A final note for the year

I feel like I've spent a good part of 2019 banging my head against a thicker than average brick wall.

Much of this feeling is tied to the controversy over the ethnogenesis of the Yamnaya people, and my often futile attempts to explain that their origin cannot be sought in what is now Iran, or, indeed, anywhere outside of Eastern Europe.

This post is my final attempt to lay out the facts in regards to this topic. Next year I'll have better things to do than to argue the bleeding obvious.

Below are two graphs from a Principal Component Analysis (PCA) based on relatively high quality ancient human genotype data from the Caucasus and surrounds. They include two typical Yamnaya individuals from burial sites north of the Caspian Sea. I made the graphs with the Vahaduo Custom PCA tool here. The relevant datasheet can be downloaded here.

Here's what I'm seeing:

- the Yamnaya individuals sit on genetic clines made up of hunter-gatherers native to the Caucasus and various parts of Eastern Europe, including a trio from the southernmost part of the Pontic-Caspian steppe (labeled Steppe_Eneolithic), with whom they form a distinct cluster

- the samples from the Caucasus and the Iranian Plateau form very different clusters, so there's no support here for the ancient Caucasus/Iranian grouping that is often haphazardly invoked in scientific literature

- there's no indication that the Yamnaya and/or Steppe_Eneolithic groups experienced recent gene flow, or, for that matter, any gene flow whatsoever, from what is now Iran.

Of course, analyses based on formal statistics suggest that the Yamnaya population harbors minor western ancestry that is missing in Steppe_Eneolithic. In fact, I was first to argue this point (see here). So let's add a couple of ancient farmers from Western Europe to my PCA to see how they affect the graphs. The relevant datasheet is available here.

Yep, the Yamnaya pair appears to be peeling away very slightly, but deliberately, from the Steppe_Eneolithic individuals towards the part of the plot occupied by the farmers.

Admittedly, I'm no Sherlock Holmes, but even with my fairly average sleuthing abilities, I'm pretty sure I know how the Yamnaya people came to be. They formed largely on the base of a population very much like Steppe_Eneolithic somewhere deep in Eastern Europe, well to the north of the Caucasus, and nowhere near the Iranian Plateau.

See also...

A note on Steppe Maykop

A note on Steppe Maykop

I'm reading a new book titled Dispersals and Diversification: Linguistic and Archaeological Perspectives on the Early Stages of Indo-European (see here). One of the chapters is authored by archeologist David Anthony, in which he makes the following claims:

A previously unknown genetic population actually was identified in Wang et al. (2019), but it was a peculiar relict-seeming group related to Paleo Siberians and American Indians (Kennewick) that had survived isolated somewhere in the Caspian steppes or perhaps in the North Caucasus Mountains. The Maykop people did admix with this previously isolated Siberian/Kennewick population in graves labeled "Steppe Maykop" in Wang et al. (2019).

But this just makes it clearer that a cultural choice motivated the Maykop people to exclude marriages with Yamnaya and pre-Yamnaya people specifically, even while exchanges of material goods, ideas, technologies continued. Neither the Maykop nor the North Caucasus/Siberian/Kennewick population can be the source of most of the CHG [Caucasus hunter-gatherer] ancestry in Yamnaya. In order to narrow down when and where CHG ancestry entered the steppes, we must widen our geographic frame beyond the Caucasus.

Unfortunately, this is way off the mark. Especially unsound is his inference that the CHG-related ancestry in the Yamnaya population may have come from beyond the Caucasus.

In fact, the chances that the Steppe Maykop people were derived from a relict Siberian/Kennewick-related group that survived into the Maykop era in the Caspian steppes or the North Caucasus are exactly zero.

The real story was surely more complicated. In my opinion, it initially involved the migration during the Eneolithic or earlier of a people rich in CHG ancestry from the southernmost steppes into the Volga Delta and surrounds, and then the back-migration during the Early Bronze Age (EBA) of their descendants with around 50% admixture from Central Asian foragers. If so, these foragers were very similar to indigenous West Siberians and also relatively closely related to Native Americans.

I don't know why such an exotic people migrated into the North Caucasus steppes to form the bulk of the Steppe Maykop population, but I'm certain they did, and one interesting possibility is that they were recruited by Maykop chiefs to create a buffer zone against hostile Yamnaya-related groups trying to push into the Caucasus, possibly from the lower Don region.

Of course, the same ancient northward migration of the CHG-rich population that may have eventually given rise to the Steppe Maykop people might also explain the deep origins of the Yamnaya people.

The key sample in all of this is VJ1001 from the Wang et al. paper. This female comes from an Eneolithic (4332-4238 calBCE) kurgan burial in the North Caucasus steppes. But despite her early date, she's genetically very similar to most Yamnaya individuals. And she's also a perfect proxy for half of the ancestry of three out of the six Steppe Maykop individuals. Here's a mixture model that I put together using the Broad MIT/Harvard software qpAdm:

RUS_Steppe_Maykop (3/6)
RUS_Eneolithic_steppe_VJ1001 0.452±0.023
RUS_Tyumen_HG 0.548±0.023
chisq 7.494
tail prob 0.874914
Full output

Indeed, these Steppe Maykop samples don't harbor any Maykop ancestry. They're simply a two-way mixture between a population closely resembling VJ1001 and another one similar to hunter-gatherers from Tyumen, West Siberia.

Importantly, a couple of Steppe Maykop-related populations were inadvertently discovered by Narasimhan et al. northeast of the Caspian Sea in what is now Kazakhstan. One of these groups is labeled Kumsay_EBA, after the location of its cemetery. It's roughly contemporaneous with Steppe Maykop and basically identical to the aforementioned Steppe Maykop trio.

KAZ_Kumsay_EBA
RUS_Eneolithic_steppe_VJ1001 0.440±0.022
RUS_Tyumen_HG 0.560±0.022
chisq 10.573
tail prob 0.646513
Full output

I suppose it's possible that Kumsay_EBA represents the migration of Steppe Maykop people into the Kazakh steppes. But even if this is true, then there had to have been an earlier migration of a group from the Kazakh steppes or West Siberia that mixed with the VJ1001-related natives of the North Caucasus steppes to give rise to Steppe Maykop.

I'm assuming that the Yamnaya-like VJ1001 and her people were the indigenous population of the North Caucasus steppes because there are no indications that they or their ancestors migrated there within any reasonable time frame from anywhere else, and certainly not from as far afield as, say, what is now Iran.

The other three Steppe Maykop individuals, who are genetic outliers in varying degrees from the main Steppe Makyop cluster, show variable levels of Maykop ancestry, with an average of about 50%. But they too harbor significant VJ1001-related ancestry. So despite the fact that there was some irregular mixing between the Maykop and Steppe Maykop peoples, this is not what created the typical Steppe Maykop genetic profile.

RUS_Steppe_Maykop_o
RUS_Eneolithic_steppe_VJ1001 0.234±0.074
RUS_Maykop_Novosvobodnaya 0.461±0.046
RUS_Tyumen_HG 0.305±0.033
chisq 7.378
tail prob 0.831667
Full output

And, of course, it should be obvious by now that the ancestry of the vast majority of Yamnaya individuals is better modeled without any input whatsoever from the Maykop or Steppe Maykop samples.

In fact, early indications are that the Yamnaya people flooded into Steppe Maykop territory from the north and completely replaced its population (see here). Despite this, in Dispersals and Diversification archeologist Kristian Kristiansen makes the following claim: "steppe Maykop expanded north, leading to the formation of the Yamnaya Culture and Proto-Indo-European". Not a chance in hell Professor.

See also...

A final note for the year

The PIE homeland controversy: August 2019 status report

Some myths die hard

An exceptional burial indeed, but not that of an Indo-European

Avalon vs Valhalla revisited

Pictured below is a new version of my Celtic vs Germanic genetic map. It's based on the same Principal Component Analysis (PCA) as the original (which can be seen here), but more focused on Northwestern Europe and produced with a different program.

To see the interactive online version, navigate to Vahaduo Custom PCA and copy paste the text from here into the empty space under the PCA DATA tab. Then press the PLOT PCA button under the PCA PLOT tab. For more guidance, refer to the screen caps here and here.

To include a wider range of populations in the key, just edit the data accordingly. For instance, to break up the ancient grouping into more specific populations, delete the Ancient: prefix in all of the relevant rows. This is what you should see:

Conversely, you can leave the ancient sample set intact and instead reorder the present-day linguistic groupings into, say, geographic groupings. To achieve this just delete all of the linguistic prefixes, such as Celtic:, Germanic:, and so on. You should end up with a datasheet like this and plot like this.

Of course, you can design your own plot by using any combination of the ancient and present-day individuals and populations that I've already run in this PCA. Their coordinates are listed here. Indeed, if you're in the possession of your own Celtic vs Germanic PCA coordinates, you can add yourself to the plot. And if you're not, see here.

It's also possible to re-process PCA data via the SOURCE tab. But I don't recommend doing this with the Celtic vs Germanic data, which are derived from a fine scale analysis and don't pack much variation. On the other hand, Global25 data are ideal for such re-processing. I made the plots below from subsets of Global25 coordinates available in a zip file here. To see how, refer to the screen caps here and here.

See also...

Modeling your ancestry has never been easier

Getting the most out of the Global25

Modeling genetic ancestry with Davidski: step by step

Big deal of 2019: ancient DNA confirms the link between Y-haplogroup N and Uralic expansions

The academic consensus is that Indo-European languages first spread into the Baltic region from the Eastern European steppes along with the Corded Ware culture (CWC) and its people during the Late Neolithic, well before the expansion of Uralic speakers into Fennoscandia and surrounds, probably from somewhere around the Ural Mountains.

On the other hand, the views that the Uralic language family is native to Northern Europe and/or closely associated with the CWC are fringe theories usually espoused by people not familiar with the topic or, unfortunately it has to be said, mentally unstable trolls.

The likely close relationship between the CWC expansion and the early spread of Indo-European languages was discussed in several papers in recent years (for instance, see here). This year, we saw the first ancient DNA paper focusing on the transition from the Bronze Age to the Iron Age in the East Baltic, including the likely first arrival of Uralic speech in what is now Estonia.

Published in Current Biology courtesy of Saag et al., the paper showed that the genetic structure of present-day East Baltic populations largely formed in the Iron Age (see here). It was during this time, the authors revealed, that the region experienced a sudden influx of Y-chromosome haplogroup N, which is today common in many Uralic speaking populations and often referred to as a Proto-Uralic marker. Little wonder then that Saag et al. linked this genetic shift in the East Baltic to the westward migrations of early Uralic speakers.

The table below, based on data from the Saag et al. paper, surely doesn't leave much to the imagination about what happened.

Unfortunately, I have to say that the genome-wide analysis in the paper was less informative than it could have been. The authors focused their attention on rather broad genetic components, and, as a result, missed an interesting fine scale distinction between their Bronze Age and Iron Age samples. The spatial maps below, based on my Global25 data for most of the ancients from Saag et al., show what I mean. The hotter the color the higher the genetic similarity between them and present-day West Eurasian populations.

Note that the Bronze Age (Baltic_EST_BA) samples are most similar to the Baltic-speaking, and thus also Indo-European-speaking, Latvians and Lithuanians, rather than the Uralic-speaking Estonians, even though they're from burial sites in Estonia. On the other hand, the Iron Age (Baltic_EST_IA) samples show strong similarity to a wider range of populations, including Estonians and many other Uralic-speaking groups.

See also...

It was always going to be this way

Fresh off the sledge

More on the association between Uralic expansions and Y-haplogroup N

Viking Age Iceland

I finally managed to get some of the Icelandic ancients from Ebenesersdóttir et al. 2018 into the Global25 datasheets (see here). Better late than never. Look for the"ISL_Viking_Age" prefix. Below is a screen cap of a Principal Component Analysis (PCA) with the new samples. It was done with an online Global25 PCA runner freely available here.

The individuals classified as unadmixed Gaels and Norse by Ebenesersdóttir et al. generally also look like it based on their Global25 coordinates.

The mixture models below, using all of the populations from the Global25 "modern pop averages scaled" datasheet, were run with an online tool freely available here. Note that the ADD DIST COL option is set to 1X. This is a useful feature for modeling the fine scale ancestry of samples that are derived from very similar populations.

See also...

They came, they saw, and they mixed

Commoner or elite?

Who were the people of the Nordic Bronze Age?

Etruscans, Latins, Romans and others

I've just added coordinates for more than 100 ancient genomes from the recently published Antonio et al. ancient Rome paper to the Global25 datasheets. Look for the population and individual codes listed here. Same links as always:

Global25 datasheet ancient scaled

Global25 pop averages ancient scaled

Global25 datasheet ancient

Global25 pop averages ancient

Thus far I've only managed to check a handful of the coordinates, so please let me know if you spot any issues. Below is a Principal Component Analysis (PCA) featuring the Etruscan and Italic speakers. I ran the PCA with an online tool specifically designed for Global25 coordinates freely available here.

Can we say anything useful about the origins of the Etruscan and early Italic populations thanks to these new genomes? Also, to reiterate my question from the last blog post, what are the genetic differences exactly between the Etruscans, early Latins, Romans and present-day Italians? Feel free to let me know in the comments below.

Update 13/11/2019: Here's another, similar PCA. This one, however, is based on genotype data, and it also highlights many more of the samples from the Antonio et al. paper. Considering these results, I'm tempted to say that the present-day Italian gene pool largely formed in the Iron Age, and that it was only augmented by population movements during later periods. The relevant datasheet is available here.

Update 13/11/2019: It seems to me that the two Latini-associated outliers show significant ancestry from the Levant, which possibly means that they're in part of Phoenician origin. These qpAdm models speak for themselves:

ITA_Ardea_Latini_IA_o
ITA_Proto-Villanovan 0.547±0.081
Levant_ISR_Ashkelon_IA2 0.453±0.081
chisq 7.573
tail prob 0.87027
Full output

ITA_Prenestini_tribe_IA_o
ITA_Proto-Villanovan 0.679±0.068
Levant_ISR_Ashkelon_IA2 0.321±0.068
chisq 7.222
tail prob 0.89033
Full output

The Proto-Villanovan singleton is also a key part of the models. Dating to the Bronze Age/Iron Age transition, she appears to be of western Balkan origin. Moreover, her steppe ancestry is probably derived directly from the Yamnaya horizon.

ITA_Proto-Villanovan
HRV_Vucedol 0.677±0.031
Yamnaya_RUS_Samara 0.323±0.031
chisq 10.397
tail prob 0.661174
Full output

The cluster made up of four early Italic speakers can be modeled with minor Proto-Villanovan-related ancestry, but, perhaps crucially, it doesn't need to be. Indeed, judging by the qpAdm output below, it's possible that almost all of its steppe ancestry came from the Bell Beaker complex, and, thus, the Corded Ware culture complex before that.

ITA_Italic_IA
Bell_Beaker_Mittelelbe-Saale 0.480±0.055
ITA_Grotta_Continenza_CA 0.411±0.042
ITA_Proto-Villanovan 0.109±0.084
chisq 10.294
tail prob 0.590205
Full output

Two out of the three available Etruscans look very similar to the Italic speakers in the above PCA plots, and yet they show a lot more Proto-Villanovan-related ancestry in my qpAdm run. The statistical fit is also relatively poor, perhaps suggesting that something important is missing.

ITA_Etruscan
Bell_Beaker_Mittelelbe-Saale 0.186±0.081
ITA_Grotta_Continenza_CA 0.283±0.064
ITA_Proto-Villanovan 0.531±0.126
chisq 17.175
tail prob 0.143143
Full output

Interestingly, the Etruscan outlier with significant North African admixture (proxied in my run by MAR_LN) doesn't need to be modeled with any Bell Beaker ancestry.

ITA_Etruscan_o
ITA_Proto-Villanovan 0.675±0.057
MAR_LN 0.325±0.057
chisq 14.864
tail prob 0.315912
Full output

Update 17/11/2019: The spatial maps below show how three groups of ancient Romans (from the Imperial, Late Antiquity and Medieval periods) compare to present-day West Eurasian populations in terms of their Global25 coordinates. The hotter the color, the higher the similarity. More here.

See also...

Getting the most out of the Global25

Modeling your ancestry has never been easier

An exceedingly simple, yet feature-packed, online tool ideal for modeling ancestry with Global25 coordinates is freely available HERE. It works offline too, after downloading the web page onto your computer. Just copy paste the coordinates of your choice under the "source" and "target" tabs, and then mess around with the buttons to see what happens. The screen caps below show me doing just that.

Another free, easy to use online tool that works with Global25 coordinates is the Principal Component Analysis (PCA) runner HERE. Below is a screen cap of me checking out one of the many PCA that it offers.

See also...

Getting the most out of the Global25

The Balkan connection

The hot topic at the moment is social inequality in Bronze Age Europe, thanks to a new paper by Mittnik et al. at Science. The full article is sitting behind an exceedingly robust paywall here.

However, the genotype dataset from the paper is freely available at the Max Planck Society's Edmond data repository here. Below is my Principal Component Analysis (PCA) of ancient West Eurasian genetic variation featuring 41 of the highest quality ancients from the new dataset. Almost all of them are from the Lech Valley in the Bavarian Alps, covering the period from the Bell Beaker culture (BBC) to the Middle Bronze Age (MBA). Two of the samples are from a mass Corded Ware culture (CWC) burial in the more northerly Tauber Valley.

I've also highlighted other ancients on the plot associated with the BBC and CWC from present-day Netherlands and Germany, respectively. The relevant PCA datasheet can be downloaded here.

Social stratification in ancient Europe is a fascinating topic, and it's an issue that I've started looking at myself (see here). However, I can't see any correlation between the inferred social standing of the individuals from the Lech and Tauber valleys and their positions in my PCA.

