A Mycenaean and an Iron Age Iranian walk into a bar...

What do they have in common? The same type of Near Eastern ancestry? From Iran? Nope, that's a joke. Obviously, they share the same type of steppe ancestry. This probably has some very important linguistic implications.

The relevant Principal Component Analysis (PCA) datasheet is available here. Below are two pairs of formal mixture models that support my inferences from the PCA.

Mycenaean
Srubnaya_MLBA 0.266±0.029
Tepecik_Ciftlik_N 0.734±0.029
taildiff: 0.588000631
Full output

Mycenaean
Minoan_Lasithi 0.790±0.023
Srubnaya_MLBA 0.210±0.023
taildiff: 0.187709803
Full output

...

Turkmenistan_IA
Namazga_CA 0.528±0.040
Srubnaya_MLBA 0.472±0.040
taildiff: 0.561330411
Full output

Turkmenistan_IA
Dzharkutan1_BA 0.530±0.037
Srubnaya_MLBA 0.470±0.037
taildiff: 0.485083377
Full output

But seriously, what's the direct link between populations like Tepecik_Ciftlik_N/Minoan_Lasithi and Namazga_CA/Dzharkutan1_BA, except some exceedingly distant farmer ancestry from the Fertile Crescent?

See also...

An early Iranian, obviously

Graeco-Aryan parallels

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

An early Iranian, obviously

Today, the part of Asia between the Caspian Sea and the Altai Mountains, known as Turan, is largely a Turkic-speaking region. But during the Iron Age it was dominated by Iranian speakers. Throughout this period it was the home of a goodly number of attested and inferred early Iranic peoples, such as the Airya, Dahae, Kangju, Massagetae, Saka and Sogdians.

Indeed, the early Iron Age Yaz II archaeological culture, located in southwestern Turan, is generally classified as an Iranian culture, and even posited to have been the Airyanem Vaejah, aka home of the Iranians, from ancient Avestan literature.

That's not to say that Iranian speakers weren't present in this part of the world much earlier. They probably were, and it's likely that we already have their genomes (see here). But the point I'm making is that Turan can't be reliably claimed to have been an Iranian realm until the Iron Age.

Ergo, any ancient DNA samples from Turan dating to the Iron Age, as opposed to, say, the Bronze Age, are very likely to be those of early Iranian speakers. One such sample is Turkmenistan_IA DA382 from Damgaard et al. 2018.

Below is a screen cap of the "time map" from homeland.ku.dk, with the slider moved to 847 BC, showing the location of the burial site where the remains of DA382 were excavated. The site is marked with the Z93 label because DA382 belongs to the Eastern European-derived Y-chromosome haplogroup R1a-Z93. Interestingly, his burial was located in close proximity to archaeological sites associated with the above mentioned and contemporaneous Yaz II culture.

DA382 didn't get much of a run in the Damgaard et al. paper, and little wonder because the authors also analyzed 73 other ancient samples. So let's take a close look at this individual's genetic structure to see whether there's anything particularly Iranian about it.

Damgaard et al. did mention that DA382 was partly of Middle to Late Bronze Age (MLBA) steppe origin. And indeed, my own mixture models using qpAdm confirm this finding with very consistent results and strong statistical fits. Here are a couple of two-way examples...

Turkmenistan_IA
Namazga_CA 0.528±0.040
Srubnaya_MLBA 0.472±0.040
taildiff: 0.561330411
Full output

Turkmenistan_IA
Dzharkutan1_BA 0.530±0.037
Srubnaya_MLBA 0.470±0.037
taildiff: 0.485083377
Full output

The fact that the MLBA Srubnaya samples from the Pontic-Caspian steppe can be used to model DA382's ancestry (alongside Bronze and Copper Age populations from Turan) with such ease shouldn't be surprising, considering the he belongs to R1a-Z93, which is the dominant Y-haplogroup in the Srubnaya and all other closely related MLBA steppe peoples.

Now, Srubnaya is generally regarded to be the proto-Iranian archaeological culture. How awesome is that considering those qpAdm fits? But, admittedly, this is just an inference, even if a robust one, based on genetic, archaeological and historical linguistics data. So apart from the fact that DA382 comes from Iron Age Turan, an Iranian-speaking realm, is there any other way to link him directly to Iranians?

