Wang et al. (2018) Suppl. data: R1b-M269 in Baltic Neolithic?


Looking for information on Novosvobodnaya samples from Wang et al. (2018) for my latest post, I stumbled upon this from the Supplementary Data 2 (download the Excel table):

Latvia_MN1.SG (ZVEJ26)

Skeletal element: petrous
Sample: Latvia_MN_dup.I4627.SG
Date: 4251-3976 calBCE
Location: Zvejnieki
mtDNA: U4a1
Y-DNA: R1b1a1a2
Coverage: 0.15
SNPs hit on autosomes: 167445

The data on Mathieson et al. (2018) is as follows:

I4627 (ZVEJ26)

Skeletal element: petrous
Origin: ThisStudy (New data; Individual first published in JonesNatureCommunications2017)
Sample: Latvia_MN
Date:4251-3976 calBCE (5280±55 BP, Ua-3639)
mtDNA: U4a1
Y-DNA: R1b1a1a(xR1b1a1a2)
Coverage: 1.77
SNPs hit on autosomes: 686273

Y-Chromosome derived SNPs: R1b1a1a:PF6475:17986687C->A; R1b1a1a:CTS3876:15239181G->C; R1b1a1a:CTS5577:16376495A->C; R1b1a1a:CTS9018:18617596C->T; R1b1a1a:FGC57:7759944G->A; R1b1a1a:L502:19020340G->C; R1b1a1a:PF6463:16183412C->A; R1b1a1a:PF6524:23452965T->C; R1b1a:A702:10038192G->A; R1b1a:FGC35:18407611C->T; R1b1a:FGC36:13822833G->T; R1b1a:L754:22889018G->A; R1b1a:L1345:21558298G->T; R1b1a:PF6249:8214827C->T; R1b1a:PF6263:21159055C->A; R1b1:CTS2134:14193384G->A; R1b1:CTS2229:14226692T->A; R1b1:L506:21995972T->A; R1b1:L822:7960019G->A; R1b1:L1349:22722580T->C; R1b:M343:2887824C->A; R1:CTS2565:14366723C->T; R1:CTS3123:14674176A->C; R1:CTS3321:14829196C->T; R1:CTS5611:16394489T->G; R1:L875:16742224A->G; R1:P238:7771131G->A; R1:P286:17716251C->T; R1:P294:7570822G->C; R:CTS207:2810583A->G; R:CTS2913:14561760A->G; R:CTS3622:15078469C->G; R:CTS7876:17722802G->A; R:CTS8311:17930099C->A; R:F33:6701239G->A; R:F63:7177189G->A; R:F82:7548900G->A; R:F154:8558505T->C; R:F370:16856357T->C; R:F459:18017528G->T; R:F652:23631629C->A; R:FGC1168:15667208G->C; R:L1225:22733758C->G; R:L1347:22818334C->T; R:M613:7133986G->C; R:M734:18066156C->T; R:P224:17285993C->T; R:P227:21409706G->C

Context of Latvia_MN1

The Middle Neolithic is known to mark the westward expansion of Comb Ware and related cultures in North-Eastern Europe.

Mathieson et al. (2017 and 2018) had this to say about the Middle Neolithic in the Baltic:

At Zvejnieki in Latvia, using 17 newly reported individuals and additional data for 5 previously reported34 individuals, we observe a transition in hunter-gatherer-related ancestry that is opposite to that seen in Ukraine. We find that Mesolithic and Early Neolithic individuals (labelled ‘Latvia_HG’) associated with the Kunda and Narva cultures have ancestry that is intermediate between WHG (approximately 70%) and EHG (approximately 30%), consistent with previous reports34–36(Supplementary Table 3). We also detect a shift in ancestry between Early Neolithic individuals and those associated with the Middle Neolithic Comb Ware complex (labelled ‘Latvia_MN’), who have more EHG-related ancestry; we estimate that the ancestry of Latvia_MN individuals comprises 65% EHG-related ancestry, but two of the four individuals appear to be 100% EHG in principal component space (Fig. 1b).

From Mathieson et al. (2018). Ancient individuals projected onto principal components defined by 777 presentday west Eurasians (shown in Extended Data Fig. 1); data include selected published individuals (faded circles, labelled) and newly reported individuals (other symbols, outliers enclosed in black circles). Coloured polygons cover individuals that had cluster memberships fixed at 100% for supervised ADMIXTURE analysis.

Other samples and errors on Y-SNP calls

The truth is, this is another sample (Latvia_MN_dup.I4627.SG) from the same individual ZVEJ26.

There is another sample used for the analysis of ZVEJ26, with the same data as in Mathieson et al. (2018), i.e. better coverage, and Y-DNA R1b1a1a(xR1b1a1a2).

Most samples in the tables from Wang et al. (2018) seem to be classified correctly, as in previous papers, but for:

  • Blätterhöhle Cave sample from Lipson et al. (2017), wrongly classified (again) as R1b1a1a2a1a2a1b2 (I am surprised no R1b-autochtonous-continuity-fan rushed to proclaim something based on this);
  • Mal’ta 1 sample from Raghavan et al. (2013) as R1b1a1a2;
  • Iron Gates HG, Schela Cladovey from Gonzalez Fortes (2017) as R1b1a1a2;
  • Oase1 from Fu (2015) as N1c1a;
  • samples from Skoglund et al. (2017) from Africa also wrongly classified as R1b1a1a2 and subclades.

It seems therefore that the poor coverage / SNPs hit on autosomes is the key common factor here for these Y-SNP calls, and so it is in the Zvejnieki MN1 duplicated sample. Anyway, if all Y-SNP calls come from the same software applied to all data, and this is going to be used in future papers, this seems to be a great improvement compared to Narasimhan et al. (2018)

EDIT (25 JUN 2018): I have been reviewing some more papers apart from Mathieson et al. (2018) and Olalde et al. (2018) to compare the reported haplogroups, and there seems to be many potential errors (or updated data, difficult to say sometimes, especially when the newly reported haplogroup is just one or two subclades below the reported one in ‘old’ papers), not only those listed above.

The sample accession number in the European Nucleotide Archive (ENA) is SAMEA45565168 (Latvia_MN1/ZVEJ26) (see here), in case anyone used to this kind of analysis wishes to repeat the Y-SNP calls on both samples.

EDIT (25 JUN 2018): Added that it is another sample with lesser coverage from the same ZVEJ26 individual.


On the Maykop – Upper Mesopotamia cultural province, distinct from the steppe


New paper (behind paywall) The Production of Thin‐Walled Jointless Gold Beads from the Maykop Culture Megalithic Tomb of the Early Bronze Age at Tsarskaya in the North Caucasus: Results of Analytical and Experimental Research, by Trifonov et al. Archaeometry (2018)

Interesting excerpts (emphasis mine):

In 1898, two megalithic tombs containing graves of a local social elite dated to the Early Bronze Age were discovered by N. I. Veselovsky near the village of Tsarskaya (modern Novosvobodnaya, Republic of Adygeya) (Fig. 1 (a)) (Baye 1900, 43–59; IAC 1901, 33–8; Sagona 2018, 281–97).

