Who's your (proto) daddy Western Europeans?

Considering the increasingly large numbers of paleogenomic samples being released online nowadays, it's no longer practical for me to try to highlight most archaeological cultures and even genetic clusters in my Principal Component Analyses (PCA) of the ancient world. Thus, from now on, I'll be focusing attention in such PCA on the main population shifts that have led to the formation of the modern-day West Eurasian gene pool and genetic substructures, like on the PCA plot below, which includes the new Lipson et al. 2017 data (available at the Reich Lab here).

The relevant PCA datasheet can be gotten here. By grouping several hundred ancient samples into just nine clusters, I'm attempting to highlight four key processes and resulting genetic shifts in Europe, the Near East and Central Asia:

- European forger populations mixing with genetically much more southern early farmers of Near Eastern origin, mostly during the Neolithic, bringing about the total disintegration of the Europe to Siberia Hunter-Gatherer cline

- "Old Europeans" being overrun and largely absorbed by Y-haplogroup R1-rich Kurgan pastoralists from the Pontic-Caspian steppe during the Eneolithic and Bronze Age, leading to the formation of at least one major new cline from the Bronze Age steppe into post-Kurgan expansion Europe

- the ancient Near East "imploding" or becoming significantly more compact in terms of genetic structure, likely due to a variety of major population expansions from the chalcolithic onwards from the eastern and western parts of the Fertile Crescent, as well as probably the Caucasus and Europe (note how the post-Neolithic western Asian cluster stretches out towards Europe)

- fully nomadic and very wide ranging pastoral and warrior cultures dominating the entire Eurasian steppe during the Iron Age, leading to the emergence of progressively more East Asian-admixed populations from west to east across the Eurasian steppe

An interesting outcome of the denser sampling from space and time in West Eurasia is that Y-haplogroup R1b, once so elusive in the ancient DNA record, is now popping up all over the place. The new Lipson et al. dataset, for instance, includes two R1b "Old Europeans" from Blatterhole in Germany dated to the Middle Neolithic. Below is the same PCA as above except with all of the ancients belonging to R1b marked with an X. The two Blatterhole samples are sitting in the largely empty space between the European/Siberian Hunter-Gatherer cline and most of the "Old Europe" cluster. The relevant PCA datasheet is available here.

So it may seem that we're back to square one in the long running effort to pinpoint the origin of Y-haplogroup R1b-L51, which encompasses almost 100% of modern-day Western European R1b lineages, and thus probably ranks as Europe's most common Y-haplogroup. But at this stage I'd say no, because R1b-L51 is a subclade of R1b-M269, of which the oldest sample comes from the Bronze Age steppe. In fact, as can be seen in the above PCA, this sample is sitting in exactly the right spot to be one of those pastoralists who overran "Old Europe", or at least a very close relative thereof.

Or am I wrong? Feel free to let me know in the comments.

I didn't bother creating a similar plot of ancient samples belonging to Y-haplogroup R1a, because, unlike R1b, this marker is still non-existent in samples from outside of Eastern Europe and Siberia dating to before the late Neolithic. And I doubt that this is simply due to a lack of the right ancient material. Moreover, the recent discovery of Y-haplogroup R1a-M417, which encompasses almost 100% of all modern-day R1a lineages on the planet, in a North Pontic steppe sample belonging to the Eneolithic Sredny Stog culture means that it's game over for the naysayers as far as the steppe origin of most modern-day R1a lineages is concerned (see here and here).

In other words, if you're still hoping to see R1a, and especially R1a-M417, pop up in non-steppe derived ancient individuals in, say, such far away places as South Asia, then you'll probably be waiting forever.

For the linguistic implications of all of this, see...

Late PIE ground zero now obvious; location of PIE homeland still uncertain, but...

Update 15/11/2017: After a couple of days of messing around with the Lipson et al. dataset, I'm certain that Late Copper Age sample Protoboleraz_LCA I2788 shows significant steppe-related admixture. This is the only sample from Lipson et al. with such an obvious signal of steppe-related input that had enough data to be analyzed individually by me with PCA and D-stats.

For the time being, amongst the best proxies for this signal appear to be Yamnaya_Samara and Samara_Eneolithic. But it's likely that the real source of the admixture is yet to enter the ancient DNA record, or at least my dataset. When it does, it'll probably be an Eneolithic pastoralist population from the North Pontic steppe.

Yamnaya_Samara also gives the best statistical fit as the single source population in qpAdm (see here). It's an important result, because it suggests that steppe peoples very similar to Yamnaya were already expanding on and out of the steppe as far back as ~3500 BCE, and perhaps a few hundred years earlier.

See also...

Migration of the Bell Beakers—but not from Iberia (Olalde et al. 2018)

Genetic and linguistic structure across space and time in Northern Europe

I feel that I need to do a double take, and demonstrate more obviously why my new PCA, the one that I introduced in the recent Tollense Valley warrior blog post (see here), should prove very useful for analyzing both genetic and ethnolinguistic links in Northern Europe between modern-day populations and ancient samples, particularly those from late prehistory to early history, which is when the main ethnolinguistic groups that today dominate Northern Europe formed. Judging by some of the reactions in the comments, not everyone was convinced, so let's try this again.

Below is a new version of the said PCA that focuses on several ancient individuals who, based on their archaeological contexts, should show strong genetic affinities to modern-day speakers of Celtic, Germanic and Slavic languages in Northern Europe. These are three Iron Age samples from what is now England, one Iron Age sample from what is now Sweden, and two Medieval samples from what is now Bohemia, Czech Republic, respectively. The relevant datasheet is available here.

And clearly these ancients do show the expected genetic affinities considering where they cluster relative to modern-day Northern Europeans in the two most significant dimensions of genetic variation. Moreover, despite the fact that the Anglo-Saxon and English Iron Age samples were all excavated from sites in eastern England, the Anglo-Saxons cluster between the English Iron Age individuals and the singleton Scandinavian Iron Age sample. This of course makes perfect sense, considering that the Anglo-Saxons were Germanic speakers with recent ancestry from very near to Scandinavia.

So everything seems in good order, and for now it's very difficult for me to consider that those Tollense Valley warriors who cluster alongside modern-day Slavic speakers on my PCA are not ethnolinguistically closer to them than to Celtic and Germanic speakers.

On the other hand, my standard PCA of West Eurasian genetic variation does a comparatively lousy job at matching ethnolinguistic origins with genetic structure, at least in Northern Europe. Note below, for instance, that the same Celtic and Germanic samples from England and Scandinavia form a tight cluster between the two Slavs from Bohemia. Hence, based on this PCA it would be very difficult, perhaps impossible, to correctly predict the ethnolinguistic ties of these ancients just by looking where they cluster relative to modern-day Germanics, Slavs and so on. Right click and open in a new tab to enlarge to the max.

But this is not surprising, because this PCA is based on a wider, more diverse range of populations, and so rather than being dominated by relatively recent, ethnolinguistic-specific genetic drift within Northern Europe, it's much more reflective of deeper, more basic genetic relationships across West Eurasia.

