Ancient North Eurasians from Siberia

September 3, 2020

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).

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

August 20, 2020

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 JuliĆ  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

July 3, 2020

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"?

December 4, 2019

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!

November 26, 2019

Google finally restored my Italianthro account:

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

November 4, 2019

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'.

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

October 8, 2019

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.

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.

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.

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.

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.

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

July 16, 2019

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

June 9, 2019

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

May 20, 2018

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.