Migration at the roof of West Asia

Click to see the full figure

The figure to the left is from The genetic prehistory of the Greater Caucasus. If you are a regular reader of this weblog, or Eurogenes, you can figure out what’s going on, and keep track of the terminology. But in 2018 I think we’re getting to the end of the line in making sense of “admixture graphs” in relation to West Eurasian population structure. The models are just getting too complicated to keep everything straight, and the distinct-populations-subject-to-pulse-admixture seems to be an assumption that may not necessarily hold.

To get a sense of what I’m talking about, the above preprint focuses on populations in and around the Caucasus region. One of the major reasons that this is important is that the Caucasus was and is to some extent a continental hinge, connecting Eastern Europe and the Pontic steppe, to the Near East. The Arab Muslims pushed north of the Caucasus, and came into conflict with the Khazars, while Cimmerians and Scythians moved south from the Pontic steppe.

The elephant in the room is the relevance to the “Indo-European controversy.” Colin Renfrew long ago posited that the Indo-European languages derive from West Asian farmers who expanded into Europe as early as ~9,000 years ago. A rival theory is that Indo-Europeans spread out of the Pontic steppe ~4,000 years ago. In 2015 two major papers suggested that the steppe was a major source of Indo-European expansion. Case closed? This preprint suggests perhaps not.

But we’ll get to that later. What do the results here show? The prose is a little hard to tease apart, but the major issues seem to be that in antiquity, or at least the period they’re focusing on, much of the gene flow seems to have been south (Near East) to the north (through the Caucasus, and out to the north slope). To some extent, we already knew this: the Yamna people of the Pontic steppe have “southern” ancestry from the Near East that earlier East European/Pontic people do not. In this preprint, the authors show that groups such as the Maykop of the north slope of the Caucasus carry Y haplogroups such as G2, and not the R1 lineages commonly found in the steppe. David W. suggests that this confirms that Near Eastern gene flow into the steppe was female-mediated.  This is plausible, but I would caution that Y chromosomes alone can be deceptive, due to the power of particular patrilineages. We’ll probably rely on the X chromosome to make a final judgment.

The plot below shows many of the relationships as a function of location and time. The green component is modal among “Iranian farmers,” the orange among “Anatolian farmers,” and the blue among “Western hunter-gatherers.”

A major aspect of this preprint is that it has to work hard to differentiate two Anatolian farmer-like signals: the first, from Anatolian farmers proper, and the second from the descendants of European farmers, who themselves are a mix of Anatolian farmers with a minority ancestry among the hunter-gatherers. The answers would probably be totally unintelligible if not for archaeology. It’s clear that the steppe people had contact with both European and Near Eastern farmers and that later East European groups that succeeded the Yamna were subject to reflux from Central Europe, and received European farmer ancestry.

Another curious nugget in their results is that there was early detection of both Ancestral North Eurasian (ANE) ancestry and, some East Eurasian gene flow (related to Han Chinese). One of their individuals carries the East Eurasian variant of EDAR, which today is only found in Finns, though it was found in reasonable frequencies among the Motala hunter-gatherers of Scandinavia. Additionally, Fu et al. 2016 found that the ancestors of Mesolithic hunter-gatherers received some gene flow from Eastern Eurasians as well (also in the supplements of Lazaridis et al. 2016).

The authors admit that there is probably population structure among ANE and undiscovered groups of East Eurasians who were traversing the Inner Asian landscape. I think this is all suggestive of some long-distance contacts, though the intensity and magnitude increased a lot with high-density societies and the mobility of pastoralism.

Much of the genetic mixing in the Near East, and to some extent in the trans-Caucasian region, seems to date to the 4th millennium. This is technically prehistory, but it is also the Uruk period. This was a phase of Mesopotamian culture expansion between 4000 and 3100 BC which resulted in replicas of Uruk style settlements as far away as Syria and southeastern Anatolia. There is even evidence of Uruk-related migration to the North Caucasus.

The Uruk experienced abrupt and sudden collapse. Uruk settlements outside of the core zone of Mesopatamia disappear.

It’s the final paragraph that warrants discussion:

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….