See also...

Tollense Valley Bronze Age warriors were very close relatives of modern-day Slavs

Ancient genomes from NE Europe suggest in tandem spread of Siberian admixture and Uralic languages into the region >3,500 ya

Tollense Valley Bronze Age warriors were very close relatives of modern-day Slavs

This is strongly suggested by the Principal Component Analysis (PCA) below, which shows that many of the Tollense Valley warriors (Welzin_BA) cluster in the Slavic-specific part of the plot. The relevant datasheet is available here.

I designed this PCA with the sole purpose of using Balto-Slavic-specific genetic drift to differentiate Slavs from Germans, except of course those Germans with a lot of Slavic ancestry, who are usually from eastern Germany and Austria. Admittedly, these Welzin_BA samples are preliminary low quality versions, but I can assure you, people who don't harbor significant Slavic ancestry never cluster in this part of the plot.

The only other ancient samples that cluster in the Slavic zone are, as expected, an early Slav from Bohemia and, interestingly, a Bronze Age individual from what is now Hungary. But we've already seen strong genetic, and indeed genealogical, links between another Hungarian Bronze Age genome and present-day Slavs (see figure 3 here).

So what's going on? Did the proto-Slavs come into existence during the Bronze Age, as opposed to the more generally accepted early Medieval Period? And did they expand from what is now Hungary? Or did they migrate there from the Baltic region? Thanks to Matt in the comments for the table below.

See also...

Genetic and linguistic structure across space and time in Northern Europe

Hilariously wrong

From a recent paper at Forensic Science International:

The most commonly found haplogroups [among Lithuanians] are R1a and N, hence it can be argued that Lithuanians originate from Pakistan/Northwest India and East China/Taiwan.

Jankauskiene et al., Population data and forensic genetic evaluation with the YfilerTM Plus PCR Amplification kit in the Lithuanian population, Forensic Science International, DOI: http://dx.doi.org/10.1016/j.fsigss.2017.10.009

For a reality check see here...

R1a: The beast among Y-haplogroups

Finngolians #2

The mad scientists are at it again. The quote below is from an American Society of Human Genetics (ASHG) talk abstract. For the whole thing see here. Now, as I've pointed out on this blog before, Finns do not have Buryat or Mongolian ancestry, or anything even closely related dating to the Middle Ages. What they do have is some sort of Siberian admixture, which has been poorly characterized to date, but is probably associated with archaeologically attested population movements across northern Eurasia during the metal ages.

We identified significant gene flow from the Buryats to the Finnish which was predicted to be occurred in 1,228 (±87) year. Moreover, 13.38% of Buryat admixture was predicted in the Finnish genome.

This sort of nonsense should never be let through peer review anywhere. It makes the ASHG and indeed population genetics look like a total joke. In fact, imagine if such sloppy inferences from population genetics are allowed to influence medical genetics work. Someone might eventually get hurt.

See also...

Finngolians #1

R1a and R1b from an early Mongolian tomb

The beast among Y-haplogroups

A lot has been written about Y-haplogroup R1a over the years. Sadly, most of it was wrong, such as its posited Pleistocene origin in the Indian subcontinent and subsequent migration to Europe.

In all likelihood, R1a was born somewhere in North Eurasia. More importantly, its R1a-M417 subclade, which encompasses almost 100% of modern-day R1a lineages, no doubt came into existence somewhere on the Pontic-Caspian (or Western) steppe in what is now Ukraine and southern Russia just 7,000-6,000 years ago.

And within a couple of thousand years it expanded in almost all directions, probably on the back of the early Indo-European dispersals (see here), to cover a massive range from Scandinavia to South Asia. It is the beast among Y-haplogroups.

The most common subclade of R1a-M417 in South Asia today is R1a-Z93, and, realistically, it couldn't have arrived there earlier than about 2,000BC. So much for the Pleistocene.

See also...

Y-haplogroup R1a and mental health

The Poltavka outlier

Yamnaya isn't from Iran just like R1a isn't from India

Ancient genomes from Neolithic North Africa (Fregel et al. 2017 preprint)

Over at bioRxiv at this LINK. The paper includes three ancient North African Y-haplogroup results: two instances of E-M35 from the Early Neolithic (5300-4800 BCE) and a singleton T-M184 from the Late Neolithic (3780-3650 BCE). Emphasis is mine:

Abstract: One of the greatest transitions in the human story was the change from hunter-gatherer to farmer. How farming traditions expanded from their birthplace in the Fertile Crescent has always been a matter of contention. Two models were proposed, one involving the movement of people and the other based on the transmission of ideas. Over the last decade, paleogenomics has been instrumental in settling long-disputed archaeological questions, including those surrounding the Neolithic revolution. Compared to the extensive genetic work done on Europe and the Near East, the Neolithic transition in North Africa, including the Maghreb, remains largely uncharacterized. Archaeological evidence suggests this process may have happened through an in situ development from Epipaleolithic communities, or by demic diffusion from the Eastern Mediterranean shores or Iberia. In fact, Neolithic pottery in North Africa strongly resembles that of European cultures like Cardial and Andalusian Early Neolithic, the southern-most early farmer culture from Iberia. Here, we present the first analysis of individuals' genome sequences from early and late Neolithic sites in Morocco, as well as Andalusian Early Neolithic individuals. We show that Early Neolithic Moroccans are distinct from any other reported ancient individuals and possess an endemic element retained in present-day Maghrebi populations, indicating long-term genetic continuity in the region. Among ancient populations, early Neolithic Moroccans share affinities with Levantine Natufian hunter-gatherers (~9,000 BCE) and Pre-Pottery Neolithic farmers (~6,500 BCE). Late Neolithic (~3,000 BCE) Moroccan remains, in comparison, share an Iberian component of a prominent European-wide demic expansion, supporting theories of trans-Gibraltar gene flow. Finally, the Andalusian Early Neolithic samples share the same genetic composition as the Cardial Mediterranean Neolithic culture that reached Iberia ~5,500 BCE. The cultural and genetic similarities of the Iberian Neolithic cultures with that of North African Neolithic sites further reinforce the model of an Iberian intrusion into the Maghreb.

Fregel et al., Neolithization of North Africa involved the migration of people from both the Levant and Europe, bioRxiv, Posted September 21, 2017, doi: https://doi.org/10.1101/191569

Ancient IBD/cM matrix analysis offer

I've had a few requests from personal genomics customers to stick their files into an Identity-by-Descent/cM matrix like the one at the link below. Also please check out the accompanying comments thread for ideas of what can be done with the output.

A Bronze Age dominion from the Atlantic to the Altai

I can do this for USD $15 per individual. The deadline for sending through the data files (which, in this run, can only be from 23andMe, Ancestry or FTDNA) is this time Tuesday.

I'll send out the results to each participant over e-mail. However, participants are encouraged to post their results in the comments thread below so that they can be discussed and analyzed further.

Update 20/09/2017: The analysis is underway. Please don't send any more data files. If there's enough interest, I'll do another run soon.

