Three key late comers in prehistoric Greece: steppe ancestry, horses and millet

A review paper at Archaeological and Anthropological Sciences posits that millet and horses arrived in what is now Greece together during the Bronze Age (see here). The author suggests that they may have been introduced via contacts with cultures to the north/northeast of Greece or directly by migrants from the Eurasian steppe. Considering the recent discovery via ancient human DNA that steppe ancestry also spread into the southern Balkans and Mycenae during the Bronze Age (see links below), I'd say the latter scenario is much more likely. I'd also add that millet and horses were probably part of an economic and cultural package expanding along with early Indo-European speakers throughout Eurasia at the time (note, for instance, how important horses are to the early Indo-European pantheon). Here's the review abstract. Emphasis is mine.

Abstract: Archaebotanical evidence for Panicum miliaceum is reviewed for prehistoric Greece including published and unpublished recent finds, providing a basis for exploring the context of the appearance of millet in Greece, the timing of its introduction and cultivation, and its significance in terms of contacts, movement of people, and cultural identity as expressed through culinary practice and food consumption. To this end, the archaeobotanical record is examined together with human isotopic, archaeozoological, and artefactual evidence. Millet is introduced to the northern part of Greece sometime during the end of the 3rd millennium bc and established as a widely used crop during the Late Bronze Age. Isotopic evidence suggests that millet consumption during the Late Bronze Age was not widespread but confined to certain regions, settlements, or individuals. Millet is suggested to reach Greece from the north after its spread westwards from China through Central Asia and the steppes of Eurasia. The timing of the introduction of millet and the horse in northern Greece coincide; the possibility therefore that they are both introduced through contacts with horse breeding cultures cultivating millet in the north and/or northeast is raised. Intensified contact networks during the Bronze Age, linking prehistoric northern Greece to central Europe and the Pontic Steppes, would have opened the way to the introduction of millet, overland via river valleys leading to the Danube, or via maritime routes, linking the Black Sea to the north Aegean. Alternatively, millet could have been introduced by millet-consuming populations, moving southwards from the Eurasian steppes.

Valamoti, S.M., Millet, the late comer: on the tracks of Panicum miliaceum in prehistoric Greece, Archaeol Anthropol Sci (2016) 8: 51. https://doi.org/10.1007/s12520-013-0152-5

See also...

Steppe invaders in the Bronze Age Balkans

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

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

The story of Y-haplogroup Q

Over at Molecular Genetics and Genomics at this LINK. I wonder how the Q1a Khvalynsk guy (see here) fits into this story?

Abstract: The human Y-chromosome has proven to be a powerful tool for tracing the paternal history of human populations and genealogical ancestors. The human Y-chromosome haplogroup Q is the most frequent haplogroup in the Americas. Previous studies have traced the origin of haplogroup Q to the region around Central Asia and Southern Siberia. Although the diversity of haplogroup Q in the Americas has been studied in detail, investigations on the diffusion of haplogroup Q in Eurasia and Africa are still limited. In this study, we collected 39 samples from China and Russia, investigated 432 samples from previous studies of haplogroup Q, and analyzed the single nucleotide polymorphism (SNP) subclades Q1a1a1-M120, Q1a2a1-L54, Q1a1b-M25, Q1a2-M346, Q1a2a1a2-L804, Q1a2b2-F1161, Q1b1a-M378, and Q1b1a1-L245. Through NETWORK and BATWING analyses, we found that the subclades of haplogroup Q continued to disperse from Central Asia and Southern Siberia during the past 10,000 years. Apart from its migration through the Beringia to the Americas, haplogroup Q also moved from Asia to the south and to the west during the Neolithic period, and subsequently to the whole of Eurasia and part of Africa.

Huang, YZ., Pamjav, H., Flegontov, P. et al., Dispersals of the Siberian Y-chromosome haplogroup Q in Eurasia, Mol Genet Genomics (2017). https://doi.org/10.1007/s00438-017-1363-8

See also...

Phylogeography of Y-haplogroup Q3-L275

Banned commentators list

People listed below aren't allowed to make posts in the comments section at this blog. If they do, their posts will be removed as soon as I see them. Moderation will be turned on when necessary to block banished commentators from returning.

Alberto

batman

Blogger

blogmaster

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Bronze

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German Dziebel

gimby20

Hayk

Helios

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Jaakko

James

Milan

Mr Snow

natsunoame

Nirjhar007

Richard Holtman

Shahanshah of Persia/Kanishka

Vincent

xyyman

I'll be updating the list regularly as more people are banned, which is highly likely. To make sure that you don't make it onto the list, think twice before you post a comment. Follow the blog rules [see HERE] and make an effort to understand the basics of what is being discussed before you join the discussion.

