I basically added information from the latest papers published, which (luckily enough for me) haven’t been too many, and I have added images to illustrate certain sections.
I have updated the PCAs by including North Caucasus samples from Wang et al. (2018), whose position I could only infer for older versions from previously published PCA graphs.
I have also added to the supplementary materials the “Tip of the Iceberg” R1b tree by Mike Walsh from the FTDNA R1b group, with permission, because some relevant genetic sections are centered on the evolution of R1b lineages, and the reader can get easily lost with so many subclades.
I have also updated maps, including some of the Y-DNA ones, and managed to finish two new maps I was working on, and I added them to the supplementary materials and to the menu above:
It is tentative because there hasn’t been any professional study or amateur attempt to date to differentiate both “steppe ancestries” in Yamna, and especially in Bell Beakers. So much for the call of professional geneticists since 2018 (see here and here) and archaeologists since 2017 (see e.g. here and here) to distinguish fine-scale population structure to be able to follow neighbouring populations which expanded with different archaeological (and thus ethnolinguistic) groups.
I think both maps are especially important today, given the current Nordicist reactionary trends arguing (yet again) for an origin of Indo-Europeans in The North™, now based on the Fearsome Tisza River hypothesis, on cephalic index values, and a few pairwise comparisons – i.e. an absolutely no-nonsense approach to the Indo-European question (LOL). At least I get to relax and sit this year out just observing how other people bury themselves and their beloved “steppe ancestry=IE” under so many new pet theories…
NOTE. Not that there is anything wrong with a northern origin of North-West Indo-European from a linguistic point of view, as I commented recently – after all, a Corded Ware origin would roughly fit the linguistic guesstimates, unlike the proposed ancestral origins in Anatolia or India. The problem is that, like many other fringe theories, it is today just based on tradition, or (even worse) ethnic, political, or personal desires, and it doesn’t make sense when all findings from disciplines involved in the Indo-European and Uralic questions are combined.
Within 20 or 30 years, when genetic genealogists (or amateur geneticists, or however you want to call them) ask why we had the opportunity since 2015 to sample as many Hungarian Yamnaya individuals as possible and we didn’t, when it is clear that the number of unscathed kurgans is diminishing every year (from an estimated 4,000 in the 20th century, of the original tens of thousands, to less than 1,500 today) the answer will not be “because this or that archaeologist or linguist was a dilettante or a charlatan‘, as they usually describe academics they dislike.
It will be precisely because the very same genetic genealogists – supposedly interested today in the origin of R1b-L151 and/or genetic marker associated with North-West Indo-Europeans – are obsessed with finding them anywhere else but for Hungary, and prefer to use their money and time to play with a few statistical tools within a biased framework of flawed assumptions and study designs, obtaining absurd results and accepting far-fetched interpretations of them, to be told exactly what they want to hear: be it the Franco-Cantabrian homeland, the Dutch or Moravian Beaker from CWC homeland, the Maykop homeland, or the Moon homeland.
Poetic justice this heritage destruction, whose indirect causes will remain written in Internet archives for everyone to see, as a good lesson for future generations.
Given my reduced free time in these months, I have decided to keep updating the text on Indo-European and Uralic migrations and/or this blog, simultaneously or alternatively, to make the most out of the time I can dedicate to this. I will add the different ‘A Song of Sheep and Horses (ASoSaH) reread’ posts to the original post announcing the books. I would be especially interested in comments and corrections to the book chapters rather than the posts, but any comments are welcome (including in the forum, where comments are more likely to stick).
Luckily enough – for those of us who want precise answers to our previous infinite models of Indo-European language expansions (viz. GAC-associated expansion, IE-speaking Old Europe, Anatolian homeland, Iran homeland, Maykop as Proto-Anatolian, Palaeolithic Continuity Theory, Celtic in the Atlantic façade, etc.) – the situation has been more clear-cut than expected: it turns out that, especially during population expansions, acute Y-chromosome bottlenecks were very common in the past, at least until the Iron Age.
Khvalynsk and Repin-Yamna expansions were no different, and that seems quite natural in hindsight, given the strong familial ties and aversion to foreigners proper of the Late Proto-Indo-European society and culture – probably not really that different from other contemporary societies, like the neighbouring Late Proto-Uralic or Trypillian ones.
During the expansion of early Khvalynsk, the most likely Indo-Anatolian culture, the society of the Don-Volga area was probably made up of different lineages including R1b-V1636, R1b-M269, R1a-YP1272, Q1a-M25, and I2a-L699 (and possibly some R1b-V88?), a variability possibly greater than that of the contemporary north Pontic area, probably a sign of this region being a sink of different east and west migrations from steppe and forest areas.