Radiocarbon dates place both tombs within the Novosvobodnaya phase of the Maykop culture, between c. 3200 and 2900 BC (Trifonov et al. 2017). Along with the human remains (one adult individual was interred in each dolmen), the tombs yielded rich funerary offerings, including artefacts made of gold, silver and semi-precious stones. (…) This paper presents results of a technical analysis of just one type of artefact, from kurgan 2 at Tsarskaya: thin-walled jointless beads made from gold.

(a) A map of the Caucasus and part of Western Asia, showing the locations of sites mentioned in the text: 1, Tsarskaya (modern Novosvobodnaya); 2, Maykop; 3, Staromyshastovskaya; 4, Andryukovskaya; 5, Psebaiskaya; 6, Inozemtsevo; 7, Kudakhurt; 8, Soyuq Bulaq; 9, Sé Girdan; 10, Tepe Gawra. (b) The string of thin-walled jointless gold beads, silver and carnelian beads from the dolmen in kurgan 2 at Tsarskaya, Western Caucasus (1898).

Ever since M. I. Rostovtzeff noted a stylistic similarity between Maykop art and Sumerian art (Rostovtzeff 1920) and M. V. Andreeva described this phenomenon within a broad cultural and chronological context (Andreeva 1977), new archaeological studies have only extended this picture of a vast cultural province that appeared between the Caucasus and the northern fringe of Western Asia (Trifonov 1987). The discovery of the Leyla-Tepe culture (Narimanov 1987) and Maykop-type kurgans in Azerbaijan (Lyonnet et al. 2008) and adjacent Iran (Muscarella 1969, 1971, 2003; Trifonov 2000) has confirmed the spatial and temporal unity of this phenomenon as a precondition for free circulation of cultural patterns and technical innovations across vast areas of the Caucasus and Western Asia. Jewellery made of gemstones and precious metals, primarily gold, was probably one such innovation.

Attempts to demarcate the historical region where the Maykop culture emerged and developed have emphasized the role of Upper Mesopotamia in the development of the Sumerian civilization and the definition of a northern centre of urbanization, independent from the centres of the south (Rothman 2002; Oats et al. 2007). The turn of the fourth millennium BC saw the development of various cultural traditions in south-east Anatolia, north-east Syria and north-west Iran; on the northern fringe, these traditions manifested themselves through the Maykop culture. Perhaps it is no coincidence that the first high-status burials containing gold and gemstone jewellery (including carnelian, turquoise and lapis lazuli) appear in these northern, rather than southern, centres in the first quarter of 4000 BC (e.g., Tepe Gawra, graves 109, 110) (Piasnall 2002). With regard to funeral rites and stylistic characteristics of jewellery pieces, these graves have many parallels with early Maykop burials (Munchaev 1975, 329; Trifonov 1987, 20).

It still remains unclear if the goldsmiths of Upper Mesopotamia mastered the technique of making thin-walled jointless beads. The gold beads from Tepe Gawra are described as spherical or ball-shaped, but their maximum diameter (5–8mm) always exceeds the length of the bore (3–4mm) (Tobler 1950, 89, 199, pl. LV, a). On the whole, these measurements are consistent with the proportions and sizes of some Maykop beads.(…)

It is quite possible that a distinctive technique of making thin-walled jointless beads from gold was a regional technological development of Maykop culture goldsmiths, within a wider tradition of Near East metalwork, as a type of production regulated by ritual beliefs (Gell 1992; Benzel 2013).

These deep-rooted Near East traditions of ritualization of the production and use of jewellery pieces made of gold, silver and gemstones in the Maykop culture, on the one hand, maintained familiar canons of ritual behaviour and, on the other, made perception of sophisticated symbolism of gemstones more difficult for neighbouring cultures with different living standards, levels of social development and value systems to understand. The jewellery traditions of the Maykop culture had no successors in the Caucasus or the adjacent steppes. In the third millennium BC, the goldsmiths of Europe and Asia had to reinvent the technique of making thin-walled jointless gold beads from scratch (Born et al. 2009).

Also interesting is Holocene environmental history and populating of mountainous Dagestan (Eastern Caucasus, Russia), by Ryabogina et al., Quaternary International (2018).

The combination of Holocene environment changes and the settlement of the territory of Dagestan.

Related excerpts, about the climate of an adjacent region of the Caucasus before, during, and after the Maykop culture:

The 7th millennium BC featured a warm and arid climate, so that time corresponds to the steppe landscapes in the final stage of the Mesolithic. It is likely that the formation of a producing economy in the mountainous zone of Dagestan gradually emerged against this background. In the Neolithic period, the area remained almost treeless, as it was still warm and quite dry. However, archaeological data indicates that long-term settlements with well-developed farming spread in the mountainous zone around 6200-5500 BC.

The beginning of increasing humidity and the appearance of deciduous forests corresponds to the early Chalcolithic period of the Eastern Caucasus. It is the most poorly studied period in the history of this region. Covering a time span of 2000 years, this period was the least saturated by archaeological sites. At the start of this period, only the stands of herdsman in the mountain zone are known, dating to the second half of the 6th millennium BC (Gadgiev, 1991). It is still not clear whether the mountains were not settled in such a favorable climatic stage. The uncertainty may be due to the fact that people have chosen other ecological niches, or it could be we simply do not have data due to the insufficient archaeological survey of the territory. It is surprising that the turn to drier climate and the reduction of deciduous forests in the inner mountainous part of Dagestan, the large, long-term settlements like Ginchi emerge with pronounced specialization in agriculture (Fig. 7 panel (2)) (Gadgiev, 1991).

After the dry climate, simultaneously with cooling, the subsequent spread of pine forests coincides with the beginning of expansion of Kura-Araxes culture from the territory of Georgia through Chechnya to the mountainous Dagestan. Debates on the impact of past climate on Kura-Araxes societies in Transcaucasus have a long history (for the comprehensive review see, for example, Connor and Kvavadze, 2014 and references therein). In general, it is clear that after 3000 BC, forest cover in most areas of the Kura-Araxes region in the Transcaucasia reached its maximum extent in the Holocene (Connor and Kvavadze, 2014). However, at the same time lakes in Central Anatolia began to dry out and Caspian Sea levels fell (Roberts et al. 2011; Leroy et al. 2013), and arid conditions were identified in mountainous Dagestan in the 4th millennium. Clearly the regional moisture balance shifted in the Eastern Caucasus only in the late 4th to early 3rd millennium BC (this study). The only available radiocarbon dating of Dagestan confirms that the agricultural settlements of the Early Bronze Age appear not in the middle of the 4th millennium BC, but in the early 3rd millennium BC; that is not earlier than the stage of increasing moistening and the appearance of pine forests.

See also:

Paleoenvironment in mid- to late Holocene in the Cis-Ural steppes, and Epigravettian in Eastern Europe

Dynamics of paleoenvironments in the Cis-Ural steppes during the mid- to late Holocene, by Khokhlova, Morgunova, Khokhlov, and Golyeva, Quaternary Research (2018), 1–15.

Interesting excerpts:

About the studied site

The Turganik settlement in the Orenburg Region constitutes part of the so-called Ivanovo microregion of cultural heritage monuments, along with the Mesolithic Starotokskaya site; an Ivanovskoye multi-layered settlement (Neolithic, Eneolithic [or Chalcolithic], Late Bronze Age); Toksky I and Toksky II settlements attributed to the Late Bronze Age (the Timber-Grave archaeological culture); an Ivanovsky ground burial dated to the Eneolithic; and the Ivanovsky kurgan cemetery of the Early Iron Age (Fig. 1).

