Yamna the likely source of modern horse domesticates; the closest lineage, from East Bell Beakers

Open access Tracking Five Millennia of Horse Management with Extensive Ancient Genome Time Series, by Fages et al. Cell (2019).

Interesting excerpts (emphasis mine):

The earliest archaeological evidence of horse milking, harnessing, and corralling is found in the ∼5,500-year-old Botai culture of Central Asian steppes (Gaunitz et al., 2018, Outram et al., 2009; see Kosintsev and Kuznetsov, 2013 for discussion). Botai-like horses are, however, not the direct ancestors of modern domesticates but of Przewalski’s horses (Gaunitz et al., 2018). The genetic origin of modern domesticates thus remains contentious, with suggested candidates in the Pontic-Caspian steppes (Anthony, 2007), Anatolia (Arbuckle, 2012, Benecke, 2006), and Iberia (Uerpmann, 1990, Warmuth et al., 2011). Irrespective of the origins of domestication, the horse genome is known to have been reshaped significantly within the last ∼2,300 years (Librado et al., 2017, Wallner et al., 2017, Wutke et al., 2018). However, when and in which context(s) such changes occurred remains largely unknown.

To clarify the origins of domestic horses and reveal their subsequent transformation by past equestrian civilizations, we generated DNA data from 278 equine subfossils with ages mostly spanning the last six millennia (n = 265, 95%) (Figures 1A and 1B; Table S1; STAR Methods). Endogenous DNA content was compatible with economical sequencing of 87 new horse genomes to an average depth-of-coverage of 1.0- to 9.3-fold (median = 3.3-fold; Table S2). This more than doubles the number of ancient horse genomes hitherto characterized. With a total of 129 ancient genomes, 30 modern genomes, and new genome-scale data from 132 ancient individuals (0.01- to 0.9-fold, median = 0.08-fold), our dataset represents the largest genome-scale time series published for a non-human organism (Tables S2, S3, and S4; STAR Methods).

genetic-affinities-horse-domesticates-pca
Genetic Affinities.
(A)
Principal Component Analysis (PCA) of 159 ancient and modern horse genomes showing at least 1-fold average depth-of-coverage. The overall genetic structure is shown for the first three principal components, which summarize 11.6%, 10.4% and 8.2% of the total genetic variation, respectively. The two specimens MerzlyYar_Rus45_23789 and Dunaujvaros_Duk2_4077 discussed in the main text are highlighted. See also Figure S7 and Table S5 for further information.
(B) Visualization of the genetic affinities among individuals, as revealed by the struct-f4 algorithm and 878,475 f4 permutations. The f4 calculation was conditioned on nucleotide transversions present in all groups, with samples were grouped as in TreeMix analyses (Figure 3). In contrast to PCA, f4 permutations measure genetic drift along internal branches. They are thus more likely to reveal ancient population substructure.

Discovering Two Divergent and Extinct Lineages of Horses

Domestic and Przewalski’s horses are the only two extant horse lineages (Der Sarkissian et al., 2015). Another lineage was genetically identified from three bones dated to ∼43,000–5,000 years ago (Librado et al., 2015, Schubert et al., 2014a). It showed morphological affinities to an extinct horse species described as Equus lenensis (Boeskorov et al., 2018). We now find that this extinct lineage also extended to Southern Siberia, following the principal component analysis (PCA), phylogenetic, and f3-outgroup clustering of an ∼24,000-year-old specimen from the Tuva Republic within this group (Figures 3, 5A and S7A). This new specimen (MerzlyYar_Rus45_23789) carries an extremely divergent mtDNA only found in the New Siberian Islands some ∼33,200 years ago (Orlando et al., 2013) (Figure 6A; STAR Methods) and absent from the three bones previously sequenced. This suggests that a divergent ghost lineage of horses contributed to the genetic ancestry of MerzlyYar_Rus45_23789. However, both the timing and location of the genetic contact between E. lenensis and this ghost lineage remain unknown.

modern-horse-domesticates-przewalski-hungary
Population modeling of the demographic changes and admixture events in extant and extinct horse lineages. The two models presented show best fitting to the observed multi-dimensional SFS in momi2. The width of each branch scales with effective size variation, while colored dashed lines indicate admixture proportions and their directionality. The robustness of each model was inferred from 100 bootstrap pseudo-replicates. Time is shown in a linear scale up to 120,000 years ago and in a logarithmic scale above.

Modeling Demography and Admixture of Extinct and Extant Horse Lineages

Phylogenetic reconstructions without gene flow indicated that IBE differentiated prior to the divergence between DOM2 and Przewalski’s horses (Figure 3; STAR Methods). However, allowing for one migration edge in TreeMix suggested closer affinities with one single Hungarian DOM2 specimen from the 3rd mill. BCE (Dunaujvaros_Duk2_4077), with extensive genetic contribution (38.6%) from the branch ancestral to all horses (Figure S7B).This, and the extremely divergent IBE Y chromosome (Figure 6B), suggest that a divergent but yet unidentified ghost population could have contributed to the IBE genetic makeup.

Rejecting Iberian Contribution to Modern Domesticates

The genome sequences of four ∼4,800- to 3,900-year-old IBE specimens characterized here allowed us to clarify ongoing debates about the possible contribution of Iberia to horse domestication (Benecke, 2006, Uerpmann, 1990, Warmuth et al., 2011). Calculating the so-called fG ratio (Martin et al., 2015) provided a minimal boundary for the IBE contribution to DOM2 members (Cahill et al., 2013) (Figure 7A). The maximum of such estimate was found in the Hungarian Dunaujvaros_Duk2_4077 specimen (∼11.7%–12.2%), consistent with its TreeMix clustering with IBE when allowing for one migration edge (Figure S7B). This specimen was previously suggested to share ancestry with a yet-unidentified population (Gaunitz et al., 2018). Calculation of f4-statistics indicates that this population is not related to E. lenensis but to IBE (Figure 7B; STAR Methods). Therefore, IBE or horses closely related to IBE, contributed ancestry to animals found at an Early Bronze Age trade center in Hungary from the late 3rd mill. BCE. This could indicate that there was long-distance exchange of horses during the Bell Beaker phenomenon (Olalde et al., 2018). The fG minimal boundary for the IBE contribution into an Iron Age Spanish horse (ElsVilars_UE4618_2672) was still important (~9.6%–10.1%), suggesting that an IBE genetic influence persisted in Iberia until at least the 7th century BCE in a domestic context. However, fG estimates were more limited for almost all ancient and modern horses investigated (median = ~4.9%–5.4%; Figure 7A).

horse-lineages-domesticates-przewalski-dom2-botai
TreeMix Phylogenetic Relationships. The tree topology was inferred using a total of ∼16.8 million transversion sites and disregarding migration. The name of each sample provides the archaeological site as a prefix, and the age of the specimen as a suffix (years ago). Name suffixes (E) and (A) denote European and Asian ancient horses, respectively. See Table S5 for dataset information. Image modified to include the likely ancestor of domesticates in a red circle, represented by Yamna, the most likely direct ancestor of the Dunaujvarus specimen.

Iron Age horses

Y chromosome nucleotide diversity (π) decreased steadily in both continents during the last ∼2,000 years but dropped to present-day levels only after 850–1,350 CE (Figures 2B and S2E; STAR Methods). This is consistent with the dominance of an ∼1,000- to 700-year-old oriental haplogroup in most modern studs (Felkel et al., 2018, Wallner et al., 2017). Our data also indicate that the growing influence of specific stallion lines post-Renaissance (Wallner et al., 2017) was responsible for as much as a 3.8- to 10.0-fold drop in Y chromosome diversity.

We then calculated Y chromosome π estimates within past cultures represented by a minimum of three males to clarify the historical contexts that most impacted Y chromosome diversity. This confirmed the temporal trajectory observed above as Byzantine horses (287–861 CE) and horses from the Great Mongolian Empire (1,206–1,368 CE) showed limited yet larger-than-modern diversity. Bronze Age Deer Stone horses from Mongolia, medieval Aukštaičiai horses from Lithuania (C9th–C10th [ninth through the tenth centuries of the Common Era]), and Iron Age Pazyryk Scythian horses showed similar diversity levels (0.000256–0.000267) (Figure 2A). However, diversity was larger in La Tène, Roman, and Gallo-Roman horses, where Y-to-autosomal π ratios were close to 0.25. This contrasts to modern horses, where marked selection of specific patrilines drives Y-to-autosomal π ratios substantially below 0.25 (0.0193–0.0396) (Figure 2A). The close-to-0.25 Y-to-autosomal π ratios found in La Tène, Roman, and Gallo-Roman horses suggest breeding strategies involving an even reproductive success among stallions or equally biased reproductive success in both sexes (Wilson Sayres et al., 2014).

Lineage is used in this paper, as in many others in genetics, as defined by a specific ancestry. I keep that nomenclature below. It should not be confused with the “lineages” or “lines” referring to Y-chromosome (or mtDNA) haplogroups.

Supporting the “archaic” nature of the Hungarian BBC horses expanding from the Pontic-Caspian steppes are:

  • Among Y-chromosome lines, the common group formed by Botai-Borly4 (closely related to DOM2), Scythian horses from Aldy Bel (Arzhani), Iron Age horses from Estonia (Ridala), horses from the Xiongnu culture (Uushgiin Uvur), and Roman horses from Autricum (Chartres).
  • Among mtDNA lines, the common group formed by Botai samples, LebyazhinkaIV NB35, and different Eurasian domesticates, including many ancient Western European ones, which reveals a likely expansion of certain subclades east and west with the Repin culture.
  • (…) DOM2 contributed 22% to the ancestor of Przewalski’s horses ca. 9.47 kya, suggesting the Holocene optimum, rather than the Eneolithic Botai culture (∼5.5 kya), as a period of population contact. This pre-Botai introgression could explain the Y chromosome topology, where Botai horses were reported to carry two different segregating haplogroups: one occupied a basal position in the phylogeny while the other was closely related to DOM2. Multiple admixture pulses, however, are known to have occurred along the divergence of DOM2 and the Botai-Borly4 lineage, including 2.3% post-Borly4 contribution to DOM2, and a more recent 6.8% DOM2 intogression into Przewalski’s horses (Gaunitz et al., 2018). Model C2 parameters accommodate all these as a single admixture pulse, likely averaging the contributions of all these multiple events.

    horse-domesticate-y-dna-mtdna
    Tip labels are respectively composed of individual sample names, their reference number as well as their age (years ago, from 2017). Red, orange, light green, green, dark green and blue refer to modern horses, ancient DOM2, Botai horses, Borly4 horses, Przewalski’s horses and E. lenensis, respectively. Black refers to wild horses not yet identified to belong to any particular cluster in absence of sufficient genome-scale data. Clades composed of only Przewalski’s horses or ancient DOM2 horses were collapsed to increase readability.