Update 22/09/2017: I've just sent out the results to the participants in the form of two text files titled "ancients_only" and "full_column". The former is a matrix of overall shared haplotype tracts in centimorgans (cM) that includes the user and 65 ancient genomes, and the latter a list of haplotype tracts, also in cM, shared between the user and well over 3000 public samples.

So what can we do with these files? For one, we can look at them, because simply eyeballing these sorts of stats can be very informative. Sorting the data in some way and calculating population averages might help with that.

The "ancients_only" file can be used for slightly more advanced analyses. For instance, below is a Neighbor joining graph produced with the Past 3 program (freely available here). I simply loaded my "ancients_only" file into Past 3, selected all of the columns and rows, and then did this: Multivariate > Clustering > Neighbor joining. Note that I cluster on the same branch as Slav_Bohemia, and this makes perfect sense considering my Polish ancestry. By the way, I dropped Oetzi from this run because he was behaving strangely, which is not unusual for low coverage genomes. Click on the image and open in a new tab for a better view.

Indeed, Past 3 can do a lot of interesting things with matrix files; anything from linear models to rotating three dimensional plots. If you'd like to repeat the linear models from my above linked to blog post, then choose the relevant two columns in your matrix and go Model > Generalized Linear Model. You should see something like this.

Moreover, a matrix with the 3000+ public samples can be gotten here and combined, in part or in whole, with your other files so that you can analyze yourself alongside a larger number of individuals.

So far so good for the Kurgan hypothesis

This is basically what I'm seeing in the ancient DNA published to date. Thus, the Kurgan hypothesis or steppe theory, which, of course, posits that the Proto-Indo-European homeland was on the Pontic-Caspian steppe, is looking really good at this stage. Indeed, unless there are some ancient DNA shocks on the way from, say, Anatolia or the Caucasus, that might buck the trend, then this one's in the bag.

See also...

"The Homeland: In the footprints of the early Indo-Europeans" time map

Genetic borders are usually linguistic borders too

Late PIE ground zero now obvious; location of PIE homeland still uncertain, but...

A homeland, but not the homeland #2

The Out-of-India Theory (OIT) challenge: can we hear a viable argument for once?

A homeland, but not the homeland #2

Back in May, in a post titled A homeland but not the homeland, I said this:

It seems increasingly likely that ancient DNA has identified a massive expansion, or a series of expansions, from Mesopotamia and/or surrounds in basically all directions dating to the Chalcolithic (ChL) and Bronze Age (BA). This phenomenon is mainly characterized by the simultaneous spread of:

- Iran_ChL-related genome-wide ancestry

- Y-haplogroup J

- South Caspian-specific mitochondrial haplogroups such as R2 and U7

In the same post I also included a list of ancient populations that showed at least two of these characteristics. I can now add two more populations to this list: the Minoans and Mycenaeans.

- Anatolia_BA, Western Turkey, 2836-1800 calBCE (Lazaridis et al. 2017)

- Egyptian mummies, Middle Egypt, 776-2 calBCE (Schuenemann et al. 2017)

- Iran_ChL, Western Iran, 4839-3796 calBCE (Lazaridis et al. 2016)

- Levant_BA, Northwestern Jordan, 2489-1966 calBCE (Lazaridis et al. 2016)

- Minoans, Crete, Greece, 2900-1700 BCE (Lazaridis et al. 2017)

- Mycenaeans, Greece, 1700-1200 BCE (Lazaridis et al. 2017)

- Sidon_BA, Southern Lebanon, 1750-1600 BCE (Haber et al. 2017)

Out of all of these groups, only the Mycenaeans are generally accepted to have been speakers of an Indo-European language. However, they differ from the others in that they harbor minor but significant ancestry from a source, or multiple sources, closely related to Yamnaya, Sinatshta and other Bronze Age peoples of the Pontic-Caspian steppe.

Possible question for the discussion in the comments: what does this say about where the Mycenaeans got their Indo-European language? Also, who wants to bet that Bronze Age samples from the Indus Valley Civilization will too make it onto my list?

See also...

Late PIE ground zero now obvious; location of PIE homeland still uncertain, but...

A Bronze Age dominion from the Atlantic to the Altai

The BEAGLE analysis that I foreshadowed a couple of days ago (see here) is finally done. The output is available for download as a matrix of shared genomic tracts in centimorgans (cM) here.

I haven't yet had a chance to look at the results in detail, but I'd say that the outcomes for the three Early Bronze Age (EBA) Afanasievo and Yamnaya individuals make a lot of sense. The high affinity that these individuals show to the Irish EBA samples is not at all surprising, but striking nonetheless. The Afanasievo people, after all, lived in the Altai Mountains deep in Asia, more than 6,000kms from Ireland.

Update 02/08/2017: Interestingly, the graphs below, based on the cM values in my coancestry matrix, suggest that upper caste Indo-Aryan-speaking Brahmins from Northern India share relatively more ancestry with the Afanasievo genome than Iranic-speakers such as Pamir Tajiks, who generally share relatively more ancestry with the younger Andronovo and Sintashta samples. The relevant datasheet is available here.

See also...

Late PIE ground zero now obvious; location of PIE homeland still uncertain, but...

Ancient herders from the Pontic-Caspian steppe crashed into India: no ifs or buts

New resource: 67 diploid ancient genomes

Published this week along with Martiniano et al. 2017, a dataset of 67 new and publicly available genomes, genotyped and imputed for 30 million markers:

Data from: The population genomics of archaeological transition in west Iberia: investigation of ancient substructure using imputation and haplotype-based methods

Martiniano R, Cassidy LM, Ó'Maoldúin R, McLaughlin R, Silva NM, Manco L, Fidalgo D, Pereira T, Coelho MJ, Serra M, Burger J, Parreira R, Moran E, Valera AC, Porfirio E, Boaventura R, Silva AM, Bradley DG

Date Published: July 28, 2017

DOI: http://dx.doi.org/10.5061/dryad.g9f5r

Keep in mind however, that this dataset is specifically designed for haplotype-based tests, like those done with Chromopainter (for more details, see S5 Text in Martiniano et al. 2017). As far as I know, it should also perform well in ADMIXTURE runs.

On the other hand, the diploid and imputed genotype calls are likely to slightly skew results in formal statistics and formal statistics-based modeling analyses. So it's best to use pseudo-haploid genomes for such tests, and/or high coverage diploid genomes if available, with 100% observed calls.

I'm about to run a quick and dirty haplotype/Principal Component analysis with this dataset using BEAGLE, mainly to check whether South Asians show greater recent genetic affinity to Afanasievo/Yamnaya over Andronovo/Sintashta (for more on this controversy, see here). It's a pity that this dataset doesn't include any genomes from Neolithic Iran, because then I'd also be able to try haplotype-based mixture models for South Asians.

By the way, I won't be using all of the 30 million markers. I've only kept the SNPs that overlap with the Harvard Medical School's 1240K SNP ancient capture array, which should mean that only a small minority of the calls in my analysis won't be real.

Update 02/08/2017: The BEAGLE run is complete and the analysis is unfolding. See post and comments here.

