Magyar tribes brought R1a-Z645, I2a-L621, and N1a-L392(xB197) lineages to the Carpathian Basin

hungarian-conquerors-turks

The Nightmare Week of “N1c=Uralic” proponents continues, now with preprint Y-chromosome haplogroups from Hun, Avar and conquering Hungarian period nomadic people of the Carpathian Basin, by Neparaczki et al. bioRxiv (2019).

Abstract:

Hun, Avar and conquering Hungarian nomadic groups arrived into the Carpathian Basin from the Eurasian Steppes and significantly influenced its political and ethnical landscape. In order to shed light on the genetic affinity of above groups we have determined Y chromosomal haplogroups and autosomal loci, from 49 individuals, supposed to represent military leaders. Haplogroups from the Hun-age are consistent with Xiongnu ancestry of European Huns. Most of the Avar-age individuals carry east Eurasian Y haplogroups typical for modern north-eastern Siberian and Buryat populations and their autosomal loci indicate mostly unmixed Asian characteristics. In contrast the conquering Hungarians seem to be a recently assembled population incorporating pure European, Asian and admixed components. Their heterogeneous paternal and maternal lineages indicate similar phylogeographic origin of males and females, derived from Central-Inner Asian and European Pontic Steppe sources. Composition of conquering Hungarian paternal lineages is very similar to that of Baskhirs, supporting historical sources that report identity of the two groups.

Interesting excerpts (emphasis mine):

All N-Hg-s identified in the Avars and Conquerors belonged to N1a1a-M178. We have tested 7 subclades of M178; N1a1a2-B187, N1a1a1a2-B211, N1a1a1a1a3-B197, N1a1a1a1a4-M2118, N1a1a1a1a1a-VL29, N1a1a1a1a2-Z1936 and the N1a1a1a1a2a1c1-L1034 subbranch of Z1936. The European subclades VL29 and Z1936 could be excluded in most cases, while the rest of the subclades are prevalent in Siberia 23 from where this Hg dispersed in a counter-clockwise migratory route to Europe (…). All the 5 other Avar samples belonged to N1a1a1a1a3-B197, which is most prevalent in Chukchi, Buryats, Eskimos, Koryaks and appears among Tuvans and Mongols with lower frequency.

haplogroup-n-pca
First two components of PCA from Hg N1a subbranch distribution in 51 populations including Avars and Conquerors. Colors indicate geographic regions. Three letter codes are given in Supplementary Table S5.

By contrast two Conquerors belonged to N1a1a1a1a4-M2118, the Y lineage of nearly all Yakut males, being also frequent in Evenks, Evens and occurring with lower frequency among Khantys, Mansis and Kazakhs.

Three Conqueror samples belonged to Hg N1a1a1a1a2-Z1936 , the Finno-Permic N1a branch, being most frequent among northeastern European Saami, Finns, Karelians, as well as Komis, Volga Tatars and Bashkirs of the Volga-Ural region.Nevertheless this Hg is also present with lower frequency among Karanogays, Siberian Nenets, Khantys, Mansis, Dolgans, Nganasans, and Siberian Tatars.

The west Eurasian R1a1a1b1a2b-CTS1211 subclade of R1a is most frequent in Eastern Europe especially among Slavic people. This Hg was detected just in the Conqueror group (K2/18, K2/41 and K1/10). Though CTS1211 was not covered in K2/36 but it may also belong to this sub-branch of Z283.

Hg I2a1a2b-L621 was present in 5 Conqueror samples, and a 6th sample form Magyarhomorog (MH/9) most likely also belongs here, as MH/9 is a likely kin of MH/16 (see below). This Hg of European origin is most prominent in the Balkans and Eastern Europe, especially among Slavic speaking groups. It might have been a major lineage of the Cucuteni-Trypillian culture and it was present in the Baden culture of the Chalcolithic Carpathian Basin.

hungarian-conquerors-y-dna
Image modified from the paper, with drawn red square around lineages of likely Ugric origin, and squares around R1a-Z93, R1a-Z283, N1a-Z1936, and N1a-M2004 samples. Y-Hg-s determined from 46 males grouped according to sample age, cemetery and Hg. Hg designations are given according to ISOGG Tree 2019. Grey shading designate distinguished individuals with rich grave goods, color shadings denote geographic origin of Hg-s according to Fig. 1. For samples K3/1 and K3/3 the innermost Hg defining marker U106* was not covered, but had been determined previously.