The ancient settlements are located at the Turganik River mouth, where the river joins the Tok River (the Samara River drainage basin). The Turganik River enters an old channel of the Tok which continues to flow due to that fact. Both valleys are wide and dissected by multiple river channels. The floodplain landscapes are mostly wet meadows with rich herb and grass vegetation, pastures, and hay fields. On both sides of the Turganik River, and farther along the right side of the Tok valley there are flat-topped elevations, with occasional forests (Chibilev, 1996). The Turganik settlement was positioned on a slightly elevated surface at the confluence of the Turganik and Tok rivers, on the right side of the valley. The settlement was inhabited in the Eneolithic and the Late Bronze Age, the fifth to fourth millennia BC.

(a and b) Location of the studied region and (c) the objects of the cultural heritage in the microregion: 1, Turganik settlement; 2, Toksky II settlement; 3, Ivanovsky dune with Ivanovsky ground cemetery; 4, Ivanovskoye II multi-layered settlement; 5, Staro-Tokskaya site; 6, Toksky I settlement; 7, Ivanovsky I kurgan cemetery.

Results and discussion

Pollen assemblages of the Atlantic optimum ~ 5500 yr BP indicate some increase in moisture supply and related afforestation of the floodplain (Lavrushin and Spiridonova, 1995). As follows from our data, the site was abandoned at that time and the no-longer-functioning cultural layer VI was gradually buried under deposits of frequent floods. According to the 14C ages obtained on archeological materials, the age of layer VI (or the second stage of the Eneolithic epoch on the Turganik settlement) may be dated to 4237–3790 cal yr BC, that is, somewhat earlier than the Holocene optimum suggested by palynologists.

Layer V shows another interval marked by increasing climate aridity and the dominance of grass steppes. As stated by the above-cited authors, the climate at the time that layer V was functioning was even dryer than during the formation of layer VI. That is confirmed by our data on the layer V composition, was formed during early Pit-Grave culture (the Early Bronze Age), in the range from 3800–3360 BC, according to the dates obtained on archeological materials (Morgunova et al., 2016b). As follows from the above, the maximum of aridity coincided with the Atlantic optimum.

It follows from the above that the Atlantic period of the Holocene was mostly characterized by arid environments; the peak of aridity fell on the early Bronze Age, the time of the early (Repino) stage of the Yamnaya culture in the Cis-Ural steppes. The Subboreal and Subatlantic periods were relatively colder and more humid, though short episodes of aridity could occur and some of them happened to be recorded in the sequence under study.

The reconstructed history of the climate changes in the Cis-Ural steppes during three intervals of the Holocene is in a good agreement with the results obtained in other regions. According to Alexandrovskiy (1996, 2000; Alexandrovskiy et al., 1999, 2004), the Atlantic period was the most arid one in the south of Russia, the subsequent intervals being comparatively wetter and colder. The extreme aridity was recorded on the Ukraine territory at the final Atlantic period, a few less arid chrono-intervals having been identified over the entire period (Kotova, 2009).

There are, however, other schemes of climate fluctuations in the central part of the Russian steppe zone; a few of them consider the Atlantic period to be humid, or even the most humid, as compared with the second half of the Holocene (Ivanov, 1992; Demkin, 1997). Also acceptable is a scenario of climatic fluctuations occurring at different times in different regions (Chendev et al., 2010). Further investigations and accumulation of empirical data would help to gain a better insight into the problem.


The ancient people inhabited the place from 5000 to 4000 BC (actually throughout the Atlantic period), when the place was not subjected to flooding. At the time of human habitation, the climate was mostly arid. Paleosols of that time are attributable to the Kastanozems (Endosalic Protosodic). They developed under grass (or herb and grass) steppes. The peak of aridity falls on the final Atlantic period. At the end of Eneolithic epoch (the fifth millennium BC) and in the Early Bronze Age (the fourth millennium BC) there were short-term but violent floods, which forced people to leave the habitable place.

During the Subboreal and Subatlantic periods of the Holocene, the climate became more humid, the floods became regular, the vegetation was dominated by meadow forbs and herbs growing on meadow-chernozem soils (Luvic Chernozem [Stagnic]), and the settlement was completely abandoned. In general, the studied sedimentary record at the Turganik archeological site reveals traceable climate change towards lower temperatures and increasing humidity in the second part of the Holocene, with occasional episodes of aridity that did not affect the general trend.

Interesting also the paper Collagen stable isotopes provide insights into the end of the mammoth steppe in the central East European plains during the Epigravettian, by Drucker et al., Quaternary Research (2018), 1-13

Location of the sites considered in this study.

About the studied site

The central East European plains are famous for their Epigravettian sites that date to around 15–12 14C ka BP (ca. 18.2–13.8 cal ka BP) and display impressive large structures made from the bones of woolly mammoth (Mammuthus primigenius; e.g., Gladkih et al., 1984; Soffer, 1985; Hoffecker, 2002). The origin of the large accumulations of mammoth remains is still a matter of debate, with the main hypotheses being the collection of natural occurrences versus active hunting (e.g., Soffer, 1985; Haynes 1989; Svoboda et al., 2005). In favor of this second scenario, studies of the mammoth remains of Yudinovo (Germonpré et al., 2008) concluded that the mammoths were hunted and, at Mezhyrich, mammoths were obtained by combined procurement via collection of carcasses and active hunting (Péan, 2015). Hunting practices were observed in older sites of the Gravettian culture in the Dnieper and Desna valleys (Demay et al., 2016).

Between ca. 22 and 12 14C ka BP (ca. 26.2–13.8 ka cal BP), the Dnieper and Desna basins correspond to the southern part of the geographical distribution of the woolly mammoth (Markova et al., 2013; Kahlke, 2014). Over time their distribution shifted northwards, while the density of the mammoth population decreased (Markova et al., 2013). According to Markova et al. (2013), the combined effect of gradual warming and growing human pressure is most likely to have had a negative impact on the mammoths, resulting in their local extinction in the Russian and Ukrainian plains around 14–12 14C kaBP (ca. 17.0–13.8 ka cal BP; Stuart et al., 2005).

Discussion and conclusion

Measured δ34S and δ15N values on bone collagen of mammoth, large canid, and fox from Mezhyrich (M), Buzhanka 2 (B), Yudinovo (Y), and Eliseevichi (E).

Humans could have taken advantage of the mammoth vulnerability, as reflected by lower δ15N values, to access animals that died naturally, collect bones, and hunt the most fragile individuals. This, with the possible assistance of domesticated dogs as hunting partners, could have countered a possible return to more suitable conditions for mammoth (Sablin and Khlopachev, 2002; Shipman, 2015). Our results confirm at least that mammoth specimens from Mezhyrich, Buzhanka 2, and, to a lesser extent, Eliseevichi were part of late mammoth populations surviving in sub-optimal conditions. They were thus most likely vulnerable to any pressure from environmental and/or human origin. Detecting further such cases among the late surviving mammoth populations using stable isotopic tracking may be a way to test if mammoth populations still had an optimal ecology or were metastable and, therefore, vulnerable to extinction. For instance, the insular Holocene population of mammoth on Saint Paul Island exhibited low and variable δ15N values, indicating suboptimal ecological conditions preceding their final disappearance (Graham et al., 2016).