During its expansion, the Khvalynsk society saw its haplogroup variability reduced, as evidenced by the succeeding expansive Repin culture:
Afanasevo, representing Pre-Tocharian (the earliest Late PIE dialect to branch off), expanded with R1b-L23 – especially R1b-Z2103 – lineages, while early Yamna expanded with R1b-L23 and I2a-L699 lineages, which suggests that these are the main haplogroups that survived the Y-DNA bottleneck undergone during the Khvalynsk expansion, and especially later during the late Repin expansion. Nevertheless, other old haplogroups might still pop up during the Repin and early Yamna period, such as the R1b-V1636 sample from Yamna in the Northern Caucasus.
It is still unclear if R1b-L23 sister clade R1b-PF7562 (formed ca. 4400 BC, TMRCA ca. 3400 BC), prevalent among modern Albanians, expanded with Yamna migrants, or if it was part of an earlier expansion of R1b-M269 into the Balkans, and represent thus Indo-Anatolian speakers who later hitchhiked the expansion of the Late PIE language from the north or west Pontic area. The early TMRCA seems to suggest an association with Repin (and therefore Yamna), rather than later movements in the Balkans.
‘Yamnaya’ or ‘steppe’ ancestry?
After the early years when population genetics relied mainly on modern Y-DNA haplogroups, geneticists and amateurs have been recently playing around with testing “ancestry percentages”, based on newly developed free statistical tools, which offer obviously just one among many types of data to achieve a proper interpretation of the past.
Today we have quite a lot Y-DNA haplogroups reported for ancient samples of more recent prehistoric periods, and they seem to offer (at least since the 2015 papers, but more evidently since the 2018 papers on Bell Beakers and Europeans, Corded Ware, or Fennoscandia among others) the most straightforward interpretation of all results published in population genomics research.
NOTE. The finding of a specific type of ancestry in one isolated 40,000-year-old sample from Tianyuan can offer very interesting information on potential population movements to the region. However, the identification of ethnolinguistic communities and their migrations among neighbouring groups in Neolithic or Bronze Age groups is evidently not that simple.
It is becoming more and more clear with each paper that the true “Yamnaya ancestry” – not the originally described one – was in fact associated with Indo-Europeans (see more on the very Yamnaya-like Yamna Hungary and early East Bell Beaker R1b samples, all of quite similar ancestry and PCA cluster before their further admixture with EEF- and CWC-like groups).
The so-called “steppe ancestry”, on the other hand, reflects the contribution of a Northern Caucasus-related ancestry to expanding Khvalynsk settlers, who spread through the steppes more than a thousand years before the expansion of Late Proto-Indo-Europeans with late Repin, and can thus be found among different groups related to the Pontic-Caspian steppes (see more on the emergence and evolution of “steppe ancestry”).
In fact, after the Yamna/Indo-European and Corded Ware/Uralic expansions, it is more likely to find “steppe ancestry” to the north and east in territories traditionally associated with Uralic languages, whereas to the south and west – i.e. in territories traditionally associated with Indo-European languages – it is more likely to find “EEF ancestry” with diminished “steppe ancestry”, among peoples patrilineally descended from Yamna settlers.
Y-DNA haplogroups, the only uniparental markers (see exceptions in mtDNA inheritance) – unlike ancestry percentages based on the comparison of a few samples and flawed study designs – do not admix, do not change, and therefore they do not lend themselves to infinite pet theories (see e.g. what David Reich has to say about R1b-P312 in Iberia directly derived from Yamna migrants in spite of their predominant EEF ancestry): their cultural continuity can only be challenged with carefully threaded linguistic, archaeological, and genetic data.
An interesting aspect of the paper, hidden among so many relevant details, is a clearer picture of how the so-called Yamnaya or steppe ancestry evolved from Samara hunter-gatherers to Yamna nomadic pastoralists, and how this ancestry appeared among Proto-Corded Ware populations.
Please note: arrows of “ancestry movement” in the following PCAs do not necessarily represent physical population movements, or even ethnolinguistic change. To avoid misinterpretations, I have depicted arrows with Y-DNA haplogroup migrations to represent the most likely true ethnolinguistic movements. Admixture graphics shown are from Wang et al. (2018), and also (the K12) from Mathieson et al. (2018).
1. Samara to Early Khvalynsk
The so-called steppe ancestry was born during the Khvalynsk expansion through the steppes, probably through exogamy of expanding elite clans (eventually all R1b-M269 lineages) originally of Samara_HG ancestry. The nearest group to the ANE-like ghost population with which Samara hunter-gatherers admixed is represented by the Steppe_Eneolithic / Steppe_Maykop cluster (from the Northern Caucasus Piedmont).
