derive some of their ancestry from migrations of farmers from the Near East, who I had assumed were derived from the same ancestral population as European hunter-gatherers. Lazaridis now realized that the ancestry of the first European farmers was distinct from European hunter-gatherers in some way. Something more complicated was going on.

[...]

Present-day Europeans and Near Easterners are mixed: they carry within them ancestry from a divergent Eurasian lineage that branched from Mal'ta, European hunter-gatherers, and East Asians before those three lineages separated from one another.

Lazaridis called this lineage "Basal Eurasian" to denote its position as the deepest split in the radiation of lineages contributing to non-Africans. The Basal Eurasians were a new ghost population, one as important as the Ancient North Eurasians, measured by the sheer number of descendant genomes they have left behind. The extent of the deviations of the Four Population Test away from the value of zero that would be expected if the populations were related by a simple tree indicates that this ghost population contributed about a quarter of the ancestry of present-day Europeans and Near Easterners. It also contributed comparable proportions of ancestry to Iranians and Indians.

[...]

After around fourteen thousand years ago, a group of hunter-gatherers spread across Europe with ancestry quite different from that of the people associated with the preceding Magdalenian culture, whom they largely displaced. Individuals living in Europe between thirty-seven thousand and fourteen thousand years ago were all plausibly descended from a common ancestral population that separated earlier from the ancestors of lineages represented in the Near East today. But after around fourteen thousand years ago, western European hunter-gatherers became much more closely related to present-day Near Easterners. This proved that new migration occurred between the Near East and Europe around this time.

[...]

The farmers in present-day Turkey expanded into Europe. [...] They mixed with local populations there and established new economies based on herding that allowed the agricultural revolution to spread into parts of the world inhospitable to domesticated crops. The different food-producing populations also mixed with one another, a process that was accelerated by technological developments in the Bronze Age after around five thousand years ago. This meant that the high genetic substructure that had previously characterized West Eurasia collapsed into the present-day very low level of genetic differentiation by the Bronze Age.

So Northern Europeans actually have a ton of the Middle Eastern ancestry that Nordicists have always looked down on. And while they still have some of their beloved European hunter-gatherer ancestry, it turns out that's Asian-related. Furthermore, large parts of their Middle Eastern and Asian/Amerindian-type ancestries arrived long after Paleolithic and Neolithic times in the Bronze Age.

Of course, the same goes for Southern Europeans, but they generally have more of the older Neolithic farmer ancestry and less of the exotic "Asiatic" ancestries. And the originators of European civilization — Minoan and Mycenaean Greeks, as well as Etruscans and Romans — were genetically like modern Southern Europeans, not like modern Northern Europeans as the Nordicists have always claimed.

So in a pissing contest of who's "more European", playing by Nordicists' own rules, Southern Europeans would have to be considered the "winners".

---------------
David Reich. Who We Are and How We Got Here: Ancient DNA and the New Science of the Human Past. Oxford: Oxford University Press, 2018.

Related: Phenotypes of Hunters and Farmers

Italianthro Restored!

Google finally restored my Italianthro account:

https://italianthro.blogspot.com/

I'll leave the new blog online but hidden from search engines and with comments disabled. If anything ever happens to the original one again, just remove the 'h' from any URL and you'll be taken there.

Italian Anthropology 2

Google disabled my Italianthro account and locked me out of it. I think it might be related to this widespread issue (which Google isn't doing shit to fix) but I never found out for sure because there is no customer support. You can fill out forms to try to unlock or restore your account, but you only get auto-generated messages. You never get a reply from a real person you can talk or ask questions to.

I don't know if that account will ever be restored, so for now I've reposted everything to this account. All of the URLs should be the same as before (i.e. post title and date) except for one letter in the name: antro now has an Italian spelling with no 'h'.

https://italiantro.blogspot.com/

Let me know if you find any broken links or other issues.

P.S. You can also access Wayback archives of the old blog (and comments), which should work forever no matter what else happens.

UPDATE 11/26/19: Italianthro Restored!

Ancient Egyptians and Nubians Were Caucasoid

Ancient DNA has confirmed that Ancient Egyptians were Middle Eastern and not Sub-Saharan African. This was already suggested by earlier genetic studies that found North Africans were the result of a back-migration from West Asia. But even before that, older craniofacial analyses of skeletal samples from all over Egypt (and Nubia) spanning all of ancient history also showed the same thing.

Here are five lines of anthropological evidence that group Ancient Egyptians and Nubians with North Africans, West/South Asians and Europeans (Caucasoids), and separate them from Sub-Saharan Africans (Negroids). Note that Horn Africans (Somalis) cluster with Caucasoids in some cases and Negroids in others, which indicates they're a racially mixed population that's not representative of either Sub-Saharan Africans or Egyptians and Nubians.

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1. Craniometric

Combined samples of Pre-Dynastic (Naqada) and Late Dynastic (Giza) Egyptians, and Bronze Age, Early Christian and Medieval Nubians, cluster with combined samples of Ancient and Modern North Africans, East Indians and Europeans.