    (A) Best maximum likelihood tree retracing the phylogenetic relationships between 270 mitochondrial genomes.

    B) Best Y chromosome maximum likelihood tree (GTRGAMMA substitution model) excluding outgroup. Node supports are indicated as fractions of 100 bootstrap pseudoreplicates. Bootstrap supports inferior to 90% are not shown. The root was placed on the tree midpoint. See also Table S5 for dataset information.

    Image modified from the paper, including a red square in archaic groups that contain the Hungarian sample, and a red circle around the most likely common ancestral stallion and mare from the Pontic-Caspian steppes.

    The paper cannot offer a detailed picture of ancient horse domestication, but it is yet another step in showing how Repin/Yamna is the most likely source of expansion of horse domesticates in Eurasia. Even more interestingly, Yamna settlers in Hungary probably expanded an ancient lineage of that horse at the same time as they spread with the Classical Bell Beaker culture. Remarkable parallels are thus found between:

    The expansion of an ancient line of horse domesticates related to Yamna Hungary/East Bell Beakers seems to be confirmed by the pre-Iberian sample from Vilars I, Els Vilars4618 2672 (ca. 700-550 BC), likely of Iberian Beaker descent, showing a lineage older than the Indo-Iranian ones, which later replaced most European lines.

    NOTE. For known contacts between Yamna and Proto-Beakers just before the expansion of East Bell Beakers, see a recent post on Vanguard Yamna groups.

    The findings of the paper confirm the expansion of the horse firstly (and mainly) through the steppe biome, mimicking the expansion of Proto-Indo-Europeans first, and then replaced gradually (or not so gradually) by lines brought to Europe during westward expansions of Bronze Age, Iron Age, and later specialized horse-riding steppe cultures. The expansion also correlates well with the known spread of animal traction and pastoralism before 2000 BC:

    animal-traction-europe
    Top image: Map with evidence of animal traction before ca. 2000 BC. Bottom image: frequency of finds of evidence for animal traction (orange), cylinder seals (purple) and potter’s wheels (green) in the 4th and 3rd millennium BC (query from the Digital Atlas of Innovations). The data points to an early peak in the expansion of this innovation at the turn of the 4th–3rd millennium BC, while direct evidence supports a radical increase from around the mid–3th millennium BC until the early 2nd millennium, coinciding with the expansion of East Bell Beakers and related European Early Bronze Age cultures. Data and image modified from Klimscha (2017).

    EDIT (3 MAY 2019): A recent reminder of these parallel developments by David Reich in Insights into language expansions from ancient DNA:

    • Yamna expansion to the west “with horses and wagons”, with a more homogeneous ancestry in modern Europeans due to later migrations from the east (and north):
    • “Descendants” of Yamna (once the culture was already “dead”), expanding to the east mainly with Corded Ware ancestry:

    Another recent open access paper on horse domestication is The horse Y chromosome as an informative marker for tracing sire lines, by Felkel et al. Scientific Reports (2019).

    Related

Fulani from Cameroon show ancestry similar to Afroasiatic speakers from East Africa

sahel-region-fulani

Open access African evolutionary history inferred from whole genome sequence data of 44 indigenous African populations, by Fan et al. Genome Biology (2019) 20:82.

Interesting excerpts (emphasis mine):

Introduction

To extend our knowledge of patterns of genomic diversity in Africa, we generated high coverage (> 30×) genome sequencing data from 43 geographically diverse Africans originating from 22 ethnic groups, representing a broad array of ethnic, linguistic, cultural, and geographic diversity (Additional file 1: Table S1). These include a number of populations of anthropological interest that have never previously been characterized for high-coverage genome sequence diversity such as Afroasiatic-speaking El Molo fishermen and Nilo-Saharan-speaking Ogiek hunter-gatherers (Kenya); Afroasiatic-speaking Aari, Agaw, and Amhara agro-pastoralists (Ethiopia); Niger-Congo-speaking Fulani pastoralists (Cameroon); Nilo-Saharan-speaking Kaba (Central African Republic, CAR); and Laka and Bulala (Chad) among others. We integrated this data with 49 whole genome sequences generated as part of the Simons Genome Diversity Project (SGDP) [14] (…)

afroasiatic-samples
Locations of samples included in this study. Each dot is an individual and the color indicates the language classification

Results and discussion

We found that the CRHG populations from central Africa, including the Mbuti from the Demographic Republic of Congo (DRC), Biaka from the CAR, and Baka, Bakola, and Bedzan from Cameroon, also form a basal lineage in the phylogeny. The other two hunter-gatherer populations, Hadza and Sandawe, living in Tanzania, group with populations from eastern Africa (Fig. 2). The two Nilo-Saharan-speaking populations, the Mursi from southern Ethiopia and the Dinka from southern Sudan, group into a single cluster, which is consistent with archeological data indicating that the migration of Nilo-Saharan populations to eastern Africa originated from a source population in southern Sudan in the last 3000 years [4, 23, 24, 25].

phylogenetic-relationship-africans
Phylogenetic relationship of 44 African and 32 west Eurasian populations determined by a neighbor joining analysis assuming no admixture. Here, the dots of each node represent bootstrap values and the color of each branch indicates language usage of each population. Human_AA human ancestral alleles

The Fulani people are traditionally nomadic pastoralists living across a broad geographic range spanning Sudan, the Sahel, Central, and Western Africa. The Fulani in our study, sampled from Cameroon, clustered with the Afroasiatic-speaking populations in East Africa in the phylogenetic analysis, indicating a potential language replacement from Afroasiatic to Niger-Congo in this population (Fig. 2). Prior studies suggest a complex history of the Fulani; analyses of Y chromosome variation suggest a shared ancestry with Nilo-Saharan and Afroasiatic populations [24], whereas mtDNA indicates a West African origin [26]. An analysis based on autosomal markers found traces of West Eurasian-related ancestry in this population [4], which suggests a North African or East African origin (as North and East Africans also have such ancestry likely related to expansions of farmers and herders from the Near East) and is consistent with the presence at moderate frequency of the −13,910T variant associated with lactose tolerance in European populations [15, 16].

Phylogenetic reconstruction of the relationship of African individuals under a model allowing for migration using TREEMIX [27] largely recapitulates the NJ phylogeny with the exception of the Fulani who cluster near neighboring Niger-Congo-speaking populations with whom they have admixed (Additional file 2: Figure S1). Interestingly, TREEMIX analysis indicates evidence for gene flow between the Hadza and the ancestors of the Ju|‘hoan and Khomani San, supporting genetic, linguistic, and archeological evidence that Khoesan-speaking populations may have originated in Eastern Africa [28, 29, 30].

afroasiatic-niger-congo-admixture
ADMIXTURE analysis of 92 African and 62 West Eurasian individuals. Each bar is an individual and colors represent the proportion of inferred ancestry from K ancestral populations. The bottom bar shows the language classification of each individual. With the increasing of K, the populations are largely grouped by their current language usage

About the Fulani, this is what the referenced study of Y‐chromosome variation among 15 Sudanese populations by Hassan et al. (2008), had to say:

  • Haplogroups A-M13 and B-M60 are present at high frequencies in Nilo-Saharan groups except Nubians, with low frequencies in Afro-Asiatic groups although notable frequencies of B-M60 were found in Hausa (15.6%) and Copts (15.2%).
  • Haplogroup E (four different haplotypes) accounts for the majority (34.4%) of the chromosome and is widespread in the Sudan. E-M78 represents 74.5% of haplogroup E, the highest frequencies observed in Masalit and Fur populations. E-M33 (5.2%) is largely confined to Fulani and Hausa, whereas E-M2 is restricted to Hausa. E-M215 was found to occur more in Nilo-Saharan rather than Afro-Asiatic speaking groups.
  • In contrast, haplogroups F-M89, I-M170, J-12f2, and JM172 were found to be more frequent in the Afro-Asiatic speaking groups. J-12f2 and J-M172 represents 94% and 6%, respectively, of haplogroup J with high frequencies among Nubians, Copts, and Arabs.
  • Haplogroup K-M9 is restricted to Hausa and Gaalien with low frequencies and is absent in Nilo-Saharan and Niger-Congo.
  • Haplogroup R-M173 appears to be the most frequent haplogroup in Fulani, and haplogroup R-P25 has the highest frequency in Hausa and Copts and is present at lower frequencies in north, east, and western Sudan.
  • Haplogroups A-M51, A-M23, D-M174, H-M52, L-M11, OM175, and P-M74 were completely absent from the populations analyzed.
fulfulde-fulani-language
Image modified from “Fulfulde Language Family Report” Author: Annette Harrison; Cartographer: Irene Tucker; SIL International 2003.

This is what David Reich will talk about in the seminar Insights into language expansions from ancient DNA:

In this talk, I will describe how the new science of genome-wide ancient DNA can provide insights into past spreads of language and culture. I will discuss five examples: (1) the spread of Indo-European languages to Europe and South Asia in association with Steppe pastoralist ancestry, (2) the spread of Austronesian languages to the open Pacific islands in association with Taiwanese aboriginal-associated ancestry, (3) the spread of Austroasiatic languages through southeast Asia in association with the characteristic ancestry type that is also represented in western Indonesia suggesting that these languages were once widespread there, (4) the spread of Afroasiastic languages through in East Africa as part of the Pastoral Neolithic farming expansion, and (5) the spread of Na-Dene languages in North America in association with Proto-Paleoeskimo ancestry. I will highlight the ways that ancient DNA can meaningfully contribute to our understanding of language expansions—increasing the plausibility of some scenarios while decreasing the plausibility of others—while emphasizing that with genetic data by itself we can never definitively determine what languages ancient people spoke.

EDIT (3 MAY 2019): Apparently, there was not much to take from the talk:

neolithic-pastoralist-africa
Pastoralist Neolithic in Africa, through a pale-green Sahelo-Sudanian steppe corridor. See full map.

This seminar (and maybe some new paper on the Neolithic expansion in Africa) could shed light on population movements that may be related to the spread of Afroasiatic dialects. Until now, it seems that Bantu peoples have been more interesting for linguistics and archaeology, and South and East Africans for anthropology.

Archaeology in Africa appears to be in its infancy, as is population genomics. From the latest publication by Carina Schlebusch, Population migration and adaptation during the African Holocene: A genetic perspective, a chapter from Modern Human Origins and Dispersal (2019):

The process behind the introduction and development of farming in Africa is still unclear. It is not known how many independent invention events there were in the continent and to which extent the various first instances of farming in northern Africa are linked. Based on the archeological record, it was proposed that at least three regions in Africa may have developed agriculture independently: the Sahara/Sahel (around 7 ka), the Ethiopian highlands (7-4 ka), and western Africa (5-3 ka). In addition to these developments, the Nile River Valley is thought to have adopted agriculture (around 7.2 ka), from the Neolithic Revolution in the Middle East (Chapter 12 – Jobling et al. 2014; Chapter 35, 37 – Mitchell and Lane 2013). From these diverse centers of origin, farmers or farming practices spread to the rest of Africa, with domesticate animals reaching the southern tip of Africa ~2 ka and crop farming ~1,8 ka (Mitchell 2002; Huffman 2007)

african-popularion-movements
Schematic representation of possible migration routes related to the expansion of herders and crop farmers during Holocene times. Arrow color indicate source populations; Brown-Eurasian, Green-western African, Blue-eastern African.

Similar to the case in Europe and the 1990s-2000s wrong haplogroup history based on the modern distribution of R1b, R1a, N, or I2, it is possible that neither of the most often mentioned haplogroups linked to the Afroasiatic expansion, E and J, were responsible for its early spread within Africa, despite their widespread distribution in certain modern Afroasiatic-speaking areas. The fact that such assessments include implausible glottochronological dates spanning up to 20,000 years for the parent language, combined with regional language continuities despite archaeological changes, makes them even more suspicious.

Similar to the case with Indo-Europeans and the “steppe ancestry” concept of the 2010s, it may be that the often-looked-for West Eurasian ancestry among Africans is the effect of recent migrations, unrelated to the Afroasiatic expansion. The results of this paper could be offering another sign of how this ancestry may have expanded only quite recently westwards from East Africa through the Sahel, after the Semitic expansion to the south:

1. From approximately 1000 BC, accompanying Nilo-Saharan peoples.

2. From approximately AD 1500, with the different population movements related to the nomadic Fulani:

sahel-nomadic-sedentary
Image from Sahel in West African History – Oxford Research Encyclopedia of African History.
  • Arguably, since the Fulani caste system wasn’t as elaborate in northern Nigeria, eastern Niger, and Cameroon, these specific groups would be a good example of the admixture with eastern populations, based on the (proportionally) huge amount of slaves they dealt with.
  • Similarly, it could be argued that the castes-based social stratification in most other territories (including Sudan) would have helped them keep a genetic make-up similar to their region of origin in terms of ancient lineages, hence similar to Chadic populations from west to east.

Reich’s assertion of the association of the language expansion with the spread of Pastoral Neolithic is still too vague, but – based on previous publications of ancient DNA in Africa and the Levant – I don’t have high hopes for a revolutionary paper in the near future. Without many samples and proper temporal transects, we are stuck with speculations based on modern distributions and scarce historical data.

fula-people-distribution
A distribution map of Fula people. Dark green: a major ethnic group; Medium: significant; Light: minor. Modified from image by Sarah Welch at Wikipedia.

About the potential genetic make-up of Cameroon before the arrival of the Neolithic, from the recent SAA 84th Annual Meeting (Abstracts in PDF):

Lipson, Mark (Harvard Medical School), Mary Prendergast (Harvard University), Isabelle Ribot (Université de Montréal), Carles Lalueza-Fox (Institute of Evolutionary Biology CSIC-UPF) and David Reich (Harvard Medical School)

[253] Ancient Human DNA from Shum Laka (Cameroon) in the Context of African Population History We generated genome-wide DNA data from four people buried at the site of Shum Laka in Cameroon between 8000–3000 years ago. One individual carried the deeply divergent Y chromosome haplogroup A00 found at low frequencies among some present-day Niger-Congo speakers, but the genome-wide ancestry profiles for all four individuals are very different from the majority of West Africans today and instead are more similar to West-Central African hunter-gatherers. Thus, despite the geographic proximity of Shum Laka to the hypothesized birthplace of Bantu languages and the temporal range of our samples bookending the initial Bantu expansion, these individuals are not representative of a Bantu source population. We present a phylogenetic model including Shum Laka that features three major radiations within Africa: one phase early in the history of modern humans, one close to the time of the migration giving rise to non-Africans, and one in the past several thousand years. Present-day West Africans and some East Africans, in addition to Central and Southern African hunter-gatherers, retain ancestry from the first phase, which is therefore still represented throughout the majority of human diversity in Africa today.

Related

N1c-L392 associated with expanding Turkic lineages in Siberia

haplogroup-n1c-tat

Second in popularity for the expansion of haplogroup N1a-L392 (ca. 4400 BC) is, apparently, the association with Turkic, and by extension with Micro-Altaic, after the Uralic link preferred in Europe; at least among certain eastern researchers.

New paper in a recently created journal, by the same main author of the group proposing that Scythians of hg. N1c were Turkic speakers: On the origins of the Sakhas’ paternal lineages: Reconciliation of population genetic / ancient DNA data, archaeological findings and historical narratives, by Tikhonov, Gurkan, Demirdov, and Beyoglu, Siberian Research (2019).

Interesting excerpts:

According to the views of a number of authoritative researchers, the Yakut ethnos was formed in the territory of Yakutia as a result of the mixing of people from the south and the autochthonous population [34].

These three major Sakha paternal lineages may have also arrived in Yakutia at different times and/ or from different places and/or with a difference in several generations instead, or perhaps Y-chromosomal STR mutations may have taken place in situ in Yakutia. Nevertheless, the immediate common ancestor(s) from the Asian Steppe of these three most prevalent Sakha Y-chromosomal STR haplotypes possibly lived during the prominence of the Turkic Khaganates, hence the near-perfect matches observed across a wide range of Eurasian geography, including as far as from Cyprus in the West to Liaoning, China in the East, then Middle Lena in the North and Afghanistan in the South (Table 3 and Figure 5). There may also be haplotypes closely-related to ‘the dominant Elley line’ among Karakalpaks, Uzbeks and Tajiks, however, limitations in the loci coverage for the available dataset (only eight Y-chromosomal STR loci) precludes further conclusions on this matter [25].

yakutia-haplogroup-n1c
17-loci median-joining network analysis of the original/dominant Elley, Unknown and Omogoy Y-chromosomal STR haplotypes with the YHRD matches from outside Yakutia populations.

According to the results presented here, very similar Y-STR haplotypes to that of the original Elley line were found in the west: Afghanistan and northern Cyprus, and in the east: Liaoning Province, China and Ulaanbaator, Northern Mongolia. In the case of the dominant Omogoy line, very closely matching haplotypes differing by a single mutational step were found in the city of Chifen of the Jirin Province, China. The widest range of similar haplotypes was found for the Yakut haplotype Unknown: In Mongolia, China and South Korea. For instance, haplotypes differing by a single step mutation were found in Northern Mongolia (Khalk, Darhad, Uryankhai populations), Ulaanbaator (Khalk) and in the province of Jirin, China (Han population).

n1c-uralic-altaic-siberia
14-loci median-joining network analysis for the original/dominant Elley (Ell), Unknown Clan
(Vil), Omogoy (Omo), Eurasian (Eur) and Xiongnu (Xuo) Y-chromosomal STR haplotypes and that for a representative ancient DNA sample (Ch0 or DSQ04) from the Upper Xiajiadian Culture
recovered from the Inner Mongolia Autonomous Region, China.

Notably, Tat-C-bearing Y-chromosomes were also observed in ancient DNA samples from the 2700-3000 years-old Upper Xiajiadian culture in Inner Mongolia, as well as those from the Serteya II site at the Upper Dvina region in Russia and the ‘Devichyi gory’ culture of long barrow burials at the Nevel’sky district of Pskovsky region in Russia. A 14-loci Y-chromosomal STR median-joining network of the most prevalent Sakha haplotypes and a Tat-C-bearing haplotype from one of the ancient DNA samples recovered from the Upper Xiajiadian culture in Inner Mongolia (DSQ04) revealed that the contemporary Sakha haplotype ‘Xuo’ (Table 2, Haplotype ID “Xuo”) classified as that of ‘the Xiongnu clan’ in our current study, was the closest to the ancient Xiongnu haplotype (Figure 6). TMRCA estimate for this 14-loci Y-chromosomal STR network was 4357 ± 1038 years or 2341 ± 1038 BCE, which correlated well with the Upper Xiajiadian culture that was dated to the Late Bronze Age (700-1000 BCE).

eurasian-n-subclades
Geographical location of ancient samples belonging to major clade N of the Y-chromosome.

NOTE. Also interesting from the paper seems to be the proportion of E1b1b among admixed Russian populations, in a proportion similar to R1a or I2a(xI2a1).

It is tempting to associate the prevalent presence of N1c-L392 in ancient Siberian populations with the expansion of Altaic, by simplistically linking the findings (in chronological order) near Lake Baikal (Damgaard et al. 2018), Upper Xiajiadian (Cui et al. 2013), among Khövsgöl (Jeong et al. 2018), in Huns (Damgaard et al. 2018), and in Mongolic-speaking Avars (Csáky et al. 2019).

However, its finding among Palaeo-Laplandic peoples in the Kola peninsula ca. 1500 BC (Lamnidis et al. 2018) and among Palaeo-Siberian populations near the Yana River (Sikora et al. 2018) ca. AD 1200 should be enough to accept the hypothesis of ancestral waves of expansion of the haplogroup over northern Eurasia, with acculturation and further expansions in the different regions since the Iron Age (see more on its potential expansion waves).

Also, a simple look at the TMRCA and modern distribution was enough to hypothesize long ago the lack of connection of N1c-L392 with Altaic or Uralic peoples. From Ilumäe et al. (2016):

Previous research has shown that Y chromosomes of the Turkic-speaking Yakuts (Sakha) belong overwhelmingly to hg N3 (formerly N1c1). We found that nearly all of the more than 150 genotyped Yakut N3 Y chromosomes belong to the N3a2-M2118 clade, just as in the Turkic-speaking Dolgans and the linguistically distant Tungusic-speaking Evenks and Evens living in Yakutia (Table S2). Hence, the N3a2 patrilineage is a prime example of a male population of broad central Siberian ancestry that is not intrinsic to any linguistically defined group of people. Moreover, the deepest branch of hg N3a2 is represented by a Lebanese and a Chinese sample. This finding agrees with the sequence data from Hallast et al., where one Turkish Y chromosome was also assigned to the same sub-clade. Interestingly, N3a2 was also found in one Bhutan individual who represents a separate sub-lineage in the clade. These findings show that although N3a2 reflects a recent strong founder effect primarily in central Siberia (Yakutia, Sakha), the sub-clade has a much wider distribution area with incidental occurrences in the Near East and South Asia.

haplogroup-n1a-M2118
Frequency-Distribution Maps of Individual Sub-clades of hg N3a2, by Ilumäe et al. (2016).

The most striking aspect of the phylogeography of hg N is the spread of the N3a3’6-CTS6967 lineages. Considering the three geographically most distant populations in our study—Chukchi, Buryats, and Lithuanians—it is remarkable to find that about half of the Y chromosome pool of each consists of hg N3 and that they share the same sub-clade N3a3’6. The fractionation of N3a3’6 into the four sub-clades that cover such an extraordinarily wide area occurred in the mid-Holocene, about 5.0 kya (95% CI = 4.4–5.7 kya). It is hard to pinpoint the precise region where the split of these lineages occurred. It could have happened somewhere in the middle of their geographic spread around the Urals or further east in West Siberia, where current regional diversity of hg N sub-lineages is the highest (Figure 1B). Yet, it is evident that the spread of the newly arisen sub-clades of N3a3’6 in opposing directions happened very quickly. Today, it unites the East Baltic, East Fennoscandia, Buryatia, Mongolia, and Chukotka-Kamchatka (Beringian) Eurasian regions, which are separated from each other by approximately 5,000–6,700 km by air. N3a3’6 has high frequencies in the patrilineal pools of populations belonging to the Altaic, Uralic, several Indo-European, and Chukotko-Kamchatkan language families. There is no generally agreed, time-resolved linguistic tree that unites these linguistic phyla. Yet, their split is almost certainly at least several millennia older than the rather recent expansion signal of the N3a3’6 sub-clade, suggesting that its spread had little to do with linguistic affinities of men carrying the N3a3’6 lineages.

haplogroup_n3a3
Frequency-Distribution Maps of Individual Subclade N3a3 / N1a1a1a1a1a-CTS2929/VL29.

It was thus clear long ago that N1c-L392 lineages must have expanded explosively in the 5th millennium through Northern Eurasia, probably from a region to the north of Lake Baikal, and that this expansion – and succeeding ones through Northern Eurasia – may not be associated to any known language group until well into the common era.

Related

Common Slavs from the Lower Danube, expanding with haplogroup E1b-V13?

late-iron-age-eastern-europe

Florin Curta has published online his draft for Eastern Europe in the Middle Ages (500-1300), Brill’s Companions to European History, Vol. 10 (2019), apparently due to appear in June.

Some interesting excerpts, relevant for the latest papers (emphasis mine):

The Archaeology of the Early Slavs

(…) One of the most egregious problems with the current model of the Slavic migration is that it is not at all clear where it started. There is in fact no agreement as to the exact location of the primitive homeland of the Slavs, if there ever was one. The idea of tracing the origin of the Slavs to the Zarubyntsi culture dated between the 3rd century BC and the first century AD is that a gap of about 200 years separates it from the Kiev culture (dated between the 3rd and the 4th century AD), which is also attributed to the Slavs. Furthermore, another century separates the Kiev culture from the earliest assemblages attributed to the Prague culture. It remains unclear as to where the (prehistoric) Slavs went after the first century, and whence they could return, two centuries later, to the same region from which their ancestors had left. The obvious cultural discontinuity in the region of the presumed homeland raises serious doubts about any attempts to write the history of the Slavic migration on such a basis. There is simply no evidence of the material remains of the Zarubyntsi, Kiev, or even Prague culture in the southern and southwestern direction of the presumed migration of the Slavs towards the Danube frontier of the Roman Empire.

Moreover, the material culture revealed by excavations of 6th- to 7th-century settlements and, occasionally, cremation cemeteries in northwestern Russia, Belarus, Poland, Moravia, and Bohemia is radically different from that in the lands north of the Danube river, which according to the early Byzantine sources were inhabited at that time by Sclavenes: no settlement layout with a central, open area; no wheel-made pottery or pottery thrown on a tournette; no clay rolls inside clay ovens; few, if any clay pans; no early Byzantine coins, buckles, or remains of amphorae; no fibulae with bent stem, and few, if any bow fibulae. Conversely, those regions have produced elements of material culture that have no parallels in the lands north of the river Danube: oval, trough-like settlement features (which are believed to be remains of above-ground, log-houses); exclusively handmade pottery of specific forms; very large settlements, with over 300 houses; fortified sites that functioned as religious or communal centers; and burials under barrows. With no written sources to inform about the names and identities of the populations living in the 6th and 7th centuries in East Central and Eastern Europe, those contrasting material culture profiles could hardly be interpreted as ethnic commonality. In other words, there is no serious basis for attributing to the Sclavenes (or, at least, to those whom early Byzantine authors called so) any of the many sites excavated in Russia, Belarus, Poland, Moravia, and Bohemia.

slavic-expansion-prague-korchak
Common Slavic expanding with Prague-Korchak from the east…or was it from the west?

Migrations

There is of course evidence of migrations in the 6th and 7th centuries, but not in the directions assumed by historians. For example, there are clear signs of settlement discontinuity in northern Germany and in northwestern Poland. German archaeologists believe that the bearers of the Prague culture who reached northern Germany came from the south (from Bohemia and Moravia), and not from the east (from neighboring Poland or the lands farther to the east). At any rate, no archaeological assemblage attributed to the Slavs either in northern Germany or in northern Poland may be dated earlier than ca. 700. In Poland, settlement discontinuity was postulated, to make room for the new, Prague culture introduced gradually from the southeast (from neighboring Ukraine). However, there is increasing evidence of 6th-century settlements in Lower Silesia (western Poland and the lands along the Middle Oder) that have nothing to do with the Prague culture. Nor is it clear how and when did the Prague culture spread over the entire territory of Poland. No site of any of the three archaeological cultures in Eastern Europe that have been attributed to the Slavs (Kolochin, Pen’kivka, and Prague/Korchak) has so far been dated earlier than the sites in the Lower Danube region where the 6th century sources located the Sclavenes. Neither the Kolochin, nor the Pen’kivka cultures expanded westwards into East Central or Southeastern Europe; on the contrary, they were themselves superseded in the late 7th or 8th century by other archaeological cultures originating in eastern Ukraine. Meanwhile, there is an increasing body of archaeological evidence pointing to very strong cultural influences from the Lower and Middle Danube to the Middle Dnieper region during the 7th century—the opposite of the alleged direction of Slavic migration.

When did the Slavs appear in those regions of East Central and Eastern Europe where they are mentioned in later sources? A resistant stereotype of the current scholarship on the early Slavs is that “Slavs are Slavonic-speakers; Slavonic-speakers are Slavs.”* If so, when did people in East Central and Eastern Europe become “Slavonic speakers”? There is in fact no evidence that the Sclavenes mentioned by the 6th-century authors spoke Slavic (or what linguists now call Common Slavic). Nor can the moment be established (with any precision), at which Slavic was adopted or introduced in any given region of East Central and Eastern Europe.** To explain the spread of Slavic across those regions, some have recently proposed the model of a koiné, others that of a lingua franca. The latter was most likely used within the Avar polity during the last century of its existence (ca. 700 to ca. 800).

*Ziółkowski, “When did the Slavs originate?” p. 211. On the basis of the meaning of the Old Church Slavonic word ięzyk (“language,” but also “people” or “nation”), Darden, “Who were the Sclaveni?” p. 138 argues that the meaning of the name the Slavs gave to themselves was closely associated with the language they spoke.

**Uncertainty in this respect dominates even in recent studies of contacts between Slavic and Romance languages (particularly Romanian), even though such contacts are presumed to have been established quite early (Paliga, “When could be dated ‘the earliest Slavic borrowings’?”; Boček, Studie). Recent studies of the linguistic interactions between speakers of Germanic and speakers of Slavic languages suggest that the adoption of place names of Slavic origin was directly linked to the social context of language contact between the 9th and the 13th centuries (Klír, “Sociální kontext”).

Avars

During the 6th century, the area between the Danube and the Tisza in what is today Hungary, was only sparsely inhabited, and probably a “no man’s land” between the Lombard and Gepid territories. It is only after ca. 600 that this area was densely inhabited, as indicated by a number of new cemeteries that came into being along the Tisza and north of present-day Kecskemét. There can therefore be no doubt about the migration of the Avars into the Carpathian Basin, even though it was probably not a single event and did not involve only one group of population, or even a cohesive ethnic group.

The number of graves with weapons and of burials with horses is particularly large in cemeteries excavated in southwestern Slovakia and in neighboring, eastern Austria. This was a region of special status on the border of the qaganate, perhaps a “militarized frontier.” From that region, the Avar mores and fashions spread farther to the west and to the north, into those areas of East Central Europe in which, for reasons that are still not clear, Avar symbols of social rank were particularly popular, as demonstrated by numerous finds of belt fittings. Emulating the success of the Avar elites sometimes involved borrowing other elements of social representation, such as the preferential deposition of weapons and ornamented belts. For example, in the early 8th century, a few males were buried in Carinthia (southern Austria) with richly decorated belts imitating those in fashion in the land of the Avars, but also with Frankish weapons and spurs. Much like in the Avar-age cemeteries in Slovakia and Hungary, the graves of those socially prominent men are often surrounded by many burials without any grave goods whatsoever.

early-avar-khaganate
Territory of the early Avar Qaganate and the location of the investigated sites in the Carpathian Basin in Csáky et al. (2019).

Carantanians

Carantania was a northern neighbor of the Lombard duchy of Friuli, which was inhabited by Slavs. According to Paul the Deacon, who was writing in the late 780s, those Slavs called their country Carantanum, by means of a corruption of the name of ancient Carnuntum (a former Roman legionary camp on the Danube, between Vienna and Bratislava). Carantanians were regarded as Slavs by the author of a report known as the Conversion of the Bavarians and Carantanians, and written in ca. 870 in order to defend the position of the archbishop of Salzburg against the claims of Methodius, the bishop of Pannonia.94 According to this text, a duke named Boruth was ruling over Carantania when he was attacked by Avars in ca. 740. He called for the military assistance of his Bavarian neighbors. The Bavarian duke Odilo (737–748) obliged, defeated the Avars, but in the process also subdued the Carantanians to his authority. Once Bavarian overlordship was established in Carantania, Odilo took with him as hostages Boruth’s son Cacatius and his nephew Chietmar (Hotimir). Both were baptized in Bavaria. During the 743 war between Odilo and Charles Martel’s two sons, Carloman and Pepin (the Mayors of the Palace in Austrasia and Neustria, respectively), Carantanian troops fought on the Bavarian side. The Bavarian domination cleared the field for missions of conversion to Christianity sent by Virgil, the new bishop of Salzburg (746–784). Many missionaries were of Bavarian origin, but some were Irish monks.

Moravians

Several Late Avar cemeteries dated to the last quarter of the 8th century are known from the lands north of the middle course of the river Danube, in what is today southern Slovakia and the valley of the Lower Morava [see image below]. By contrast, only two cemeteries have so far been found in Moravia (the eastern part of the present-day Czech Republic), along the middle and upper course of the Morava and along its tributary, the Dyje. In both Dolní Dunajovice and Hevlín, the latest graves may be dated by means of strap ends and belt mounts with human figures to the very end of the Late Avar period. (…)

The archaeological evidence pertaining to burial assemblages dated to the early 9th century is completely different. Shortly before or after 800, all traces of cremation—with or without barrows—disappear from the valley of the Morava river and southwestern Slovakia, two regions in which cremation had been the preferred burial rite during the previous centuries. This dramatic cultural change has often been interpreted as a direct influence of both Avar and Frankish burial rites, but it coincides in time with the adoption of Christianity by local elites. In spite of conversion, however, the representation of status through furnished burial continued well into the 9th century. Unlike Avar-age sites in Hungary and the surrounding regions, many men were buried in 9th-century Moravia together with their spurs, in addition to such weapons as battle axes, “winged” lance heads, or swords with high-quality steel blades of Frankish production.

morvaian-sites
Relevant Moravian sites mentioned in Curta’s new book.

When the Magyars inflicted a crushing defeat on the Bavarians at Bratislava (July 4, 907), the fate of Moravia was sealed as well. Moravia and the Moravians disappear from the radar of the written sources, and historians and archaeologists alike believe that the polity collapsed as a result of the Magyar raids.

Magyars

(…) although there can be no doubt about the relations between Uelgi and the sites in Hungary attributed to the first generations of Magyars, those relations indicate a migration directly from the Trans-Ural lands, and not gradually, with several other stops in the forest-steppe and steppe zones of Eastern Europe. In the lands west of the Ural Mountains, the Magyars are now associated with the Kushnarenkovo (6th to 8th century) and Karaiakupovo (8th to 10th century) cultures, and with such burial sites as Sterlitamak (near Ufa, Bashkortostan) and Bol’shie Tigany (near Chistopol, Tatarstan).* However, the same problem with chronology makes it difficult to draw the model of a migration from the lands along the Middle Volga. Many parallels for the so typically Magyar sabretache plates found in Hungary are from that region. They have traditionally been dated to the 9th century, but more recent studies point to the coincidence in time between specimens found in Eastern Europe and those from Hungary.

* Ivanov, Drevnie ugry-mad’iary; Ivanov and Ivanova, “Uralo-sibirskie istoki”; Boldog et al., “From the ancient homelands,” p. 3; Ivanov, “Similarities.” Ivanov, “Similarities,” p. 562 points out that the migration out of the lands along of the Middle Volga is implied by the disappearance of both cultures (Kushnarenkovo and Karaiakupovo) in the mid-9th century. For the Kushnarenkovo culture, see Kazakov, “Kushnarenkovskie pamiatniki.” For the Karaiakupovo culture, see Mogil’nikov, “K probleme.”

Given that the Magyars are first mentioned in relation to events taking place in the Lower Danube area in the 830s, the Magyar sojourn in Etelköz must have been no longer than 60 years or so—a generation. (…)

arrival-of-hungarians-feszty-slavs
A detail of the Arrival of the Hungarians, Árpád Feszty’s and his assistants’ vast (1800 m2) cyclorama, painted to celebrate the 1000th anniversary of the Magyar conquest of Hungary, now displayed at the Ópusztaszer National Heritage Park in Hungary. This specific detail is probably based on the account on The Annals of Fulda, which narrates under the year 894 that the Hungarians crossed the Danube into Pannonia where they “killed men and old women outright and carried off the young women alone with them like cattle to satisfy their lusts and reduced the whole” province “to desert”.

It has become obvious by now that one’s impression of the Magyars as “Easterners” and “steppe-like” was (and still is) primarily based on grave finds, while the settlement material is considerably more aligned with what is otherwise known from other contemporary settlement sites in Central and Southeastern Europe. The dominant feature on the 10th- and 11th-century settlements in Hungary is the sunken-floored building of rectangular plan, with a stone oven in a corner. Similarly, the pottery resulting from the excavation of settlement sites is very similar to that known from many other such sites in Eastern Europe. Moreover, while clear changes taking place in burial customs between ca. 900 and ca. 1100 are visible in the archaeological record from cemeteries, there are no substantial differences between 10th- and the 11th-century settlements in Hungary. (…)

As a matter of fact, the increasing quantity of paleobotanical and zooarchaeological data from 10th-century settlements strongly suggests that the economy of the first generations of Magyars in Hungary was anything but nomadic. To call those Magyars “half-nomad” is not only wrong, but also misleading, as it implies that they were half-way toward civilization, with social changes taking place that must have had material culture correlates otherwise visible in the burial customs.

Comments

The origin of “Slavs” (i.e. that of “Slavonic” as a language, whatever the ancestral Proto-Slavic ethnic make-up was) is almost as complicated as the origin of Albanians, Basques, Balts, or Finns. Their entry into history is very recent, with few reliable sources available until well into the Middle Ages. If you add our ignorance of their origin with the desire of every single researcher or amateur out there to connect them to the own region (or, still worse, to all the regions where they were historically attested), we are bound to find contradictory data and a constantly biased selection of information.

Furthermore, it is extremely complicated to connect any recent population to its ancestral (linguistic) one through haplogroups prevalent today, and just absurd to connect them through ancestral components. This, which was already suspected for many populations, has been confirmed recently for Basques in Olalde et al. (2019) and will be confirmed soon for Finns with a study of the Proto-Fennic populations in the Gulf of Finland.

NOTE. Yes, the “my parents look like Corded Ware in this PCA” had no sense. Ever. Why adult people would constantly engage in that kind of false 5,000-year-old connections instead of learning history – or their own family history – escapes all comprehension. But if something is certain about human nature, is that we will still see nativism and ancestry/haplogroup fetishism for any modern region or modern haplogroups and their historically attested ethnolinguistic groups.

balto-slavic-pca
Genetic structure of modern Balto-Slavic populations within a European context according to the three genetic systems. Image from Kushniarevich et al. (2015)

As you can see from my maps and writings, I prefer neat and simple concepts: in linguistics, in archaeology, and in population movements. Hence my aversion to this kind of infinite proto-historical accounts (and interpretations of them) necessary to ascertain the origins of recent peoples (Slavs in this case), and my usual preference for:

  • Clear dialectal classifications, whether or not they can be as clear cut as I describe them. The only thing that sets Slavic apart from other recent languages is its connection with Baltic, luckily for both. Even though this connection is disputed by some linguists, and the question is always far from being resolved, a homeland of Proto-Balto-Slavic would almost necessarily need to be set to the north of the Carpathian Mountains in the Bronze Age (or at least close to them).
  • NOTE. A dismissal of a connection with Baltic would leave Slavic a still more complicated orphan, and its dialectal classification within Late PIE more dubious. Its union with Balto-Slavic locates it close to Germanic, and thus as a Bronze Age North-West Indo-European dialect close to northern Germany. So bear with me in accepting this connection, or enter the linguistic hell of arguing for Indo-Slavonic of R1a-Z93 mixed with Temematic….

  • A priori “pots = people” assumption, which may lead to important errors, but fewer than the usual “pots != people” of modern archaeologists. The traditional identification of the Common Slavic expansion with the Prague-Korchak culture – however undefined this culture may be – has clear advantages: it may be connected (although admittedly with many archaeological holes) with western cultures expanding east during the Bronze Age, and then west again after the Iron Age, and thus potentially also with Baltic.
  • A simplistic “haplogroup expansion = ethnolinguistic expansion”, which is quite useful for prehistoric migrations, but enters into evident contradictions as we approach the Iron Age. Common Slavs may be speculatively (for all we know) associated with an expansion of recent R1a-M458 lineages – among other haplogroups – from the east, and possibly Balto-Slavic as an earlier expansion of older subclades from the west, as I proposed in A Clash of Chiefs.
r1a-m458-underhill-2015
Modern distribution of R1a-M458, after Underhill et al. (2015).

NOTE. The connection of most R1a-Z280 lineages is more obviously done with ancient Finno-Ugric peoples, as it is clear now (see here and here).

Slavs appeared first in the Danube?

No matter what my personal preference is, one can’t ignore the growing evidence, and it seems that Florin Curta‘s long-lasting view of a Danubian origin of expansion for Common Slavic, including its condition as a lingua franca of late Avars, won’t be easy to reject any time soon:

1) Theories concerning Chernyakhov as a Slavic homeland will apparently need to be fully rejected, due to the Germanic-like ancestry that will be reported in the study by Järve et al. (2019).

EDIT (3 MAY 2019). From their poster Shift in the genetic landscape of the western Eurasian Steppe not due to Scythian dominance, but rather at the transition to the Chernyakhov culture (Ostrogoths) (download PDF):

(…) the transition from the Scythian to the Chernyakhov culture (~2,100–1,700 cal BP) does mark a shift in the Ponto-Caspian genetic landscape. Our results agree well with the Ostrogothic origins of the Chernyakhov culture and support the hypothesis that Scythian dominance was cultural rather than achieved through population replacement.

scythians-chernyakhov-ostrogoths-jarve
PCA of novel and published ancient samples from Scythian/Sarmatian and related groups on the background of modern samples presented as population medians. Δ – ref. 1, ○ – ref. 2, □ – ref. 3, ◊ – this study. Embedded are the locations of some of the samples. Notice the wide cluster formed by the three samples, from Hungarian Scythians in the west to steppe-like peoples in the east.

2) Therefore, unless Przeworsk shows the traditionally described mixture of populations in terms of ancestry and/or haplogroups, it will also be a sign of East Germanic peoples expanding south (and potentially displacing the ancestors of Slavs in either direction, east or south).

It would seem we are stuck in a Danubian vs. Kievan homeland for Common Slavs, then:

3) About the homeland in the Kiev culture, two early Avar females from Szólád have been commented to cluster “among Modern Slavic populations” based on some data in Amorim et al. (2018).

Rather than supporting an origin of Slavs in common with modern Russians, Poles, and Ukranians as observed in the PCA, though, the admixture of AV1 and AV2 (ca. AD 540-640) paradoxically supports an admixture of Modern Slavs of Eastern Europe in common with early Avar peoples (an Altaic-speaking population) and other steppe groups with an origin in East Asia… So this admixture would actually support a western origin of the Common Slavs with which East Asian Avars may have admixed, and whose descendants are necessarily sampled at later times.

pca-medieval-avar-longobards
Procrustes transformed PCA of medieval ancient samples against POPRES imputed SNP dataset. AV1 and Av2 samples have been circled in red. Color coding of medieval samples is same as in Figs 1 and 2. Two letter and three codes for POPRES samples: AL=Albania, AT=Austria, BA=Bosnia-Herzegovina, BE=Belgium, BG=Bulgaria, CH=Switzerland, CY=Cyprus, CZ=Czech Republic, DE=Germany, DK=Denmark, ES=Spain, FI=Finland, FR=France, GB=United Kingdom, GR, Greece, HR=Croatia, HU=Hungary, IE=Ireland, IT=Italy, KS=Kosovo, LV=Latvia, MK=Macedonia, NO=Norway, NL=Netherlands, PL=Poland, PT=Portugal, RO=Romania, SM=Serbia and Montenegro, RU=Russia, Sct=Scotland, SE=Sweden, SI=Slovenia, SK=Slovakia, TR=Turkey, UA=Ukraine.

4) Favouring Curta’s Danubian origin (or even an origin near Bohemia) at the moment are thus:

  • The “western” cluster of Early Slavs from Brandýsek, Bohemia (ca. AD 600-900).
  • Two likely Slavic individuals from Usedom, in Mecklenburg-Vorpommern (AD 1200) show hg. R1a-M458 and E1b-M215 (Freder 2010).
  • An early West Slav individual from Hrádek nad Nisou in Northern Bohemia (ca. AD 1330) also shows E1b-M215 (Vanek et al. 2015).
  • One sample from Székkutas-Kápolnadülő (SzK/239) among middle or late Avars (ca. AD 650-710), a supposed Slavonic-speaking polity, of hg. E1b-V13.
  • Two samples from Karosc (K1/13, and K2/6) among Hungarian conquerors (ca. AD 895-950), likely both of hg. E1b-V13, probably connected to the alliance with Moravian elites.
  • Possibly a West Slavic sample from Poland in the High Middle Ages (see below).

A later Hungarian sample (II/53) from the Royal Basilica, where King Béla was interred, of hg. E1b1, supports the importance of this haplogroup among elite conquerors, although its original relation to the other buried individuals is unknown.

NOTE. You can see all ancient samples of haplogroup E to date on this Map of ancient E samples, with care to identify the proper subclades related to south-eastern Europe. About the ancestral origin of the haplogroup in Europe, you may read Potential extra Iberomaurusian-related gene flow into European farmers, by Chad Rohlfsen.

Even assuming that the R1a sample reported from the late Avar period is of a subclade typically associated with Slavs (I know, circular reasoning here), which is not warranted, we would have already 6 E1b1b vs. 1-2 R1a-M458 in populations that can be actually assumed to represent early Slavonic speakers (unlike many earlier cultures potentially associated with them), clearly earlier than other Slavic-speaking populations that will be sampled in eastern Europe. It is more and more likely that Early Slavs are going to strengthen Curta’s view, and this may somehow complicate the link of Proto-Slavic with eastern European BA cultures like Trzciniec or Lusatian.

NOTE. I am still expecting a clear expansion associated with Prague-Korchak, though, including a connection with bottlenecks based on R1a-M458 in the Middle Ages, whether the expansion is eventually shown to be from the west (i.e. Bohemia -> Prague -> Korchak), or from the east (i.e. Kiev -> Korchack -> Prague), and whether or not this cultural community was later replaced by other ‘true’ Slavonic-speaking cultures through acculturation or population movements.

slavic-origins
Common theories on Slavic origins.. After “The Early Slavs. Culture and Society in Early Medieval Europe” by P. M. Barford, Cornell University Press (2001). Image by Hxseek at Wikipedia.

5) Back to Przeworsk and the “north of the Carpathians” homeland (i.e. between the Upper Oder and the Upper Dniester), but compatible with Curta’s view: Even if Common Slavic is eventually evidenced to be driven by small migrations north and south of the Danube during the Roman Iron Age, before turning into a mostly “R1a-rich” migration or acculturation to the north in Bohemia and then east (which is what this early E1b-V13 connection suggests), this does not dismiss the traditional idea that Late Bronze Age – Iron Age central-eastern Europe was the Proto-Slavic homeland, i.e. likely the Pomeranian culture disturbed by the East Germanic migrations first (in Przeworsk), and the migrations of steppe nomads later (around the Danube).

Even without taking into account the connection with Baltic, the relevance of haplogroup E1b-V13 among Early Slavs may well be a sign of an ancestral population from the northern or eastern Carpathian region, supported by the finding of this haplogroup among the westernmost Scythians. The expansion of some modern E1b-CTS1273 lineages may link Slavic ancestrally with the Lusatian culture, which is an eastern (very specific) Urnfield culture group, stemming from central-east Europe.

An important paper in this respect is the upcoming Zenczak et al., where another hg. E1b1 will be added to the list above: such a sample is expected from Poland (from Kowalewko, Maslomecz, Legowo or Niemcza), either from the Roman Iron Age or Early Middle Ages, close to an early population of likely Scandinavian origin (eight I1 samples), apart from other varied haplogroups, with little relevance of R1a. Whether this E-V13 sample is an Iron Age one (justifying the bottleneck under E-V13 to the south) or, maybe more likely, a late one from the Middle Ages (maybe supporting a connection of the Gothic/Slavic E1b bottleneck with southern Chernyakhov or further west along the Danube) is unclear.

The finding of south-eastern European ancestry and lineages in both, Early Slavs and East Germanic tribes* suggests therefore a Slavonic homeland near (or within) the Przeworsk culture, close to the Albanoid one, as proposed based on topohydronymy. This may point to a complex process of acculturation of different eastern European populations which formed alliances, as was common during the Iron Age and later periods, and which cannot be interpreted as a clear picture of their languages’ original homeland and ancestral peoples (in the case of East Germanic tribes, apparently originally expanding from Scandinavia under strong I1 bottlenecks).

* Iberian samples of the Visigothic period in Spain show up to 25% E1b-V13 samples, with a mixture of haplogroups including local and foreign lineages, as well as some more E1b-V13 samples later during the Muslim period. Out of the two E1b samples from Longobards in Amorim et al. (2018), only SZ18 from Szólád (ca. AD 412-604) is within E1b-V13, in a very specific early branch (SNP M35.2), further locating the expansion of hg. E1b-V13 near the Danube. Samples of haplogroup J (maybe J2a) or G2a among Germanic tribes (and possibly in Poland’s Roman Iron Age / Early Middle Ages) are impossible to compare with early Hungarian ones without precise subclades.

east-slavic-expansion
East Slavic expansion in topo-hydronymy. Image from (Udolph 1997, 2016).

I already interpreted the earlier Slavic samples we had as a sign of a Carpathian origin and very recent bottlenecks under R1a lineages among Modern Slavs:

The finding of haplogroup E1b1b-M215 in two independent early West Slavic individuals further supports that the current distribution of R1a1a1b1a-Z282 lineages in Slavic populations is the product of recent bottlenecks. The lack of a precise subclade within the E1b1b-M215 tree precludes a proper interpretation of a potential origin, but they are probably under European E1b1b1a1b1-L618 subclade E1b1b1a1b1a-V13 (formed ca. 6100 BC, TMRCA ca. 2800 BC), possibly under the mutation CTS1273 (formed ca. 2600 BC, TMRCA ca. 2000 BC), in common with other ancient populations around the Carpathians (see below §viii.11. Thracians and Albanians). This gross geographic origin would support the studies of the Common Slavic homeland based on toponymy (Figure 66), which place it roughly between the Upper Oder and the Upper Dniester, north of the Carpathians (Udolph 1997, 2016).

EDIT (8 APR 2019): Another interesting data is the haplogroup distribution among Modern Slavs and neighbouring peoples (see Wikipedia). For example, the bottleneck seen in Modern Albanians, under Z5017 subclade, also points to an origin of the expansion of E1b-V13 subclades among multiethnic groups around the Lower Danube coinciding with the Roman Iron Age, given the estimates for the arrival of Proto-Albanian close to the Latin and Greek linguistic frontier.

Remarkable is also its distribution among Rusyns, East Slavs from the Carpathians not associated with the Kievan Rus’, isolated thus quite soon from East Slavic expansions to the east. They were reported to show ca. 35% hg. E1b-V13 globally in FTDNA, with a frequency similar to or higher than R1a, in common with South Slavic peoples*, reflecting thus a situation similar to the source of East Slavs before further R1a-based bottlenecks (and/or acculturation events) to the east:

* Although probably due in part to founder effects and biased familial sampling, this should be assumed to be common to all FTDNA sampling, anyway.

rusyns-map
Map showing the full geographic extent of the Rusyn people in Central Europe, prior to World War I (Carpatho Rusyn Society).

Repeating what should be already evident: in complex organizations and/or demographically dense populations (more common since the Iron Age), we can’t expect language change to happen in the same way as during the known Neolithic or Chalcolithic population replacements, be it in Finland, Hungary, Iberia, or Poland. For example, no matter whether Romans (2nd c. BC) brought some R1b-U152 and other Mediterranean lineages to Iberia; Germanic peoples entering Hispania (AD 5th c.) were of typically Germanic lineages or not; Muslims who spoke mainly Berber (AD 8th c.) and were mainly of hg. E1b-M81 (and J?) brought North African ancestry; etc. the language or languages of Iberia changed (or not) with the political landscape: neither with radical population replacements (or full population continuity), nor with the dominant haplogroups’ ancestral language.

Y-chromosome haplogroups are, in those cases, useful for ascertaining a more recent origin of the population. Like the finding of certain R1a-Z645, I2a-L621 & N-L392 lineages among Hungarians shows a recent origin near the Trans-Urals forest-steppes, or the finding of I1, R1b-U106 & E1b-V13 among Visigoths shows a recent origin near the Danube, the finding of Early Slavs (ca. AD 6th-7th c.) originally with small elite groups of hg. R1a-M458 & E1b-V13 from the Lower/Middle Danube – if strengthened with more Early Slavic samples, with Slavonic partially expanding as a lingua franca in some regions – is not necessarily representative of the Proto-Slavic community, just as it is clearly not representative of the later expansion of Slavic dialects. It would be representative, though, of the same processes of acculturation repeated all over Eurasia at least since the Iron Age, where no genetic continuity can be found with ancestral languages.

Related

Scytho-Siberians of Aldy-Bel and Sagly, of haplogroup R1a-Z93, Q1b-L54, and N

iron-age-sakas-aldy-bel-scythians

Recently, a paper described Eastern Scythian groups as “Uralic-Altaic” just because of the appearance of haplogroup N in two Pazyryk samples.

This simplistic identification is contested by the varied haplogroups found in early Altaic groups, by the early link of Cimmerians with the expansion of hg. N and Q, by the link of N1c-L392 in north-eastern Europe with Palaeo-Laplandic, and now (paradoxically) by the clear link between early Mongolic expansion and N1c-L392 subclades.

A new paper (behind paywall) offers insight into the prevalent presence of R1a-Z93 among eastern Scytho-Siberian groups (most likely including Samoyedic speakers in the forest-steppes), and a new hint to the westward expansion of haplogroups Q and N (probably coupled with the so-called “Siberian ancestry”) from the east with different groups of Iron Age steppe nomads:

Genetic kinship and admixture in Iron Age Scytho-Siberians, by Mary et al. Human Genetics (2019).

Interesting excerpts (emphasis mine):

From an archeological and historical point of view, the term “Scythians” refers to Iron Age nomadic or seminomadic populations characterized by the presence of three types of artifacts in male burials: typical weapons, specific horse harnesses and items decorated in the so-called “Animal Style”. This complex of goods has been termed the “Scythian triad” and was considered to be characteristic of nomadic groups belonging to the “Scythian World” (Yablonsky 2001). This “Scythian World” includes both the Classic (or European) Scythians from the North Pontic region (7th–3th century BC) and the Southern Siberian (or Asian) populations of the Scythian period (also called Scytho-Siberians). These include, among others, the Sakas from Kazakhstan, the Tagar population from the Minusinsk Basin (Republic of Khakassia), the Aldy-Bel population from Tuva (Russian Federation) and the Pazyryk and Sagly cultures from the Altai Mountains.

mtdna-scytho-siberians
Proportions of Scythian mtDNA haplogroups. Western (blue) and eastern (pink) Eurasian lineages are equally distributed in the Arzhan Scytho-Siberian sample. The U5a2a1 haplogroup shared between the two Scythian groups studied is in bold

In this work, we first aim to address the question of the familial and social organization of Scytho-Siberian groups by studying the genetic relationship of 29 individuals from the Aldy-Bel and Sagly cultures using autosomal STRs. (…) were obtained from 5 archeological sites located in the valley of the Eerbek river in Tuva Republic, Russia (Fig. 1). All the mounds of this archeological site were excavated but DNA samples were not collected from all of them. 14C dates mainly fall within the Hallstatt radiocarbon calibration plateau (ca. 800–400 cal BC) where the chronological resolution is poor. Only one date falls on an earlier segment of calibration curve: Le 9817–2650 ± 25 BP, i.e. 843–792 cal BC with a probability of 94.3% (using the OxCal v4.3.2 program). This sample (Bai-Dag 8, Kurgan 1, grave 10) is not from one of the graves studied but was used to date the kurgan as a whole.

Y-chromosome haplogroups were first assigned using the ISOGG 2018 nomenclature. In order to improve the precision of haplogroup definition, we also analyzed a set of Y-chromosome SNP (Supplementary Table 2). Nine samples belonged to the R1a-M513 haplogroup (defined by marker M513) and two of these nine samples were characterized as belonging to the R1a1a1b2-Z93 haplogroup or one of its subclades. Six samples belonged to the Q1b1a-L54 haplogroup and five of these six samples belonged to the Q1b1a3-L330 subclade. One sample belonged to the N-M231 haplogroup.

haplogroups-scythian-siberians

The distribution of these haplogroups in the population must be confronted with the prevalence of kinship among the samples. Although five individuals belonged to haplogroup Q1b1a3-L330, three of them (ARZ-T18, ARZ-T19 and ARZ-T20) were paternally related (Fig. 2). It must, therefore, be considered that haplogroup Q1b1a3-L330 is present in three independent instances (given that the remaining two instances exhibit no close familial relationship with other samples or one another). All five were buried on the Eki-Ottug 1 archaeological site (although in two different kurgans).

In the same way, although two groups, of two and three individuals, shared haplotypes belonging to the R1a-M513 haplogroup, these groups likely include a father/son pair (ARZ-T2 and ARZ-T12). Therefore, among nine R1a-M513 men, we found six independent haplotypes, one being present in two independent instances. All R1a-M513 haplotypes, however, including those attributed to the R1a1a1b2-Z93 subclade, only differed by one-step mutations, across 5 loci at most. All R1a-M513 individuals were buried on the same site, Eki-Ottug 2, in a single Kurgan.

y-haplogroups-r1a-n-q1b

Haplogroup R1a-M173 was previously reported for 6 Scytho-Siberian individuals from the Tagar culture (Keyser et al. 2009) and one Altaian Scytho-Siberian from the Sebÿstei site (Ricaut et al. 2004a), whereas haplogroup R1a1a1b2-Z93 (or R1a1a1b-S224) was described for one Scythian from Samara (Mathieson et al. 2015) and two Scytho-Siberians from Berel and the Tuva Republic (Unterländer et al. 2017). On the contrary, North Pontic Scythians were found to belong to the R1b1a1a2 haplogroup (Krzewińska et al. 2018), showing a distinction between the two groups of Scythians. (…) The absence of R1b lineages in the Scytho-Siberian individuals tested so far and their presence in the North Pontic Scythians suggest that these 2 groups had a completely different paternal lineage makeup with nearly no gene flow from male carriers between them.

The seven other male individuals studied in this work were found to carry Eastern Eurasian Y haplogroups Q1b1a and one of its subclades (n = 6) and N (n = 1). Haplogroup Q1b1a-L54 was previously described in four males from the Bronze Age in the Altai Mountains (Hollard et al. 2014, 2018) and was clearly associated with Siberian populations (Regueiro et al. 2013).

The N-M231 haplogroup emerged from haplogroup K in Southern Asia around 21,000 years BCE, maybe in Southern China (Shi et al. 2013; Ilumäe et al. 2016). Previous studies attested to its presence in samples from Neolithic and Bronze Age in China (Li et al. 2011; Cui et al. 2013). Waves of northwestern expansion of this haplogroup are described as beginning during the Paleolithic period (Derenko et al. 2006; Shi et al. 2013) but traces of this expansion in archeological samples were reported only in two Scytho-Siberian males from the Altai (Pilipenko et al. 2015).

The sample of haplogroup N comes from the Aldy-Bel culture (ARZ-T15), from the Eerbek site, but has no radiocarbon date. All Q1b-L330 samples come from the Sagly culture, and three are paternally related. The other Q1b-L54 sample is from other tombs in one kurgan at Aldy Bel.

It seems that – exactly as expected – different waves of steppe nomads brought different lineages at a time (the Iron Age) when many regions incorporated different eastern lineages without necessarily changing language. Just like the expansion of N among Ugrians and Samoyeds, and N1c among Finno-Permic peoples, and like many other lineages expanding with federation-like groups in eastern, central, and western Europe

Related

How the genocidal Yamnaya men loved to switch cultures

yamnaya-expansion-bell-beaker

After some really interesting fantasy full of arrows, it seems Kristiansen & friends are coming back to their most original idea from 2015, now in New Scientist’s recent clickbait Story of most murderous people of all time revealed in ancient DNA (2019):

Teams led by David Reich at Harvard Medical School and Eske Willerslev at the University of Copenhagen in Denmark announced, independently, that occupants of Corded Ware graves in Germany could trace about three-quarters of their genetic ancestry to the Yamnaya. It seemed that Corded Ware people weren’t simply copying the Yamnaya; to a large degree they actually were Yamnayan in origin.

If you think you have seen that movie, it’s because you have. They are at it again, Corded Ware from Yamna, and more “steppe ancestry” = “more Indo-European. It seems we haven’t learnt anything about “Steppe ancestry” since 2015. But there’s more:

Genocidal peoples who “switch cultures”

Burial practices shifted dramatically, a warrior class appeared, and there seems to have been a sharp upsurge in lethal violence. “I’ve become increasingly convinced there must have been a kind of genocide,” says Kristian Kristiansen at the University of Gothenburg, Sweden.

The collaboration revealed that the origin and initial spread of Bell Beaker culture had little to do – at least genetically – with the expansion of the Yamnaya or Corded Ware people into central Europe. “It started in It is in that region that the earliest Bell Beaker objects – including arrowheads, copper daggers and distinctive Bell-shaped pots – have been found, in archaeological sites carbon-dated to 4700 years ago. Then, Bell Beaker culture began to spread east, although the people more or less stayed put. By about 4600 years ago, it reached the most westerly Corded Ware people around where the Netherlands now lies. For reasons still unclear, the Corded Ware people fully embraced it. “They simply take on part of the Bell Beaker package and become Beaker people,” says Kristiansen.

The fact that the genetic analysis showed the Britons then all-but disappeared within a couple of generations might be significant. It suggests the capacity for violence that emerged when the Yamnaya lived on the Eurasia steppe remained even as these people moved into Europe, switched identity from Yamnaya to Corded Ware, and then switched again from Corded Ware to Bell Beaker.

Notice what Kristiansen did there? Yamnaya men “switched identities” into Corded Ware, then “switched identities” into Bell Beakers…So, the most aggresive peoples who have ever existed, exterminating all other Europeans, were actually not so violent when embracing wholly different cultures whose main connection is that they built kurgans (yes, Gimbutas lives on).

NOTE. By the way, just so we are clear, only Indo-Europeans are “genocidal”. Not like Neolithic farmers, or Palaeolithic or Mesolithic populations, or more recent Bronze Age or Iron Age peoples, who also replaced Y-DNA from many regions…

yamnaya-corded-ware-bell-beaker

In fact, there is much stronger evidence that these Yamnaya Beakers were ruthless. By about 4500 years ago, they had pushed westwards into the Iberian Peninsula, where the Bell Beaker culture originated a few centuries earlier. Within a few generations, about 40 per cent of the DNA of people in the region could be traced back to the incoming Yamnaya Beakers, according to research by a large team including Reich that was published this month. More strikingly, the ancient DNA analysis reveals that essentially all the men have Y chromosomes characteristic of the Yamnaya, suggesting only Yamnaya men had children.

“The collision of these two populations was not a friendly one, not an equal one, but one where the males from outside were displacing local males and did so almost completely,” Reich told New Scientist Live in September. This supports Kristiansen’s view of the Yamnaya and their descendants as an almost unimaginably violent people. Indeed, he is about to publish a paper in which he argues that they were responsible for the genocide of Neolithic Europe’s men. “It’s the only way to explain that no male Neolithic lines survived,” he says.

So these unimaginably violent Yamnaya men had children exclusively with their Y chromosomes…but not Dutch Single Grave peoples. These great great steppe-like northerners switched culture, cephalic index…and Y-chromosome from R1a (and others) to R1b-L151 to expand Italo-Celtic From The West™.

It’s hilarious how (exactly like their latest funny episode of PIE from south of the Caucasus) this new visionary idea copied by Copenhagen from amateur friends (or was it the other way around?) had been already rejected before this article came out, in Olalde et al. (2019), and that “Corded Ware=Indo-European” fans have become a parody of themselves.

What’s not to love about 2019 with all this back-and-forth hopping between old and new pet theories?

NOTE. I would complain (again) that the obsessive idea of the Danes is that Denmark CWC is (surprise!) the Pre-Germanic community, so it has nothing to do with “steppe ancestry = Indo-European” (or even with “Corded Ware = Indo-European”, for that matter), but then again you have Koch still arguing for Celtic from the West, Kortlandt still arguing for Balto-Slavic from the east, and – no doubt worst of all – “R1a=IE / R1b=Vasconic / N1c=Uralic” ethnonationalists arguing for whatever is necessary right now, in spite of genetic research.

So prepare for the next episode in the nativist and haplogroup fetishist comedy, now with western and eastern Europeans hand in hand: Samara -> Khvalynsk -> Yamnaya -> Bell Beaker spoke Vasconic-Tyrsenian, because R1b. Wait for it…

Vanguard Yamnaya groups

On a serious note, interesting comment by Heyd in the article:

A striking example of this distinction is a discovery made near the town of Valencina de la Concepción in southern Spain. Archaeologists working there found a Yamnaya-like kurgan, below which was the body of a man buried with a dagger and Yamnaya-like sandals, and decorated with red pigment just as Yamnaya dead were. But the burial is 4875 years old and genetic information suggests Yamnaya-related people didn’t reach that far west until perhaps 4500 years ago. “Genetically, I’m pretty sure this burial has nothing to do with the Yamnaya or the Corded Ware,” says Heyd. “But culturally – identity-wise – there is an aspect that can be clearly linked with them.” It would appear that the ideology, lifestyle and death rituals of the Yamnaya could sometimes run far ahead of the migrants.

NOTE. I have been trying to find which kurgan is this, reviewing this text on the archaeological site, but didn’t find anything beyond occasional ochre and votive sandals, which are usual. Does some reader know which one is it?

yamna-expansion-bell-beakers
Yamna expansion and succeeding East Bell Beaker expansion, without color on Bell Beaker territories. Notice vanguard Yamna groups in blue where East Bell Beakers later emerge. See original image with Bell Beaker territories.

Notice how, if you add all those vanguard Yamna findings of Central and Western Europe, including this one from southern Spain, you begin to get a good idea of the territories occupied by East Bell Beakers expanding later. More or less like vanguard Abashevo and Sintashta finds in the Zeravshan valley heralded the steppe-related Srubna-Andronovo expansions in Turan…

It doesn’t seem like Proto-Beaker and Yamna just “crossed paths” at some precise time around the Lower Danube, and Yamna men “switched cultures”. It seems that many Yamna vanguard groups, probably still in long-distance contact with Yamna settlers from the Carpathian Basin, were already settled in different European regions in the first half of the 3rd millennium BC, before the explosive expansion of East Bell Beakers ca. 2500 BC. As Heyd says, there are potentially many Yamna settlements along the Middle and Lower Danube and tributaries not yet found, connecting the Carpathian Basin to Western and Northern Europe.

These vanguard groups would have more easily transformed their weakened eastern Yamna connections with the fashionable Proto-Beaker package expanding from the west (and surrounding all of these loosely connected settlements), just like the Yamna materials from Seville probably represent a close cultural contact of Chalcolithic Iberia with a Yamna settlement (the closest known site with Yamna traits is near Alsace, where high Yamna ancestry is probably going to be found in a Bell Beaker R1b-L151 individual).

This does not mean that there wasn’t a secondary full-scale migration from the Carpathian Basin and nearby settlements, just like Corded Ware shows a secondary (A-horizon?) migration to the east with R1a-Z645. It just means that there was a complex picture of contacts between Yamna and European Chalcolithic groups before the expansion of Bell Beakers. Doesn’t seem genocidal enough for a popular movie, tho.

Related

ASoSaH Reread (I): Y-DNA haplogroups among Indo-Europeans (apart from R1b-L23)

eneolithic-early-admixture-steppe-ancestry

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).

This is mainly a reread of iv.2. Indo-Anatolians and vi.1. Disintegrating Indo-Europeans.

Indo-Anatolians and Late Indo-Europeans

I have often written about R1b-L23 as the majority haplogroup among Late Proto-Indo-Europeans (see my predictions for 2018 and my summary of 2018), but always expected other haplogroups to pop up somewhere along the way, in Khvalynsk, in Repin, in Yamna, and in Bell Beakers (see e.g. the post on common fallacies of R1a/IE-fans).

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.

y-dna-khvalynsk
Y-DNA samples from Khvalynsk and neighbouring cultures. See full version here.

Y-DNA haplogroups

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.

chalcolithic-early-y-dna
Y-DNA samples from Yamnaya and neighbouring cultures. See full version here.

‘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.

PCA-caucasus-steppe-all
Yamnaya (Indo-European peoples) and their evolution in the steppes, together with North Pontic (eventually Uralic) peoples.Notice how little Indo-European ancestry changes from Khvalynsk (Indo-Anatolian) to Yamna Hungary (North-West Indo-Europeans) Image modified from Wang et al. (2018). See more on the evolution of “steppe ancestry”.

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.

Related

“Steppe ancestry” step by step: Khvalynsk, Sredni Stog, Repin, Yamna, Corded Ware

dzudzuana_pca-large

Wang et al. (2018) is obviously a game changer in many aspects. I have already written about the upcoming Yamna Hungary samples, about the new Steppe_Eneolithic and Caucasus Eneolithic keystones, and about the upcoming Greece Neolithic samples with steppe ancestry.

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.

anatolia-neolithic-steppe-eneolithic
Image modified from Wang et al. (2018). Marked are in orange: equivalent Steppe_Maykop ADMIXTURE; in red, approximate limit of Anatolia_Neolithic ancestry found in Yamna populations; in blue, Corded Ware-related groups. “Modelling results for the Steppe and Caucasus cluster. Admixture proportions based on (temporally and geographically) distal and proximal models, showing additional Anatolian farmer-related ancestry in Steppe groups as well as additional gene flow from the south in some of the Steppe groups as well as the Caucasus groups.”

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.

PCA-caucasus-steppe-samara

This is what Steppe_Maykop looks like, different from Steppe_Eneolithic:

steppe-maykop-admixture

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:

PCA-khvalynsk-steppe

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.

PCA-sredni-stog-steppe

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.

steppe-ancestry-admixture-sredni-stog

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.

afanasevo-admixture

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:

PCA-repin-yamna

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:

yamnaya-admixture

5. Corded Ware

Corded Ware represents a quite homogeneous expansion of a late Sredni Stog population, compatible with the traditional location of Proto-Corded Ware peoples in the steppe-forest/forest zone of the Dnieper-Dniester region.

PCA-latvia-ln-steppe

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:

sintashta-poltavka-andronovo-admixture

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.

steppe-ancestry-admixture-latvia

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.

Conclusion

Yamna and Corded Ware show a similar “steppe ancestry” due to convergence. I have said so many times (see e.g. here). This was clear long ago, just by looking at the Y-chromosome bottlenecks that differentiate them – and Tomenable noticed this difference in ADMIXTURE from the supplementary materials in Mathieson et al. (2017), well before Wang et al. (2018).

This different stock stems from (1) completely different ancestral populations + (2) different, long-lasting Y-chromosome bottlenecks. Their similarities come from the two neighbouring cultures admixing with similar populations.

If all this does not mean anything, and each lab was going to support some pre-selected archaeological theories from the 1960s or the 1980s, coupled with outdated linguistic models no matter what – Anthony’s model + Ringe’s glottochronological tree of the early 2000s in the Reich Lab; and worse, Kristiansen’s CWC-IE + Germano-Slavonic models of the 1940s in the Copenhagen group – , I have to repeat my question again:

What’s (so much published) ancient DNA useful for, exactly?

See also

Related