The results of δ34S analyses showed no differences among mammoth according to the site but possibly a forage range partitioning between mammoth and coexisting large ungulates. Thus, variability in the mobility pattern for the mammoth between the high and low δ15N groups, such as migratory versus sedentary individuals, is not supported so far. We consider that rapid environmental modifications over time, probably not detectable through radiocarbon dating, can be a valid alternative explanation. Combined with direct competition with other large herbivores, such as the horse, and hunting of the most vulnerable individuals, the loss of their optimal habitat was likely to be the driving factor behind the local extinction of the mammoth in the central East European plains.


Ancient genomes from North Africa evidence Neolithic migrations to the Maghreb

BioRxiv preprint now published (behind paywall) Ancient genomes from North Africa evidence prehistoric migrations to the Maghreb from both the Levant and Europe, by Fregel et al., PNAS (2018).

NOTE. I think one of the important changes in this version compared to the preprint is the addition of the recent Iberomaurusian samples.

Abstract (emphasis mine):

The extent to which prehistoric migrations of farmers influenced the genetic pool of western North Africans remains unclear. Archaeological evidence suggests that the Neolithization process may have happened through the adoption of innovations by local Epipaleolithic communities or by demic diffusion from the Eastern Mediterranean shores or Iberia. Here, we present an analysis of individuals’ genome sequences from Early and Late Neolithic sites in Morocco and from Early Neolithic individuals from southern Iberia. We show that Early Neolithic Moroccans (∼5,000 BCE) are similar to Later Stone Age individuals from the same region and possess an endemic element retained in present-day Maghrebi populations, confirming a long-term genetic continuity in the region. This scenario is consistent with Early Neolithic traditions in North Africa deriving from Epipaleolithic communities that adopted certain agricultural techniques from neighboring populations. Among Eurasian ancient populations, Early Neolithic Moroccans are distantly related to Levantine Natufian hunter-gatherers (∼9,000 BCE) and Pre-Pottery Neolithic farmers (∼6,500 BCE). Late Neolithic (∼3,000 BCE) Moroccans, in contrast, share an Iberian component, supporting theories of trans-Gibraltar gene flow and indicating that Neolithization of North Africa involved both the movement of ideas and people. Lastly, the southern Iberian Early Neolithic samples share the same genetic composition as the Cardial Mediterranean Neolithic culture that reached Iberia ∼5,500 BCE. The cultural and genetic similarities between Iberian and North African Neolithic traditions further reinforce the model of an Iberian migration into the Maghreb.

Ancestry inference in ancient samples from North Africa and the Iberian Peninsula. PCA analysis using the Human Origins panel (European, Middle Eastern, and North African populations) and LASER projection of aDNA samples.

Relevant excerpts:

FST and outgroup-f3 distances indicate a high similarity between IAM and Taforalt. As observed for IAM, most Taforalt sample ancestry derives from Epipaleolithic populations from the Levant. However, van de Loosdrecht et al. (17) also reported that one third of Taforalt ancestry was of sub-Saharan African origin. To confirm whether IAM individuals show a sub-Saharan African component, we calculated f4(chimpanzee, African population; Natufian, IAM) in such a way that a positive result for f4 would indicate that IAM is composed both of Levantine and African ancestries. Consistent with the results observed for Taforalt, f4 values are significantly positive for West African populations, with the highest value observed for Gambian and Mandenka (Fig. 3 and SI Appendix, Supplementary Note 10). Together, these results indicate the presence of the same ancestral components in ∼15,000-y old and ∼7,000-y-old populations from Morocco, strongly suggesting a temporal continuity between Later Stone Age and Early Neolithic populations in the Maghreb. However, it is important to take into account that the number of ancient genomes available for comparison is still low and future sampling can provide further refinement in the evolutionary history of North Africa.

Genetic analyses have revealed that the population history of modern North Africans is quite complex (11). Based on our aDNA analysis, we identify an Early Neolithic Moroccan component that is (i) restricted to North Africa in present-day populations (11); (ii) the sole ancestry in IAM samples; and (iii) similar to the one observed in Later Stone Age samples from Morocco (17). We conclude that this component, distantly related to that of Epipaleolithic communities from the Levant, represents the autochthonous Maghrebi ancestry associated with Berber populations. Our data suggests that human populations were isolated in the Maghreb since Upper Paleolithic times. Our hypothesis is in agreement with archaeological research pointing to the first stage of the Neolithic expansion in Morocco as the result of a local population that adopted some technological innovations, such as pottery production or farming, from neighboring areas.

By 3,000 BCE, a continuity in the Neolithic spread brought Mediterranean-like ancestry to the Maghreb, most likely from Iberia. Other archaeological remains, such as African elephant ivory and ostrich eggs found in Iberian sites, confirm the existence of contacts and exchange networks through both sides of the Gibraltar strait at this time. Our analyses strongly support that at least some of the European ancestry observed today in North Africa is related to prehistoric migrations, and local Berber populations were already admixed with Europeans before the Roman conquest. Furthermore, additional European/ Iberian ancestry could have reached the Maghreb after KEB people; this scenario is supported by the presence of Iberian-like Bell-Beaker pottery in more recent stratigraphic layers of IAM and KEB caves. Future paleogenomic efforts in North Africa will further disentangle the complex history of migrations that forged the ancestry of the admixed populations we observe today.

Ancestry inference in ancient samples from North Africa and the Iberian Peninsula. (B) ADMIXTURE analysis using the Human Origins dataset (European, Middle Eastern, and North African populations) for modern and ancient samples (K = 8). (D) Detail of ADMIXTURE analysis using the Human Origins dataset (European, Middle Eastern, North African, and sub-Saharan African populations) for modern and ancient samples, including Taforalt.

Also, from the main author’s Twitter account:

I just realized that the paragraph with information on data availability is missing! Sequence data in the European Nucleotide Archive (PRJEB22699). Consensus mtDNA sequences are available at the National Center of Biotechnology Information (Accession Numbers MF991431-MF991448).

I find it hard to believe that this genetic continuity from Upper Palaeolithic to Late Neolithic could be representative of an autochthonous development of Afroasiatic. An important population movement – likely more than one – must be found in ancient DNA influencing North-Central and North-East Africa, probably during the time of the Green Sahara corridor.

See here:

Bayesian estimation of partial population continuity by using ancient DNA and spatially explicit simulations


Open access Bayesian estimation of partial population continuity by using ancient DNA and spatially explicit simulations, by Silva et al., Evolutionary Applications (2018).