If, perhaps, you're frustrated that your pet theory isn't working out, then come up with a better pet theory. But whatever you do, don't troll here. Also please note, discussions with banned commentators are forbidden, and those who break this rule will also be banned.

Female mobility and exogamy as the main drivers of foreign admixture during the Late Neolithic/Early Bronze Age shift in Central Europe (Knipper et al. 2017)

The paper is still embargoed. I'll update this post after I've read it. The press release is here. Who wants to bet that this is exactly what happened on the Eneolithic/Early Bronze Age steppe?

Corina Knipper et al., Female exogamy and gene pool diversification at the transition from the Final Neolithic to the Early Bronze Age in central Europe, PNAS (2017). www.pnas.org/cgi/doi/10.1073/pnas.1706355114

Update 05/0/2017: I just had a quick look at the paper. It's not as comprehensive as I had hoped, because it lacks Y-chromosome and genome-wide data. But anyway, the authors do make a strong case for male patrilocality and abundant female exogamy at the sites in question. Below is the paper abstract. Emphasis is mine:

Human mobility has been vigorously debated as a key factor for the spread of bronze technology and profound changes in burial practices as well as material culture in central Europe at the transition from the Neolithic to the Bronze Age. However, the relevance of individual residential changes and their importance among specific age and sex groups are still poorly understood. Here, we present ancient DNA analysis, stable isotope data of oxygen, and radiogenic isotope ratios of strontium for 84 radiocarbon-dated skeletons from seven archaeological sites of the Late Neolithic Bell Beaker Complex and the Early Bronze Age from the Lech River valley in southern Bavaria, Germany. Complete mitochondrial genomes documented a diversification of maternal lineages over time. The isotope ratios disclosed the majority of the females to be nonlocal, while this is the case for only a few males and subadults. Most nonlocal females arrived in the study area as adults, but we do not detect their offspring among the sampled individuals. The striking patterns of patrilocality and female exogamy prevailed over at least 800 y between about 2500 and 1700 BC. The persisting residential rules and even a direct kinship relation across the transition from the Neolithic to the Bronze Age add to the archaeological evidence of continuing traditions from the Bell Beaker Complex to the Early Bronze Age. The results also attest to female mobility as a driving force for regional and supraregional communication and exchange at the dawn of the European metal ages.

See also...

A plausible model for the formation of the Yamnaya genotype

Corded Ware women more mobile than their men (Sjögren et al. 2016)

R1b-V88: out of the Balkans and into Africa?

Late last year I tentatively suggested that R1b-PF6279/V88, also known as R1b1a2, and formerly as R1b1c, may have entered Africa from Iberia, rather than from the Near East as is generally accepted, because of its presence in an Early Neolithic sample from Iberia (see here).

This is now looking a lot more plausible due to the recent discovery that the eastern Balkans was home to Mesolithic foragers belonging to R1b-PF6279/V88 (see page 122 here and discussion here) and indeed a pre-Neolithic R1b hotspot (see Mathieson et al. 2017 and González-Fortes et al. 2017). Note also the continued absence of R1b in the growing selection of prehistoric samples from Anatolia and the Levant.

So here's a theory that I think is worth considering for the time being: R1b-V88, and perhaps even R1b, originated in an as yet unsampled Balkan population, dating to the Upper Paleolithic and ancestral to the so called Villabruna cluster (see here), that eventually contributed ancestry to all present-day Europeans, as well as many present-day Asians and, via an Iberian route, Africans. Is there any archaeological evidence for the existence of such an Upper Paleolithic group in the Balkans?

The pseudo-steppe theory: last line of defense against the inevitable

A popular tactic used by those none too pleased with the presence of Bronze Age steppe or Yamnaya-related ancestry in South Asians is to claim that this ancestry isn't actually from the steppe. Thus, they call it "pseudo-steppe" ancestry.

This theory is based on the assumption that a population very similar to Yamnaya formed independently in South Asia, or at least as part of a cline of such ancestry running from South Asia to the Eastern European steppe.

At best, this is highly speculative and, at worst, insane; a last ditch attempt to counter mounting genetic evidence backing the Aryan Invasion Theory (AIT), or, if you're politically correct, Aryan Migration Theory (AMT). But it gets traction for the time being, including, unfortunately, among some Indian scientists (see here), simply because of the lack of ancient DNA data points for Central and South Asia.

However, we do have Iran_HotuIIIb, a Mesolithic or Neolithic forager from the Hotu Cave near the southeastern coast of the Caspian Sea in modern-day Iran. This location is not in Central Asia per se, but pretty damn close by anyone's standards, because it's next door to the Turkmenistan border.