Steppe_Eneolithic samples, of R1b1 lineages, are probably expanded Khvalynsk peoples, showing thus a proximate ancestry of an Early Eneolithic ghost population of the Northern Caucasus. Steppe_Maykop samples represent a later replacement of this Steppe_Eneolithic population – and/or a similar population with further contribution of ANE-like ancestry – in the area some 1,000 years later.
This is what Steppe_Maykop looks like, different from Steppe_Eneolithic:
NOTE. This admixture shows how different Steppe_Maykop is from Steppe_Eneolithic, but in the different supervised ADMIXTURE graphics below Maykop_Eneolithic is roughly equivalent to Eneolithic_Steppe (see orange arrow in ADMIXTURE graphic above). This is useful for a simplified analysis, but actual differences between Khvalynsk, Sredni Stog, Afanasevo, Yamna and Corded Ware are probably underestimated in the analyses below, and will become clearer in the future when more ancestral hunter-gatherer populations are added to the analysis.
2. Early Khvalynsk expansion
We have direct data of Khvalynsk-Novodanilovka-like populations thanks to Khvalynsk and Steppe_Eneolithic samples (although I’ve used the latter above to represent the ghost Caucasus population with which Samara_HG admixed).
We also have indirect data. First, there is the PCA with outliers:
Second, we have data from north Pontic Ukraine_Eneolithic samples (see next section).
Third, there is the continuity of late Repin / Afanasevo with Steppe_Eneolithic (see below).
3. Proto-Corded Ware expansion
It is unclear if R1a-M459 subclades were continuously in the steppe and resurged after the Khvalynsk expansion, or (the most likely option) they came from the forested region of the Upper Dnieper area, possibly from previous expansions there with hunter-gatherer pottery.
Supporting the latter is the millennia-long continuity of R1b-V88 and I2a2 subclades in the north Pontic Mesolithic, Neolithic, and Early Eneolithic Sredni Stog culture, until ca. 4500 BC (and even later, during the second half).
Only at the end of the Early Eneolithic with the disappearance of Novodanilovka (and beginning of the steppe ‘hiatus’ of Rassamakin) is R1a to be found in Ukraine again (after disappearing from the record some 2,000 years earlier), related to complex population movements in the north Pontic area.
NOTE. In the PCA, a tentative position of Novodanilovka closer to Anatolia_Neolithic / Dzudzuana ancestry is selected, based on the apparent cline formed by Ukraine_Eneolithic samples, and on the position and ancestry of Sredni Stog, Yamna, and Corded Ware later. A good alternative would be to place Novodanilovka still closer to the Balkan outliers (i.e. Suvorovo), and a source closer to EHG as the ancestry driven by the migration of R1a-M417.
The first sample with steppe ancestry appears only after 4250 BC in the forest-steppe, centuries after the samples with steppe ancestry from the Northern Caucasus and the Balkans, which points to exogamy of expanding R1a-M417 lineages with the remnants of the Novodanilovka population.
4. Repin / Early Yamna expansion
We don’t have direct data on early Repin settlers. But we do have a very close representative: Afanasevo, a population we know comes directly from the Repin/late Khvalynsk expansion ca. 3500/3300 BC (just before the emergence of Early Yamna), and which shows fully Steppe_Eneolithic-like ancestry.
Compared to this eastern Repin expansion that gave Afanasevo, the late Repin expansion to the west ca. 3300 BC that gave rise to the Yamna culture was one of colonization, evidenced by the admixture with north Pontic (Sredni Stog-like) populations, no doubt through exogamy:
This admixture is also found (in lesser proportion) in east Yamna groups, which supports the high mobility and exogamy practices among western and eastern Yamna clans, not only with locals:
We don’t have a comparison with Ukraine_Eneolithic or Corded Ware samples in Wang et al. (2018), but we do have proximate sources for Abashevo, when compared to the Poltavka population (with which it admixed in the Volga-Ural steppes): Sintashta, Potapovka, Srubna (with further Abashevo contribution), and Andronovo:
The two CWC outliers from the Baltic show what I thought was an admixture with Yamna. However, given the previous mixture of Eneolithic_Steppe in north Pontic steppe-forest populations, this elevated “steppe ancestry” found in Baltic_LN (similar to west Yamna) seems rather an admixture of Baltic sub-Neolithic peoples with a north Pontic Eneolithic_Steppe-like population. Late Repin settlers also admixed with a similar population during its colonization of the north Pontic area, hence the Baltic_LN – west Yamna similarities.