Abstract (emphasis mine):

The retrieval of ancient DNA from osteological material provides direct evidence of human genetic diversity in the past. Ancient DNA samples are often used to investigate whether there was population continuity in the settlement history of an area. Methods based on the serial coalescent algorithm have been developed to test whether the population continuity hypothesis can be statistically rejected by analysing DNA samples from the same region but of different ages. Rejection of this hypothesis is indicative of a large genetic shift, possibly due to immigration occurring between two sampling times. However, this approach is only able to reject a model of full continuity model (a total absence of genetic input from outside), but admixture between local and immigrant populations may lead to partial continuity. We have recently developed a method to test for population continuity that explicitly considers the spatial and temporal dynamics of populations. Here we extended this approach to estimate the proportion of genetic continuity between two populations, by using ancient genetic samples. We applied our original approach to the question of the Neolithic transition in Central Europe. Our results confirmed the rejection of full continuity, but our approach represents an important step forward by estimating the relative contribution of immigrant farmers and of local hunter‐gatherers to the final Central European Neolithic genetic pool. Furthermore, we show that a substantial proportion of genes brought by the farmers in this region were assimilated from other hunter‐gatherer populations along the way from Anatolia, which was not detectable by previous continuity tests. Our approach is also able to jointly estimate demographic parameters, as we show here by finding both low density and low migration rate for pre‐Neolithic hunter‐gatherers. It provides a useful tool for the analysis of the numerous aDNA datasets that are currently being produced for many different species.

A) Different zones defined for computing proportions of ancestry in Central Europeans 4,500 BP. B) Schematic representation of various population contributions. C) Mean proportions of ancestry from the various PHG zones (A+B+C+D) in Central European populations from zone A at the end of the Neolithic transition 4,500 BP, computed for autosomal and mitochondrial markers.

Relevant excerpts:

Our results are in general accordance with two distinct ancestry components that have previously been detected at the continental scale by Lazaridis, Patterson et al. (2014): the “early European farmer” (EEF), which corresponds here to the NFA from Anatolia (zone C in Figure 3), and the “West European hunter-gatherer” (WHG), which corresponds here to the PHG from zones A and B in Figure 3. Notably, the contribution of an Ancient North Eurasians (ANE) component is not included in our model as we did not consider potential post-Neolithic immigration waves, which could have contributed to the modern European genetic pool, such as the wave that came from the Pontic steppes and was associated with the Yamnaya culture (Haak, Lazaridis et al. 2015). Without considering the ANE ancestry component, our estimate of the autosomal genetic contribution of Early farmers to the gene pool of Central European populations (25%) tends to be lower than the EEF ancestry estimated in most modern Western European populations, but is of the same order than the estimations in modern Estonians and in the ancient Late Neolithic genome “Karsdorf” from Germany (Lazaridis, Patterson et al. 2014, Haak, Lazaridis et al. 2015). Note that the contribution of hunter-gatherers to Neolithic communities appears to be variable in different regions of Europe (Skoglund, Malmstrom et al. 2012, Brandt, Haak et al. 2013, Lazaridis, Patterson et al. 2014), while we computed an average value for Central Europe. Moreover, we computed the ancestry of the two groups at the end of the Neolithic period while previous studies estimated it in modern times. Finally, previous studies used molecular information to directly estimate admixture proportions, while we use molecular information to estimate the model parameters and, then, we computed the expected genetic contributions of both groups using the best parameters, without using molecular information during this second step. Model assumptions may thus influence the inferences on the relative genetic contribution of both groups. In particular, we made the assumption of a uniform expansion of NFA with constant and similar assimilation of PHG over the whole continent but spatio-temporally heterogeneous environment, variable assimilation rate and long distance dispersal may have played an important role. The effects of those factors should be investigated in future studies.

The Proto-Indo-European – Euskarian hypothesis


Another short communication by Juliette Blevins has just been posted, A single sibilant in Proto-Basque: *s, *Rs, *sT and the phonetic basis of the sibilant split:

Blevins (to appear) presents a new reconstruction of Proto-Basque, the mother of Basque and Aquitanian, based on standard methods in historical linguistics: the comparative method and the method of internal reconstruction. Where all previous reconstructions of Proto-Basque assume a contrast between two sibilants, *s, a voiceless apical sibilant, and *z a voiceless laminal sibilant (Martinet 1955; Michelena 1977; Lakarra 1995; Trask 1997), this proposal is unique in positing only a single sibilant *s. Under this account, all instances of Common Basque /z/ are derived from *s. More specifically, in syllable coda, *Rs > *Rz (R a sonorant) while in the syllable onset, *sT > *zT (T an oral stop). The true split of *s into /s/ vs. /z/ occurs when clusters like *Rz or *zT are further simplified to /z/.

In this talk, internal evidence for a single sibilant, *s, in Proto-Basque is presented, and sound changes underlying the sibilant split are examined within the context of Evolutionary Phonology (Blevins 2004, 2006, 2015, 2017). Similar sound changes are identified in other languages with similar cluster types (e.g. Kümmel 2007:232), and the phonetic basis of the sibilant split is informed by recent studies of sibilant retraction (e.g. Stevens and Harrington 2016; Stuart-Smith et al. 2018).

Blevins, already known for her previous work on the Basque language, was known internationally for her recent controversial proposal of a genetic relationship between Proto-Indo-European and Basque. Apparently, a book with her full model, Advances in Proto-Basque Reconstruction with evidence for the Proto-Indo-European-Euskarian Hypothesis, will be published by Routledge soon.

I was never convinced, not just about a genetic connection, but about the very possibility of discovering it if there was any, mainly because such a link would be quite old, and Basque is known to have been greatly influenced by surrounding IE prestige languages for millennia until it was first attested in the 16th century. Internal reconstruction can only avail a gross reconstruction of few aspects up to a certain point in time, probably not very far beyond the Pre-Roman period, and that only thanks to the available Aquitanian inscriptions.

There are indeed certain known migrations that could be linked with a pan-European population movement, the most likely one for this hypothesis being the Villabruna cluster (the Villabruna sample itself being of haplogroup R1b pre-P297), and especially the expansion of R1b-V88 lineages, found widespread in Europe from west to east, from Mesolithic Iberia to Khvalynsk.

This haplogroup is also found in Sardinia, which may be connected to the expansion of V88 subclades (which I have speculatively proposed could be linked to Afro-Asiatic) into Africa through Italy and the Green Sahara; although it could also be linked to a speculative Vasconic-Iberian – Palaeo-Sardo group.

Without knowing the exact Pre-Proto-Indo-European stage at which Blevins would place the Basque separation, it is difficult to know how it could fit within any macro-language proposal – and thus potential ancestral population expansion.

If you are interested in this hypothesis, I suggest Koch’s controversial paper of 2013 Is Basque an Indo-European Language? (PDF), appeared in JIES 41 (1 & 2)….And of course the many papers rejecting it in the same volume. You also have Forni’s writings supporting this association.

Seeing how many Basque nationalists (obviously obsessed with racial purity) are still rooting for an autochthonous Palaeolithic origin of R1b lineages (especially P312) linked with the Basque language and dat huge Vasconic Western Europe; and now, after Olalde & Mathieson 2018, how some are also suggesting a Neolithic link with Sardinians and the Neolithic expansion, for lack of further modern genetic differences with other Western Europeans… I wonder how a lot of people inclined to believe this nonsense today, and mentally linking Vasconic with haplogroup R1b, will be paradoxically necessarily tied precisely to this kind of macro-family proposals in the future.


Proto-Indo-European homeland south of the Caucasus?