But instead of tackling this let’s focus on the paper that came out of the Willerslev group, The first horse herders and the impact of early Bronze Age steppe expansions into Asia. This is a final manuscript in Science. That means it was probably written before The Genomic Formation of South and Central Asia. When it comes to South Asia, the results from the two publications are consanant. There is no conflict.*

More interesting are the results in West Asia, and the linguistic supplement. In the authors note that tablets now indicate an Indo-Aryan presence in Syria ~1750 BC. Second, Assyrian merchants record Indo-European Hittite, or Nesili (the people of Nesa), as early as ~2500 BC.

As suggested in earlier work Hittite remains don’t suggest steppe influence. David W. says:

The apparent lack of steppe ancestry in five Hittite-era, perhaps Indo-European-speaking, Anatolians was interpreted in Damagaard et al. 2018 as a major discovery with profound implications for the origin of the Anatolian branch of Indo-European languages.

But I disagree with this assessment, simply because none of these Hittite-era individuals are from royal Hittite, or Nes, burials. Hence, there’s a very good chance that they were Hattians, who were not of Indo-European origin, even if they spoke the Indo-European Hittite language because it was imposed on them.

The main aspect I’d bring up with this is that in other areas steppe ancestry has spread deeply and widely into the population, including non-Indo-European ones. It is certainly possible that the sample is not needed enough to pick up the genuinely Hittite elite, but I probably lean to the likelihood that the steppe signal won’t be found. It seems that the Anatolian languages were already diversified by ~2000 BC, and perhaps earlier. Linguists have long suggested that they are the outgroup to other Indo-European languages, though this could just be a function of their isolation among highly settled and socially complex populations.

Two alternative models present themselves for these results. The Anatolian Indo-European languages expanded through elite diffusion,  part of the same general migrations that emerged out of the Yamna culture ~3000 BC. The lack of a steppe signal may be due to sampling bias, as David W. suggested, or, more likely in my opinion, simple dilution of the signal. Second, the steppe migrations were one part of a broader palette of population movements and cultural diffusions, and the Anatolian Indo-Europeans are basal to the efflorescence of the steppe derived branches.

The evidence of the explosion of Indo-Aryans in the years after 2000 BC in West and South Asia, as well as the expansion of Iranians across vast swaths of Inner Asia during the same period, suggest to me that Indo-Iranians are most definitely part of the steppe pulse. The connection to the Sintashta charioteers presents itself, and, connections to the Uralic languages indicates incubation in the trans-Volga region.

In West Asia, the Indo-Aryans crashed themselves against the most advanced civilizations of their time. Like the Bulgars, and unlike the Hittites, Indo-Aryan Mitanni was totally absorbed by their non-Indo-European Hurrian substrate. Indo-Aryan linguistic influence was preserved in their names, their gods, and in particular words relating to chariots. And yet in 2017’s Continuity and Admixture in the Last Five Millennia of Levantine History from Ancient Canaanite and Present-Day Lebanese Genome Sequences, the authors observe:

We next tested a model of the present-day Lebanese as a mixture of Sidon_BA and any other ancient Eurasian population using qpAdm. We found that the Lebanese can be best modeled as Sidon_BA 93% ± 1.6% and a Steppe Bronze Age population 7% ± 1.6% (Figure 3C; Table S6). To estimate the time when the Steppe ancestry penetrated the Levant, we used, as above, LD-based inference and set the Lebanese as admixed test population with Natufians, Levant_N, Sidon_BA, Steppe_EMBA, and Steppe_MLBA as reference populations. We found support (p = 0.00017) for a mixture between Sidon_BA and Steppe_EMBA which has occurred around 2,950 ± 790 ya (Figure S13B).

This needs to be more explored. The admixture could have come from many sources. I am curious about the frequency of R1a1a-z93 among modern-day Syrians and Lebanese.

For me these arguments can only be resolved with a deeper understanding of linguistic evolution. The close relationship of Indo-Aryan and Iranian languages is obvious to any speaker of either of these languages (I can speak some Bengali). A divergence in the range of 4 to 5 thousand years before the present seems most likely to me. But the relationship of the other Indo-European languages is much less clear.

One of the arguments in Peter Bellwood’s First Farmers is that the Indo-European languages exhibit a “rake-like” topology with the exception of Indo-Iranian, which forms a clear clade. To him and others in his camp, this argues for deep divergences very early in time.