Brace et al. "Clines and clusters versus 'Race:' a test in ancient Egypt and the case of a death on the Nile". Am J Phys Anthro, 1993.

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2. Cranial Non-metric

Pre-Dynastic Egyptians from Naqada (#59), 26th-30th Dynasty Egyptians from Gizeh (#60), 12th-13th Dynasty Nubians from Kerma (#61), and Early Christian or Christian Nubians (#62) cluster with South Asians (#44) and several European groups: Greeks (#48), Scandinavians (#51 and #52) and Germans (#53). [NOTE that Somalis are up with Sub-Saharan Africans (#63)]


Hanihara et al. "Characterization of biological diversity through analysis of discrete cranial traits". Am J Phys Anthro, 2003.

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3. Dental Metric

Pre-Dynastic and 12th-29th Dynasty Egyptians cluster with Afghans and North Indians on the edge of a larger cluster of Europeans and West Asians. [NOTE that here again, Somalis show Sub-Saharan affinities and don't cluster with Ancient Egyptians.]


Hanihara and Ishida. "Metric dental variation of major human populations". Am J Phys Anthro, 2005.

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4. Dental Non-metric

12th Dynasty (Lisht), Roman/Byzantine (El Hesa), and Byzantine (Kharga) Egyptians, and Pharonic, Meroitic, X-group and Christian Nubians, cluster with other North Africans and Europeans (Poundbury, England).


Joel D. Irish. "Diachronic and synchronic dental trait affinities of late and post-pleistocene peoples from North Africa". Homo, 1998b.

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5. Prognathism

Ancient Egyptians from Badari, Pre-Dynastic Egyptians from Naqada, and 26th-30th Dynasty Egyptians from Gizeh, as well as 12th-13th Dynasty Nubians from Kerma and Early Christian or Christian Nubians, all cluster with Europeans and West/South Asians on the negative end of the prognathism scale.


Tsunehiko Hanihara. "Frontal and Facial Flatness of Major Human Populations". Am J Phys Anthro, 2000.

Catherine Zeta Jones Is NOT Greek

The ancestry of British actress Catherine Zeta Jones is Welsh and Irish, but people keep claiming she's part Greek, and it was just repeated in a Jeopardy question. The source is an old CNN interview she did with Larry King which was transcribed online like this:

Zeta is my grandmother's name, who is of north Greek origin. But Zeta is a Greek name. And everyone thinks that I just put Zeta in to spice up Catherine Jones. And that's completely untrue.

But there's a disclaimer at the beginning that says it's a "rush transcript" that "may not be in its final form". If she were saying that her grandmother was Greek, there would be no "But..." after the first sentence. That only makes sense if she's denying having Greek ancestry, which is what she actually said:

Zeta is my grandmother's name, who is of not Greek origin. But Zeta is a Greek name. And everyone thinks that I was, just put Zeta in to spice up Catherine Jones. And that's completely untrue. [...] I've always been called that. And in my school in Wales, Jones is a very, very popular name in Wales. And so there were like three Catherine Joneses in my class, so they always called me Zeta.

It's thought that her last name is "Zeta-Jones", but she added the hyphen later. "Zeta" was originally her middle name and her grandmother's first name, and this is where it actually came from:

"I've been performing since I was 11 years old," said Catherine, named for her maternal grandmother Catherine Fair and her paternal grandmother Zeta Jones (Zeta was the name of a ship that her great-grandfather had sailed on).

Many people have also claimed, with no evidence, that she's Hispanic or even half black because they believe Nordicist stereotypes and can't accept the fact that some Northern Europeans are naturally darker and more "exotic" looking. A sane commenter responds:

Whenever a N.European has darker hair, eyes or complexion, somebody (predictably) uses a stereotype of another ethnic group to suggest they're an "outsider" to their own heritage. Despite popular stereotypes, differences in appearance are normal within any one group due to expected genetic variation within families and their older tribal histories. This is the norm--and not the exception. Anyone proud of their heritage won't like racists taking that from them.

Catherine Zeta Jones is just a dark Brit. That's all.

What "Olive-Skinned" Really Means

Most people either think that "olive" skin tone is the same as "tan" or "dark", or that it refers to some kind of "nonwhite" or "mixed" ethnicity ranging from the Mediterranean region to Latin America. But it's actually one of several skin undertones that have nothing to do with what race or shade someone is.

When shopping for foundation, you've probably heard the terms "cool," "warm," or "neutral" to describe how a shade will look on skin. Those terms refer to your skin's undertone and are used to determine which foundation shade will match it the best.

Cool, warm, or neutral undertones are the colors that come through your skin from underneath the surface to affect its overall hue. It's not about how light or dark your skin is; people of all skin colors, from very fair to deep, can have cool, warm, or neutral undertones. Here's what each of these terms means:

Cool: Hints of bluish, pink, or a ruddy complexion.