User Camulogène Rix at Anthrogenica posted an interesting excerpt of Reich’s new book in a thread on ancient DNA studies in the news (emphasis mine):

Ancient DNA available from this time in Anatolia shows no evidence of steppe ancestry similar to that in the Yamnaya (although the evidence here is circumstantial as no ancient DNA from the Hittites themselves has yet been published). This suggests to me that the most likely location of the population that first spoke an Indo-European language was south of the Caucasus Mountains, perhaps in present-day Iran or Armenia, because ancient DNA from people who lived there matches what we would expect for a source population both for the Yamnaya and for ancient Anatolians. If this scenario is right the population sent one branch up into the steppe-mixing with steppe hunter-gatherers in a one-to-one ratio to become the Yamnaya as described earlier- and another to Anatolia to found the ancestors of people there who spoke languages such as Hittite.

The thread has since logically become a trolling hell, and it seems not to be working right for hours now.

Reich’s proposal based on ancestral components to explain the formation of a people and language is a continuation of their emphasis on ancestry to explain cultures and languages. It seems quite interesting to see this happen again, given their current trend to surreptitiously modify their previous ‘Yamnaya ancestry’ concept and Yamnaya millennia-long R1a-R1b community (that supposedly explains a Yamna -> Corded Ware -> Bell Beaker migration) to a more general ‘steppe people’ sharing a ‘steppe ancestry’ who spoke a ‘steppe language’.

Interesting arrows of dispersal of steppe ancestry, from Yamna -> Corded Ware -> Bell Beaker, from David Reich’s new book (yes, from 2018, number one bestseller in

This new idea based on ancestral components suffers thus from the same essential methodological problems, which equate it – yet again – to pure speculation:

  1. It is a conclusion based on the genomic analysis of few individuals from distant regions and different periods, and – maybe more disturbingly – on the lack of steppe ancestry in the few samples at hand.
  2. Wait, what? Steppe ancestry? So they are trying to derive potential genetic connections among specific prehistoric cultures with a poorly depicted genetic sketch, based on previous flawed concepts (instead of on anthropological disciplines), which seems a rather long stretch for any scientist, whether they are content with seeing themselves as barbaric scientific conquerors of academic disciplines or not. In other words, statistics is also science (in fact, the main one to assert anything in almost any scientific field), and you cannot overcome essential errors (design, sampling, hypothesis testing) merely by using a priori correct statistical methods. Results obtained this way constitute a statistical fallacy.

  3. Even if the sampling and hypothesis testing were fine, to derive anthropological models from genomic investigation is completely wrong. Ancestral component ≠ population.
  4. To include not only potential migrations, but also languages spoken by these potential migrants? It’s sad that we have a need to repeat it, but if ancestral component ≠ population, how could ancestral component = language?

The Proto-Indo-European-speaking community

This is what we know about the formation of a Proto-Indo-European community (i.e. a community speaking a reconstructible Proto-Indo-European language) in the Pontic-Caspian steppe, which is based on linguistic reconstruction and guesstimates, tracing archaeological cultures backwards from cultures known to have spoken ancient (proto-)languages, and helping both disciplines with anthropological models (for which ancient genomics is only helping select certain details) of migration or – rarely – cultural diffusion:

NOTE. The following dates are obviously simplified. Read here a more detailed linguistic assessment based on phonology.

Most likely Pre-Proto-Anatolian migration with Suvorovo-Novodanilovka chiefs in the North Pontic steppe and the Balkans.
  • ca. 5000 BC. Early Proto-Indo-European (or Indo-Uralic) spoken probably during the formation and development of a loose Early Khvalynsk – Sredni Stog I cultural-historical community over the Pontic-Caspian steppe region, whose indigenous population probably had mainly Caucasus hunter-gatherer ancestry.
  • ca. 4500 BC. Khvalynsk probably speaking Middle Proto-Indo-European expands, most likely including Suvorovo-Novodanilovka chiefs into the North Pontic steppe, and probably expanding R1b-M269 lineages for the first time.
  • ca. 4000 BC. Separated communities develop, including North Pontic cultures probably gradually dominated by R1a-Z645 (potentially speaking Proto-Uralic); and Khvalynsk (and Repin) cultures probably dominated by R1b-L23 lineages, most likely developing a Late Proto-Indo-European already separated from Proto-Anatolian.
  • ca. 3500 BC. A Proto-Corded Ware population dominated by R1a-Z645 expands to the north, and slightly later an early Yamna community develops from Late Khvalynsk and Repin, expanding to the west of the Don River, and to the east into Afanasevo. This is most likely the period of reduction of variability and expansion of subclades of R1a-Z645 and R1b-L23 that we expect to see with more samples.
  • ca. 3000 BC. Expansion of Corded Ware migrants in northern Europe, and Yamna migrants along the Danube and into the Balkans, with further reduction and expansion of certain subclades.
  • ca. 2500 BC. Expansion of Bell Beaker migrants dominated by R1b-L51 subclades in Europe, and late Corded Ware migrants in east Yamna expanding R1a-Z93 subclades.

All these events are compatible with language reconstruction in mainstream European schools since at least the 1980s, supported by traditional archaeological research of the past 20 years, and is being confirmed with Genomics.

For those willingly lost in a myriad of new dreams boosted by the shallow comment contained in David Reich’s paragraph on CHG ancestry, even he does not doubt that the origin of Late Proto-Indo-European lies in Yamna, to the north of the Caucasus, based on Anthony’s (2007) account:

Both images from the book, posted by Twitter user Jasper at

NOTE: By the way, David Anthony, one of the main sources of information for Reich’s group, never considered Corded Ware to have received Yamna migrants, and althought he changed his model due to the conclusions of the 2015 papers, he has recently changed his model again to adapt it to the inconsistencies found in phylogeography.

CHG ancestry and PIE homeland south of the Caucasus

As for the potential origins of CHG ancestry in early Proto-Indo-European speakers, I already stated clearly my opinion quite recently. They may be attributed to:

Just to be clear, an expansion of Proto-Anatolian to the south, through the Caucasus, cannot be discarded today. It will remain a possibility until Maykop and more Balkan Chalcolithic and Anatolian-speaking samples are published.

However, an original Early Proto-Indo-European community south of the Caucasus seems to me highly unlikely, based on anthropological data, which should drive any conclusion. From what I could read, here are the rather simplistic arguments used:

  • Gimbutas and Maykop: Maykop was thought to be (in Gimbutas’ times) a rather late archaeological culture, directly connected to a Transcaucasian Copper Age culture ca. 2400-2300 BC. It has been demonstrated in recent years that this culture is substantially older, and even then language guesstimates for a Late PIE / Proto-Anatolian would not fit a migration to the north. While our ignorance may certainly be used to derive far-fetched conclusions about potential migrations from and to it, using Gimbutas (or any archaeological theory until the 1990s) today does not make any sense. Still less if we think that she favoured a steppe homeland.

NOTE. It seems that the Reich Lab may have already access to Maykop samples, so this suggested Proto-Indo-European – Maykop connection may have some real foundation. Regardless, we already know that intense contacts happened, so there will be no surprise (unless Y-DNA shows some sort of direct continuity from one to the other).