We identified potential relatives within Conqueror cemeteries but not between them. The uniform paternal lineages of the small Karos3 (19 graves) and Magyarhomorog (17 graves) cemeteries approve patrilinear organization of these communities. The identical I2a1a2b Hg-s of Magyarhomorog individuals appears to be frequent among high-ranking Conquerors, as the most distinguished graves in the Karos2 and 3 cemeteries also belong to this lineage. The Karos2 and Karos3 leaders were brothers with identical mitogenomes 11 and Y-chromosomal STR profiles (Fóthi unpublished). The Sárrétudvari commoner cemetery seems distinct from the others, containing other sorts of European Hg-s. Available Y-chromosomal and mtDNA data from this cemetery suggest that common people of the 10th century rather represented resident population than newcomers. The great diversity of Y Hg-s, mtDNA Hg-s, phenotypes and predicted biogeographic classifications of the Conquerors indicate that they were relatively recently associated from very diverse populations.

Surprising about the Hungarian conquerors – although in line with the historical accounts – is the varied patrilineal origin of clans, including Q1a, G2a2b, I1, E1b1b, R1b, J1, or J2 – some of which (depending on specific lineages) may have appeared earlier in the Carpathian Basin or south-eastern Europe.

However, out of the 27 conqueror elite samples, 17 are of haplogroups most likely related to Ugric populations beyond the Urals: R1a-Z645, I2-L621, and two specific N1a-L392 lineages (see below). In fact, there are three high-ranking conqueror elites of hg. I2-L621 (one of them termed a “leader”, brother to an unpublished leader of Karos3, and all of them possibly family), one of hg. R1a-Z280, one of hg. R1a-Z93 (which should be added to the Árpáds), and one of hg. N1a-Z1936, which gives a good idea of the ruling class among the elite Ugric settlers.

NOTE. The Q1a sample is also likely to be found in the mixed population of the West Siberian forest-steppes, since it was found in Mesolithic-Neolithic samples from eastern Europe to Lake Baikal, and in Bronze Age Siberian groups, although admittedly it may have formed part of an Avar Transtisza group, or even earlier Hunnic or Scythian groups along the steppes. Without precise subclades it’s impossible to know.

arrival-of-hungarians-arpad
The seven chieftains of the Hungarians, detail of Arrival of the Hungarians, from 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. Image from Wikipedia.

I2a-L621

I2a-L621 (xS17250) or I2a1b2 in the old nomenclature, is found in 6 early conquerors (including one leader), on a par with R1a and N samples. This haplogroup is found widely distributed in ancient samples, due to its early split (formed ca. 9200 BC, TMRCA ca. 4500 BC) and expansion, probably with Neolithic populations. I can’t seem to find samples of this early haplogroup from the Carpathian Basin, as mentioned in the text, although it wouldn’t be strange, because it appears also in Neolithic Iberia, and in modern populations from western Europe.

Nevertheless, I2a-L621 samples seem to be concentrated mainly in Mesolithic-Neolithic cultures of Fennoscandia, and appeared also in Sikora et al. (2017) in a sample of the High Middle Ages from Sunghir (ca. AD 1100-1200), probably from the Vladimir-Suzdalian Rus’, in a region where clearly tribes of Volga Finns were being assimilated at the time. The reported SNP call by Genetiker is A16681 (see Yfull), deep within I2a-CTS10228. It is possibly also behind a modern Saami from Chalmny Varre (ca. AD 1800) of hg. I2a in Lamnidis et al. (2018).

Lacking precise subclades from Hungarian conquerors this is pure speculation, but modern samples may also point to I2a-CTS10228 (formed ca. 3100 BC, TMRCA ca. 1800 BC) as a Finno-Ugric lineage in common with R1a, which must have expanded to the Urals and beyond with eastern Corded Ware groups or (more likely) succeeding cultures. This is in line with the association of certain I2a lineages with modern Uralic peoples or populations from their historical regions in eastern Europe, and linked thus to the most likely homeland of Uralians in the eastern European forests:

uralic-groups-haplogroup-r1a
Additional file 6: Table S5. Y chromosome haplogroup frequencies in Eurasia. Modified by me: in bold haplogroup N1c and R1a from Uralic-speaking populations, with those in red showing where R1a is the major haplogroup. Observe that all Uralic subgroups – Finno-Permic, Ugric, and Samoyedic – have some populations with a majority of R1a, and also of I lineages. Data from Tambets et al. (2018).

R1a-Z645

Regarding the important question of the ethnic makeup of Ugric populations stemming from the Urals, the most interesting (and expected) data is the presence of R1a-Z645 lineages among high-ranking conquerors, in particular four R1a-Z280 subclades proper of Finno-Ugrians.

This proves that, in line with the old split and expansion of R1a-CTS1211 (formed ca. 2600 BC, TMRCA ca. 2400 BC), and its finding in Bronze Age Fennoscandian samples, only some late R1a-Z280 (xZ92) lineages (see Z280 on YFull) may show a clear identification with early acculturated Uralic speakers, with the main early acculturated Balto-Slavic R1a haplogroup remaining R1a-M458.

I recently hypothesized this late connection of Slavs with very specific R1a-Z280 (xZ92) lineages based on analyses of modern populations (like Slovenians), because the connection of ancient Finno-Ugrians with modern Z92 samples was already evident:

(…) subclades of hg. R1a1a1b1a2-Z280 (xR1a1a1b1a2a-Z92) seem to have also been involved in early Slavic expansions, like R1a1a1b1a2b3a-CTS3402 (formed ca. 2200 BC, TMRCA ca. 2200 BC), found among modern West, South, and East Slavic populations and in Fennoscandia, prevalent e.g. among modern Slovenians which points to a northern origin of its expansion (Maisano Delser et al. 2018).

This finding also supports the expected shared R1a-Z280 lineages among ancient Finno-Ugric populations, as predicted from the study of modern Permic and Ugric peoples in Dudás et al. (2019).

r1a-z282-z280-z2125-distribution
Modified image, from Underhill et al. (2015). Spatial frequency distributions of Z282 (green) and Z93 (blue) affiliated haplogroups. Notice the distribution of R1a-Z280 (xZ92), i.e. R1a-M558, compared to the ancient Finno-Ugric distribution.

Furthermore, while we don’t have precise R1a-Z93 lineages to compare with the new Hunnic sample reported, we already know that some archaic R1a-Z2124 subclades stem from the forest-steppe areas of the Cis- and Trans-Urals, and the two newly reported R1a-Z93 Hungarian conqueror elites, like those of the Árpád dynasty, probably belong to them.

There is an obvious lack of continuity in specific paternal lineages among the Hunnic, the Avar, and the Conqueror periods, which makes any simplistic identification of all R1a-Z93 lineages as stemming from Avars, Huns, or the Iron Age Pontic-Caspian steppes clearly flawed. Comparing R1a-Z93 in Hungarian Conquerors with Huns is like comparing them with samples of the Srubna or earlier periods… Similarly, comparing the Hunnic R1b-U106 or the early Avar I1 to later Hungarian samples is not warranted without precise subclades, because they most likely correspond to different Germanic populations: Goths among Huns, then Longobards, then likely peoples descended from Franks and Irish Monks (the latter with R1b-P312).

N1a-L392

Second behind R1a subclades are, as expected, N1a-L392 (N1c in the old nomenclature).

Avars are dominated by a specific N1a-L392 subclade, N1a-B197, as we recently discovered in Csáky et al. (2019).

Hungarian conquerors show three N1a-Z1936 subclades, which is known to stem from the northern Ural region, including the Arctic (likely Palaeo-Laplandic peoples) and cross-stamped cultures of the northern Eurasian forests.

haplogroup_n3a4
Frequency-Distribution Maps of Individual Subclade N3a4 / N1a1a1a1a2-Z1936, probably with the Samic (first) and Fennic (later) expansions into Paleo-Lakelandic and Palaeo-Laplandic territories.

On the other hand, the two N1a-M2118 lineages are more clearly associated with Palaeo-Siberian populations east of the Urals, but became incorporated into the Ugric stock in the Trans-Urals region probably in the same way as N1a-Z1936, by infiltration from (and acculturation of) hunter-gatherers of forest and taiga cultures.

NOTE. You can read more about the infiltration of N1a lineages in the recent post Corded Ware—Uralic (IV): Hg R1a and N in Finno-Ugric and Samoyedic expansions, and in the specific sections for each Uralic group in A Clash of Chiefs.

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

Conclusion

The picture offered by the paper on Hungarian Conquerors, while in line with historical accounts of multi-ethnic tribes incorporating regional lineages, shows nevertheless patrilineal clans clearly associated with Uralic peoples, in a distribution which could have been easily inferred from ancient Trans-Uralian forest-steppe cultures and modern samples (even regarding I2a-L621).

In spite of this, there is a great deal of discussion in the paper about specific N1a subclades in Hungarian conquerors, while the presence of R1a-Z280 (among early Magyar elites!) is interpreted, as always, as recently acculturated Slavs. This is sadly coupled with the simplistic identification of I2a-L621 as of local origin around the Carpathians.

The introduction of the paper to the history of Hungarians is also weird, for example giving credibility to the mythic accounts of the Árpád dynasty’s origin in Attila, which is in line, I guess, with what the authors intended to support all along, i.e. the association of Magyars with Turks from the Eurasian steppes, which they are apparently willing to achieve by relating them to haplogroup R1a-Z93

The conclusion is thus written to appease modern nation-building myths more than anything else, like many other papers before it:

It is generally accepted that the Hungarian language was brought to the Carpathian Basin by the Conquerors. Uralic speaking populations are characterized by a high frequency of Y-Hg N, which have often been interpreted as a genetic signal of shared ancestry. Indeed, recently a distinct shared ancestry component of likely Siberian origin was identified at the genomic level in these populations, modern Hungarians being a puzzling exception36. The Conqueror elite had a significant proportion of N Hgs, 7% of them carrying N1a1a1a1a4-M2118 and 10% N1a1a1a1a2-Z1936, both of which are present in Ugric speaking Khantys and Mansis. At the same time none of the examined Conquerors belonged to the L1034 subclade of Z1936, while all of the Khanty Z1936 lineages reported in 37 proved to be L1034 which has not been tested in the 23 study. Population genetic data rather position the Conqueror elite among Turkic groups, Bashkirs and Volga Tatars, in agreement with contemporary historical accounts which denominated the Conquerors as “Turks”. This does not exclude the possibility that the Hungarian language could also have been present in the obviously very heterogeneous, probably multiethnic Conqueror tribal alliance.

So, back to square one, and new circular reasoning: If ancient populations from north-eastern Europe believed to represent ancient Finno-Ugrians are of R1a-Z645 lineages, it’s because they were not Finno-Ugric speakers. If ancient and modern populations known to be of Finno-Ugric language show clear connections with R1a-Z645, it’s because they are “multi-ethnic”.

The only stable basis for discussion in genetic papers, apparently, is the own making of geneticists, with their traditional 2000s “R1a=Indo-European” and “N1c=Uralic”, coupled with national beliefs. It does not matter how many predictions based on that have been proven wrong, or how many predictions based on the Corded Ware = Uralic expansion have been proven right.

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

R1a-Z280 and R1a-Z93 shared by ancient Finno-Ugric populations; N1c-Tat expanded with Micro-Altaic

Two important papers have appeared regarding the supposed link of Uralians with haplogroup N.

Avars of haplogroup N1c-Tat

Preprint Genetic insights into the social organisation of the Avar period elite in the 7th century AD Carpathian Basin, by Csáky et al. bioRxiv (2019).

Interesting excerpts (emphasis mine):

After 568 AD the Avars settled in the Carpathian Basin and founded the Avar Qaganate that was an important power in Central Europe until the 9th century. Part of the Avar society was probably of Asian origin, however the localisation of their homeland is hampered by the scarcity of historical and archaeological data.

Here, we study mitogenome and Y chromosomal STR variability of twenty-six individuals, a number of them representing a well-characterised elite group buried at the centre of the Carpathian Basin more than a century after the Avar conquest.

The Y-STR analyses of 17 males give evidence on a surprisingly homogeneous Y chromosomal composition. Y chromosomal STR profiles of 14 males could be assigned to haplogroup N-Tat (also N1a1-M46). N-Tat haplotype I was found in four males from Kunpeszér with identical alleles on at least nine loci. The full Y-STR haplotype I, reconstructed from AC17 with 17 detected STRs, is rare in our days. Only nine matches were found among haplotypes in YHRD database, such as samples from the Ural Region, Northern Europe (Estonia, Finland), and Western Alaska (Yupiks). We performed Median Joining (MJ) network analysis using N-Tat haplotypes with ten shared STR loci (Fig. 3, Table S9). All modern N-Tat samples included in the network had derived allele of L708 as well. Haplotype I (Cluster 1 in Fig. 3) is shared by eight populations on the MJ network among the 24 identical haplotypes. Cluster 1 represents the founding lineage, as it is described in Siberian populations, because this haplotype is shared by the most populations and it is more diverse than Cluster 2.