Below is a map that I put together to illustrate the absurdity of the pseudo-steppe theory. Note that Iran_HotuIIIb is located between Yamnaya and present-day Brahmins from northern India, so if you're a proponent of the pseudo-steppe theory you'd expect him to pack quite a bit of Yamnaya-like ancestry, right? Well, he shows a slither, but much less than the Brahmins. How is this possible geographically and temporally, without a migration from the steppe to India some time after Iran_HotuIIIb was alive? Realistically, it's not.

The ancestry proportions on the map are based on an easily reproducible ADMIXTURE analysis. The test was run in unsupervised mode, but I designed the dataset to help the algorithm flesh out the so called Early Bronze Age steppe (Steppe_EBA) component. The full Q output is available in this spreadsheet.

In fact, there are several ways to show that present-day South Asians share relatively recent and direct ancestry with Bronze Age Eastern Europeans, including, for example, with a haplotype test (see here). But I'd say that a well designed ADMIXTURE analysis is an especially effective way of doing it, because both a strength and weakness of ADMIXTURE is that it's sensitive to ethnic-specific genetic drift. Thus, the so called ancestral populations that it infers are often just exaggerated signals of relative inbreeding, isolation, and/or rapid expansions experienced by founders of ethnic groups.

Clearly, my Steppe_EBA component is a signal of a relatively small, young founder population expanding rapidly across and out of the Eurasian steppe during the Bronze Age. That's because it peaks at extreme levels in genetically and archaeologically closely related Bronze Age steppe populations, such as Afanasievo, Poltavka and Yamnaya, and their recent descendants, such as Andronovo, Corded Ware and Srubnaya. In other words, ADMIXTURE has probably managed to pick up the genetic signal of a Bronze Age ethnic group.

This signal also shows up at relatively high frequencies in many geographically disparate Indo-European speakers, such as the Indian Brahmins and Icelanders. That's obviously good news for those of us who favor the Eastern European steppe as the Proto-Indo-European homeland.

Among the other ancients, those that plausibly can't have ancestry from the EBA steppe because they're too old, it's only seen at high levels in such populations as Eastern European Hunter-Gatherers (EHG), Caucasus Hunter-Gatherers (CHG), and Ancient North Eurasians (ANE). In other words, populations that contributed in a big way to the formation of the Bronze Age peoples of the Eastern European steppe.

It's important to understand also that my ADMIXTURE analysis isn't anything particularly original. Iran_HotuIIIb shows essentially the same behavior in this standard Principal Component Analysis (PCA) of West Eurasian populations, sitting just south of present-day South Central Asians (SC_Asia). This, of course, is out of whack with geography, and, again, can only be explained by significant gene flow from the north into South Central Asia after Iran_HotuIIIb's time.

See also...

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

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

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

The focus turns to the Kazakh steppe

As we wait for more ancient data to be released, here are a couple of interesting quotes from an interview with Harvard's David Reich at Lapham's Quarterly (for the whole thing see here). Emphasis is mine:

I spent the morning corresponding with an archaeologist in Kazakhstan. We’re trying to do genetics in the Bronze Age and the pre–Bronze Age variants in Kazakhstan.

...

We’d like to characterize the genetic variation in Kazakhstan because it’s a key connection point between the regions south of the Hindu Kush mountains, Iran and India in the south, the steppes in Russia in the north, in China in the East; it’s all mixing up there. And we’re trying to understand the connections. It’s a place of movement and migration, and understanding the genetics is interesting. So that was my morning, and then I spent some hours talking with colleagues today in the laboratory about ancient farmers of Israel and Iran that we have some data from.

It looks like this interview was done a while ago, because he's talking about samples from Iran and Israel that have already been published, so I'd say the Kazakhstan data will be part of the upcoming ancient DNA paper on South Asia.

See also...

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?

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

Genetic and archaeological continuity from Khvalynsk to Yamnaya

Over a year ago, using the D-stats/nMonte method of mixture modeling (see here), I noticed that Yamnaya did not appear to be simply a two-way mixture between Eastern European and Caucasus Hunter-Gatherers (EHG and CHG, respectively), but the result of a much more complex process:

Using the most plausible reference samples currently available - almost all of them older than Yamnaya, and thus unlikely to skew the results with Yamnaya admixture - reveals the following models for the two Yamnaya sets from Kalmykia and Samara, respectively.

Yamnaya_Kalmykia
Khvalynsk 57.7
Kotias 28.3
Hungary_EN 12.9
Ulchi 1.1
AfontovaGora3 0
Anatolia_Neolithic 0
Karelia_HG 0
Loschbour 0
MA1 0
Motala_HG 0

distance%=1.9125 / distance=0.019125

Yamnaya_Samara
Khvalynsk 56.75
Kotias 26.4
Hungary_EN 10.85
Karelia_HG 4.4
Loschbour 1.6
AfontovaGora3 0
Anatolia_Neolithic 0
MA1 0
Motala_HG 0
Ulchi 0

distance%=2.1354 / distance=0.021354

Very interesting but hardly surprising. Essentially what we're seeing there is potentially very strong genetic continuity from the Eneolithic to the Early Bronze Age on the Pontic-Caspian Steppe. In other words, from Khvalynsk to Yamnaya.

However, at some point between the Eneolithic and the Early Bronze Age, the steppes saw a major influx of extra CHG, represented by the ~27% of Kotias-related admixture. Considering the relevant uniparental data, with lots of Y-HG R1b and no Y-HG J among Yamnaya males, I'd say this CHG came with women.

Also, the relatively high admixture related to early Hungarian Plain farmers (Hungary EN) is a fairly curious detail that has not been reported before. If real, it probably represents gene flow from the Neolithic and/or Chalcolithic Balkans to the Pontic-Caspian Steppe. Again, in all likelihood it mostly came with women, perhaps from Tripolye-Cucuteni and/or Varna communities.

The reason I mention this now is because I can reproduce basically the same model using the updated qpAdm methodology described recently in Lazaridis et al. 2017, which relies on a relatively large number (≥16) of ancient genomes/populations as outgroups (see here), and, in my experience, causes many formerly successful models to fail miserably (the taildiff dives from >0.05 to <0.05). Note that in my dataset Khvalynsk is now labeled Samara_Eneolithic, Kotias as CHG, and Hungary_EN as Hungary_N.

Yamnaya_Kalmykia
CHG 0.334±0.044
Hungary_N 0.115±0.031
Samara_Eneolithic 0.550±0.032
taildiff: 0.419775785
chisq: 13.368
Full output

Yamnaya_Samara
CHG 0.267±0.040
Hungary_N 0.130±0.027
Samara_Eneolithic 0.603±0.030
taildiff: 0.300777879
chisq: 15.106
Full output

Here's a formerly successful model in which Steppe_EMBA (a grouping which includes Afanasievo, Poltavka, Russia_EBA and Yamnaya) is posited as a mixture between EHG and Chalcolithic farmers from the Zagros Mountains in what is now Iran. It clearly fails when I use CHG as one of the outgroups.

Steppe_EMBA
EHG 0.544±0.020
Iran_ChL 0.456±0.020
taildiff: 0.00279643007
chisq: 31.553
Full output

vs.

Steppe_EMBA
CHG 0.310±0.034
Hungary_N 0.121±0.023
Samara_Eneolithic 0.568±0.025
taildiff: 0.50194795
chisq: 12.316
Full output

Now, tight statistical fits are great, but they don't always reflect reality, especially when fine scale genetic structure is being tested. So does my model have any support from archeology? In other words, does archaeological data show continuity between Khvalynsk and Yamnaya (Pit-Grave culture)? According to Morgunova and Turetskij 2016 it does. Emphasis is mine:

Abstract: The aim of the paper is to provide the research results concerning the Pit-Grave culture sites of the south Ural region, which is a part of the Volga-Ural interfluve. The Pit-Grave culture developed mostly out of the Khvalynsk Eneolithic culture at the turn of the 5th–4th millennium cal BC. People of the Sredny Stog and forest-steppe Eneolithic cultures from the Middle Volga region also influenced the Pit-Grave culture. The paper considers the radiocarbon data (more than 120 dates), specifies the periodization of the Pit-Grave culture of the Volga-Ural interfluve, singles out the three stages of its development. The chronology of the culture is determined 3900–2300 cal BC. The authors provide new information about the Pit-Grave economy. Paleopedology, palynology, anthropology, metallography, ceramic technical, and technological analyses were used together with archaeological methods to make a more detailed description of the culture.

...

A number of steppe Eneolithic features remained at the Repin stage. The cultural continuity between the Pit-Grave, Khvalynsk, and Sredny Stog Eneolithic cultures was proved by the following features: skeletons in crouched supine position with bent legs to the left or to the right, heads at the eastern sector of burials, ochre coverage with high or low density, multiple burials, egg-shaped ceramics with neck and crushed shell impurity. Technical and technological analysis of pottery was another evidence demonstrating the pottery continuity between the Khvalynsk and Repin traditions (Vasilyeva 2002; Salugina 2005). Big soil burial grounds were substituted by individual burials under the barrow. The spread of local production copper articles was a distinctive feature of the Pit-Grave culture. This was the phenomenon, which archaeologists consider to be the beginning of the Early Bronze Age in steppe of Eastern Europe.

Morgunova N. and Turetskij M., Archaeological and natural scientific studies of Pit-Grave culture barrows in the Volga-Ural interfluve, Estonian Journal of Archaeology, Vol. 20, Issue 2, doi: 10.3176/arch.2016.2.02

See also...

Another look at the genetic structure of Yamnaya

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 (see here).

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...

A homeland, but not the homeland #3

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