Yamnaya-related migrations into Iberia: infiltration rather than invasion (Martiniano et al. 2017)

The Martiniano et al. preprint that appeared at bioRxiv more than two months ago was published at PLoS Genetics today (see here). The paper packs a lot of supplementary information that wasn't included with the preprint. Below is the press release about the paper from the Public Library of Science (PLoS). Emphasis is mine.

The genomes of individuals who lived on the Iberian Peninsula in the Bronze Age had minor genetic input from Steppe invaders, suggesting that these migrations played a smaller role in the genetic makeup and culture of Iberian people, compared to other parts of Europe. Daniel Bradley and Rui Martiniano of Trinity College Dublin, in Ireland, and Ana Maria Silva of University of Coimbra, Portugal, report these findings July 27, 2017 in PLOS Genetics.

Between the Middle Neolithic (4200-3500 BC) and the Middle Bronze Age (1740-1430 BC), Central and Northern Europe received a massive influx of people from the Steppe regions of Eastern Europe and Asia. Archaeological digs in Iberia have uncovered changes in culture and funeral rituals during this time, but no one had looked at the genetic impact of these migrations in this part of Europe. Researchers sequenced the genomes of 14 individuals who lived in Portugal during the Neolithic and Bronze Ages and compared them to other ancient and modern genomes.

In contrast with other parts of Europe, they detected only subtle genetic changes between the Portuguese Neolithic and Bronze Age samples resulting from small-scale migration. However, these changes are more pronounced on the paternal lineage. "It was surprising to observe such a striking Y chromosome discontinuity between the Neolithic and the Bronze Age, such as would be consistent with a predominantly male-mediated genetic influx" says first author Rui Martiniano. Researchers also estimated height from the samples, based on relevant DNA sequences, and found that genetic input from Neolithic migrants decreased the height of Europeans, which subsequently increased steadily through later generations.

The study finds that migration into the Iberian Peninsula occurred on a much smaller scale compared to the Steppe invasions in Northern, Central and Northwestern Europe, which likely has implications for the spread of language, culture and technology. These findings may provide an explanation for why Iberia harbors a pre-Indo-European language, called Euskera, spoken in the Basque region along the border of Spain and France. It has been suggested that Indo-European spread with migrations through Europe from the Steppe heartland; a model that fits these results.

Daniel Bradley says "Unlike further north, a mix of earlier tongues and Indo-European languages persist until the dawn of Iberian history, a pattern that resonates with the real but limited influx of migrants around the Bronze Age."

Martiniano R, Cassidy LM, Ó'Maoldúin R, McLaughlin R, Silva NM, Manco L, et al. (2017) The population genomics of archaeological transition in west Iberia: Investigation of ancient substructure using imputation and haplotype-based methods. PLoS Genet 13(7): e1006852. https://doi.org/10.1371/journal.pgen.1006852

See also...

New resource: 67 diploid ancient genomes

Late PIE ground zero now obvious; location of PIE homeland still uncertain, but...

The Out-of-India Theory (OIT) challenge: can we hear a viable argument for once?

Recent weeks have seen a rash of activity from OIT proponents defending their "truth", largely as a response to a news feature in The Hindu on new genetic evidence backing the Aryan Invasion or Migration Theory (AIT/AMT). A few examples:

Genetics Might Be Settling The Aryan Migration Debate, But Not How Left-Liberals Believe

Genetics and the Aryan invasion debate

Propagandizing the Aryan Invasion Debate: A Rebuttal to Tony Joseph

Here We Go Again: Why They Are Wrong About The Aryan Migration Debate This Time As Well

The problematics of genetics and the Aryan issue

Too early to settle the Aryan migration debate?

The people who wrote these articles are able to string sentences together in a reasonable way, but apart from that, their efforts are clumsy at best. Not only do they not appear to completely understand what they're attempting to debunk, but they also fail to offer an OIT that realistically incorporates new findings from ancient and modern-day DNA.

AIT/AMT is now firmly backed by ancient DNA from Eastern Europe and high resolution modern-day DNA from South Asia. To quote myself from a week ago:

During the past couple of years ancient DNA has revealed the presence of Y-chromosome haplogroup R1a in Eastern European remains dated to the Mesolithic, Neolithic, Eneolithic and Bronze Age. Moreover, the Bronze Age remains, packed in ancestry derived from Eastern European hunter-gatherers (or EHG) and totally lacking any sort of South Asian admixture, belong to R1a-Z645, which is the ancestral clade of by far the most common varieties of R1a in Europe and South Asia today: R1a-Z282 and R1a-Z93, respectively. And on top of that, South Asians, especially those speaking Indo-European languages, show significant admixture derived from EHG.

The conclusion from this data is self-evident: during the Bronze Age R1a-Z645 became a very important Y-chromosome lineage in Europe and quickly moved to South Asia, in all likelihood on the back of the Indo-European expansion.

Pre-Indo-European Eastern Europe and South Asia were not the same world; they were world's apart. Thus, you will never read anything like this, no matter how much ancient DNA from South Asia is sequenced:

During the past couple of years ancient DNA has revealed the presence of Y-chromosome haplogroup R1a in South Asian remains dated to the Mesolithic, Neolithic, Eneolithic and Bronze Age. Moreover, the Bronze Age remains, packed in ancestry derived from South Asian hunter-gatherers, and totally lacking any sort of European admixture, belong to R1a-Z645, which is the ancestral clade of by far the most common varieties of R1a in Europe and South Asia today: R1a-Z282 and R1a-Z93, respectively. And on top of that, Europeans, especially those speaking Indo-European languages, show significant admixture derived from South Asian hunter-gatherers.

So, OIT proponents, what counter-arguments can you offer? And can you come up with a new vision for OIT that coherently takes into account ancient DNA from Eastern Europe?

However, to ensure that the debate is a fruitful one not derailed regularly by anti-AIT/pro-OIT red herrings, let's take care of the most obvious of these red herrings now. I reserve the right to delete any comments that attempt to go down these tired, irrelevant avenues without a very good excuse for doing so.

You: So and so found Y-haplogroup P* and other basal clades upstream of R1a in Papuans, therefore R1a and Indo-Europeans are from South Asia. Me: Nonsense. R1 and R1a are found in the remains of Eastern European Mesolithic foragers. Were these individuals recently arrived Indo-European-speakers from South Asia? Try harder.

You: It doesn't matter that Eastern European Mesolithic foragers belonged to R1a, because the most common form of R1a in the world is R1a-M417, and if it originated in India then OIT is a reality. Me: But what are the chances realistically that R1a-M417 is from India or South Asia, considering that prehistoric European samples, with absolutely no signals of ancestry from South Asia, belong to both M417+ and M417- lineages? In fact, Europe is the most likely homeland of R1a-M417.

You: India has incredible diversity in R1a, therefore it's the R1a and Indo-European homeland. Me: No it doesn't. India, and indeed, South Asia as a whole are dominated by one fairly young subclade: R1a-Z93. Europe is home to three different subclades that show up at perceptible frequencies: R1a-Z282, found throughout much of the continent; R1a-L664, mostly confined to Northwestern Europe; and R1a-Z93, mostly confined to far Eastern Europe.

You: Many unique Indian ethnic groups are yet to be tested genetically. They may show surprising results, including new subclades of R1a. Me: If you dig hard enough, you'll always find some exceptions to the rule. But how do you know where the ancestral lineages of such exceptions in South Asia were during, say, the Neolithic? What makes you think they were in South Asia? To prove that South Asia is the homeland of its by far most dominant R1a subclade, R1a-Z93, then at the very least you need to show that other, closely and distantly related subclades, are also found at perceptible frequencies in whole regions of South Asia, and therefore that they have some sort of history there. Otherwise we can safely assume that R1a-Z93 and the few exceptions to the R1a-Z93 rule in South Asia are relative latecomers from somewhere else.

You: But we have no ancient DNA from South Asia yet, and it may produce a huge shock. Me: For you yes, but not for me. What are the chances realistically that R1a was present among both European and South Asian foragers? I'd say practically zero. Feel free to raise it to a few per cent to make yourself feel better, but we both know the hard reality.

You: Ancient DNA from South Asia might show that Northern India was home to a population very similar to Yamnaya, and if so, then the Yamnaya-related ancestry in modern-day Indians is native to India. Me: There's no logic behind this. Yamnaya and other closely related Bronze Age groups were very specific mixtures of Mesolithic foragers and Neolithic farmers living in Eastern Europe and surrounds. There's absolutely no reason to assume that such unique mixtures would also form independently in South Asia, or even outside of Europe's generally accepted borders.

You: Bronze Age Europeans who belonged to R1a also carried southern admixture from Iran, or maybe even India. Me: In prehistoric samples, R1a is always highly correlated with Eastern European Hunter-Gatherer (EHG) ancestry, so positing that it also arrived in Europe with a southern population makes no sense. And why would this southern ancestry be from Iran or India? Why not the Caucasus? We know from ancient DNA that the type of southern ancestry that these ancient Europeans carried has been sitting in the Caucasus since the Upper Paleolithic. Moreover, they lack South Caspian- and South Asian-specific markers such as mtDNA haplogroup U7. How were such markers purged from their gene pool if they or their recent ancestors arrived in Europe from Iran or India?

You: Chickens and mice came from South Asia, therefore Indo-Europeans came from South Asia too. Me: Bullshit. Do better or go away.

Does anyone want to claim that I don't know what I'm talking about? Or perhaps that I'm just putting out Eurocentric propaganda? If you don't understand my arguments, and that they're indeed very solid arguments, then there's no hope for you. Go and find a new hobby or profession, because you're not cut out for this.

OK, now what we have the formalities out of the way, who wants to have a go at salvaging OIT in the comments? Don't be shy.

See also...

Ancient herders from the Pontic-Caspian steppe crashed into India: no ifs or buts

R1a-M417 from Eneolithic Ukraine!!!11

Late PIE ground zero now obvious; location of PIE homeland still uncertain, but...

On the Mesolithic colonization of Scandinavia (Günther et al. 2017 preprint)

Over at bioRxiv at this LINK. The main takeaway point from this preprint is that Scandinavia was a more happening place than most of the rest of Europe during the Mesolithic, because at the time it was the meeting place between two relatively divergent forager groups, West European hunter-gatherers (WHG) and East European hunter-gatherers (EHG), that entered the peninsula from different directions, the southwest and northeast, respectively, and mixed to form Scandinavian hunter-gatherers (SHG). Other key points:

- EHG probably dispersed across Scandinavia in a counter-clockwise direction via an ice-free route along the Atlantic coast in what is now Norway, because SHG samples from northern and western Scandinavia show more EHG ancestry than those from southern and eastern Scandinavia

- at least 17% of the SNPs that are common in SHG are not found in present-day Europeans, suggesting that a large part of European variation has been lost since the Mesolithic

- although it's unlikely that SHG made a significant contribution to the present-day Northern European gene pool, some gene-variants common in SHG that appear to be associated with metabolic, cardiovascular, developmental and psychological traits are carried at high frequencies by present-day Northern Europeans, especially compared to present-day Southern Europeans, probably due to strong selective pressures specific to northern latitudes in Europe

- SHG is inferred to have had fair skin and varied blue to light-brown eye color, which makes sense considering that it was a mixture of apparently fair-skinned/brown-eyed EHG and dark-skinned/blue-eyed WHG, except that the frequencies of blue-eyed variants and one fair-skinned variant in SHG are much higher than expected from its EHG/WHG mixture ratios, again pointing to strong selective pressures specific to northern latitudes in Europe acting upon certain gene-variants

- a 3D computer generated facial reconstruction of an SHG female based on data from a very high (57x) coverage genome sequence looks, at least to me, like a fairly typical present-day Northern European woman (see Figure S9.1 in the supp info here), though I suspect that the result might be biased in some way, simply because it's impossible to know whether variants associated with specific facial traits in present-day Northern Europeans were also associated with the same facial traits in SHG.

Citation...

Günther et al., Genomics of Mesolithic Scandinavia reveal colonization routes and high-latitude adaptation, bioRxiv, Posted July 17, 2017, doi: https://doi.org/10.1101/164400

North European admixture in the Han Chinese (Charleston et al. 2017 preprint)

Over at bioRxiv at this LINK. Emphasis is mine. The estimated date of the North European-related admixture signal is probably much too late. These sorts of estimates always look way off. And I doubt that it's largely the result of the Silk Road, which linked China to the Near East and Mediterranean rather than to Northern Europe. More likely it reflects gene flow from the Pontic-Caspian steppe in Eastern Europe during the Bronze and Iron ages, via the Afanasievo, Andronovo, and other closely related steppe peoples (see here).

Abstract: As are most non-European populations around the globe, the Han Chinese are relatively understudied in population and medical genetics studies. From low-coverage whole-genome sequencing of 11,670 Han Chinese women we present a catalog of 25,057,223 variants, including 548,401 novel variants that are seen at least 10 times in our dataset. Individuals from our study come from 19 out of 22 provinces across China, allowing us to study population structure, genetic ancestry, and local adaptation in Han Chinese. We identify previously unrecognized population structure along the East-West axis of China and report unique signals of admixture across geographical space, such as European influences among the Northwestern provinces of China. Finally, we identified a number of highly differentiated loci, indicative of local adaptation in the Han Chinese. In particular, we detected extreme differentiation among the Han Chinese at MTHFR, ADH7, and FADS loci, suggesting that these loci may not be specifically selected in Tibetan and Inuit populations as previously suggested. On the other hand, we find that Neandertal ancestry does not vary significantly across the provinces, consistent with admixture prior to the dispersal of modern Han Chinese. Furthermore, contrary to a previous report, Neandertal ancestry does not explain a significant amount of heritability in depression. Our findings provide the largest genetic data set so far made available for Han Chinese and provide insights into the history and population structure of the world's largest ethnic group.

...