  • Gamkrelidze & Ivanov: they argued for an Armenian homeland (and are thus at the origin of yet another autochthonous continuity theory), but they did so to support their glottalic theory, i.e. merely to support what they saw as favouring their linguistic model (with Armenian being the most archaic dialect). The glottalic theory is supported today – as far as I know – mainly by Kortlandt, Jagodziński, or (Nostraticist) Bomhard, but even they most likely would not need to argue for an Armenian homeland. In fact, their support of a Graeco-Aryan group (also supported by Gamkrelidze & Ivanov) would be against this, at least in archaeological terms.
  • Colin Renfrew and the Anatolian homeland: This conceptual umbrella of language spreading with farming everywhere has changed so much and so many times in the past 20 years, with so many glottochronological and archaeological estimates circulating, that you can support anything by now using them. Mostly used today for abstract models of long-lasting language contacts, cultural diffusion, and constellation analogies. Anyway, he strives to keep up-to-date information to revise the model, that much is certain:
  • Glottochronology, phylogenetic trees, Swadesh list analysis, statistical estimates, psychics, pyramid power, and healing crystals: no, please, no.
Science Magazine
“A first line of evidence comes from linguistic analysis based on quantitative lexical data, which returned a tree compatible with the Anatolian hypothesis

In principle, unlike many other recent autochthonous continuity theories, I doubt there can be much racial-based opposition anywhere in the world to an origin of Proto-Indo-European in the Middle East, where the oldest civilizations appeared – apart, obviously, from modern Northeast and Northwest Caucasian, Kartvelian, or Semitic speakers, who may in turn have to revisit their autochthonous continuity theories radically…

Nevertheless, it is obvious that prehistoric (and many historic) migrations are signalled by the reduction in variability and expansion of certain Y-DNA haplogroups, and not just by ancestral components. That is generally accepted, although the reasons for this almost universal phenomenon are not always clear.

In fact, Proto-Anatolian and Common Anatolian speakers need not share any ancestral component, PCA cluster, or any other statistical parameter related to steppe populations, not even the same Y-DNA haplogroups, given that approximately three thousand years might have passed between their split from an Indo-Hittite community and the first attested Anatolian-speaking communities…We must carefully follow their tracks from Anatolia ca. 1500 BC to the steppe ca. 4500 BC, otherwise we risk creating another mess like the Corded Ware one.

In my opinion, the substantial contribution of EHG ancestry and R1a-M417 lineages to the Pontic-Caspian steppe (probably ca. 6500 BC) from Central or East Eurasia is the most recent sizeable genomic event in the region, and thus the best candidate for the community that expanded a language ancestral to Proto-Indo-European – whether you call it Pre-Proto-Indo-European, Pre-Indo-Uralic, or Eurasiatic, depending on your preferences.

An early (and substantial) contribution of CHG ancestry in Khvalynsk relative to North Pontic cultures, if it is found with new samples, may actually be a further proof of the Caucasian substrate of Proto-Indo-European proposed by Kortlandt (or Bomhard) as contributing to the differentiation of Middle PIE from Uralic. Genomics could thus help support, again, traditional disciplines in accepting or rejecting academic controversial theories.


In the case of an Early PIE (or Indo-Uralic) homeland, genomic data is scarce. But all traditional anthropological disciplines point to the Pontic-Caspian steppe, so we should stick to it, regardless of the informal suggestion written by a renown geneticist in one paragraph of a book conceived as an introduction to the field.

It seems we are not learning much from the hundreds of peer-reviewed, statistically (superficially, at least) sound genetic papers whose anthropological conclusions have been proven wrong by now. A lot of people should be spending their time learning about the complex, endless methods at hand in this kind of research – not just bioinformatics – , instead of fruitlessly speculating about wild unsubstantiated proposals.

As a final note, I would like to remind some in the discussion, who seem to dismiss the identification of CHG with Proto-Indo-European by supporting a “R1a-R1b” community for PIE, of their previous commitment to ancestral components in identifying peoples and languages, and thus their support to Reich’s (and his group’s) fundamental premises.

You cannot have it both ways. At least David Reich is being consistent.


The preferred northwest passage to Scandinavia

Pontus Skoglund writes (and shares publicly) his perspective on early postglacial migrations of hunter-gatherers into Scandinavia, in Northwest Passage to Scandinavia (Nat. Ecol. Evol.): an initial migration from the south and a second coastal migration north of the Scandinavian ice sheet.

He sums up the recently published Open Access paper Population genomics of Mesolithic Scandinavia: Investigating early postglacial migration routes and high-latitude adaptation, by Günther, Malmström , Svensson, Omrak, et al. PLoS Biol (2018) 16(1): e2003703, based on preprint at BioRxiv Genomics of Mesolithic Scandinavia reveal colonization routes and high-latitude adaptation (2017).


Scandinavia was one of the last geographic areas in Europe to become habitable for humans after the Last Glacial Maximum (LGM). However, the routes and genetic composition of these postglacial migrants remain unclear. We sequenced the genomes, up to 57× coverage, of seven hunter-gatherers excavated across Scandinavia and dated from 9,500–6,000 years before present (BP). Surprisingly, among the Scandinavian Mesolithic individuals, the genetic data display an east–west genetic gradient that opposes the pattern seen in other parts of Mesolithic Europe. Our results suggest two different early postglacial migrations into Scandinavia: initially from the south, and later, from the northeast. The latter followed the ice-free Norwegian north Atlantic coast, along which novel and advanced pressure-blade stone-tool techniques may have spread. These two groups met and mixed in Scandinavia, creating a genetically diverse population, which shows patterns of genetic adaptation to high latitude environments. These potential adaptations include high frequencies of low pigmentation variants and a gene region associated with physical performance, which shows strong continuity into modern-day northern Europeans.

The ice sheet distribution – which did not improve nuch for thousands of years – was clearly the greatest barrier for potential migrations in the region.

Baltischer Süßwassersee Vorläufer der Ostsee vor 12.000 Jahren, by Juschki and Koyos at Wikipedia

See also:

Population replacement in Early Neolithic Britain, and new Bell Beaker SNPs


New (copyrighted) preprint at BioRxiv, Population Replacement in Early Neolithic Britain, by Brace et al. (2018).

Abstract (emphasis mine):

The roles of migration, admixture and acculturation in the European transition to farming have been debated for over 100 years. Genome-wide ancient DNA studies indicate predominantly Anatolian ancestry for continental Neolithic farmers, but also variable admixture with local Mesolithic hunter-gatherers. Neolithic cultures first appear in Britain c. 6000 years ago (kBP), a millennium after they appear in adjacent areas of northwestern continental Europe. However, the pattern and process of the British Neolithic transition remains unclear. We assembled genome-wide data from six Mesolithic and 67 Neolithic individuals found in Britain, dating from 10.5-4.5 kBP, a dataset that includes 22 newly reported individuals and the first genomic data from British Mesolithic hunter-gatherers. Our analyses reveals persistent genetic affinities between Mesolithic British and Western European hunter-gatherers over a period spanning Britain’s separation from continental Europe. We find overwhelming support for agriculture being introduced by incoming continental farmers, with small and geographically structured levels of additional hunter-gatherer introgression. We find genetic affinity between British and Iberian Neolithic populations indicating that British Neolithic people derived much of their ancestry from Anatolian farmers who originally followed the Mediterranean route of dispersal and likely entered Britain from northwestern mainland Europe.

Also, Genetiker has updated Y-SNP calls from new data published from the Harvard group.

The R1b lineages that expanded from (Yamna->) East Bell Beakers -> Western Europe are more and more clearly of R1b-L151 subclades, as expected.