NOTE. A direct admixture with west Yamna populations through exogamy by the ancestors of this Baltic population cannot be ruled out yet (without direct access to more samples), though, because of the contacts of Corded Ware with west Yamna settlers in the forest-steppe regions.
A similar case is found in the Yamna outlier from Mednikarovo south of the Danube. It would be absurd to think that Yamna from the Balkans comes from Corded Ware (or vice versa), just because the former is closer in the PCA to the latter than other Yamna samples. The same error is also found e.g. in the Corded Ware → Bell Beaker theory, because of their proximity in the PCA and their shared “steppe ancestry”. All those theories have been proven already wrong.
NOTE. A similar fallacy is found in potential Sintashta→Mycenaean connections, where we should distinguish statistically that result from an East/West Yamna + Balkans_BA admixture. In fact, genetic links of Mycenaeans with west Yamna settlers prove this (there are some related analyses in Anthrogenica, but the site is down at this moment). To try to relate these two populations (separated more than 1,000 years before Sintashta) is like comparing ancient populations to modern ones, without the intermediate samples to trace the real anthropological trail of what is found…Pure numbers and wishful thinking.
The genetic formation of Central and South Asian populations has been unclear because of an absence of ancient DNA. To address this gap, we generated genome-wide data from 362 ancient individuals, including the first from eastern Iran, Turan (Uzbekistan, Turkmenistan, and Tajikistan), Bronze Age Kazakhstan, and South Asia. Our data reveal a complex set of genetic sources that ultimately combined to form the ancestry of South Asians today. We document a southward spread of genetic ancestry from the Eurasian Steppe, correlating with the archaeologically known expansion of pastoralist sites from the Steppe to Turan in the Middle Bronze Age (2300-1500 BCE). These Steppe communities mixed genetically with peoples of the Bactria Margiana Archaeological Complex (BMAC) whom they encountered in Turan (primarily descendants of earlier agriculturalists of Iran), but there is no evidence that the main BMAC population contributed genetically to later South Asians. Instead, Steppe communities integrated farther south throughout the 2nd millennium BCE, and we show that they mixed with a more southern population that we document at multiple sites as outlier individuals exhibiting a distinctive mixture of ancestry related to Iranian agriculturalists and South Asian hunter-gathers. We call this group Indus Periphery because they were found at sites in cultural contact with the Indus Valley Civilization (IVC) and along its northern fringe, and also because they were genetically similar to post-IVC groups in the Swat Valley of Pakistan. By co-analyzing ancient DNA and genomic data from diverse present-day South Asians, we show that Indus Periphery-related people are the single most important source of ancestry in South Asia — consistent with the idea that the Indus Periphery individuals are providing us with the first direct look at the ancestry of peoples of the IVC — and we develop a model for the formation of present-day South Asians in terms of the temporally and geographically proximate sources of Indus Periphery-related, Steppe, and local South Asian hunter-gatherer-related ancestry. Our results show how ancestry from the Steppe genetically linked Europe and South Asia in the Bronze Age, and identifies the populations that almost certainly were responsible for spreading Indo-European languages across much of Eurasia.
NOTE. The supplementary material seems to be full of errors right now, because it lists as R1b-M269 (and further subclades) samples that have been previously expressly said were xM269, so we will have to wait to see if there are big surprises here. So, for example, samples from Mal’ta (M269), Iron Gates (M269 and L51), and Latvia Mesolithic (L51), a Deriivka sample from 5230 BC (M269), Armenia_EBA (Z2103)…Also, the sample from Yuzhnyy Oleni Ostrov is R1a-M417 now.
EDIT (1 APR 2018): The main author has confirmed on Twitter that they have used a new Y Chr caller that calls haplogroups given the data provided, and depending on the coverage tried to provide a call to the lowest branch of the tree possible, so there are obviously a lot of mistakes – not just in the subclades of R. A revision of the paper is on its way, and soon more people will be able to work with the actual samples, since they say they are releasing them.
Nevertheless, since it is subclades (and not haplogroups) the apparent source of gross errors, for the moment it seems we can say with a great degree of confidence that:
New samples of East Yamna / Poltavka are of haplogroup R1b-L23.
Afanasevo is confirmed to be dominated by R1b-M269.
With lesser confidence in precise subclades, we find that:
A sample from Hajji Firuz in Iran ca. 5650 BC, of subclade R1b-Z2103, may confirm Mesolithic R1b-M269 lineages from the Caucasus as the source of CHG ancestry to Khvalynsk/Yamna, and be thus the reason why Reich wrote about a potential PIE homeland south of the Caucasus. (EDIT 11 APR 2018) The sample shows steppe ancestry, therefore the date is most likely incorrect, and a new radiocarbon dating is due. It is still interesting – depending on the precise subclade – for its potential relationship with IE migrations into the area.
New samples of East Yamna / Poltavka are of haplogroup R1b-Z2103.
Afanasevo migrants are mainly of haplogroup R1b-Z2103.
The Darra-e Kur sample, ca. 2655, of haplogroup R1b-L151, without a clear cultural adscription, may be the expected sign of Afanasevo migrants (Pre-Proto-Tocharian speakers) expanding a Northern Indo-European (in contrast with a Southern or Graeco-Aryan) dialect, in a region closely linked with the later desert mummies in the Tarim Basin. Its early presence there would speak in favour of a migration through the Inner Asian Mountain Corridor previous to the one caused by Andronovo migrants.
Sintashta shows a mixed R1b-Z2103 / R1a-Z93 society.
Later Indo-Iranian migrations are apparently dominated by R1a-Z2123, an early subclade of R1a-Z93, also found in Srubna.
R1b is also seen later in BMAC (ca. 1487 BC), although its subclade is not given.
There is also a sample of R1a-Z283 subclade in the eastern steppe (ca. 1600 BC). What may be interesting about it is that it could mark one of the subclades not responsible for the expansion of Balto-Slavic (or responsible for it with the expansion of Srubna, for those who support an Indo-Slavonic branch related Sintashta-Potapovka).
A sample of R1b-U106 subclade is found in Loebanr_IA ca. 950 BC, which – together with the sample of Darra-e Kur – is compatible with the presence of L51 in Yamna.
NOTE. Errors in haplogroups of previously published samples make every subclade of new samples from the supplementary table questionable, but all new samples (safe for the Darra_i_Kur one) were analysed and probably reported by the Reich Lab, and at least upper subclades in each haplogroup tree seem mostly coherent with what was expected. Also, the contribution of Iranian Farmer related (a population in turn contributing to Hajji Firuz) to Khvalynsk in their sketch of the genetic history may be a sign of the association of R1b-M269 lineages with CHG ancestry, although previous data on precise R1b subclades in the region contradict this. (EDIT 11 APR 2018) The sample of Hajji Firuz is most likely much younger than the published date, hence its younger subclade may be correct. No revision or comment on this matter has been published, though.
Also, it seems that the Corded Ware culture appears now irrelevant for Late Proto-Indo-European migrations. Observe:
Our results also shed light on the question of the origins of the subset of Indo-European languages spoken in India and Europe (45). It is striking that the great majority of Indo-European speakers today living in both Europe and South Asia harbor large fractions of ancestry related to Yamnaya Steppe pastoralists (corresponding genetically to the Steppe_EMBA cluster), suggesting that “Late Proto-Indo-European”—the language ancestral to all modern Indo- European languages—was the language of the Yamnaya (46). While ancient DNA studies have documented westward movements of peoples from the Steppe that plausibly spread this ancestry to Europe (5, 31), there has not been ancient DNA evidence of the chain 488 of transmission to South Asia. Our documentation of a large-scale genetic pressure from Steppe_MLBA groups in the 2nd millennium BCE provides a prime candidate, a finding that is consistent with archaeological evidence of connections between material culture in the Kazakh middle-to-late Bronze Age Steppe and early Vedic culture in India (46).
NOTE. If they correct the haplogroups soon, I will update the information in this post. Unless there is a big surprise that merits a new one, of course.
EDIT (1 APR 2018): Multiple minor edits to the original post.
EDIT (2 APR 2018): While I and other simple-minded people were only looking to confirm our previous theories using Y-DNA haplogroups, and are content with wildly speculating over the consequences if some of those strange (probably wrong) ones were true, intelligent people are using their time for something useful, interpreting the results of the investigation as described in the paper, to offer a clearer picture of Indo-Iranian migrations for everyone:
Featured image, from the article: “A Tale of Two Subcontinents. The prehistory of South Asia and Europe are parallel in both being impacted by two successive spreads, the first from the Near East after 7000 BCE bringing agriculturalists who mixed with local hunter-gatherers, and the second from the Steppe after 3000 BCE bringing people who spoke Indo-European languages and who mixed with those they encountered during their migratory movement. Mixtures of these mixed populations then produced the rough clines of ancestry present in both South Asia and in Europe today (albeit with more variable proportions of local hunter-gatherer-related ancestry in Europe than in India), which are (imperfectly) correlated to geography. The plot shows in contour lines the time of the expansion of Near Eastern agriculture. Human movements and mixtures, which also plausibly contributed to the spread of languages, are shown with arrows.”