It is hard to deny that the steppe migrations between 4 and 5 thousand years ago had something to do with the distribution of modern Indo-European languages. But, it is harder to falsify the model that there were earlier Indo-European migrations, perhaps out of the Near East, that preceded these. Only a deeper understanding of linguistic evolution, and multidisciplinary analysis of regional substrates will generate the clarity we need.

* I’m going to skip the Botai angle in this post.

Is American genetic diversity enough?


In the nearly 20 years since the draft of the human genome was complete,* we’ve moved on to bigger and better things. In particular, researchers are looking to diversify their panels of human genetic diversity, because of differences between groups matter. You can’t just substitute them for each other genetically.

There have been efforts to diversify the population panels recently, but that prompts the question whether American population coverage is sufficient. My first thought is that the genetic diversity in the USA is probably getting us 90% of the way there. Consider Spencer’s comment about Queens, it’s the most ethnically diverse large conurbation in the country.

There are some gaps though. In Who We Are David Reich points out the distinctiveness of Indian population genetics. The subcontinent has lots of large census populations which have drifted upward deleterious alleles due to long-term endogamy. And, many of these populations don’t have a strong representation in the Diaspora.

In contrast, much of the rest of the world is panmictic enough that an American panel can pick up most of the variation. American Chinese are skewed toward Guandong and Fujian, but a substantial number of people from other parts of China have arrived in the last generation. Regional structure is not so strong that you’ll miss out on too much, aside from very rare variants which are more extended pedigree scale rather than population scale.

There are small populations such as Hadza, Khoikhoi, and Pygmies in Africa which are probably going to be missed by American population panels, but the total census size of these groups is pretty low (for comparison, there are 1 million Pulayar Dalits in the state of Kerala alone). Much of the rest of Africa is West African variation well represented in African Americans, and Bantu and Nilotic variation probably captured my immigrant communities.

I’d propose supplementing American genetic diversity with sampling Cape Coloureds in South Africa.

* No discussions about how the genome isn’t totally complete. I know that.

Are Turks Armenians under the hood?

Benedict Anderson’s Imagined Communities: Reflections on the Origin and Spread of Nationalism is one of those books I haven’t read, but should. In contrast, I have read Azar Gat’s Nations, which is a book-length counterpoint to Imagined Communities. To take a stylized and extreme caricature, Imagined Communities posits nations to be recent social and historical constructions, while Nations sees them as primordial, and at least originally founded on on ties of kinships and blood.

The above doesn’t capture the subtlety of  Gat’s book, and I’m pretty sure it doesn’t capture that of Anderson’s either. But, those are the caricatures that people take away and project in public, especially Anderson’s (since Gat’s is not as famous).

When it comes to “imagined communities” I recently have been thinking how much that of modern Turks fits into the framework well. Though forms of pan-Turkic nationalism can be found as earlier as 9th-century Baghdad, the ideology truly emerges in force in the late 19th century, concomitantly with the development of a Turkish identity in Anatolia which is distinct from the Ottoman one.

The curious thing is that though Turkic and Turkish identity is fundamentally one of language and secondarily of religion (the vast majority of Turkic peoples are Muslim, and there are periods, such as the 17th century when the vast majority of Muslims lived in polities ruled by people of Turkic origin*), there are some attempts to engage in biologism. This despite the fact that the physical dissimilarity of Turks from Turkey and groups like the Kirghiz and Yakut is manifestly clear.

Several years ago this was made manifestly clear in the paper The Genetic Legacy of the Expansion of Turkic-Speaking Nomads across Eurasia. This paper clearly shows that Turkic peoples across Eurasia have been impacted by the local genetic substrate. In plainer language, the people of modern-day Turkey mostly resemble the people who lived in Turkey before the battle of Manzikert and the migration of Turkic nomads into the interior of the peninsula in the 11th century A.D. Of course, there is some genetic element which shows that there was a migration of an East Asian people into modern day Anatolia, but this component in the minority one.**

Sometimes the Turkish fascination with the biological comes out in strange ways, Turkish genealogy database fascinates, frightens Turks. Much of the discussion has to do with prejudice against Armenians and Jews. But the reality is that most Turks at some level do understand that they are descended from Greeks, Armenians, Georgians, etc.