Quite interesting are the early samples from Poland, of R1b1a1a2a2-Z2103 and R1b1a1a2a1a-L151 lineages – , which may point (different to the more homogeneous L151 distribution in Western Europe) to a mix in both original (east-west) Yamna groups. This could tentatively be used to explain the Graeco-Aryan influence that some linguists see in Balto-Slavic (or its superstrate).

That link would then be quite early, to account for an influence during the Yamna settlements in Hungary, before its expansion as East Bell Beakers, but we haven’t seen a clearly differentiated subgroup (yet) in Archaeology, Anthropology, or Genomics within the Hungarian Yamna/East Bell Beaker community, so I am not convinced. It could be just that different scattered subclades mixed with the general L151 population pop up (following old Yamna lineages, or having being added along the way), as expected in an expansion over such a great territory – as if some scattered samples of R1a, I1, I2, J, etc. were found.

We need more early samples from south-eastern Europe and the steppe during the Chalcolithic to ascertain the composition and migration paths of the different Yamna settlers.

Other interesting findings are the early (Proto-)Bell Beaker samples of haplogroup R1b with no steppe ancestry from Spain – which some autochthonous continuists wanted to believe was a proof of some kind – , which are actually R1b-V88, a haplogroup known to have expanded throughout Europe quite early. In fact, this subclade has been recently shown to have most likely expanded through the Green Sahara region, and is potentially linked to the expansion of Afro-Asiatic.

See also:

Genetic prehistory of the Baltic Sea region and Y-DNA: Corded Ware and R1a-Z645, Bronze Age and N1c


Open Access The genetic prehistory of the Baltic Sea region, by Mittnik et al., Nature Communications 9: 442 (2018), based on preprint The Genetic History of Northern Europe, at BioRxirv.

As you can see, it follows my predictions in terms of haplogroups, and sadly the same trend to substitute ‘Yamna’ for ‘steppe’ while keeping linguistic interpretations unchanged…

Important excerpts for the Indo-European question (emphasis mine):

Mesolithic to Neolithic

In the archaeological understanding, the transition from Mesolithic to Neolithic in the Eastern Baltic region does not coincide with a large-scale population turnover and a stark shift in economy as seen in Central and Southern Europe. Rather, it is signified by a change in networks of contacts and the use of pottery, among other material, cultural and economic changes. Our results suggest continued admixture between groups in the south of the Eastern Baltic region, who are more closely related to WHG, and northern or eastern groups, more closely related to EHG. Neolithic social networks from the Eastern Baltic to the River Volga could also explain similarities of the hunter-gatherer pottery styles, although morphologically analogous ceramics might also have developed independently due to similar functionality. The genetic evidence for a change in networks and possibly even a large-scale population movement is most pronounced in the Middle Neolithic in individuals attributed to the CCC. The distribution of this culture overlaps in the north with the Narva culture and extends further north to Finland and Karelia. Its spread in the Eastern Baltic is linked with a significant change in imported raw materials, artefacts, and the appearance of village-like settlements15.

Neolithic to Chalcolithic

We see a further population movement into the regions surrounding the Baltic Sea with the CWC in the Late Neolithic that was accompanied by the first evidence of extensive animal husbandry in the Eastern Baltic. The presence of ancestry from the Pontic-Caspian Steppe among Baltic CWC individuals without the genetic component from north-western Anatolian Neolithic farmers must be due to a direct migration of steppe pastoralists that did not pick up this ancestry in Central Europe. It suggests import of the new economy by an incoming steppe-like population independent of the agricultural societies that were already established to the south and west of the Baltic Sea. The presence of direct contacts to the steppe could lend support to a linguistic model that sees an early branching of Balto-Slavic from a Proto-Indo-European language, for which the west Eurasian steppe was proposed as a homeland. However, as farmer ancestry is found in later Eastern Baltic individuals, it is likely that considerable individual mobility and a network of contact throughout the range of the CWC facilitated its spread eastward, possibly through exogamous marriage practices. Conversely, the appearance of mitochondrial haplogroup U4 in the Central European Late Neolithic after millennia of absence could indicate female gene-flow from the Eastern Baltic, where this haplogroup was present at high frequency.

PCA and ADMIXTURE analysis reflecting Late Neolithic in Northern European prehistory. a Principal components analysis of 1012 present-day West Eurasians (grey points, modern Baltic populations in dark grey) with 294 projected published ancient and ancient North European samples introduced in this study (marked with a red outline). b Ancestral components in ancient individuals estimated by ADMIXTURE (k = 11)
Zoomed-in version of the European Late Neolithic PCA.

So, we see that no farmer ancestry is found in the Baltic (unlike in Western Yamna), that PCA of Late Neolithic is closer to Corded Ware samples from Europe (or to earlier samples from the region) and not to Yamna, as suggested at first by the Zvejnieki individual.

There obviously was exogamy – which may in fact justify the findings in PCA close to Yamna (like the Zvejnieki sample), although researchers obviate that.

Also, as expected, no R1b-M269 in the Baltic (during the Corded Ware period), most are R1a with the majority showing subclade R1a-Z645 (and others poor SNP coverage), which support the reduction in haplogroup diversity to this very subclade during the expansion of Corded Ware peoples, as I predicted it would happen.

Bronze Age

Local foraging societies were, however, not completely replaced and contributed a substantial proportion to the ancestry of Eastern Baltic individuals of the latest LN and Bronze Age. This ‘resurgence’ of hunter-gatherer ancestry in the local population through admixture between foraging and farming groups recalls the same phenomenon observed in the European Middle Neolithic and is responsible for the unique genetic signature of modern-day Eastern Baltic populations.

We suggest that the Siberian and East Asian related ancestry in Estonia, and Y-haplogroup N in north-eastern Europe, where it is widespread today, arrived there after the Bronze Age, ca. 500 calBCE, as we detect neither in our Bronze Age samples from Lithuania and Latvia. As Uralic speaking populations of the Volga-Ural region show high frequencies of haplogroup N, a connection was proposed with the spread of Uralic language speakers from the east that contributed to the male gene pool of Eastern Baltic populations and left linguistic descendants in the Finno-Ugric languages Finnish and Estonian. A potential future direction of research is the identification of the proximate population that contributed to the arrival of this eastern ancestry into Northern Europe.

I predicted that haplogroup N arrived probably to the region west of the Urals with the Sejma-Turbino phenomenon, and that it expanded quite late, probably through founder effects. A late arrival to the region leaves obviously (safe for these researchers and others working with old ideas) only the Corded Ware culture (represented by steppe admixture and mainly haplogroup R1a-Z645) as the vector of expansion of Uralic languages, which show obviously a dialectalization process and regional expansion much older than 500 BC…

It is funny to see how people keep trying to identify R1a with ‘Yamnaya’, now ‘steppe’, but always Indo-European (an ethnolinguistic term, mind you) supposedly because of the ‘Yamnaya’ (now ‘steppe’) admixture, but the only ‘mark’ of Uralic languages for the same researchers in the same paper using this very concept is nevertheless, paradoxically, haplogroup N, with an assumption explicitly based on prevalence in modern populations

This admixture vs. haplogroup question for language and culture identification in genetic papers is really gettting messed up with new data, now in a contortionist-like way…

Images and text: Content of the paper is licensed under CC-by 4.0.

See also: