Steppe Eneolithic and the Origins of Yamnaya Ancestry

Steppe Eneolithic and the Origins of Yamnaya Ancestry

In 2019, Wang et al. published three ancient DNA samples from the northern Caucasus foothill steppe region. These samples were labeled Steppe Eneolithic. In genetic studies, the term “Steppe” is usually used to describe DNA profiles related or similar to the Yamnaya genetic profile.

These Steppe Eneolithic samples were remarkable because they already displayed a genetic profile similar to Yamnaya, while being older than the Yamnaya culture itself. At the time, however, they did not receive the attention they deserved, largely because they were not associated with a well-known archaeological culture.

In 2024, Lazaridis et al. proposed a new term for this population: the Caucasus–Lower Volga (CLV) cline, and added additional samples with similar genetic profiles. Genetic modeling showed that Yamnaya populations derived most of their genome-wide ancestry from these CLV groups.

Despite this important progress, two crucial Y-DNA markers were still not securely linked to the CLV population: R1b-M269 and R1a-M417, the two major lineages that later expanded widely across Eurasia and are often associated with the spread of Indo-European languages.

New Evidence

The study by Ghalichi et al. (2024) helped fill this gap. Two new cases of R1b-M269 were discovered north of the Caucasus, predating the Yamnaya culture. These individuals had ancestry similar to other Steppe Eneolithic groups and were dated to approximately 3800 BCE.

It is worth noting that an older R1b-M269 sample (I33307) from Kalmykia, dated to around 3700 BCE, had already been reported. However, its genetic profile is identical to Core Yamnaya, which raises the possibility that the radiocarbon date may be somewhat inaccurate. In Lazaridis et al., this sample was classified as part of the Russia_CaspianInland_EBA_Yamnaya group, which is why it is not included in the list of Steppe Eneolithic Y-DNA samples (4700–3000 BCE).
(See the map for the geographic area.)

Steppe Eneolithic Y-DNA Distribution (4700–3000 BCE)

• R1b-V1636 — 9 samples

• R1b-M269 — 2 samples

• R1a — 1 sample

• I2-L699 — 1 sample

• J2a — 1 sample

• J2b2a — 1 sample

Another possible early R1b-M269 case (around 4500 BCE) was discovered in a Varna culture burial in Bulgaria. Its autosomal profile clearly shows Steppe Eneolithic–related ancestry, indicating that it was not related to local Neolithic farmers but rather represents a migrant from the Caucasus–Lower Volga region.

The Varna culture itself was unusual, characterized by a highly hierarchical social structure. Its graves contained large quantities of gold artifacts, among the earliest known examples of gold metallurgy.

The Role of J2b2a-L283

Another important finding in the Steppe Eneolithic context is J2b2a-L283. Its autosomal profile shows a stronger Caucasus-related component, which is understandable. Nevertheless, it clearly belonged to the emerging Eneolithic kurgan populations, from which both the Yamnaya culture and the Corded Ware culture would develop several centuries later.

The Question of R1a-M417

So far, R1a-M417 has not yet been identified in the Steppe Eneolithic / CLV context. However, one of the oldest R1a-M198 samples (closely related to M417) was found in the Middle Volga region, suggesting that M417 may eventually be discovered within the CLV or Steppe Eneolithic populations as well. Currently, one of the oldest known R1a-M417 individuals comes from a Yamnaya layer in the Balkans.

Implications for Yamnaya Origins

Based on these findings, I remain skeptical of the hypothesis that Yamnaya originated directly from the Sredny Stog culture in Ukraine, as sometimes suggested in discussions.

The highest diversity of R1b-Z2103 lineages that are positive for Z2103 but negative for Z2106 is found in the Near East, suggesting that the region where Z2103 began its expansion was likely close to the Caucasus. Additional ancient DNA samples will hopefully clarify this question in the future.

Note

The Eneolithic Y-DNA list above does not include samples from Steppe Maykop, which have a different origin related to Central Asian ancestry.

In addition, one R1b-V1636 sample is dated to around 2800 BCE, slightly later than the 3000 BCE cutoff used here. However, its autosomal profile indicates that it belonged to the preceding Steppe Eneolithic population, rather than to Core Yamnaya.

See also

• Qpadm models about the Core Yamnaya origins.
• The Yamnaya culture and it's legacy.

Qpadm models about the Core Yamnaya origins.

We have one sample from north Caucasus PG2004 which is similar to so called BP group of Caucasus Lower Volga cline.

Modeling Yamnaya as a three-way mixture of CLV_BP, Aknashen and Ukraine Neolithic hunter gatherers was very easy, as in the paper.