Well, he's very similar in terms of overall genetic structure to some of the least Turkic-admixed Iranian speakers still living in Turan, and might well be ancestral to them.

For instance, below is a Principal Component Analysis (PCA) featuring a wide range of ancient and present-day West Eurasian samples. Note that, in line with the qpAdm models, DA382 clusters about half-way between the populations of the MLBA steppe and pre-Kurgan expansion Turan, and amongst present-day Yaghnobi and Pamiri Tajiks. In fact, he clusters at the apex of a southeast > northwest cline made up of Tajiks that appears to be pulling towards Europeans.

Needless to say, Tajiks, especially Pamiri Tajiks, also pack a lot of Srubnaya-related ancestry. I've talked about this plenty of times at this blog (for instance, see here). But what happens if I try to model Pamiri and Yaghnobi Tajiks with DA382?

Tajik
Turkmenistan_IA 0.892±0.023
Han 0.108±0.023
taildiff: 0.794566182
Full output

Wow, it's an awesome fit! My mind's made up: DA382 was probably an Iranian speaker and, more specifically, an Eastern Iranian speaker. Who disagrees and why? Feel free to let me know in the comments (unless you're banned, in which case, f*ck off).

See also...

A Mycenaean and an Iron Age Iranian walk into a bar...

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

New PCA featuring Botai horse tamers, Hun and Saka warriors, and many more...

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

Click HERE to view the interactive time map, and give it some time to load if you're on a slow connection. Use the slider tool to explore different time periods from 6385 BC to 1 BC. It's still a work in progress, so feel free to let the author, Mikkel Nørtoft, know if anything should be added, tweaked and/or generally improved.

Below is a screen cap of the map with the slider moved to 3618 BC. Note the sheep in the North Pontic steppe and the wheels just west of it. Wink, wink, nudge, nudge...

See also...

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

SMBE 2018 abstracts

Abstracts from the upcoming SMBE 2018 conference (8-12 July) are now available HERE. Below are a few that I found interesting. Emphasis is mine. Feel free to post your own favorites in the comments.

The first Epipaleolithic Genome from Anatolia suggests a limited role of demic diffusion in the Advent of Farming in Anatolia

Feldman et al.

Anatolia was home to some of the earliest farming communities, which in the following millennia expanded into Europe and largely replaced local hunter-gatherers. The lack of genetic data from pre-farming Anatolians has so far limited demographic investigations of the Anatolian Neolithisation process. In particular, it has been unclear whether farming was adopted by indigenous hunter-gatherers in Central Anatolia or imported by settlers from earlier farming centers. Here we present the first genome-wide data from an Anatolian Epipaleolithic hunter-gatherer who lived ~15,000 years ago, as well as from Early Neolithic individuals from Anatolia and the Levant. By using a comparative dataset of modern and ancient genomes, we estimate that the earliest Anatolian farmers derive over 90 percent of their ancestry from the local Epipaleolithic population, indicating a high degree of genetic continuity throughout the Neolithic transition. In addition, we detect two distinct waves of gene flow during the Neolithic transition: an earlier one related to Iranian/Caucasus ancestry and a later one linked to the Levant. Finally, we observe a genetic link between Epipaleolithic Near-Easterners and post-glacial European hunter-gatherers that suggests a bidirectional genetic exchange between Europe and the Near East predating 15,000 years ago. Our results suggest that the Neolithisation model in Central Anatolia was demographically similar to the one previously observed in the southern Levant and in the southern Caucasus-Iran highlands, further supporting the limited role of demic diffusion during the early spread of agriculture in the Near East, in contrast to the later Neolithisation of Europe.

...

Demographic processes in Estonia from Bronze Age through Iron Age to Medieval times

Metspalu et al.

N3 and R1a are the two most common Y chromosome haplogroups among modern Estonians. R1a appears with Corded Ware culture but the arrival of hg N has not been determined. To this end we have extracted and studied aDNA from teeth of 18 individuals bracketing the changes in the material culture in the end of the Bronze and early Iron Age. We find N3 in Iron Age but not in Bronze Age. Due to the small sample size we cannot refute the existence of hg N in the latter. In genome wide analyses the Bronze Age and especially Iron Age samples appear very similar to modern Estonians implying population continuity. Christianization (13 cc AD) established a new elite of West European origin, which presumably had an impact on the genetic structure of the local population. To investigate this we extracted DNA from teeth of 35 individuals, who have been uncovered from both rural (considered local Estonian population) and town (likely of West European origin) cemeteries of Estonia. We compared the low coverage genomes with each other and with relevant modern and ancient Estonian and other European populations. We find that there is a clear discontinuity between the elite and common people, where the former group genetically with modern German samples and the latter with modern Estonians. We do find three individuals of mixed genetic ancestry. But importantly we do not see a steady shift of either local population strata, which suggests limited contact between the elite and the common people.

...

Genetic transition in the Swiss Late Neolithic and Early Bronze Age

Furtwaengler et al.

Recent studies have shown that the beginning of the Neolithic period as well as final stages of the Neolithic were marked by major genetic turnovers in European populations.The transition from hunter-gatherers to agriculturalists and farmers/farming in the 6 th millennium BP coincided with a human migration from the Near East. In the 3 rd millennium BP a second migration into Central Europe occurred originating from the Pontic steppe linked to the spread of the Corded Ware Complex ranging as far southwest as modern day Switzerland. These genetic processes are well studied for example for the Middle-Elbe-Saale region in Eastern Germany, however, little is known from the regions that connect Central and Southern Europe. Here, we investigate genomic data from 69 individuals from the Swiss Plateau and Southern Germany that span the transition of the Neolithic to the Bronze Age (5500 to 4000 BP). Our results show a similar genetic process as reported for the Middle-Elbe-Saale region suggesting that the migration from the Pontic steppe reached all the way into the Swiss plateau. The high quality of the ancient genomic data also allowed an analysis of core families within multiple burials, the determination and qualification of different ancestry components and the determination of the migration route taken by the ancestors of the Late Neolithic populations in this region. This study presents the first comprehensive genome wide dataset from Holocene individuals from the Swiss plateau and provides the first glimpse into the genetic history of this genetically and linguistically diverse region.

...

Genome-Wide Ancient DNA Portrays the Forming of the Finnish Population Along a 1400-Year Transect

Majander et al.

The Finnish population has long been a subject of interest for the fields of medical and population genetics, due to its isolation-affected genetic structure and the associated unique set of inherited diseases. Recent advances in ancient DNA techniques now enable the in-depth investigation of Finland's demographic past: the impact of migrations, trade and altering livelihood practices. Here we analyse genome-wide data from over 30 individuals, representing ten archaeological burial sites from southern Finland, that span from the 5th to 19th century. We find the historical individuals to differ genetically from Finns today. Comparing them with surrounding ancient and modern populations, we detect a transition from genotypes generally connected with prehistoric hunter-gatherers, and specifically resembling those of the contemporary Saami people, into a more East-Central European composition, associated with the established agricultural lifestyle. Starting from the Iron Age and continuing through the Early Medieval period, this transition dates remarkably late compared to the respective changes in most regions of Europe. Our results suggest a population shift, presumably related to Baltic and Slavic influences, also manifested in the archaeological record of the local artefacts from the late Iron Age. Our observations also agree with the archaeological models of relatively recent and gradual adoption of farming in Finland.

...

Population migration and dairy pastoralism on the Bronze Age Mongolian steppe

Warinner et al.

The steppe belt that extends across Eurasia was the primary corridor of Eneolithic and Bronze Age migrations that reshaped the genetics of Europe and Asia and dispersed the Indo-European language family. Beginning in the Eneolithic, a new and highly mobile pastoralist society formed on the Western Steppe. These steppe herders expanded both westwards, contributing to the Corded Ware culture of Eastern and Central Europe, and eastwards, contributing to the mobile pastoralist Afanasevo, Sintashta, Andronovo, and Okunevo cultures in Central Asia. The eastern extent of this Western Steppe herder expansion is not well defined. Here we investigate genome-wide ancestry data obtained from 20 Late Bronze Age (16th-9th century BCE) khirigsuur burials from Khovsgol, Mongolia and further investigate evidence for dairy pastoralism by LC-MS/MS analysis of dental calculus. Overall, we observe limited Western Steppe gene flow into Late Bronze Age Mongolia, but adoption of Western ruminant dairying by ca. 1500 BCE.

...

The Transition to Farming in Eneolithic (Copper Age) Ukraine was Largely Driven by Population Replacement

Schmidt et al.

The transition to a farming-based economy during the Neolithic happened relatively late in southeastern Europe. Material changes occurred through pottery manufacture, but it wasn't until the sixth millennium BCE that farming was adopted by the Cucuteni-Trypillian archaeological complex (4800-3000 BCE). In many parts of Europe, early farmers who were descended from Anatolian migrants slowly admixed with local hunter-gatherers over the course of the Neolithic. In Eastern Europe and the Balkans, this process may have been more complex since early farmers would likely have admixed with local groups prior to spreading into continental Europe. Studies from the Baltic and Estonia suggest little genetic input from early farmers or continuous admixture with hunter-gatherers. Here, we investigate the impact of Trypillian migrations into Ukraine through the analyses of 19 ancient genomes (0.6 to 2.1X coverage) from the site of Verteba Cave. Ceramic typology and radiocarbon dating of the cave indicate continuous occupation from the Mesolithic to the Medieval Period, with peak occupation coinciding with the middle to late Tripolye. We show that the Trypillians replaced local Ukrainian Neolithic cultures. Also, hunter-gatherers contributed very little ancestry to the Trypillians, who are genetically indistinct from early Neolithic farmers. The one exception is a female that has mostly steppe-related ancestry. Direct radiocarbon dating of this individual places her in the the Middle Bronze Age (3545 years before present). Her lack of farmer ancestry suggests abrupt population replacement resulting perhaps from inter-group hostilities or plague that spread through Europe during the Late Neolithic.

See also...

Ahead of the pack

Genetic borders are usually linguistic borders too

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

How relevant is Arslantepe to the PIE homeland debate?

Below is an abstract of a presentation from the recent 11th International Congress on the Archaeology of the Ancient Near East (see here). It was discussed on at least a couple of DNA forums, and hailed by some as potentially pivotal to the Proto-Indo-European (PIE) homeland debate. Emphasis is mine:

Palaeogenetic and Anthropological Perspectives on late Chalcolithic and Early Bronze Age Arslantepe

Skourtanioti, Eirini - Max Planck Institute for the Science of Human History, Jena
Selim, Erdal Yilaz - Department of Anthropology, Hacettepe University, Ankara

While Anatolia was highlighted as the genetic origin of early Neolithic European farmers, the genetic substructure in Anatolia itself as well as the demographic and cultural changes remain unclear. In eastern Anatolia, the archaeological record reflects influences from North-Central Anatolia, the northeastern sectors of Fertile Crescent and the Caucasus, and suggests that some of these were brought along with the movement of people. Central to this question is the archaeological site of Arslantepe (6 th -1 st millennium BC), strategically located at the Upper Euphrates, the nexus of all three regions. Arslantepe also developed one of the first state societies of Anatolia along with advanced metal-technologies. Archaeological research suggests that conflicts with surrounding groups of pastoralists affiliated to the Caucasus might have contributed to the collapse of its palatial system at the end of the Chalcolithic period (4 th millennium BC). To test if these developments were accompanied by genetic changes, we generated genome-wide data from 18 ancient individuals spanning from the Late Chalcolithic period to the Early Bronze Age of Arslantepe. Our results show no evidence for a major genetic shift between the two time periods. However, we observe that individuals from Arslantepe are very heterogeneous and differentiated from other ancient western and central Anatolians in that they have more Iran/Caucasus related ancestry. Our data also show evidence for an ongoing but also recent confluence of Anatolian/Levantine and Caucasus/Iranian ancestries, highlighting the complexity of the Chalcolithic and Bronze Age periods in this region.

Actually, I think it's likely that the good people at Max Planck will try to weave these results into their southern PIE homeland theory.

If so, they'll probably claim that since there's no discernible steppe ancestry in any of the 18 ancient Anatolians, then it's unlikely that there were any migrations from the Pontic-Caspian steppe into Anatolia during the Chalcolithic and Bronze Age periods. Ergo, they're likely to conclude that the PIE homeland was located south of the steppe, perhaps in Transcaucasia or what is now Iran, because this is where some of the Arslantepe ancients appear to source a lot of their recent ancestry from. But that would be a mistake.

First of all, the best way to link Arslantepe to the Proto-Indo-Europeans is probably via the massive Arslantepe Royal Tomb, which shows a close archaeological relationship with kurgan burials in the North Caucasus. Here's an abstract from a paper on the topic from 2011 (freely available here). Emphasis is mine:

The first appearance of the Kurgan funerary tradition in the Northern Caucasus (Majkop‐Novosvobodnaya) dates to the second half of the fourth millennium and records an impressive display and accumulation of wealth in the grave goods (mainly metal objects) which stress the emergence of radical social transformations in the communities of the region. But kurgans also signal a new approach to territory and a different conception of the landscape. The Arslantepe Royal Tomb (in the Upper Euphrates Valley, eastern Anatolia), which is dated to 3100-2900 B.C., shows that far-reaching influences from the Northern Caucasus were already crossing the Greater Caucasus range and that they were being assimilated by the Anatolian power groups. The set of traits that the Arslantepe Royal Tomb shares with the funerary representations of the northern Caucasian Kurgans (ritual, grave goods and eventually the location chosen for the burial) was the result of a symbolic and ideological selection performed by a local community. It aimed at legitimizing and justifying the current historical and political contingencies and the emergence of new images and power rules. What can be grasped of the general sense and cultural values of the phenomenon of the northern Caucasian Kurgans by means of an interpretation performed by an external (and distant) community?

So we can still argue that the PIE homeland was on the steppe, with Anatolian languages splitting from the rest of PIE in the North Caucasus, and then making their way south to Anatolia along with people of fully Caucasian genetic structure. As far as I can tell, that's essentially the theory proposed recently by Damgaard et al. in their linguistic supplement (see here).

But I'm not a huge fan of this solution. Clearly, the archaeological and genetic changes at Arslantepe from the Chalcolithic to the Bronze Age were in large part caused by the arrival of groups associated with the Kura-Araxes archaeological culture, and they were probably speakers of early Hurra-Urutian languages. If there were any migrants directly from the North Caucasus at Arslantepe, then I'd say that they probably spoke Caucasian languages.

Indo-Europeans certainly arrived at Arslantepe during the Late Bronze Age, with the expansion of the Hittite Empire into the region. But it's impossible to say how much, if any, PIE ancestry they brought with them, because obviously language shifts are a complex phenomenon, and this is especially true in state societies, like the Hittite Empire, in which cultural and political domination can lead to widespread language change without any accompanying gene flow.

See also...

Ahead of the pack

Genetic borders are usually linguistic borders too

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

Friendly Yeniseian steppe pastoralists

For most people the Proto-Indo-European (PIE) homeland debate isn't just about language, but also, or even more so, about things like ancestry, politics, racism, and ethnic pride.

I don't want to get into all the dirty details in this post, but, for instance, many of those who argue vehemently against a steppe homeland seem to really hate the idea that their ancestors were, at some level, dominated by a bunch of sheep and cow herders from an obscure part of Eastern Europe. Why? I'm pretty sure because they find this humiliating.

Hence, PIE homeland debates are generally very emotional and often degenerate into shouting matches. So what would happen if we assumed that those sheep herders weren't Indo-Europeans? Let's say, for the sake of argument, that they spoke extinct Yeniseian languages.

Well, nothing really...

We still have them expanding in a big way out of the Pontic-Caspian steppe. And the fact that their impact was mostly male mediated, especially at the two ends of its range, in Iberia and India, suggests that they weren't just friendly pastoralists looking for new grazing fields.

In fact, in India, steppe ancestry, including Y-chromosome haplogroup R1a, is much more pronounced in the upper castes than in the rest of society. But who knows what that means, right? Maybe those friendly "Yeniseian" steppe pastoralists just got lucky or something?

But that's OK, perhaps this will always remain a mystery? In any case, does anyone know if there's any sort of Yeniseian substrate in Celtic or Mycenaean? Haha.

See also...

"Heavily sex-biased" population dispersals into the Indian Subcontinent (Silva et al. 2017)

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

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

Ahead of the pack

Eurogenes Blog January 2018 (see here):

Yamnaya and other similar Eneolithic/Bronze Age herder groups from the Eurasian steppe were mostly a mixture of Eastern European Hunter-Gatherers (EHG) and Caucasus Hunter-Gatherers (CHG). But they also harbored minor ancestry from at least one, significantly more westerly, source that pulled them away from the EHG > CHG north/south genetic cline.

...

It's interesting and, I'd say, important to note that the West Asian reference groups produce amongst the worst statistical fits (in bold text). What this suggests is that Yamnaya did not harbor extra West Asian ancestry on top of its CHG input.

...

Rather, Blatterhole_MN is simply the best proxy in this analysis for the non-CHG/EHG ancestry in Yamnaya, and the important question is why?

Considering also the presence at the top of the list of Koros_HG (which includes Hungary_HG I1507), Germany_MN and Vinca_MN, the likely answer is its high ratio of Western European Hunter-Gatherer (WHG) ancestry.

...

So is the missing piece of the Yamnaya puzzle a population with roughly equal ratios of Early Neolithic (EN) and WHG ancestries from the Carpathian Basin or surrounds? Quite possibly. But let's wait and see what happens when I add the ancient groups from the Balkans and North Pontic steppe from the forthcoming Mathieson et al. 2018 to this analysis.

And now Wang et al. May 2018 (see here).

In principal component space Eneolithic individuals (Samara Eneolithic) form a cline running from EHG to CHG (Fig. 2D), which is continued by the newly reported Eneolithic steppe individuals.

...

However, PCA results also suggest that Yamnaya and later groups of the West Eurasian steppe carry some farmer related ancestry as they are slightly shifted towards ‘European Neolithic groups’ in PC2 (Fig. 2D) compared to Eneolithic steppe.

...

Importantly, our results show a subtle contribution of both Anatolian farmer-related ancestry and WHG-related ancestry (Fig.4; Supplementary Tables 13 and 14), which was likely contributed through Middle and Late Neolithic farming groups from adjacent regions in the West. A direct source of Anatolian farmer-related ancestry can be ruled out (Supplementary Table 15).

...

We find that Yamnaya individuals from the Volga region (Yamnaya Samara) have 13.2±2.7% and Yamnaya individuals in Hungary 17.1±4.1% Anatolian farmer-related ancestry (Fig.4; Supplementary Table 18)– statistically indistinguishable proportions. Replacing Globular Amphora by Iberia Chalcolithic, for instance, does not alter the results profoundly (Supplementary Table 19). This suggests that the source population was a mixture of Anatolian farmer-related ancestry and a minimum of 20% WHG ancestry, a profile that is shared by many Middle/Late Neolithic and Chalcolithic individuals from Europe of the 3 rd millennium BCE analysed thus far.

Strikingly similar, don't you think? However, I'm not implying that they copied me. The point I'm making is that I predicted this outcome ahead of anyone else, and was able to demonstrate it without some of the key ancient samples that Wang et al. had access to. Indeed, kudos to them for finding and successfully sequencing those all important new Eneolithic steppe samples.

Moreover, these results mean that it's no longer plausible to argue that the Yamnaya population, by and large, formed due to recent gene flow from south of the Caucasus, let alone from what is now Iran, into the Pontic-Caspian steppe. Obviously, this is a major problem for anyone arguing that the Proto-Indo-European (PIE) homeland may have been located somewhere south of the Caucasus, such as Paul Heggarty of the Max Planck Institute for the Science of Human History (for instance, see here). But hey, never mind the facts when you have an awesome theory, right?

See also...

Yamnaya: home-grown

Genetic borders are usually linguistic borders too

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