To interrogate this further I decided to look at a data set of Greeks, Turks, Armenians, Georgians, and a few other groups, including Yakuts, who are the most northeastern of Turkic peoples. The SNP panel was >200,000, and I did some outlier pruning. Additionally, I didn’t have provenance on a lot of the Greeks, except some labeled as from Thessaly. I therefore just split those up with “1” being closest to the Thessaly sample and “3” the farthest.

First, let’s look at the PCA.

Read More

Mesolithic and Neolithic, Of Cheddar and Bread


It’s been a big week for “Cheddar Man” and the science around him. I already talked about the issue blog-wise for my day job. Additionally, Spencer and I did a podcast on the topic (if you haven’t, please subscribe and leave positive reviews and ratings on iTunes and Stitcher; next we’ll post our conversation with Chris Stringer, don’t miss it!).

So at this point I’ll put some other thoughts here that are “big picture.”

Cheddar Man may have been black but probably wasn’t

Much of the media is focused on the predicted pigmentation of Cheddar Man. That is, dark. Back when the La Brana Western Hunter-Gatherer results came in with the same finding, several population genomics people pointed out that it might not be valid to predict their phenotype based on modern training sets.

Here are some thoughts:

  • Cheddar Man and the WHG in general were probably darker than modern Northern Europeans. There is detectable selection in modern Europeans for pigmentation alleles down to the present, and Northern Europeans are the palest people in the world. And, pigmentation is polygenic, but it’s not hyperpolygenic. That’s why GWAS and early selection tests picked up pigmentation loci as hits so often.
  • Cheddar Man and the WHG in general were probably not as dark as tropical people. The only people who live(d) at very high latitudes who were very darkly complected were Tasmanian Aboriginals and Australian Aboriginals (Melbourne is at the same latitude south as Lisbon is north). In contrast, we see that Khoisan are brown, sometimes rather lightly so, while the peoples of non-European heritage who live in high latitudes are not dark-skinned, though they are not as light-skinned as Europeans.

We don’t have a time machine, so we won’t know with finality. But, it seems that pigmentation pathways are finite, and eventually we can probably be more confident if Cheddar Man had a genetic architecture that would lead to fewer and smaller melanocytes.

The First Farmers replaced WHG to a great extent in Britain

The preprint that came out with the Cheddar Man documentary really focused mostly on the Neolithic farmers. The data set was large, and it emphasized that the discontinuity between the farmers, who were EEF from Anatolian stock (modern Sardinians are their best proxies), the hunter-gatherers. WHG is genetically homogeneous, so they couldn’t reject the proposition that there was no admixture of British hunter-gatherers into the farmer population Basically, the thesis that Peter Bellwood outlined in First Farmers is well supported by these results. The farmers brought agricullture, and pushed aside or absorbed the hunter-gatherers.

It is notable to me that they found more hunter-gatherer ancestry (possibly) in eastern and northern populations, but not much in farmers from Wales. Additionally, though they couldn’t be definitive about it, the EEF settlers of Britain seem to have more affinities with the Western Mediterranean populations than the Central European ones. This suggests that perhaps the farmers arrived by sea or coast-hugging from the south and west, rather than from the south and east.

The arrival of farming to Britain was different

Farmers came to Britain later than to the continent. The shift from hunter-gatherer to farming was rapid. One model for why there was lack of admixture is that the farming cultural package was fully adapted to Northern Europe by the time they began settling the island. In contrast, on the mainland farmers were changing a Middle Eastern lifestyle into something that could take root in cold northern climes where there were already local residents.

Sometimes cultural and ecological changes drive rapid expansions of human populations

Today Europe, and much of Western Eurasia, is characterized by isolation by distance dynamics between populations. What you see in the transition from the Mesolithic to the Neolithic, and later with the arrival of metal age populations (Bell Beakers), is that populations can turnover fast, and that rapid expansion and growth can result in homogeneity across huge distances and then sharp continuities across cultural divides. The classical example of this is that hunter-gatherers and farmers in Central Europe did not exchange much in the way of genes for centuries, and their between population variance accounted for ~10% of their pooled variance (this is what you see comparing Han and Europeans). Additionally, WHG and EEF are both relatively homogeneous, at least before the latter began to absorb WHG at different fractions across its range. WHG descends from a late Pleistocene expansion, after the Last Glacial Maximum. Similarly, the EEF expanded rapidly from its Anatolian point of origin.

Britons didin’t become Britons genetically until the Bronze Age

Ten years ago many people thought that Cheddar Man and his people were the ancestors of most of the people who lived in Britain today. At the same time as this preprint came out, the Bell Beaker paper was officially published. We now know that Britain went through two massive demographic transitions in less than 2,000 years, with on the order of a 90% replacement in a few centuries both times.

Why? Was this typical? Those are for a later post….

The “Finns” are probably an Iron Age intrusion into the East Baltic

One of the first things I wrote at length about historical population genetics, in late 2002, happened to be a rumination on the Y chromosomal phylogeography of Finnic peoples. At the time there was debate as to the provenance of the N1c Y chromosomal haplotype (this is the haplotype of the Rurikids by the way). Just as R1b is ubiquitous in Western Europe, and R1a in Eastern Europe (and to some extent in Indo-Iranian lands), N1c has an extensive distribution in the northern zone of Eurasia.

The question at the time was whether N1c was from Europe and in particular the Finnic peoples, or, whether it was from Siberia.

NJ Tree of Pairwise Fst

Today we have many of the questions resolved. At this point, we know that the Finns, Sami, and Estonians, all exhibit evidence of gene flow from a Siberian-like population. This is clear on any genome-wide analyses. Though this is very much a minority component, even among the Sami, because it is genetically very different from the Northern European background, it is clear on any analysis.

Ancient DNA has also established the likelihood that this Siberian-like element is relatively new to the Baltic region. In a recent paper, The genetic prehistory of the Baltic Sea region:

We suggest that the Siberian and East Asian related ancestry in Estonia, and Y-haplogroup N in north-eastern Europe, where it is widespread today, arrived there after the Bronze Age, ca. 500 calBCE, as we detect neither in our Bronze Age samples from Lithuania and Latvia.

This is not the only ancient DNA paper that shows this. Of course, sampling is imperfect, and perhaps they’ve missed pockets of ancient Finnic peoples. But the most thorough analysis of Mesolithic hunter-gatherers in Scandinavian does not pick them up either, Population genomics of Mesolithic Scandinavia: Investigating early postglacial migration routes and high-latitude adaptation. Populations, such as the Comb Ceramic Culture, which have been identified as possible ancestors of the modern Finnic culture and ethnicity, lack the distinctive Siberian-like component.

At the SMBE 2017, I saw a poster which had results that were sampled from Finland proper, and distinctive ancestry of Siberian-like peoples was present in an individual who lived after 500 AD. This means that in all likelihood the circumpolar Siberian population which introduced this new element into the East Baltic arrived in the period between 500 BC and 500 AD.

Someone with more knowledge of paleoclimatology and archaeology needs to comment at this point. Something happened in this period, and it probably left a big ethno-linguistic impact. But I don’t know enough detail to say much (the Wikipedia entries are out of date or don’t illuminate).

I will add when I run Treemix Finns get the Siberian gene flow you’d expect. But the Lithuanians get something from the Finns. Since the Lithuanians have appreciable levels of N1c, that is not entirely surprising to me (the basal flow from the Yakut/European region to Belorussians may be more CHG/ANE).

Additionally, I will note that on a f-3 test Lithuanians have nearly as high a z-score (absolute) as Swedes (i.e., Finn; Swede/Lithuanian, Yakut), indicating that the predominant Northern European ancestry isn’t necessarily Scandinavian, as much as something between Lithuanian-like and Swedish-like (on Admixture tests the Finns do seem to have less EEF than Swedes, and Lithuanians probably the least of all among non-Finn peoples).

Addendum: I should note here that the genetics is getting clearer, but I have no great insight into the ethno-linguistic aspect. Perhaps the Siberian-like people did not introduce Finnic languages into the Baltic. Perhaps that was someone else. But I doubt it. That being said, though the Siberian-like component adds great distinctiveness to the Finns, it is important to add that by and large Finns are actually generic (if highly drifted) Northern Europeans.

 

Function & phylogeny, where the twain shall diverge


Every few years I get asked about Nuristanis and Kalash. The reason is that these people are often white. By white, I mean that some Nuristanis and Kalash are fair-skinned, blonde-haired, and blue-eyed. Entering “Nuristani” into Google images returns some very white faces. And you have weird news stories about ‘white’ Taliban, because non-locals don’t realize that some Nuristanis look like Northern Europeans.

Since the vast majority of people who look like white Northern Europeans are white Northern Europeans, many people assume that the Nuristanis and Kalash must have some kinship to white Northern Europeans. More precisely, many have spread the legend that these people have some relationship to the soldiers of Alexander the Great (even though Macedonians are Southern European…details).

As it turns out, they do have some kinship to Europeans…but not inordinately more than any of the other peoples of the region. The TreeMix plot at the top of this post shows that Greeks are far closer to Iranian Jews than they are to Kalash. In fact, the Kalash clearly have a non-trivial proportion of “Ancestral South Indian” South Asian ancestry.

Because of their high genetic drift (they’re endogamous and kind of inbred) a lot of population genetic analyses are a bit more difficult with the Kalash samples that are out there. But their genetic affinities are clear:

Table S4 show highly significant evidence (p value < 10−10) in the Kalash when using Armenia and Chamar as surrogates. Eight other pairings of surrogates give p values < 10−5. In all cases, the surrogate pairs include one group from South Asia (Chamar, Kol) and the other from West Eurasia (Armenia, Adygei, Brahui, Hungarians, Palestinians, Tuscans), consistent with admixture from a West Eurasian source.

Chamar are a Dalit caste of Northern India if you don’t know.

So what’s going on with the Kalash and Nuristanis? Appreciable frequencies of alleles which are correlated with traits like blue-eyes are found amongst them. Though the frequencies are much lower than in Northern Europeans. Very white looking Nuristanis and Kalash may be highly salient to photographers and the Western media, but it turns out most Nuristanis and Kalash look West Asian, with a minority who are dark-skinned enough that their South Asian ancestry is also quite clear.

This disjunction between appearance and ancestry should not surprise us. There has been a lot of recent change in physical appearance across populations over the last 10,000 years. Europeans themselves have changed in appearance. Similarly, other populations have as well. Some of them look similar to Europeans due to happenstance or convergence.

Another case are the Ainu of Japan. Though as an unadmixed group they no longer exist, old photos show some of them exhibiting an appearance not typical of East Asians. This led early anthropologists to posit that the Ainu were a “lost white race.” And yet to my knowledge, no European ancestry is found in Hokkaido, or in the Tohoku region, where Ainu-like people lived down to the early medieval period.

The moral of the story: don’t judge the contents of the book by its cover.

From the eastern edge of the sea of grass

A new preprint, Mitogenomic data indicate admixture components of Asian Hun and Srubnaya origin in the Hungarian Conquerors, throws a rather large sample of medieval mtDNA samples at the question of the ethnogenesis of the Magyar people. The context here is that Maygars speak a non-Indo-European language, with a distant relationship to the Finnic ones, but genetically are not much distinguished from their neighbors.

But the differences are not non-zero, ‘best of breed’ methods can pick up a small fraction of exotic ancestry.

The results above are based on samples from cemeteries of the Magyar elite. These are people who identified with a steppe confederacy which coalesced in the 9th century in the Volga region, and in 895 conquered the region of Central Europe which we now term Hungary. In the 10th century, the Magyar tribes were an alien and predatory force within Europe, pagan and exotic. They were, in fact, the latest in a long time of mobile Central Asian horsemen, going back to the Scythian, Sarmatians, Huns, and Avars.

What the data from these results confirm is that a substantial proportion of the maternal lineages of the Hungarian nobility were East Eurasian. Their genetic profiles, in fact, resembled Scythians who had mixed with Altaic peoples, even if they were not descended from those people. What this suggests is that the Magyar conquest elite was part of a broader landscape of ethnolinguistically distinct, but interconnected, steppe confederations. The preprint gives great space to the likelihood that a substantial proportion of the Magyar elite was in fact Turkic in origin.

This leads to one of the major assertions of the paper: the distinctive linguistic identity of modern Hungarians is a legacy of the Hunnic period, and not the Magyar conquest. From what I can tell they didn’t have older DNA, so I don’t know how they came to this conclusion, except that the conquest Magyars were small in number, and seem to have been decimated over time (Mongol invasion, the battle of Mohacs).