One finding from our analysis of admixture signals that most likely fit a one-pulse admixture model is our observation of admixture from Northern European populations to the Northwestern provinces of China (Gansu, Shaanxi, Shanxi), but not other parts of China. Previous analysis of the HGDP data, based on patterns of haplotype sharing among 10 Han Chinese from Northern China, estimated a single pulse of ~6% West Eurasian ancestry among the Northern Han Chinese. The estimated date of admixture was around 1200 CE. This signal is also observed among the Tu people, an ethnic minority also from Northwestern China; the authors attributed this signal to contact through the Silk Road (Hellenthal et al. 2014). We estimate a lower bound of admixture proportion due to Northern Europeans at approximately 2%-5%, with an admixture date of about 26 +/-3 generations for Gansu, and 47 +/-3 generations for Shaanxi [Table S8]. Using a generation time of about 26-30 years (Moorjani et al. 2016), these estimates correspond to admixture events occurring at around 700 CE and 1300 CE, respectively, corresponding roughly to the Tang and Yuan dynasty in China. However, these estimated dates should be interpreted with caution, as both the violation of a single pulse admixture model and the additional noise in inter-­marker LD estimates due to low coverage data could bias the estimates.

Charleston et al.,A comprehensive map of genetic variation in the world's largest ethnic group - Han Chinese, bioRxiv, Posted July 13, 2017, doi: https://doi.org/10.1101/162982

See also...

Late PIE ground zero now obvious; location of PIE homeland still uncertain, but...

Indian confirmation bias

In a largely fact free but obfuscation rich comment piece at The Hindu, Indian scientists Gyaneshwer Chaubey and Kumarasamy Thangaraj ask whether it's too early to settle the Aryan migration debate. See here.

No, it's not too early. It's game over chaps, and has been for a while.

During the past couple of years ancient DNA has revealed the presence of Y-chromosome haplogroup R1a in Eastern European remains dated to the Mesolithic, Neolithic, Eneolithic and Bronze Age. Moreover, the Bronze Age remains, packed in ancestry derived from Eastern European hunter-gatherers (or EHG) and totally lacking any sort of South Asian admixture, belong to R1a-Z645, which is the ancestral clade of by far the most common varieties of R1a in Europe and South Asia today: R1a-Z282 and R1a-Z93, respectively. And on top of that, South Asians, especially those speaking Indo-European languages, show significant admixture derived from EHG.

The conclusion from this data is self-evident: during the Bronze Age R1a-Z645 became a very important Y-chromosome lineage in Europe and quickly moved to South Asia, in all likelihood on the back of the Indo-European expansion. Yet, in spite of this, Gyaneshwer and Kumarasamy make the following claim in their article.

Moreover, there is evidence which is consistent with the early presence of several R1a branches in India (our unpublished data).

Potentially powerful stuff, you might say. But hang on, what are Gyaneshwer and Kumarasamy seeing in their data that could possibly reverse the current reality about R1a? Did they find R1a in South Asian remains from the Mesolithic and Neolithic? Or perhaps they've uncovered South Asian Bronze Age remains that belong to R1a-Z645 and lack any signals of ancestry from Eastern Europe?

This is impossible. The ancient DNA from Eastern Europe says so. That's because pre-Indo-European Eastern Europe and South Asia were not the same world; they were world's apart. Thus, you will never read anything like this, no matter how much ancient DNA from South Asia is sequenced:

During the past couple of years ancient DNA has revealed the presence of Y-chromosome haplogroup R1a in South Asian remains dated to the Mesolithic, Neolithic, Eneolithic and Bronze Age. Moreover, the Bronze Age remains, packed in ancestry derived from South Asian hunter-gatherers, and totally lacking any sort of European admixture, belong to R1a-Z645, which is the ancestral clade of by far the most common types of R1a in Europe and South Asia today: R1a-Z282 and R1a-Z93, respectively. And on top of that, Europeans, especially those speaking Indo-European languages, show significant admixture derived from South Asian hunter-gatherers.

See also...

The resistance crumbles

Ancient herders from the Pontic-Caspian steppe crashed into India: no ifs or buts

The Out-of-India Theory (OIT) challenge: can we hear a viable argument for once?

Working topology for Eurasian population structure

Here's my new "basic" qpGraph topology that I'll be using to test phylogenetic and mixture models for Eurasians. I think it reconciles a few key findings from recent scientific literature. Please note that since my main interest is post-Neolithic prehistory of West Eurasia, and in particular the early Indo-European expansions, I don't want to make this model unnecessarily complex by adding "dead end" Upper Paleolithic genomes.

But I welcome ideas on how to improve and make use of this topology, so if, say, adding Ust_Ishim helps, then let's do it. The ancient samples featured in the above graph are listed here and the graph file is available here. Feel free to post your own versions of the graph file in the comments and I'll run them as soon as possible. But please remember to label the samples correctly at all times.

Update 13/07/2017: Thanks to Matt in the comments, here's a neater version of the same model, with a lower (highest) Z score and slightly different mixture coefficients. It includes a couple of zero edges, which are generally undesirable, but these might disappear when more populations are added to the topology. The graph file is available here.

Armenian confirmation bias

Current Biology recently published a paper by Margaryan and Derenko et al. titled Eight Millennia of Matrilineal Genetic Continuity in the South Caucasus. I wasn't going to bother calling out the authors on their, unfortunately I have to say, rather dubious claim, but then I saw this ScienceNordic article enthusiastically attempting to drive home their misguided point, so a few words are now in order.

“It’s basically the same female population in the region over the past 8,000 years. It’s very surprising considering the many waves of migration and cultural shifts,” says lead-author Ashot Margaryan from the Centre for GeoGenetics at the National History Museum of Denmark, University of Copenhagen.

Genetics have remained constant for 8,000 years in world’s melting pot

I'm at a loss as to why Ashot Margaryan is very surprised. I'm not even mildly surprised. Why? Let's take a closer look at what we're dealing with here:

- the authors sequenced just 52 full mitogenomes to represent 8,000 years of prehistory and early history in the South Caucasus

- they lumped all of these sequences together into an "Ancient" sample set as if they were from a single time slice (I know, pretty crazy)

- they then ran a few complex models on this neither here nor there sample set, and concluded that it resembled the maternal gene pool of present-day Armenians.

Well, duh, present-day Armenians are more or less the end product of the population history of the last eight thousand years in what is now Armenia and surrounds. Is anyone still as surprised about this as Ashot? Surely not.

Obviously, the problem here is that the authors have mistaken their none too surprising outcome to mean that the South Caucasus has not experienced any major upheavals in its maternal gene pool over the past 8,000 years, which, if actually true, would indeed be very surprising, and even shocking.

But the haplogroup assignments of the 52 mitogenomes are reported in the spreadsheet here, and just by eyeballing these results, I can tell that they suggest an influx of foreign ancestry, probably from the Pontic-Caspian steppe, to the South Caucasus after the Early Bronze Age (EBA). Note, for instance, the presence of what appear to be typically steppe haplogroups U4a, U2e1e and U5a1b in the samples dated to the Middle Bronze Age (MBA), Late Bronze Age (LBA) and Early Iron Age (EIA), respectively.