BP group 66.5%

Aknashen 19.5%

UNHG. 14.0%

p value 0.17

It didn't require from me any extra effort to find the correct "settings" (outgroups). Adding western Ukraine Trypllia farmers didn't made the model better. Aknashen was still wanted while Trypllia not. I tried various ways to make Trypllia wanted, but it didn't work. Finally, I removed Aknashen and forced the model to rely solely in Trypllia. I got a p value lower than 0.05. So practically a failure. Nikitin 2024 dedicates a special chapter to this subject. Their conclusion is that Yamnaya probably do have some Trypllia but it's very low.

My conclusions are the same. Aknashen ancestry in Yamnaya is real. It is supported by Y DNA of preceding period, and I am sure that the scrupulous analysis of mtDNA will show the same result. Finally, if I had access to the Nalchik farmer data then the percentage would be twice higher.

Then I tried the same in G25 and here a surprise. The raw unscaled models behaved in the same manner as qpadm albeit with a different proportion. But the scaled G25 showed a clear a preference for Trypllia. I said in this group many times that tools and settings matters. And this an excellent example why the use of scaled models can lead to misleading results. I want to remind that scaled coordinates are artificially altered numbers.

Keeping this in mind let's see what means CLV-steppe ancestry in Bronze Age Anatolia that Lazaridis 2024 supposedly has found but Lazaridis 2022 didn't detected. I don't have Cayonu files to reproduce their models in qpadm but what we know from G25 behavior is that:

+ When Anatolia N and CHG are used in models as source they mask the steppe ancestry and show a lower number. This was done in Lazaridis 2022 which not only didn't find any steppe in BA Anatolia but even in Van Urartu.

+ When a Mesopotamian or Levantine and Iran Neo are used as sources and CHG and Anatolia_N are ommitted then this exaggerates the steppe ancestry in northwest Asia. In Lazaridis 2024 they used Cayonu Neolithic which was a north Mesopotamian population. This exaggerated the steppe ancestry in Bronze Age Anatolia where Hittite lived. If the same Cayonu was used for Minoans, they would find steppe even in Minoans and probably Alalakh Hurrians and Semites also. Which doesn't make sense. Uniparental markers do NOT support such high level of CLV-steppe ancestry in BA Anatolia. Quite contrary they speak about very low or virtual absence of it. Just one R1b-V1636 in the midst of more than dozen local haplotypes.

My conclusions are that based on this genetic data it is not possible to consider IE homeland issue fully solved. We are very close it. And the broad picture. But the exact details are not still there, and linguistics also can be helpful.

What is needed now is to have more than 50 Y DNA from Bronze Age Anatolia. Also, a large number of Y DNA is needed from South Caucasus and historic Armenia LC-EBA period to see what happened to those Late Chalcolithic migrants from north. How much they left an impact and why their autosomes were diluted after the Areni C. It is also important to have samples from LC kurganic burials like Aknalich and Soyuq Bulaq.

See also

• When the tools and settings matters. Part 1 and Part 2

The Kurgan builders.

The Caucasus Lower Volga cline (CLV) having both south Caucasian and Eastern European foragers ancestry is a genetic term. The material cultures behind this term were variable but they had one important common feature. Virtually all the samples from CLV cline were found from kurgans. Kurgan is an artificial mound, a tumulus on top of the grave. A kurgan could harbor a single or multiple graves. Given that it requires a lot of manpower it was usually built for elite persons. Smaller kurgans also existed. Other prominent features of kurganic burials in Pontic Caspian steppe were the red ochre, the raised knee position etc. The origins of this tradition are uncertain, and the dates of the oldest kurgans debated but what is well known now that it expanded with Eneolithic (is equal to Chalcolithic) pastoralists (4500-3500bc) who had the CLV ancestry, replacing older local hunter's flat grave tradition. Besides kurgans, we know that those people had a patrilocal and exogamic culture.

Kurganic people moved to north, toward middle Volga region and the Khvalynsk culture (after 4500bc) emerged there. In middle Volga region various branches of haplogroup Q1 were integrated into early steppe pastoralist communities. Q-L939 the branch of Georgian Bagrationi. Q-YP1669, probably Q-F26062 and some others to be found.

They also moved toward the Balkanian peninsula where they mixed with local European farmers where various kurganic cultures emerged (Cernavoda, Usatovo, Suvorovo). This migration has been proposed to be the source of Anatolian languages. Currently there is no evidence that they reached deep into Anatolia. Nevertheless it's possible that some obscure ethnicities like Mysians in Turkish Thrace and north west Anatolia were derived from those migrants. Although a later Yamnayan origin is also possible for them. Yamnayans themselves also had kurgans.