Nevertheless, the PCA is interesting in that it highlights considerable genetic heterogeneity within the Lech Valley BBC population. Indeed, how is this heterogeneity even possible, if, as per Mittnik et al., ancient DNA "has shown that the spread of the BBC throughout continental Europe did not involve large-scale migrations"?

Below is another version of my PCA, but this time focusing on three males: Lech Valley Beakers UNTA58_68Sk1 and WEHR_1192SkA, as well as ALT_4 from the aforementioned mass CWC grave in the Tauber Valley. Note that UNTA58_68Sk1 and WEHR_1192SkA represent genetically the most southern and northern, respectively, Lech Valley BBC samples that had enough data to be run in my analysis. I chose to focus on males because they carry the Y-chromosome, which can be informative about male-mediated ancient population expansions.

The PCA outcomes for these individuals are generally in line with their results in other types of genetic analyses, including those based on formal statistics. For instance, compared to the other two, ALT_4 harbors excess early steppe herder ancestry, UNTA58_68Sk1 excess early European farmer ancestry, and WEHR_1192SkA excess European hunter-gatherer ancestry. Moreover...

- UNTA58_68Sk1 shows a non-local isotopic signature and belongs to Y-haplogroup G2a, a marker essentially missing from BBC populations north of the Alps, and is best modeled as a two-way mixture between Bronze Age populations from the Balkans and the Pontic-Caspian steppe (see here), which probably means that he was a migrant to the Lech Valley from south of the Alps

- importantly, UNTA58_68Sk1 is not an isolated case, at least in the sense that several other BBC individuals from Bavaria, Bohemia, Hungary and Poland show varying ratios of Balkan-related ancestry, although almost all of these people are women

- WEHR_1192SkA is very similar to Bell Beakers from the northern Netherlands with whom he shares the R1b-P312 Y-haplogroup, suggesting that he was part of a population that moved into the Lech Valley from potentially as far away as the North Sea coast

- although ALT_4 probably shares the R1b-L51 Y-haplogroup with WEHR_1192SkA and many other BBC and Bronze Age individuals from the Bavarian Alps and surrounds, this can't be used as evidence of significant local genetic continuity after the CWC period, especially considering the comparatively eastern genome-wide structure of ALT_4.

Of course, archeological data suggest that the BBC was influenced in some important ways by the Copper and Bronze Age cultures of the Balkans and Carpathian Basin. So much so, in fact, that Marija Gimbutas, author of The Civilization of the Goddess, believed that the BBC originated in the Balkans from a synthesis of the local Vucedol culture and the intrusive Yamnaya culture from the Pontic-Caspian steppe.

Considering the ancient DNA evidence, however, the main demographic center of the early BBC could not have been south of the Alps.

Rather, it appears that early BBC and even CWC groups from north of the Alps moved into the Balkans and Carpathian Basin, where they may have established contacts with the local elites. If so, this might explain the significant southern cultural influences on the BBC, but limited accompanying genetic impact. This scenario also has support from archeological data (for instance, see here).

See also...

Is Yamnaya overrated?

The Boscombe Bowmen

Single Grave > Bell Beakers

Y-haplogroup R1a and mental health

I've updated my map of pre-Corded Ware culture R1a samples with a couple of new entries from Central and South Asia (the original is still here). However, before any of you get overly excited, please note that these samples aren't older than the Corded Ware culture. The reason I added them to my map is to counter the ongoing absurd claims online that South Asian R1a isn't derived from European R1a.

Just in case the map can't be viewed in all of its glory in some devices, here's what the fine print says:

The oldest example of R1a in ancient DNA from Central Asia is dated to 2132-1940 calBCE (ID I3770, Narasimhan 2019). Moreover, this sequence is closely related to much older R1a samples from Central, Eastern and Northern Europe, and phylogenetically nested within their diversity. Thus, it must surely represent a population expansion from Europe to Central Asia. Indeed, it's also associated with the Bronze Age Andronovo archeological culture, which is usually seen as an offshoot of the Corded Ware culture (CWC) of Late Neolithic Europe. The vast majority of present-day R1a lineages in Central Asia are closely related to that of I3770, and so must also ultimately derive from Europe.

The oldest instance of R1a in ancient DNA from South Asia is dated to just 1044-922 calBCE (ID I12457, Narasimhan 2019). This sequence, as well as the vast majority of present-day South Asian R1a lineages, are closely related to much older R1a samples from Central, Eastern and Northern Europe, and phylogenetically nested within their diversity. Thus, they must surely represent a population expansion from Europe to South Asia via Central Asia, in all likelihood during the Bronze Age. Even if R1a existed in South Asia before the Bronze Age, which is extremely unlikely, because it's found in samples from indigenous European hunter-gatherers, the vast majority of present-day R1a lineages in South Asia must be ultimately from Europe.

The idea that most, if not all, South Asian R1a is derived from European R1a seriously scares a lot of people. This is obvious in many online discussions on the topic. I suspect they're so frightened by it because, in their minds, it has the potential to encourage discrimination and even racism, perhaps by re-defining the colonization of much of the world by European nations in the recent past as the natural order of things?

In any case, clearly we're dealing with some sort of mass phobia here. I've got advice for those of you suffering from this problem: if you're honestly worried that the geographic provenance and expansion history of some Y-haplogroup is going to negatively impact on your life in any meaningful way, then it's time to find yourself a quality mental health professional. All the best with that.

See also...

The mystery of the Sintashta people

The Poltavka outlier

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

Commoner or elite?

I recently started looking at the correlations between Y-chromosome haplogroups and social standing in ancient Europe, and was surprised by what I learned about the five currently sampled prehistoric Scandinavians belonging to Y-haplogroup R1b. I certainly wasn't expecting to uncover these stories about a mass human sacrifice, a bog body, and an Arctic circle warrior:

- The earliest Scandinavian in the ancient DNA record belonging to R1b comes from a grave site in what is now northern Norway (VK531, Margaryan et al. 2019). This individual has a genome-wide profile similar to that of local Mesolithic hunter-gatherers, but is dated to just ~2,400 BCE. During this time, Scandinavia was dominated by a "new" population associated with the Battle-Axe culture (BAC), with high levels of ancestry from the steppes of Eastern Europe. Since VK531 wasn't buried with any BAC grave goods, and indeed with no grave goods at all, it's possible that he may have been from a remnant forager population that was displaced and ultimately forced into extinction.

- R1b-U106 is today by far the most common R1b subclade in Scandinavia, but it's not yet clear how it managed to attain this status. Was it perhaps through elite dominance? The earliest ancient individual belonging to R1b-U106 is dated to 2275-2032 calBCE and comes from a Late Neolithic, likely post-BAC burial ground in what is now Sweden (RISE98, Lilla Beddinge, grave 49, southern skeleton, Allentoft et al. 2015). However, RISE98 wasn't buried in any way that would suggest he was an individual of high social standing. In fact, he was found in a mass grave, along with two other adults and two infants, possibly representing a human sacrifice. The only artefact in the grave was a bone needle. More details are available here.

- During the Nordic Bronze Age it became customary for Scandinavian elites to be laid to rest in richly furnished barrows, while commoners were buried in flat graves with few or no offerings. Human remains recovered from a "commoner" flat grave cemetery dated to the Early Bronze Age near the present-day city of Aalborg, northern Denmark, included the skeleton of a male belonging to Y-haplogroup R1b-M269 (RISE47, grave 3, skeleton 8, Allentoft et al. 2015). Keep in mind, however, that this might have been another case of an ancient Scandinavian R1b-U106 if not for missing data. A flint dagger was found alongside one of the skeletons in this cemetery, but RISE47 wasn't accompanied by any grave goods (see here).

- One of the most amazing archeological discoveries made in Scandinavia is the Trundholm Sun Chariot. Found in a peat bog on the island of Zealand, Denmark, in 1902, it's thought to be an Indo-European religious artefact dating back to the Nordic Bronze Age; a representation of a horse pulling the sun and perhaps also the moon in a spoked wheel chariot. Another important discovery in a peat bog near Trundholm dating to the Nordic Bronze Age was the body of a man belonging to R1b-M269 (RISE276, Trundholm mose II, bog find 1940, Allentoft et al. 2015). However, chances are slim that RISE276 was a charioteer or, say, a spiritual guru who accidentally drowned in the bog. Most Danish bog bodies are thought to have belonged to sacrificial victims or executed criminals.

- Interestingly, the earliest likely Scandinavian warrior belonging to R1b, and also R1b-U106, is from an early Iron Age burial in present-day northwestern Norway (VK418, Margaryan et al. 2019). This site isn't quite as far north as the grave of the above mentioned VK531, but it's still well within the Arctic circle. Apparently, VK418 was buried with some impressive weapons, potentially of "eastern origin", including a shield, spearheads and a sword. Who knows, he may even have been an elite warrior for his time and place?

The other two main Scandinavian Y-haplogroups, I1a and R1a, haven't yet been found in prehistoric Nordic remains from such, shall we say, depressing burials. That's not to say, of course, that they won't be sooner or later. RISE175, from Allentoft et al. 2015, is currently the only individual who fits the bill as a representative of the Nordic Bronze Age elite. He was buried in a barrow grave in what is now southwest Sweden and probably belongs to Y-haplogroup I1a. That's not much to go on, but perhaps it's a sign of things to come?

See also...

Isotopes vs ancient DNA in prehistoric Scandinavia

Who were the people of the Nordic Bronze Age?

They came, they saw, and they mixed

Roopkund Lake dead

Fifteen of the Roopkund Lake samples from the Harney et al. paper published today at Nature Communications made it into the Global25 datasheets. Look for the prefix IND_Roopkund here...

Global25 datasheet ancient scaled

Global25 pop averages ancient scaled

Global25 datasheet ancient

Global25 pop averages ancient

Their genotypes are freely available in a ~590K SNP dataset via the Reich Lab here. I might be able to run more of the samples at some point if and when they're released in a dataset with more SNPs.

In any case, much like everyone else, I don't have a clue how those Mediterranean migrants ended up in the Himalayas back in the 1800s, but I do know where they came from. Most appear to have been from Crete, while others from mainland Greece. However, one of the individuals that I was able to analyze with the Global25 was almost certainly an Anatolian Greek. Below are a couple of Principal Component Analyses (PCA) based on the Global25 data. The relevant datasheet is available here.

I don't yet have a strong opinion about the origins of the earlier, typically South Asian Roopkund dead. They may have been visitors from all over India, or members of different castes from northern India. A PCA with six of these individuals can be seen here and the relevant datasheet gotten here. Any thoughts? Feel free to share them in the comments below.

Update 23/08/2019: A new ~1240K SNP genotype dataset with the Roopkund Lake samples is now available here. More markers means that I can produce more accurate PCA and run almost twice as many of the samples. I've updated all of the datasheets accordingly. The links are the same.

See also...

Getting the most out of the Global25

A surprising twist to the Shirenzigou nomads story

The Poltavka outlier

A surprising twist to the Shirenzigou nomads story

Remember those potentially Afanasievo-derived and Tocharian-related Shirenzigou nomads from the Ning et al. paper? Well, in my opinion, they're probably neither. The genotypes and other data for these Iron Age individuals from the eastern Tian Shan are available here.

Below are a few successful and not so successful qpAdm mixture models for them. Note that I tried to use a wide range of relevant "right pops", but also retain a lot of markers, specifically to be able to discriminate between different types of steppe and steppe-derived sources of gene flow (refer to the full output). Admittedly, the Shirenzigou nomads can be modeled with Afanasievo-related ancestry, but...

CHN_Shirenzigou_IA
KAZ_Botai 0.161±0.023
KAZ_Wusun 0.490±0.023
NPL_Mebrak_2125BP 0.349±0.019
chisq 5.793
tail prob 0.926172
Full output

CHN_Shirenzigou_IA
KAZ_Botai 0.143±0.022
NPL_Mebrak_2125BP 0.295±0.019
Saka_Tian_Shan 0.562±0.024
chisq 6.796
tail prob 0.870794
Full output

CHN_Shirenzigou_IA
KAZ_Botai 0.185±0.023
NPL_Mebrak_2125BP 0.428±0.021
RUS_Sintashta_MLBA 0.270±0.026
TJK_Sarazm_En 0.117±0.027
chisq 11.351
tail prob 0.414345
Full output

CHN_Shirenzigou_IA
KAZ_Botai 0.032±0.027
KAZ_Zevakinskiy_LBA 0.567±0.025
NPL_Mebrak_2125BP 0.401±0.019
chisq 15.157
tail prob 0.232961
Full output

CHN_Shirenzigou_IA
NPL_Mebrak_2125BP 0.452±0.031
RUS_Afanasievo 0.435±0.025
RUS_Okunevo_BA 0.114±0.049
chisq 19.808
tail prob 0.0708003
Full output

CHN_Shirenzigou_IA
NPL_Mebrak_2125BP 0.409±0.031
RUS_Okunevo_BA 0.173±0.050
Yamnaya_RUS_Caucasus 0.418±0.026
chisq 20.453
tail prob 0.0589872
Full output

CHN_Shirenzigou_IA
NPL_Mebrak_2125BP 0.464±0.033
RUS_Okunevo_BA 0.104±0.053
Yamnaya_RUS_Samara 0.432±0.027
chisq 27.189
tail prob 0.0072566
Full output

Both the Wusun and Saka are generally accepted to have been the speakers of Indo-Iranian languages. So it's possible that the Shirenzigou nomads were Indo-Iranian speakers too, or at least derived from such peoples.

Surprisingly, NPL_Mebrak_2125BP was the key to obtaining the best statistical fits. This is a trio of samples, roughly contemporaneous with the Shirenzigou nomads, from a burial site high up in the Himalayas in what is now Nepal (see here).

To be honest, I'm not quite sure why the Himalayan ancients work so well in my models. Perhaps they're just a really good proxy for an Iron Age population from the northern edge of the Tibetan Plateau?

By the way, most of the Shirenzigou nomads made it into the latest Global25 datasheets (see here). They can be analyzed in a variety of ways described in this blog post: Getting the most out of the Global25. Below is a screen cap of me comparing the effectiveness of Afanasievo, Sintashta and Wusun samples as proxies for the steppe ancestry in the Shirenzigou nomads with an online tool freely available here. As expected, the algorithm picks Sintashta ahead of Afanasievo, and the Wusun ahead of both.

See also...

They mixed up Huns with Tocharians

Some myths die hard

The mystery of the Sintashta people

Did South Caspian hunter-fishers really migrate to Eastern Europe?

The idea that most of the Near Eastern-related ancestry in the ancient populations of the Pontic-Caspian (PC) steppe is, one way or another, sourced from the territory of present-day Iran is a fairly popular one nowadays (for instance, see here). It might turn out to be correct, once there are enough relevant samples to test it properly, but in my opinion the chances of this are slim.

My skepticism is based on literally hours of analyses with the currently available ancients from the Caucaso-Caspian region, like, for instance, the admixture graphs below featuring foragers and early farmers from Russia, Georgia and Iran. The relevant qpGraph and dot files are available here.

Note that the further I move away from Eastern Europe in these graphs when looking for the source of the southern ancestry in the Eneolithic population from the southernmost part of the PC steppe (Piedmont_En), the more difficult it is for me to create a statistically sound model. What might this tell us about the provenance of this so called southern ancestry?

See also...

The PIE homeland controversy: August 2019 status report

Some myths die hard

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

The PIE homeland controversy: August 2019 status report

Archeologist David Anthony has a new paper on the Indo-European homeland debate titled Archaeology, Genetics, and Language in the Steppes: A Comment on Bomhard. It's part of a series of articles dealing with Allan R. Bomhard's "Caucasian substrate hypothesis" in the latest edition of The Journal of Indo-European Studies. It's also available, without any restrictions, here.

Any thoughts? Feel free to share them in the comments below. Admittedly, I found this part somewhat puzzling (emphasis is mine):

It was the faint trace of WHG, perhaps 3% of whole Yamnaya genomes, that identified this admixture as coming from Europe, not the Caucasus, according to Wang et al. (2018). Colleagues in David Reich’s lab commented that this small fraction of WHG ancestry could have come from many different geographic places and populations.

I think that's highly optimistic. It really should be obvious by now thanks to archeological and ancient genomic data, including both uniparental and genome-wide variants, that the Yamnaya people were practically entirely derived from Eneolithic populations native to the Pontic-Caspian (PC) steppe. So, in all likelihood, this was also the source of their minor WHG ancestry.

Indeed, they clearly weren't some mishmash of geographically, culturally and genetically disparate groups that had just arrived in Eastern Europe, but the direct descendants of closely related and already significantly Yamnaya-like peoples associated with long-standing PC steppe archeological cultures such as Khvalynsk and Sredny Stog. I discussed this earlier this year, soon after the Wang et al. paper was published:

On Maykop ancestry in Yamnaya

I hope I'm wrong, but I get the feeling that the scientists at the Reich Lab are finding this difficult to accept, because it doesn't gel with their theory that archaic Proto-Indo-European (PIE) wasn't spoken on the PC steppe, but rather south of the Caucasus, and that late or rather nuclear PIE was introduced into the PC steppe by migrants from the Maykop culture who were somehow involved in the formation of the Yamnaya horizon.

Inexplicably, after citing Wang et al. on multiple occasions and arguing against any significant gene flow between Maykop and Yamnaya groups, Anthony fails to mention Steppe Maykop. But the Steppe Maykop people are an awesome argument against the idea that there was anything more than occasional mating between the Maykop and Yamnaya populations, because they were wedged between them, and yet clearly distinct from both, with a surprisingly high ratio of West Siberian forager-related ancestry (see here and here).

Despite all the talk lately about the potential cultural, linguistic and genetic ties between Maykop and Yamnaya, including claims that the latter possibly acquired its wagons from the former, my view is that the Steppe Maykop and Yamnaya wagon drivers may have competed with each other and eventually clashed in a big way. Indeed, take a look at what happens after Yamnaya burials rather suddenly replace those of Steppe Maykop just north of the Caucasus around 3,000 BCE.

Yamnaya_RUS_Caucasus
RUS_Progress_En_PG2001 0.808±0.058
RUS_Steppe_Maykop 0.000
UKR_Sredny_Stog_II_En_I6561 0.192±0.058
chisq 13.859
tail prob 0.383882
Full output

Yep, total population replacement with no significant gene flow between the two groups. Apparently, as far as I can tell, there's not even a hint that a few Steppe Maykop stragglers were incorporated into the ranks of the newcomers. Where did they go? Hard to say for now. Maybe they ran for the hills nearby?

Intriguingly, Anthony reveals a few details about new samples from three different Eneolithic steppe burial sites associated with the Khvalynsk culture:

The Reich lab now has whole-genome aDNA data from more than 30 individuals from three Eneolithic cemeteries in the Volga steppes between the cities of Saratov and Samara (Khlopkov Bugor, Khvalynsk, and Ekaterinovka), all dated around the middle of the fifth millennium BC.

...

Most of the males belonged to Y-chromosome haplogroup R1b1a, like almost all Yamnaya males, but Khvalynsk also had some minority Y-chromosome haplogroups (R1a, Q1a, J, I2a2) that do not appear or appear only rarely (I2a2) in Yamnaya graves.

As far as I can tell, he suggests that they'll be published in the forthcoming Narasimhan et al. paper. If so, it sounds like the paper will have many more ancient samples than its early preprint that was posted at bioRxiv last year.

For me the really fascinating thing in regards to these new samples is how scarce Y-haplogroup R1a appears to have been everywhere before the expansion by the putative Indo-European-speaking steppe ancestors of the Corded Ware culture (CWC) people. It's basically always outnumbered by other haplogroups wherever it's found prior to about 3,000 BCE, even on the PC steppe. But then, suddenly, its R1a-M417 subclade goes BOOM! And that's why I call it...

The beast among Y-haplogroups

At this stage, I'm not sure how to interpret the presence of Y-haplogroup J in the Khvalynsk population. It may or may not be important to the PIE homeland debate. Keep in mind that J is present in two foragers from Karelia and Popovo, northern Russia, dated to the Mesolithic period and with no obvious foreign ancestry. So it need not have arrived north of the Caspian as late as the Eneolithic with migrants rich in southern ancestry from the Caucasus or what is now Iran. In other words, for the time being, the steppe PIE homeland theory appears safe.

Update 20/12/2019: A note on Steppe Maykop

See also...

Is Yamnaya overrated?

The PIE homeland controversy: January 2019 status report

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

They mixed up Huns with Tocharians

I don't yet have the genomes from the recent Ning et al. paper on the Iron Age nomads from the Shirenzigou site in the eastern Tian Shan. But I do have most of the previously published data featured in the paper, including the Damgaard et al. 2018 Hun and Saka samples from the western Tian Shan.

After reading the Ning et al. paper between the lines and running a few analyses of my own, it's clear to me that most of the supposedly Tocharian-related Shirenzigou individuals actually share a very close relationship with the Tian Shan Huns, and indeed may have been their ancestors.

For instance, Ning et al. found that a large part of the ancestry of the Shirenzigou ancients could be modeled with the Tian Shan Huns, which was an anachronistic approach because the former are older than the latter. They also found that Ulchi-related ancestry was a key part of the genetic structure of eight out of the ten Shirenzigou individuals, and this likewise appears to be an important part of the genetic structure of the Tian Shan Huns.

Note the strong statistical fits in the Global25/nMonte and qpAdm mixture models below, respectively, which characterize these Huns as a two-way mixture between the Ulchi and the earlier Tian Shan Saka. And keep in mind that the Saka also harbor significant Ulchi-related ancestry.

Hun_Tian_Shan
Saka_Tian_Shan,92
Ulchi,8
distance%=1.2553

Hun_Tian_Shan
Saka_Tian_Shan 0.928±0.009
Ulchi 0.072±0.009
chisq 4.409
tail prob 0.992464
Full output

Moreover, the Shirenzigou males belong to Y-haplogroups Q1a and R1b (two instances of each), and they share the latter with one of the Tian Shan Huns. Judging by the data from the relevant BAM files, it's also possible that the Shirenzigou males share a very rare subclade of R1b with the Hun, defined by the PH155 mutation (see here). The Y-haplogroup assignments for the other Tian Shan Huns end at R and R1, but that's almost certainly due to missing data.

On the other hand, two Tian Shan Sakas belong to Y-haplogroup R1a but none to R1b, which fits with the pattern from currently available ancient DNA that R1a was more common than R1b in Saka-related groups, such as the Scythians and Sarmatians (see here).

This is all very interesting, because the Huns replaced the Saka in the western Tian Shan, and, considering their R1b and excess Ulchi-related ancestry, very likely moved into the region from the direction of Shirenzigou. Indeed, in my opinion a strong argument can now be made that the Iron Age population from the Shirenzigou region took part in the formation of the Hunnic confederacy.

So where does that leave the theory presented by Ning et al. that the Shirenzigou ancients may have been closely related, and perhaps even ancestral, to the Tocharians, simply because they packed a lot of Yamnaya-related and possibly proto-Tocharian Afanasievo ancestry, and were living close to the Tarim Basin, where Tocharian languages were subsequently first attested?

I'm not sure, but I now find it difficult to reconcile this theory with the fact that they were closely related, and probably ancestral, to the Tian Shan Huns. As far as I'm aware, Huns cannot be linked to Tocharians in any meaningful way.

Of course it's possible that different Afanasievo-derived groups were living in the Tarim Basin and surrounds, and, as some merged with new populations pushing into the region from the east and adopted non-Indo-European languages, others retained their Tocharian speech and eventually split into communities speaking Tocharian A, B and apparently also C (see here).

But this has to be demonstrated directly with ancient DNA from archeological sites where Tocharian languages were attested. Till then, I'll keep thinking that Ning et al. wrote a paper about Tocharians that really should've been a paper about Huns.

Here's a famous wall painting of Tocharian princes from the cave of the sixteen sword-bearers in the Tarim Basin, dated to 432–538 AD. They don't look like guys with a lot of Ulchi-related admixture to me, but I might be wrong. Feel free to let me know what you think in the comments below.

Update 08/17/2019: The Shirenzigou nomads are now in my dataset. Below are a few successful and not so successful qpAdm mixture models for them. Note that I tried to use a wide range of relevant "right pops", but also retain a lot of markers, specifically to be able to discriminate between different types of steppe and steppe-derived sources of gene flow (refer to the full output). Admittedly, the Shirenzigou nomads can be modeled with Afanasievo-related ancestry, but...

CHN_Shirenzigou_IA
KAZ_Botai 0.161±0.023
KAZ_Wusun 0.490±0.023
NPL_Mebrak_2125BP 0.349±0.019
chisq 5.793
tail prob 0.926172
Full output

CHN_Shirenzigou_IA
KAZ_Botai 0.143±0.022
NPL_Mebrak_2125BP 0.295±0.019
Saka_Tian_Shan 0.562±0.024
chisq 6.796
tail prob 0.870794
Full output

CHN_Shirenzigou_IA
KAZ_Botai 0.185±0.023
NPL_Mebrak_2125BP 0.428±0.021
RUS_Sintashta_MLBA 0.270±0.026
TJK_Sarazm_En 0.117±0.027
chisq 11.351
tail prob 0.414345
Full output

CHN_Shirenzigou_IA
KAZ_Botai 0.032±0.027
KAZ_Zevakinskiy_LBA 0.567±0.025
NPL_Mebrak_2125BP 0.401±0.019
chisq 15.157
tail prob 0.232961
Full output

CHN_Shirenzigou_IA
NPL_Mebrak_2125BP 0.452±0.031
RUS_Afanasievo 0.435±0.025
RUS_Okunevo_BA 0.114±0.049
chisq 19.808
tail prob 0.0708003
Full output

CHN_Shirenzigou_IA
NPL_Mebrak_2125BP 0.409±0.031
RUS_Okunevo_BA 0.173±0.050
Yamnaya_RUS_Caucasus 0.418±0.026
chisq 20.453
tail prob 0.0589872
Full output

CHN_Shirenzigou_IA
NPL_Mebrak_2125BP 0.464±0.033
RUS_Okunevo_BA 0.104±0.053
Yamnaya_RUS_Samara 0.432±0.027
chisq 27.189
tail prob 0.0072566
Full output

Both the Wusun and Saka are generally accepted to have been the speakers of Indo-Iranian languages. So it's possible that the Shirenzigou nomads were Indo-Iranian speakers too, or at least derived from such peoples.

Surprisingly, NPL_Mebrak_2125BP was the key to obtaining the best statistical fits. This is a trio of samples, roughly contemporaneous with the Shirenzigou nomads, from a burial site high up in the Himalayas in what is now Nepal (see here).

To be honest, I'm not quite sure why the Himalayan ancients work so well in my models. Perhaps they're just a really good proxy for an Iron Age population from the northern part of the Tibetan Plateau? By the way, most of the Shirenzigou nomads made it into the latest Global25 datasheets (see here).

See also...

The mystery of the Sintashta people

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

Viking invasion at bioRxiv

A new preprint featuring hundreds of Viking Age genomes has appeared at bioRxiv [LINK]. Titled Population genomics of the Viking world, it looks like a solid effort overall, although I'm skeptical about its conclusions. I might elaborate on that in the comments below, but I'll have a lot more to say on the topic if and when I get to check out the ancient genomes with my own tools. Details about the new samples, including their Y-chromosome haplogroup assignments, are available here. Below is the abstract, emphasis is mine:

The Viking maritime expansion from Scandinavia (Denmark, Norway, and Sweden) marks one of the swiftest and most far-flung cultural transformations in global history. During this time (c. 750 to 1050 CE), the Vikings reached most of western Eurasia, Greenland, and North America, and left a cultural legacy that persists till today. To understand the genetic structure and influence of the Viking expansion, we sequenced the genomes of 442 ancient humans from across Europe and Greenland ranging from the Bronze Age (c. 2400 BC) to the early Modern period (c. 1600 CE), with particular emphasis on the Viking Age. We find that the period preceding the Viking Age was accompanied by foreign gene flow into Scandinavia from the south and east: spreading from Denmark and eastern Sweden to the rest of Scandinavia. Despite the close linguistic similarities of modern Scandinavian languages, we observe genetic structure within Scandinavia, suggesting that regional population differences were already present 1,000 years ago. We find evidence for a majority of Danish Viking presence in England, Swedish Viking presence in the Baltic, and Norwegian Viking presence in Ireland, Iceland, and Greenland. Additionally, we see substantial foreign European ancestry entering Scandinavia during the Viking Age. We also find that several of the members of the only archaeologically well-attested Viking expedition were close family members. By comparing Viking Scandinavian genomes with present-day Scandinavian genomes, we find that pigmentation-associated loci have undergone strong population differentiation during the last millennia. Finally, we are able to trace the allele frequency dynamics of positively selected loci with unprecedented detail, including the lactase persistence allele and various alleles associated with the immune response. We conclude that the Viking diaspora was characterized by substantial foreign engagement: distinct Viking populations influenced the genomic makeup of different regions of Europe, while Scandinavia also experienced increased contact with the rest of the continent.

Margaryan et al., Population genomics of the Viking world, bioRxiv, posted July 17, 2019, doi: https://doi.org/10.1101/703405

See also...

They came, they saw, and they mixed

Who were the people of the Nordic Bronze Age?

Asiatic East Germanics