Citation...

Margaryan and Derenko et al., Eight Millennia of Matrilineal Genetic Continuity in the South Caucasus, Current Biology 27, 1–6 July 10, 2017, DOI: 10.1016/j.cub.2017.05.087

Out-of-India chickens coming home to roost

Razib has posted a spacious but none-too-technical review of the ongoing Aryan Invasion Theory (AIT) controversy, along with some personal anecdotes and predictions about how ancient DNA from South Asia might shape the debate in the near future (see here).

It should be a useful guide to the topic for those of you who aren't quite as excited about reading about my latest adventures with qpGraph as many of the regulars in the comments here.

One thing that I'd perhaps add to Razib's post is that the ancient DNA record now boasts Late Neolithic Yamnaya-like Corded Ware Culture individuals from the East Baltic region that belong to Y-haplogroup R1a-Z645. And that's usually as far as their lineages go (see here).

This is important, because the Z645 mutation is directly and recently ancestral to the pair of likely post-Neolithic mutations that define the two R1a subclades most common in Europe and South Asia today: Z282 and Z93, respectively.

So not only are the "European" R1a-Z282 and "South Asian" R1a-Z93 relatively young sister clades, but their ancestral clade has now been found in ancient samples from Northeastern Europe that probably predate their appearance by only a few generations, if that. Of course, the upshot of all of this is that R1a-Z93 could not have originated very far from the East Baltic, which makes South Asia look about as likely as the homeland of this subclade as the goddamn moon. Conversely, it makes AIT look very plausible indeed.

However, granted, this might seem very confusing to anyone who hasn't been studying the R1a topology for years, and perhaps better left out of the more mainstream debates on AIT for the sake of simplicity. By the way, I found this part of Razib's post especially intriguing:

One scientist who holds to the position that most South Asian ancestry dates to the Pleistocene argued to me that we don’t know if ancient Indian samples from the northwest won’t share even more ancestry than the Iranian Neolithic and Pontic steppe samples. In other words, ANI was part of some genetic continuum that extended to the west and north. This is possible, but I do not find it plausible.

I suspect that this scientist's rather fanciful suggestion (which really flies in the face of very solid models based on ancient genomic data from Europe and surrounds) is a hint of the direction that the debate will take right after the publication of ancient genomes from South Asia. Because when that happens, obfuscators like this guy (usually hopeless Out-of-India proponents) will either have to concede defeat and quit the debate, or ramp up their obfuscations to spectacular new highs.

And please don't mistake my confidence on this issue for bluff and bluster. It's not exactly the best kept secret out there that ancient samples from India and Pakistan are now ready, and...oops I probably can't say more than that for now. Pity.

See also...

Ancient herders from the Pontic-Caspian steppe crashed into India: no ifs or buts

Indian confirmation bias

Iron Age nomads vs Bronze Age herders: Sarmatians and Yamnaya in qpGraph

If we are to take these qpGraph models fairly literally, and I don't see why not, since they're very tight fits overall, then the early Sarmatians from what is now Pokrovka, Russia, derived almost 80% of their ancestry from Yamnaya or a very closely related group, while the rest of their ancestry came from a source that was a ~50/50 mixture between Han-like East Asians and a population closely related to Neolithic and Chalcolithic farmers from what is now Iran.

This topology also tests for the same Iran Neolithic/Chalcolithic-related input in Yamnaya, and I think it's very important to note that the relevant admixture edges (D7 to D9) are 0%, which suggests that Yamnaya did not harbor this type of ancestry. I didn't bother testing for East Asian-related admixture in Yamnaya in the same way, because it never shows such signals in other analyses.

The clearly more complex ancestry of the Sarmatians is probably best explained by the fact that they belonged to a true nomadic warrior culture, and indeed one that managed to spread its influence across vast stretches of Eurasia. So these two Sarmatian individuals, both from Unterlander et al. 2017, may have had recent ancestors from as far afield as Central Asia and Siberia. On the other hand, Yamnaya was a semi-nomadic pastoralist population, and although also highly mobile and prone to long-distance expansions, probably not as mobile as the Sarmatians.

Update 30/06/2017: Interestingly, adding Siberian Upper Paleolithic genome MA1 to the topology in the main model slightly shifts the admixture coefficients for Yamnaya, resulting in an arguably more accurate outcome in which it's modeled as a 50/50 mixture of populations closely related to Eastern European and Caucasus Mesolithic foragers.

See also...

Ancient herders from the Pontic-Caspian steppe crashed into India: no ifs or buts

Matters of geography

The steppe north of the Black Sea in Ukraine has basically always been considered a part of Europe, and just over 100 years ago some guy with a map decided that the steppe between the eastern coast of the Black Sea in Russia and the Ural River in western Kazakhstan should also be Europe.

So nowadays, right or wrong, it's generally accepted that the entire steppe region west of the Ural River, known as the Pontic-Caspian steppe, is in Eastern Europe. Here's a map courtesy of Wikipedia showing how the official boundary between Eastern Europe and Asia has shifted since the 18th century.

But this decision wasn't entirely arbitrary, because the current boundary between Eastern Europe and Asia by and large follows several major geographic barriers, including the Caucasus Mountains, the Caspian Sea and the Ural Mountains. It'd be hard to argue that these barriers haven't had a profound impact across the ages on the character of Europe and its people, and this has probably been known for well over a couple hundred years.

For instance, if we're to trust the most common interpretations of the works of ancient geographers like Hecataeus and Herodotus, then their worlds in some important ways resembled the typical Principal Component Analysis (PCA) of West Eurasian genetic variation. And it seems that they had a pretty good idea where both the strong continental boundaries and fuzzy areas were located.

Below, on the geographic map inspired by Herodotus, Europa or Europe is delineated from much of Asia by the Black Sea, the Caucasus Mountains and the Caspian Sea, while on the genetic map, most European and Asian populations form two, more or less parallel, clusters fairly cleanly separated by empty space (this was first noted in Lazaridis et al. 2013). Indeed, this empty space is the work of the Black Sea, the Caucasus Mountains and the Caspian Sea acting as rather effective barriers to gene flow between Eastern Europe and Asia (see Yunusbayev et al. 2012).

However, on the genetic map, the Iranic Scythians of the Asian steppes straddle my somewhat arbitrary red line separating Europa and Asia, and this is echoed on the Herodotus map by Iranic and related peoples like the Massagetae and Issedones, who inhabit the seemingly undefined part of the world between Europa and Asia east of the Caspian Sea (Mare Caspium).

Nothing really ground breaking, but pretty cool stuff.

On a related note, I've seen the term "mainland Europe" used recently in at least one of the big ancient DNA papers to describe the part of Europe west of the Pontic-Caspian steppe. It seems that the authors wanted to underline the fairly stark genetic difference that existed between most of Europe and the steppe just prior to the expansion of Yamnaya and related steppe herder groups that initiated the formation of the present-day European gene pool.