The new "finding" in Lazaridis 2024 paper is about the migration of those kurganic CLV people to south Caucasus. I use brackets because the presence of steppe ancestry in Areni cave was known since 2016. But only now it got an interpretation. This migration apparently occurred at 4300BC a period known as late Chalcolithic which is associated with Chaff faced ware. Chaff ware was not from steppe but had a local Neolithic origin. A syncretic culture emerged. A variant of which is known as Leila tepe culture in what is now Azerbaijan. We have already seen that those chaff groups moved to west toward Anatolia. The R1b-V1636 is associated with this event. We have now three cases of V1636 stretched from Aintab to Sevan basin.

The new proposal of Lazaridis is that they were the IE Anatolians and based on this they propose that the CLV is the place were Indo-Europeans emerged. This is the main difference of this paper from the Lazaridis 2022 in which they placed the homeland of PIE ( or Indo-Anatolians ) in south of Caucasus.

My next post will be about the Indo European origins based on the linguistics and how those known migrations fits into the great picture. Also I will discuss the real number of steppe ancestry in Bronze Age Anatolia and will show that the current data size is very small for having a good understanding of the situation.

** The first picture is a reconstruction of a kurgan .They are usually eroded over time. This one reconstructed. The second picture is the tumulus of Lydian king Alyattes. This is one of the largest known kurgans.

The Yamnaya culture and it's legacy

Yamnaya was an Early Bronze Age culture (3300-2700BC) that expanded in Pontic Caspian steppe in East Europe. It is remarkable due to its homogenous population which contrasts to the diversity of the preceding Eneolithic (means Chalcolithic) cultures. In just 1000 years genes associated with Yamnaya culture reached what is now Portugal and western Mongolia, Britain and South-Central Asia. The scale of this expansion can be considered as one of the most important demographic events in West Eurasia after the invention of farming in Fertile Crescent. In 2022 Lazaridis et al. announced that all modern living IE nations languages can be derived from the Yamnaya.

In 2015 the first ancient DNA of Yamnayans was published and it became clear that they do not directly descend from Eastern European hunter gatherers. Roughly the half of their ancestry was from south, and the "Armenian like" term was used to label this southern part. Then in late 2015 the Caucasian hunter gatherers (CHG) were discovered. In 2016 Iranian Zagros Neolithic farmers were published and since then both those populations were used to model the Yamnayans.

Despite those statistical models the formation of Yamnayans gene pool was remaining an obscure subject. Finally in April 2024 two large papers dedicated to this subject provided more clarity. Various Neolithic and Eneolithic groups living in East Europe had both shared ancestry and differences. One group who was living from North Caucasus foothills to Lower Volga (site Berezhnovka ) was labeled as "Caucasus Lower Volga cline". CLV had both southern Caucasian hunters and farmers (Aknashen) ancestry.

I will dedicate a special topic to describe what is CLV but at this stage it's sufficient to say that Yamnayans had up to 80% of their ancestry from CLV. Not only Yamnayans but all Kurganic cultures (Sredni Stog, Khvalynsk, Cernabida, Usatovo) of eastern Europe and Balkans had the CLV ancestry mixed with various local European populations. For the case of Yamnayans the second part of their ancestry was the Ukraine hunter gatherers (UNHG), which made them similar to Sredni Stog. Even though the most important Y DNA's were missing in Sredni Stog which could mean that Proto Yamnaya's exact immediate geographic location is not found yet but shouldn't be outside from Don - Volga - Caucasus triangle. With more sampling we will learn the exact origin of the two most successful lineages. R1b-M269 and R1a-M417.

The samples from the new paper are not available yet but we have two samples from the forementioned CLV cline. They are from the Progress - 2 site in the North Caucasus (4500-4000BC, see the last map for the location). I used them to model various Yamnaya samples and got the same result as in Lazaridis 2024. More than 75-80% of ancestry in regular Yamnaya samples is from the Progress / CLV. Some outliers have extra ancestries which is normal given their location.

You may wonder how "Hittites from east, from historic Armenia" and "Yamnaya from CLV" are connected to each other? After all there is no evidence that Hittites and other Anatolian speakers descend from Yamnaya culture. What is the common shared ancestry between IE Anatolians and Yamnaya?

Another topic will be dedicated to this subject.