Warm: Skin skews yellow, sallow, peachy, or golden.

Neutral: Has no obvious overtones of pink or sallow skin, but rather the skin's natural color is more evident.

[...]

Does your skin look somewhat ashen or gray? You might have the wild card of the bunch — olive skin — which is a combination of the natural neutral, slightly yellow undertone everyone has plus the greenish ashen hue that's unique to olive skin. Olive skin tone is very specific, but is not neutral, as some tend to call it.

Here's what the 4 different undertones would look like on light untanned skin:


And here are people of different races and shades with 3 of the undertones:

Indo-Europeans Were Likely from the Near East

A new book and a new study on ancient DNA both say that the homeland of Indo-European languages was most likely south of the Caucasus Mountains in the Near East, and not in the Russian Steppe from where many later spread out. This would explain why the earliest IE languages were in Anatolia, and why IE-speakers in SE Europe have higher "Caucasus/Iran" ancestry and lower "Russian/Siberian" ancestry.

While the genetic findings point to a central role for the Yamnaya in spreading Indo-European languages, tipping the scales definitively in favor of some variant of the steppe hypothesis, those findings do not yet resolve the question of the homeland of the original Indo-European languages, the place where these languages were spoken before the Yamnaya so dramatically expanded. Anatolian languages known from four-thousand-year-old tablets recovered from the Hittite Empire and neighboring ancient cultures did not share the full wagon and wheel vocabulary present in all Indo-European languages spoken today. Ancient DNA available from this time in Anatolia shows no evidence of steppe ancestry similar to that in the Yamnaya (although the evidence here is circumstantial as no ancient DNA from the Hittites themselves has yet been published). This suggests to me that the most likely location of the population that first spoke an Indo-European language was south of the Caucasus Mountains, perhaps in present-day Iran or Armenia, because ancient DNA from people who lived there matches what we would expect for a source population both for the Yamnaya and for ancient Anatolians. If this scenario is right, the population sent one branch up into the steppe—mixing with steppe hunter-gatherers in a one-to-one ratio to become the Yamnaya as described earlier—and another to Anatolia to found the ancestors of people there who spoke languages such as Hittite.

David Reich. Who We Are and How We Got Here: Ancient DNA and the New Science of the Human Past. New York: Pantheon Books, 2018.

The insight that the Caucasus mountains served not only as a corridor for the spread of CHG/Neolithic Iranian ancestry but also for later gene-flow from the south also has a bearing on the postulated homelands of Proto-Indo-European (PIE) languages and documented gene-flows that could have carried a consecutive spread of both across West Eurasia. Perceiving the Caucasus as an occasional bridge rather than a strict border during the Eneolithic and Bronze Age opens up the possibility of a homeland of PIE south of the Caucasus, which itself provides a parsimonious explanation for an early branching off of Anatolian languages. Geographically this would also work for Armenian and Greek, for which genetic data also supports an eastern influence from Anatolia or the southern Caucasus. A potential offshoot of the Indo-Iranian branch to the east is possible, but the latest ancient DNA results from South Asia also lend weight to an LMBA spread via the steppe belt. The spread of some or all of the proto-Indo-European branches would have been possible via the North Caucasus and Pontic region and from there, along with pastoralist expansions, to the heart of Europe. This scenario finds support from the well attested and now widely documented 'steppe ancestry' in European populations, the postulate of increasingly patrilinear societies in the wake of these expansions (exemplified by R1a/R1b), as attested in the latest study on the Bell Beaker phenomenon.

Wang et al. "The genetic prehistory of the Greater Caucasus". bioRxiv, 2018.

Eurasian Origin and Back-Migration of L3 and DE

This was already suggested a few years back by Farrell et al. (2013) and Shi Yan et al. (2013), and now a new study is saying the same thing again.

Background: After three decades of mtDNA studies on human evolution the only incontrovertible main result is the African origin of all extant modern humans. In addition, a southern coastal route has been relentlessly imposed to explain the Eurasian colonization of these African pioneers. Based on the age of macrohaplogroup L3, from which all maternal Eurasian and the majority of African lineages originated, that out-of-Africa event has been dated around 60-70 kya. On the opposite side, we have proposed a northern route through Central Asia across the Levant for that expansion. Consistent with the fossil record, we have dated it around 125 kya. To help bridge differences between the molecular and fossil record ages, in this article we assess the possibility that mtDNA macrohaplogroup L3 matured in Eurasia and returned to Africa as basic L3 lineages around 70 kya.

Results: The coalescence ages of all Eurasian (M,N) and African L3 lineages, both around 71 kya, are not significantly different. The oldest M and N Eurasian clades are found in southeastern Asia instead near of Africa as expected by the southern route hypothesis. The split of the Y-chromosome composite DE haplogroup is very similar to the age of mtDNA L3. A Eurasian origin and back migration to Africa has been proposed for the African Y-chromosome haplogroup E. Inside Africa, frequency distributions of maternal L3 and paternal E lineages are positively correlated. This correlation is not fully explained by geographic or ethnic affinities. It seems better to be the result of a joint and global replacement of the old autochthonous male and female African lineages by the new Eurasian incomers.

Conclusions: These results are congruent with a model proposing an out-of-Africa of early anatomically modern humans around 125 kya. A return to Africa of Eurasian fully modern humans around 70 kya, and a second Eurasian global expansion by 60 kya. Climatic conditions and the presence of Neanderthals played key roles in these human movements.

Cabrera et al. "Carriers of mitochondrial DNA macrohaplogroup L3 basic lineages migrated back to Africa from Asia around 70,000 years ago". bioRxiv, 2017.

Genetic Continuity in Greece

Greek women through the ages: Minoan, Cycladic, Mycenaean, Classical, Modern  (SOURCE)

A recent study suggested based on modern samples that Greeks hadn't changed much since ancient times. Now that's been proven with ancient DNA from Bronze Age Minoan and Mycenaean Greeks. Their ancestry is mostly Anatolian Neolithic farmer with some Caucasus/Iran and less Russian/Siberian admixture (the last two related to the spread of Indo-European languages), they had dark hair and dark eyes, and there was no difference between the elites and common people. They're genetically closest to modern Southeastern Europeans, and not to Northern Europeans or Africans as different people have claimed.

The origins of the Bronze Age Minoan and Mycenaean cultures have puzzled archaeologists for more than a century. We have assembled genome-wide data from 19 ancient individuals, including Minoans from Crete, Mycenaeans from mainland Greece, and their eastern neighbours from southwestern Anatolia. Here we show that Minoans and Mycenaeans were genetically similar, having at least three-quarters of their ancestry from the first Neolithic farmers of western Anatolia and the Aegean, and most of the remainder from ancient populations related to those of the Caucasus and Iran. However, the Mycenaeans differed from Minoans in deriving additional ancestry from an ultimate source related to the hunter–gatherers of eastern Europe and Siberia, introduced via a proximal source related to the inhabitants of either the Eurasian steppe or Armenia. Modern Greeks resemble the Mycenaeans, but with some additional dilution of the Early Neolithic ancestry. Our results support the idea of continuity but not isolation in the history of populations of the Aegean, before and after the time of its earliest civilizations.

[...]

The elite Mycenaean individual from the ‘royal’ tomb at Peristeria in the western Peloponnese did not differ genetically from the other three Mycenaean individuals buried in common graves.

[...]

Other proposed migrations, such as settlement by Egyptian or Phoenician colonists, are not discernible in our data, as there is no measurable Levantine or African influence in the Minoans and Mycenaeans, thus rejecting the hypothesis that the cultures of the Aegean were seeded by migrants from the old civilizations of these regions.

[...]

Phenotype prediction from genetic data has enabled the reconstruction of the appearance of ancient Europeans who left no visual record of their pigmentation. By contrast, the appearance of the Bronze Age people of the Aegean has been preserved in colourful frescos and pottery, depicting people with mostly dark hair and eyes. We used the HIrisPlex tool (Supplementary Information section 4) to infer that the appearance of our ancient samples matched the visual representations (Extended Data Table 2), suggesting that art of this period reproduced phenotypes naturalistically.

We estimated the fixation index, F_ST, of Bronze Age populations with present-day West Eurasians, finding that Mycenaeans were least differentiated from populations from Greece, Cyprus, Albania, and Italy (Fig. 2), part of a general pattern in which Bronze Age populations broadly resembled present-day inhabitants from the same region (Extended Data Fig. 7).

The modern Greek samples used in the study for comparison are not the best. Thessaloniki is in the north of the country, the Coriell database doesn't provide specific origins, and there are only two Cretans. I believe that if Mycenaeans were compared to their descendants in the Peloponnese, continuity would be nearly perfect instead of just very strong.


Lazaridis et al. "Genetic origins of the Minoans and Mycenaeans". Nature, 2017.

Related: Ancient Roman DNA, Ancient-to-Modern Genetic Distances

Reconstruction of an Ancient Egyptian

A 3,500-year-old noble Egyptian called Nebiri has been brought back to life through modern forensics.

Scientists have reconstructed the face of the ancient mummy, and discovered he had a prominent nose, wide jaw, straight eyebrows and moderately thick lips.

[...]

Nebiri is thought to have been a member of the Egyptian elite who served as the Chief of the Stables, looking after royal horses, during the reign of Thutmoses III, a pharaoh from the 18th Dynasty of ancient Egypt.

His remains were discovered in the Valley of the Queens in Luxor in 1904, but as the tomb has been plundered, just his head and jars containing his organs remained.

Researchers from the University of Turin have now used a range of facial reconstruction techniques to produce an impressive facial approximation.

To reconstruct his face, the researchers used a mixture of computer modelling and anthropological research.

The team then used a computer programme to start to build up a picture of the Egyptian's face.

[...]

Speaking to Live Science, Raffaella Bianucci, who led the study, said: 'He was between 45 [and] 60 years old when he died.

Shivali Best. "The face of Nebiri revealed: Scientists reconstruct the head of the ancient Egyptian 'Chief of Stables', 3,500 years after he died of heart failure". MailOnline – Science & Tech, 11:01 BST, 20 June 2017.

Genetics of Peloponnesean Greeks

This new study refutes Nordicist and Afrocentrist claims of population replacement in Ancient Greece, showing that modern Peloponnesean Greeks are most closely related to other Southern Europeans, and far from both Slavic and non-European groups. They're also distinct from Greek-speaking populations in Asia Minor, only partly overlapping with those on the Aegean coast nearest to Greece.

Peloponnese has been one of the cradles of the Classical European civilization and an important contributor to the ancient European history. It has also been the subject of a controversy about the ancestry of its population. In a theory hotly debated by scholars for over 170 years, the German historian Jacob Philipp Fallmerayer proposed that the medieval Peloponneseans were totally extinguished by Slavic and Avar invaders and replaced by Slavic settlers during the 6th century CE. Here we use 2.5 million single-nucleotide polymorphisms to investigate the genetic structure of Peloponnesean populations in a sample of 241 individuals originating from all districts of the peninsula and to examine predictions of the theory of replacement of the medieval Peloponneseans by Slavs. We find considerable heterogeneity of Peloponnesean populations exemplified by genetically distinct subpopulations and by gene flow gradients within Peloponnese. By principal component analysis (PCA) and ADMIXTURE analysis the Peloponneseans are clearly distinguishable from the populations of the Slavic homeland and are very similar to Sicilians and Italians. Using a novel method of quantitative analysis of ADMIXTURE output we find that the Slavic ancestry of Peloponnesean subpopulations ranges from 0.2 to 14.4%. Subpopulations considered by Fallmerayer to be Slavic tribes or to have Near Eastern origin, have no significant ancestry of either. This study rejects the theory of extinction of medieval Peloponneseans and illustrates how genetics can clarify important aspects of the history of a human population.

Figure 2: Genetic similarity of Peloponneseans, Sicilians and Italians. PCA analysis of several European populations. (a) Notice the north to south distribution of the populations and that the Peloponneseans are placed to the far right of the graph and overlap with the Sicilians. (b) PCA analysis of Southern European populations illustrating the close relationship between Peloponneseans, Sicilians and Italians (TSI is an Italian population). (c) Network analysis illustrating the high connectivity between the Peloponnesean populations as well as between the Peloponneseans, the Sicilians and the Italians. Notice the distance between Peloponneseans and the Slavic, and Near Eastern populations. Peloponneseans are connected with the Near Eastern populations through Crete and Dodecanese.

Figure 3: Testing the theory of replacement of medieval Peloponneseans by Slavs and Asia Minor settlers. (a) PCA analysis shows the broad separation of Peloponneseans from four populations of the Slavic homeland (Ukrainians, Polish, Russians and Belarusians). (b) PCA comparisons of the Peloponneseans with three Greek-speaking Asia Minor populations shows only partial overlap with the population of the Asia Minor Aegean coast.

Supplementary Figure 2: Comparisons of Peloponneseans with non-European populations. PCA analysis of Peloponneseans and A. Near East. B. Caucasus. C. North Africa. D. East Africa. E. Arabia. F. West Siberia populations.

Stamatoyannopoulos et al. "Genetics of the peloponnesean populations and the theory of extinction of the medieval peloponnesean Greeks". Eur J Hum Genet, 2017.

Related: Genetic Continuity in Greece

Scythians Had East Asian Ancestry

Nordicists have often claimed that Iron Age Scythians were blonde, blue-eyed "Aryans" most similar to modern Northern Europeans, but ancient DNA analysis shows they were a mix of Yamnaya people from the Russian Steppe (who were mostly brunet) and East Asian Mongoloids.

During the 1st millennium before the Common Era (BCE), nomadic tribes associated with the Iron Age Scythian culture spread over the Eurasian Steppe, covering a territory of more than 3,500 km in breadth. To understand the demographic processes behind the spread of the Scythian culture, we analysed genomic data from eight individuals and a mitochondrial dataset of 96 individuals originating in eastern and western parts of the Eurasian Steppe. Genomic inference reveals that Scythians in the east and the west of the steppe zone can best be described as a mixture of Yamnaya-related ancestry and an East Asian component. Demographic modelling suggests independent origins for eastern and western groups with ongoing gene-flow between them, plausibly explaining the striking uniformity of their material culture. We also find evidence that significant gene-flow from east to west Eurasia must have occurred early during the Iron Age.

[...]

Since the PCA of west Eurasia in Fig. 4 does not allow one to examine the ancient samples in relation to contemporary East Asian populations, we also carried out PCA of all 2,345 modern individuals in the Human Origins dataset, onto which we also projected the ancient individuals (Fig. 5). It is evident from this PCA that ancestry of the Iron Age samples falls on a continuum between present-day west Eurasians and eastern non-Africans, which is in concordance with the mitochondrial haplogroup analyses. The eastern Scythians display nearly equal proportions of mtDNA lineages common in east and west Eurasia, whereas in the western Scythian groups, the frequency of lineages now common in east Eurasia is generally lower, even reaching zero in four samples of the initial Scythian phase of the eight to sixth century BCE (group #1 in Fig. 2), and reaches 18–26% during later periods (sixth to second century BCE; #2 and #3) (Supplementary Table 7).

The Scythian samples are in black:

Figure 5 | Principal component analysis. PCA of ancient individuals (according colours see legend) projected on modern individuals of the Human Origins dataset (grey). Iron Age Scythians are shown in black; CHG, Caucasus hunter-gatherer; LNBA, late Neolithic/Bronze Age; MN, middle Neolithic; EHG, eastern European hunter-gatherer; LBK_EN, early Neolithic Linearbandkeramik; HG, hunter-gatherer; EBA, early Bronze Age; IA, Iron Age; LBA, late Bronze Age; WHG, western hunter-gatherer.

Unterlander et al. "Ancestry and demography and descendants of Iron Age nomads of the Eurasian Steppe". Nature Communications, 2017.

First Ancient Egyptian Genomes

This is a talk that will be given at the 82nd annual meeting of the Society for American Archaeology in Vancouver, BC, Canada from March 29–April 2, 2017. New genome-wide data show that Ancient Egyptians had less Sub-Saharan African ancestry than Modern Egyptians and were closely related to Middle Easterners, which will surely upset Afrocentrists. More ancient genomes from earlier periods should follow soon.

[203] Ancient Egyptian Mummy Genomes Suggest an Increase of Sub-Saharan African Ancestry in Post-Roman Periods

Krause, Johannes (Max Planck Institute—SHH), Verena Schuenemann (Institute for Archaeological Sciences, University of Tübingen), Alexander Peltzer (Department for Archaeogenetics, Max Planck Inst), Wolfgang Haak (Department for Archaeogenetics, Max Planck Inst) and Stephan Schiffels (Department for Archaeogenetics, Max Planck Inst)

Egypt, located on the isthmus of Africa, is an ideal region to study historical population dynamics due to its geographic location and documented interactions with ancient civilizations in Africa, Asia, and Europe. Particularly, in the first millennium BCE Egypt endured foreign domination leading to growing numbers of foreigners living within its borders possibly contributing genetically to the local population. Here we mtDNA and nuclear DNA from mummified humans recovered from Middle Egypt that span around 1,300 years of ancient Egyptian history from the Third Intermediate to the Roman Period. Our analyses reveal that ancient Egyptians shared more Near Eastern ancestry than present-day Egyptians, who received additional Sub-Saharan admixture in more recent times. This analysis establishes ancient Egyptian mummies as a genetic source to study ancient human history and offers the perspective of deciphering Egypt’s past at a genome-wide level.

Hopefully a deeper transect, into the Old Kingdom, Early Dynastic, and Predynastic, is to follow. My prediction (I would be happy to be wrong) is that this DNA came from tooth or bone — I think mummified soft tissue has mostly been a source of disappointment. Differential relatedness of modern Copts and non-Copts to the ancient samples would be something to look out for.

If any of these talks is going to really upset some people, it’ll be this one.

LINK

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UPDATE 06/01/17: The study that goes with this talk is out now, and the Ancient Egyptian samples cluster with ancient Neolithic Levantines, and modern Bedouin, Palestinian and Lebanese Arabs.

ABSTRACT: Egypt, located on the isthmus of Africa, is an ideal region to study historical population dynamics due to its geographic location and documented interactions with ancient civilizations in Africa, Asia and Europe. Particularly, in the first millennium BCE Egypt endured foreign domination leading to growing numbers of foreigners living within its borders possibly contributing genetically to the local population. Here we present 90 mitochondrial genomes as well as genome-wide data sets from three individuals obtained from Egyptian mummies. The samples recovered from Middle Egypt span around 1,300 years of ancient Egyptian history from the New Kingdom to the Roman Period. Our analyses reveal that ancient Egyptians shared more ancestry with Near Easterners than present-day Egyptians, who received additional sub-Saharan admixture in more recent times. This analysis establishes ancient Egyptian mummies as a genetic source to study ancient human history and offers the perspective of deciphering Egypt’s past at a genome-wide level.

Figure 4 | Principal component analysis and genetic clustering of genome-wide DNA from three ancient Egyptians. (a) Principal Component Analysis-based genome-wide SNP data of three ancient Egyptians, 2,367 modern individuals and 294 previously published ancient genomes, (b) subset of the full ADMIXTURE analysis (Supplementary Fig. 4).

Schuenemann et al. "Ancient Egyptian mummy genomes suggest an increase of Sub-Saharan African ancestry in post-Roman periods". Nature Communications, 2017.

Natufians NOT Sub-Saharan African

This new ancient DNA study refutes the Afrocentrist claim that Natufians were from Sub-Saharan Africa or had Negroid admixture.

Craniometric analyses have suggested that the Natufians may have migrated from north or sub-Saharan Africa, a result that finds some support from Y chromosome analysis which shows that the Natufians and successor Levantine Neolithic populations carried haplogroup E, of likely ultimate African origin, which has not been detected in other ancient males from West Eurasia (Supplementary Information, section 6). However, no affinity of Natufians to sub-Saharan Africans is evident in our genome-wide analysis, as present-day sub-Saharan Africans do not share more alleles with Natufians than with other ancient Eurasians (Extended Data Table 1).

Lazaridis et al. "The genetic structure of the world's first farmers". bioRxiv, 2016.

Related: Natufians NOT Source of European Neolithic

Phenotypes of Hunters and Farmers

Europeans are descended from prehistoric hunter-gatherers and farmers. Here's what we know about the origins and physical appearance of these two populations from anthropology (Coon 1939, Pinhasi 2012) and genetics (Lipson 2012, Lazaridis 2014), along with representations of what they might have looked like:

Mesolithic Hunters had broad faces, dark skin, light eyes and were intermediate between Western and Eastern Eurasians. So to represent them I chose a Uralic Norwegian Lapp that I darkened and gave blue eyes.

Neolithic Farmers had narrow faces, light skin, dark eyes and were Western Eurasian, closest to modern Sardinians. So I chose an untanned, long-faced Mediterranean soccer player from Sardinia to represent them.


Of course, they didn't all look exactly the same. We know that there was diversity and overlap in some of their traits (Gamba 2014). But in general, the phenotypic variation we see in Europe today is the result of waves of settlements by these two distinct types from Siberia and the Middle East since ancient times, and the mixing that occurred between them in different proportions (Haak 2015), plus selective pressures favoring further depigmentation in some places.


Related: Who's Really "More European"?

AIMs Overestimate Admixture

AIMs are a subset of SNPs chosen for their informativeness about ancestry and often used by geneticists instead of genome-wide data to save time and money. However, according to Galanter et al. (2010), this can lead to errors and overestimations of admixture, especially when the panel of AIMs is very small:

Ancestry informative markers (AIMs) have been used as a cost-effective way to estimate individual ancestral proportions in admixed populations such as African Americans and Latinos. [...] We compared differences in ancestry estimated with different size AIMs panels with ancestry estimated from genomewide markers. [...] There was an inverse correlation between the number of AIMs used to estimate ancestry and mean and standard deviation of the error in ancestry estimation. Using AIMs, African ancestry was consistently overestimated, while the major ancestral component (European in Puerto Ricans and Native American in Mexicans) was systematically underestimated. Using 300 or fewer AIMs consistently produced a standard deviation of ancestry estimation error of 10% or greater. [...] There is both systematic bias resulting in overestimation of African ancestry (and underestimation of other continental ancestry) and random error. Such error is inversely proportional to the number of AIMs used.

Bauchet et al. (2007) found that even larger panels of AIMs, while somewhat more accurate, still lead to a loss of structure, and therefore an overestimation of admixture, compared with using the full SNP data set:

Using <1,200 EuroAIMs of the type available in this panel gradually leads to loss of consistent structure and a corresponding increase in misclassification of individual origins (fig. 7C).

While the number of AIMs used is clearly a big factor in the accuracy level of results, another problem is that AIMs may not even be as informative about ancestry as they claim, according to Bolnick et al. (2007):

Furthermore, some of the most "informative" AIMs involve loci that have undergone strong selection, which makes it unclear whether these markers indicate shared ancestry or parallel selective pressures (such as similar environmental exposures in different geographic regions) or both.

Hopefully, all this criticism will get more notice, and geneticists will stop trying to cut corners by using these inferior markers for quantifying individual ancestry.

Related: Overestimated Admixture in Brisighelli (2012)

European Ancestry of Ashkenazi Jews

According to a new genetic study, Ashkenazi Jews are mostly European on their maternal side, and that admixture comes from Western and Central Europe when diaspora males from the Levant arrived first in Rome and found wives among local Italian women who converted to Judaism. They then migrated further west and north and acquired other European admixture in the same way, before finally heading east. Intermixing slowed after that because they have very little Slavic or Turkic (Khazar) admixture. Their paternal side remains mostly Near Eastern. This all fits well with findings from anthropology some 75 years ago.

Overall, it seems that at least 80% of Ashkenazi maternal ancestry is due to the assimilation of mtDNAs indigenous to Europe, most likely through conversion. The phylogenetic nesting patterns suggest that the most frequent of the Ashkenazi mtDNA lineages were assimilated in Western Europe, ~2 ka or slightly earlier. Some in particular, including N1b2, M1a1b, K1a9 and perhaps even the major K1a1b1, point to a north Mediterranean source. It seems likely that the major founders were the result of the earliest and presumably most profound wave of founder effects, from the Mediterranean northwards into central Europe, and that most of the minor founders were assimilated in west/central Europe within the last 1,500 years. The sharing of rarer lineages with Eastern European populations may indicate further assimilation in some cases, but can often be explained by exchange via intermarriage in the reverse direction.

The Ashkenazim therefore resemble Jewish communities in Eastern Africa and India, and possibly also others across the Near East, Caucasus and Central Asia, which also carry a substantial fraction of maternal lineages from their 'host' communities. Despite widely differing interpretations of autosomal data, these results in fact fit well with genome-wide studies, which imply a significant European component, with particularly close relationships to Italians. As might be expected from the autosomal picture, Y-chromosome studies generally show the opposite trend to mtDNA (with a predominantly Near Eastern source) with the exception of the large fraction of European ancestry seen in Ashkenazi Levites.

Evidence for haplotype sharing with non-Ashkenazi Jews for each of the three main haplogroup K founders may imply a partial common ancestry in Mediterranean Europe for Ashkenazi and Spanish-exile Sephardic Jews, but may also, at least in part, be due to subsequent gene flow, especially into Bulgaria and Turkey, both of which witnessed substantial immigration from Ashkenazi communities in the fourteenth and fifteenth centuries. Gene flow could have been substantial in some cases—ongoing intermarriage is likely when these communities began living in closer proximity after the Spanish exile. A partial common ancestry for all European Jews—both Ashkenazi and Sephardic—is again strongly supported by the autosomal results.

Jewish communities were already spread across the Graeco-Roman and Persian world >2,000 years ago. It is thought that a substantial Jewish community was present in Rome from at least the mid-second century BCE, maintaining links to Jerusalem and numbering 30,000-50,000 by the first half of the first century CE. By the end of the first millennium CE, Ashkenazi communities were historically visible along the Rhine valley in Germany. After the wave of expulsions in Western Europe during the fifteenth century, they began to disperse once more, into Eastern Europe.

These analyses suggest that the first major wave of assimilation probably took place in Mediterranean Europe, most likely in the Italian peninsula ~2 ka, with substantial further assimilation of minor founders in west/central Europe. There is less evidence for assimilation in Eastern Europe, and almost none for a source in the North Caucasus/Chuvashia, as would be predicted by the Khazar hypothesis—rather, the results show strong genetic continuities between west and east European Ashkenazi communities, albeit with gradual clines of frequency of founders between east and west.

Costa et al. "A substantial prehistoric European ancestry amongst Ashkenazi maternal lineages". Nature Communications, 2013.

The Jews have been left to the end because they do not as a whole fit into any single racial classification heretofore outlined. Historically the Jews of the Biblical period in Palestine were a Semitic-speaking people composed of various Mediterranean strains which had blended together at the time of the formation of the Jewish nation. These Mediterranean strains must have included a small Mediterranean type comparable to the present Yemeni Arabs; a taller, longer-faced strain with a tendency to nasal convexity, as is found among Irano-Afghan peoples today; and a straight-nosed, presumably Atlanto-Mediterranean element contributed by the Philistines.

The Jews began their expansion from Palestine as early as the time of the Babylonian Captivity; at this time they settled Mesopotamia in large numbers, and from there began an expansion into central Asia of which colonies still remain. In the Hellenistic period they migrated into Asia Minor and the Black Sea region, as well as into Egypt; these emigrants became Hellenistic Jews. Under the Romans they settled in Italy, France, and Spain, with especial concentrations in Spain and in the cities of the Rhineland. The Jews expelled from Spain in 1492 and during previous expulsions became the Sephardim, whose descendants are to be found in various countries bordering on the Mediterranean, especially Morocco, the Salonika region of what is now Greece, and Turkey. The Rhineland Jews, persecuted at the time of the First Crusade, moved eastward into Poland, the Ukraine and other central European countries, and met there and absorbed a group of Hellenistic Jews moving westward, among whom were some who had lived among the Turkish Khazars in the Crimea and elsewhere. The two groups blended and the Germanic speech of the more numerous western element prevailed. The modern Yiddish-speaking Ashkenazim are the descendants of this amalgamated body. Racially they preserve to a large measure their Mediterranean character, altered partly by Alpine admixture which has in many cases produced Dinaricization. This Alpine, as well as some Nordic, admixture was probably obtained largely in France and Germany before their departure eastward. The most persistent Palestinian Mediterranean traits which the Jews preserve is a narrowness of the face. The Jewish facial expression, by which many Jews may be distinguished, is a cultural and not a genetic character.

Carleton Coon. The Races of Europe. New York: MacMillan, 1939.