I can see why they did this, but to my mind they got things backwards. That's because the term mainland implies the opposite of island and/or peninsula, and of course the part of Europe west of the Pontic-Caspian steppe is a relatively narrow strip of land surrounded by water, so it's a peninsula. Let's visualize these two models on a map of Europe courtesy of Wikipedia:

I understand that my model might result in heart palpitations for some readers, especially those from Western Europe, who generally see their part of Europe as core Europe, but I feel that it makes good sense from a purely geographic POV.

See also...

Max Planck scientists: on a mission against geography

Genetic borders are usually linguistic borders too

Ancient herders from the Pontic-Caspian steppe crashed into India: no ifs or buts

It's now more than obvious that South Asia experienced an almighty pulse of admixture from an Early Bronze Age (EBA) population originally from somewhere on the Pontic-Caspian steppe in Eastern Europe. This is fairly easy to demonstrate thanks to ancient DNA from Europe and West Asia. One way of doing it is with the qpGraph algorithm.

Moreover, the widespread presence of Y-chromosome haplogroup R1a in South Asia is, at least in large part, linked to this event, because:

- Mesolithic Eastern European foragers belonging to basal clades of R1a do not show any South Asian or even Near Eastern ancestry, so it's likely that R1a is native to Eastern Europe and surrounds

- If R1a is native to Eastern Europe then it can't also be native to South Asia, which is not only thousands of miles away, but also ecologically a different world

- The most common R1a subclades in the world today, R1a-M417 and one of its main daughter branches R1a-Z93, appear in Late Neolithic and Bronze Age European pastoralist groups (Corded Ware, Srubnaya and closely related peoples) that harbor high levels of Eastern European forager ancestry and no signs of South Asian admixture

- Practically 100% of the R1a in South Asia today belongs to the R1a-Z93 subclade, which, based on full Y-chromosome sequencing data, looks like it began expanding rapidly only during the EBA, eventually making its way to South Asia, and this is in line with the available ancient DNA evidence

- In South Asia, R1a and ancient steppe admixture peak in groups that speak Indo-European, including Indo-Aryan, languages, suggesting that both are genetic signals of the Indo-European expansions into the Indian subcontinent

So we're now at a stage where anyone with at least moderate thinking capacity, whose mind isn't poisoned by extreme bias, has to agree that there was a rather large movement of people from the Eurasian steppes into South Asia during the Bronze Age. No ifs or buts.

Ancient DNA from South Asia is on the way. It might throw up a few surprises and force a new model of how the Indo-Europeans and R1a got to South Asia, but it won't turn things upside down. In other words, don't expect the Out-of-India or "indigenous Aryans" theory to suddenly come into the picture as a viable alternative to the Aryan Invasion Theory (AIT), occasionally presented as the more politically correct Aryan Migration Theory (AMT).

Many Indians still don't get this, or rather they refuse to get it, which is very frustrating, especially if you're a regular in the comments section here. But admittedly it can also be very entertaining.

Last week The Hindu published an interesting piece on the latest developments in South Asian population genetics that were making the AIT, or at least AMT, look like a sure thing:

How genetics is settling the Aryan migration debate

Soon after came this peculiarly titled retort in the Swarajya online magazine, in which unfortunately it's impossible to find a single coherent argument:

Genetics Might Be Settling The Aryan Migration Debate, But Not How Left-Liberals Believe

Generally hilarious stuff, except the parts where the author abuses blogger Razib Khan for moving with the latest genetic data and arguing in favor of the Aryan expansion into India (see here and here).

So what are we to expect when the first big paper with ancient DNA from South Asia comes out, probably in the next few months? For starters, accusations of racism and maybe even hate speech against anyone who claims that the results support AIT or AMT, or anything even close. And lots of shouting and carrying on. But also a lot more comic relief.

See also...

Descendants of ancient European (fair?) maidens in Central Asia's highlands

The Out-of-India Theory (OIT) challenge: can we hear a viable argument for once?

Late PIE ground zero now obvious; location of PIE homeland still uncertain, but...

qpGraph open thread

I managed to put together a simple qpGraph model for the Kalash using present-day populations. It's largely based on the model for the Paniya by Nakatsuka et al. (see Supplementary Figure 5. here). The graph and pops files for my model can be downloaded here and here, respectively. I'm now working on a more complex model for the Kalash that includes ancient genomes from Eastern Europe and West Asia.

I'm willing take a few requests for qpGraph models in the comments below. Please note, however, that these requests will have to be accompanied by graph and pops files, and the graph files must be correctly set out; if they don't work, then they don't work, and you won't get your graph. On the other hand, you only need to supply pops files with the correct populations and I'll do the rest.

See also...

Ancient herders from the Pontic-Caspian steppe crashed into India: no ifs or buts

A homeland, but not the homeland

It seems increasingly likely that ancient DNA has identified a massive expansion, or a series of expansions, from Mesopotamia and/or surrounds in basically all directions dating to the Chalcolithic (ChL) and Bronze Age (BA). This phenomenon is mainly characterized by the simultaneous spread of:

- Iran_ChL-related genome-wide ancestry

- Y-haplogroup J

- South Caspian-specific mitochondrial haplogroups such as R2 and U7

At least two of these characteristics are shared by five groups that have appeared in the Near Eastern and African ancient DNA record as probable post-Neolithic newcomers, at least in part, at their respective sampling sites:

- Anatolia_BA, Western Turkey, 2836-1800 calBCE (Lazaridis et al. 2017)

- Egyptian mummies, Middle Egypt, 776-2 calBCE (Schuenemann et al. 2017)

- Iran_ChL, Western Iran, 4839-3796 calBCE (Lazaridis et al. 2016)

- Levant_BA, Northwestern Jordan, 2489-1966 calBCE (Lazaridis et al. 2016)

- Sidon_BA, Southern Lebanon, 1750-1600 BCE (Haber et al. 2017)

I'm confident that many more such groups will soon be added to the ancient DNA record, probably including Levant_ChL from the upcoming Harney et al. 2017 (a teaser of the paper can be seen here). Below, a map of Mesopotamia courtesy of Wikipedia.

It's an interesting and important question who these likely Mesopotamian migrants and their descendants were in terms of linguistic affinities. It seems that they left a massive genetic imprint on the Near East and much of North Africa, and perhaps also Central Asia and Southeastern Europe, so they probably also left some sort of linguistic legacy.

Obviously, it's highly improbable that most of them were Indo-European speakers. So if most of them weren't Indo-Europeans, then the phenomenon I'm describing here can't be related to the Proto-Indo-European (PIE) expansion. Forget the idea of an West Asian linguistic hot spot spewing out different, distantly related language families, including Indo-European, via the migrations of closely related Iran_ChL-like populations over a span of a few thousand years; it's plain stupid.

So who were they?

See also...

A homeland, but not the homeland #2

Late PIE ground zero now obvious; location of PIE homeland still uncertain, but...

Steppe admixture in Mycenaeans, lots of Caucasus admixture already in Minoans (Lazaridis et al. 2017)