And at the end here is the list of Y DNA found in Yamnaya at this stage. R1b-Z2103 (west Asia Balkans), R1b-L51 (west Europe), R1a-M417 (Asia - Europe), I2-L699, J2b-L283 (west Balkans), R1b-PF7562 (Balkans Mycenean ) Q-L939 and isolated cases of J1 and R1b-V1636.

See also

• Population dynamics in North Caucasus

Hittites from east.

Before commenting the deeper origins of Indo Europeans proposed in the new Lazaridis 2024 paper I want to speak about the origins of Anatolians. Since the advent of paleogenetic it became clear that they can't be native to plain Anatolia. They could be either from east or from north-west. Since 2018 the accumulating data was giving clear preference to the eastern origins. From historic Armenia. But a doubt was still persisting, because the Balkanian route was not fully investigated. In 2023 Penske et al. has found a migration from steppe to eastern Balkans before Yamna around 4000BC but they didn't even discuss the possibility that they could be related to Hittites. It's because their autosomes were rapidly diluted having a limited impact in Bulgaria yet alone in Anatolia. Also the Y DNA associated to this migration the I2-L699 was missing from inland ancient Anatolian samples.

While a R1b-V1636 was found in Bronze Age Gaziantep quite close temporaly and geographically to the kingdom Armi mentioned in Eblaite texts around 2500bc. Hittites names have been discovered in this kingdom recently.

The V1636 was popular in CLV cline stretching from Caucasus to Lower Volga and was found in ancient Malatya and Sevan basin also.

Besides this R1b other haplotypes found in Bronze Age Anatolia also show self-evident eastern affiliation.

The J1-Z1828 who's subbranch BY94 can be an Iron Age Luwian lineage like the J2- L70 which itself is derived from the L25 a haplotype from east.

A J2-Z6065 which was found in Masis Blue in Neolithic Armenia. We have also bunch of of J2-M67 and G2-M406, even though this latter is related to Hattic people so it can't be interpreted as a prove of Hittites from east.

In conclusion all ancient Y DNA that we have in inland Anatolia do have an eastern origin, from historic Armenia and around. One of them is further from north of Caucasus region. Their autosomes are also shifted to east compared to the preceding periods.

One exception is the I2-P78 from MBA Yassi Tepe near Izmir, which is clearly from Mycenean Greece, so this person was almost certainly a Greek people representative who probably settled in western Anatolian coasts after the 2000BC. They are mentioned in Hittite texts as Ahhiyawa.

In sum we can now say with high certitude that Indo-European languages were spoken in Armenian Highlands at minimum the last 6300 years. The 4300BC is not a random number it's the date of the start of Late Chalcolithic associated with Chaff faced ware which had a tremendous impact on southern Anatolia. But not much on northern Anatolia where Hattic related people were settled since more earlier times. Probably since 5800BC.

But that would require another thread which I will do the next time.

PS a proposed location of Armi. Another location is in the northeast of Ebla.

See also

• One possible way to know the language(s) spoken in Kur-Araxian culture is to examine the onomastics in those places where there was continuity

The dramatic expansion of Yamna is related to the introducing of food producing in Steppe

The dramatic expansion of Yamnaya is related to the introducing of food producing in Steppe. Suddenly large grasslands became pastures for herding, which itself resulted in dramatic increase of population.

Farming was invented in Near East, Fertile Crescent. See in maps. This period is known as Neolithic and starts at 9500 BC. The oldest evidence of crop raising is found in Diyarbekir region of Turkey. Farmers moved to different directions. They had diverse haplogroups and almost certainly were speaking different languages.

 Levantine farmers moved to Africa. Farmers from Anatolia moved to Europe. Iranian farmers moved to Central Asia and India (this latter is disputed). And farmers from historic Armenia moved to Caucasus and then farming in the form of pastoralism appear in Steppe, North of Caucasus.

For IE puzzle there are two main options. 

1. PIE was the language of farmers living in historic Armenia who moved to north mixed with locales and expanded there as Yamna.

2. PIE was the language of Steppic hunters who learned farming (borrowed terminology) and metallurgy from Near Eastern farmers. And then expanded. 

There is a third option also that farmers of west Ukraine who themselves were from Anatolia affected Steppe. But this option is not anymore popular after the paleogenetic findings.

In both cases we would have Near Eastern terminology in IE. Some examples.

• gud in Sumerian means ox, bull and we have kov, cow in IE languages. *gʷṓws in PIE
• hawirni means lamb in Hurrian, *Hovi in PIE sheep
• *tawr in Semitic and *táwros in PIE. 
• urud in Sumerian means copper, and *h₁rewdʰ is the red color in PIE.

Also, numbers like six, seven and word wine but this latter has been proposed to be from IE to Kartvelian.