Nine males share N-Tat haplotype II (on a minimum of eight detected alleles), all of them buried in the Danube-Tisza Interfluve. We found 30 direct matches of this N-Tat haplotype II in the YHRD database, using the complete 17 STR Y-filer profile of AC1, AC12, AC14, AC15, AC19 samples. Most hits came from Mongolia (seven Buryats and one Khalkh) and from Russia (six Yakuts), but identical haplotypes also occur in China (five in Xinjiang and four in Inner Mongolia provinces). On the MJ network, this haplotype II is represented by Cluster 2 and is composed of 45 samples (including 32 Buryats) from six populations (Fig. 3).

y-str-haplogroup-n-mongolian-ugrians
Median Joining network of 162 N-Tat Y-STR haplotypes Allelic information of ten Y-STR loci were used for the network. Only those Avar samples were included, which had results for these ten Y-STR loci. The founder haplotype I (Cluster 1) is shared by eight populations including three Mongolian, three Székely, three northern Mansi, two southern Mansi, two Hungarian, eight Khanty, one Finn and two Avar (AC17, AC26) chromosomes. Haplotype II (Cluster 2) includes 45 haplotypes from six populations studied: 32 Buryats, two Mongolians, one Székely, one Uzbek, one Uzbek Madjar, two northern Mansi and six Avars (AC1, AC12, AC14, AC15, AC19 and KSZ 37). Haplotype III (indicated by a red arrow) is AC8. Information on the modern reference samples is seen in Table S9.

A third N-Tat lineage (type III) was represented only once in the Avar dataset (AC8), and has no direct modern parallels from the YHRD database. This haplotype on the MJ network (see red arrow in Fig. 3) seems to be a descendent from other haplotype cluster that is shared by three populations (two Buryat from Mongolia, three Khanty and one Northern Mansi samples). This haplotype cluster also differs one molecular step (locus DYS393) from haplotype II. We classified the Avar samples to downstream subgroup N-F4205 within the N-Tat haplogroup, based on the results of ours and Ilumäe et al.18 and constructed a second network (Fig. S4). The N-F4205 network results support the assumption that the N-Tat Avar samples belong to N-F4205 subgroup (see SI chapter 1d for more details).

Based on our calculation, the age of accumulated STR variance (TMRCA) within N-Tat lineage for all samples is 7.0 kya (95% CI: 4.9 – 9.2 kya), considering the core haplotype (Cluster 1) to be the founding lineage. Y haplogroup N-Tat was not detected by large scale Eurasian ancient DNA studies but it occurs in late Bronze Age Inner Mongolia and late medieval Yakuts, among them N-Tat has still the highest frequency.

Two males (AC4 and AC7) from the Transtisza group belong to two different haplotypes of Y-haplogroup Q1. Both Q1a-F1096 and Q1b-M346 haplotypes have neither direct nor one step neighbour matches in the worldwide YHRD database. A network of the Q1b-M346 haplotype shows that this male had a probable Altaian or South Siberian paternal genetic origin.

EDIT (5 APR 2019): The paper offers an interesting late sample before the arrival of Hungarian conquerors, although we don’t know which precise lineage the sample belongs to:

One sample in our dataset (HC9) comes from this population, and both his mtDNA (T1a1b) and Y chromosome (R1a) support Eastern European connections. (…) Furthermore, we excluded sample HC9 from population-genetic statistical analyses because it belongs to a later period (end of 7th – early 9th centuries)

Apparently, then, results are consistent with what was already known from studies of modern populations:

According to Ilumäe et al. study, the frequency peak of N-F4205 (N3a5-F4205) chromosomes is close to the Transbaikal region of Southern Siberia and Mongolia, and we conclude that most Avar N-Tat chromosomes probably originated from a common source population of people living in this area, completely in line with the results of Ilumäe et al.

haplogroup_n1
Geographic-Distribution Map of hg N3 from Ilumäe et al.

Finno-Ugrians share haplogroup R1a-Z280

Another paper, behind paywall, Genetic history of Bashkirian Mari and Southern Mansi ethnic groups in the Ural region, by Dudás et al. Molecular Genetics and Genomics (2019).

Interesting excerpts (emphasis mine):

Y‑chromosome diversity

The most frequent haplogroups of the Bashkirian Maris were N1b-P43 (42%), R1a-Z280 (16%), R1a-Z93 (16%), N1c-Tat (13%), and J2-M172 (7%). Furthermore, subgroup R1b-M343 accounted for 4% and I2a-P37 covered 2% of the lineages. None of the Mari N1c Y chromosomes belonged to the N1c subgroups investigated (L1034, VL29, Z1936).

In the case of the Southern Mansi males, the most frequent haplogroups were N1b-P43 (33%), N1c-L1034 (28%) and R1a-Z280 (19%). The frequencies of the remaining haplogroups were as follows: R1a-M458 (6%), I1-L22 (3%), I2a-P37 (3%), and R1b-P312 (3%). The haplotype and haplogroup diversities of the Bashkirian Mari group were 0.9929 and 0.7657, whereas these values for the Southern Mansi were 0.9984 and 0.7873, respectively. The results show that, in both populations, haplotypes are much more diverse than haplogroups.

bashkir-mari-southern-mansi
Haplogroup frequencies of the Bashkirian Mari and the Southern Mansi ethnic groups in Ural region

Genetic structure

(..) the studied Bashkirian Mari and Southern Mansi population groups formed a compact cluster along with two Khanty, Northern Mansi, Mari, and Estonian populations based on close Fst-genetic distances (< 0.05), with nonsignificant p values (p > 0.05) except for the Estonian population. All of these populations belong to the Finno-Ugric language family. Interestingly, the other Mansi population studied by Pimenoff et al. (2008) (pop # 38) was located a great distance from the Southern Mansi group (0.268). In addition, the Bashkir population (pop # 6) did not show a close genetic affinity to the Bashkirian Mari group (0.194), even though it is the host population. However, the Russian population from the Eastern European region of Russia (pop # 49) showed a genetic distance of 0.055 with the Southern Mansi group. All Hungarian speaking populations (pops 13, 22, 23, 24, 50, and 51) showed close genetic affinities to each other and to the neighbouring populations, but not to the two studied populations.

y-dna-hungarians-ugric-mansi
Multidimensional scaling (MDS) plot constructed on Fstgenetic distances of Y haplogroup frequencies of 63 populations compared. The haplogroup frequency data used for population comparison together with references are seen in Online Resource 2 (ESM_2). Pairwise Fst-genetic distances and p values between 63 populations were calculated as shown in Online Resource 3 (ESM_3) Fig. 4 Multidimensional scaling (MDS) plot constructed on Rstgenetic distances of 10 STR-based Y haplotype frequencies of 21 populations compared. Image modified to include labels of modern populations.

Phylogenetic analysis

Median-joining networks were constructed for:

N-P43 (earlier N1b):

(…) TMRCA estimates for this haplogroup were made for all P43 samples (n = 157) 8.7 kya (95% CI 6.7–10.8 kya), for the N-P43 Asian.

N1c-Tat:

(…) 75% of Buryats belonged to Haplotype 2, indicating that the Buryats studied by us is a young and isolated population (Bíró et al. 2015). Bashkirian Mari samples derive from Haplotype 2 via Haplotype 3 (see dark purple circles on the top of Fig. 6a). Haplotype 3 contained six males (2 Buryat, 1 Northern Mansi, and 3 Khanty samples from Pimenoff et al. 2008). The biggest Bashkirian Mari haplotype node (3 Mari samples) was positioned three mutational steps away from Haplotype 1 and the remaining Mari samples can be derived from this haplotype. Southern Mansi haplotypes were scattered within the network except for two, which formed a smaller haplotype node with two Northern Mansi and two Khanty samples from Pimenoff et al. (2008).

n1c-n-tat-uralic-ugric
Median-Joining Networks (MJ) of 153 N-Tat (a) and 26 N-L1034 (b) haplotypes constructed. The circle sizes are proportional to the haplotype frequencies. The smallest area is equivalent to one individual. For N-Tat network, we used data from Southern Mansi (n = 11), Bashkirian Mari (n = 6) samples with Hungarian (n = 12), Hungarian speaking Székely (n = 6), Northern Mansi (n = 14), Mongolian (n = 16), Buryat (n = 44), Finnish (n = 13), Uzbek Madjar (n = 2), Uzbek (n = 3), Khanty (n = 4) populations studied earlier by us (Fehér et al. 2015; Bíró et al. 2015) and Khanty (n = 18) and Mansi (n = 4) studied by Pimenoff et al. (2008)

R1a-Z280 haplotypes, shared by Maris, Mansis, and Hungarians, hence ancient Finno-Ugrians:

The founder R1a-Z280 haplotype was shared by four samples from four populations (1 Bashkirian Mari; 1 Southern Mansi; 1 Hungarian speaking Székely; and 1 Hungarian), as presented in Fig. 7 (Haplotype 1). Haplotype 2 included five males (3 Bashkirian Mari and 2 Hungarian), as it can be seen in Fig. 7. Haplotype 4 included two shared haplotypes (1 Bashkirian Mari and one Hungarian speaking Csángó). The remaining two Bashkirian Mari haplotypes differ from the founder haplotype (Haplotype 1) by two mutational steps via Hungarian or Hungarian and Bashkirian Mari shared haplotypes. Beside Haplotype 1, the remaining Southern Mansi haplotypes were shared with Hungarians (Haplotype 5 or turquoise blue and red-coloured circles above Haplotype 7) or with Hungarians and Hungarian speaking Székely group (Haplotypes 3, 5, and 6). Haplotype 7 included ten Hungarian speakers (Hungarian, Székely, and Csángó). One Hungarian and one Uzbek Khwarezm shared haplotype can be found in Fig. 7 as well (red and white-coloured circle). All the other haplotypes were scattered in the network. The age of accumulated STR variation within R1a-Z280 lineage for 93 samples is estimated to be 9.4 kya (95% CI 6.5–12.4 kya) considering Haplotype 1 (Fig. 7) to be the founder.

r1a-z280-ugrians
Median-Joining Networks (MJ) of 93 R1a-Z280 haplotypes constructed. The circle sizes are proportional to the haplotype frequencies. The smallest area is equivalent to one individual. We used haplotype data from Bashkirian Mari (n = 7), Southern Mansi (n = 7), Hungarian (n = 52), Hungarian speaking Székely (n = 11), Hungarian speaking Csángó (n = 10), Uzbek Ferghana (n = 2), Uzbek Tashkent (n = 1), Uzbek Khwarezm (n = 1) and Northern Mansi (n = 2) populations

R1a-Z93 as isolated lineages among Permic and Ugric populations:

Figure 8 depicts an MJ network of R1a-Z93* samples using 106 haplotypes from the 14 populations (Fig. 8). All of the Bashkirian Mari samples (7 haplotypes) formed a very isolated branch and differed from the one Hungarian haplotype (Fig. 8, see Haplotype 1) by seven mutational steps as well from two Uzbek Tashkent samples (see Haplotype 3). Another Hungarian sample shared two haplotypes of Uzbek Khwarezm samples in Haplotype 4. This haplotype can be derived from Haplotype 3 (Uzbek Tashkent). Haplotype 2 included one Hungarian and one Khakassian male. The remaining three Hungarian haplotypes are outliers in the network and are not shared by any sample. The other population samples included in the network either form independent clusters such as Altaians, Khakassians, Khanties, and Uzbek Madjars or were scattered in the network. The age of accumulated STR variation (TMRCA) within R1a-Z93* lineage for 106 samples is estimated as 11.6 kya (95% CI 9.3–14.0 kya) considering an Armenian haplotype (Fig. 8, “A”) to be the founder and the median haplotype.

r1a-z93-ugrians
Median-Joining Networks (MJ) of 106 R1a-Z93 haplotypes constructed. The circle sizes are proportional to the haplotype frequencies. The smallest area is equivalent to one individual. We used the next haplotype data: 7 Bashkirian Mari, 6 Khanty, 4 Uzbek Madjar, 5 Uzbek Ferghana, 9 Uzbek Tashkent, 7 Uzbek Khwarezm, 2 Mongolian, 2 Buryat, 6 Hungarian samples tested by us for this study or published earlier (Bíró et al. 2015) and populations (3 Armenian; 3 Afghan Tajik;
16 Altaian; 24 Khakassian; 12 Kyrgyz) from Underhill et al. (2015)

Comments

The results of modern populations for N (especially N1c) subclades show really wide clusters and ancient TMRCA, consistent with their known ancient and wide distribution in northern and eastern Eurasian groups, and thus with infiltration of different lineages with eastern nomads (and northern Arctic populations) coupled with later bottlenecks, as well as acculturation of groups.

EDIT (2 APR): Interesting is the specific subclade to which ancient Mongolic-speaking Avars belong (information from Yfull) N1c-F4205 (TMRCA ca. 500 BC), subclade of N1c-Y6058 (formed ca. 2800 BC, TMRCA ca. 2800 BC). This branch also gives the “European” branch N1c-CTS10760 (formed ca. 2800 BC, TMRCA ca. 2100 BC), and is subclade of a branch of N1c-L392 (formed ca. 4400 BC, TMRCA ca. 2800 BC). A northern expansion of N1c-L392 is probably represented by its branch N1c-Z1936 (formed ca. 2800, TMRCA ca. 2100 BC), the most likely candidate to appear in the Kola Peninsula in the Bronze Age as the Palaeo-Laplandic population (see here). Read more about potential routes of expansion of haplogroup N.

On the other hand, R1a-Z280 lineages form a tight cluster connecting Permic with Ugric groups, with R1a-Z93 showing early isolation (probably) between Cis-Urals and Trans-Urals regions. While both Corded Ware lineages in Finno-Ugrians are most likely related to the Abashevo expansion through Seima-Turbino and the Andronovo-like Horizon (and potentially later Eurasian expansions), a plausible hypothesis would be that Finno-Ugrians are related to an expansion of R1a-Z283 haplogroups (we already knew about the Finno-Permic connection), while the ancient connection between Permians and Hungarians with R1a-Z93 would correspond to this haplogroup’s potentially tighter link with an early Samoyedic split.

I don’t think that an explosive expansion of eastern Corded Ware groups of R1a-Z645 lineages will show a clear-cut division of haplogroups among Eastern Uralic groups, though, and culturally I doubt we will have such a clear image, either (similar to how the explosive expansion of Bell Beakers cannot be easily divided by regional/language group into R1b-L151 subclades before the known bottlenecks). Relevant in this regard are the known Z93 samples from the Árpád dynasty.

Nevertheless, this data may represent a slightly more recent wave of R1a-Z280 lineages linked to the expansion of Ugric into the Trans-Uralian region, after their split from Finno-Permic, still in close contact with Indo-Iranians in Poltavka and Sintashta-Potapovka, evident from the early and late Indo-Iranian borrowings, during a common period when Samoyedic had already separated.

Such a “Z283 over Z93” layer in the Trans-Urals (and Cis-Urals?) forest-steppes would be similar to the apparent replacement of Z284 by Z282 in the Eastern Baltic during the Bronze Age (possibly with the second or Estonian Battle Axe wave or, much more likely during later population movements). Such an early R1a-Z93 split could potentially be supported also by the separation into bottlenecks under “Northern” (R1a-Z283) Finno-Ugric-speaking Abashevo-related groups and “Southern” (R1a-Z93) acculturated Indo-Iranian-speaking Abashevo migrants developing Sintashta-Potapovka admixing with Poltavka R1b-Z2103 herders.

r1a-z282-z280-z2125-distribution
Modified image, from Underhill et al. (2015). Spatial frequency distributions of Z282 (green) and Z93 (blue) affiliated haplogroups.. Notice the potential Finno-Ugric-associated distribution of Z282 (especially R1a-M558, a Z280 subclade), the expansion of R1a-Z2123 subclades with Central Asian forest-steppe groups.

Conclusion

Let’s review some of the most common myths about Hungarians (and Finno-Ugrians in general) repeated ad nauseam, side by side with my assertions:

❌ N (especially N1c-Tat) in ancient and modern samples represent the True Uralic™ N1c peoples including Magyar tribes? Nope.

✅ Ancient N (especially N1c-Tat) lineages among Uralic populations expanded relatively recently, and differently in different regions (including eastern steppe nomads and northern arctic populations) not associated with a particular language or language group? Yep (read the series on Corded Ware = Uralic expansion).

❌ Modern Hungarian R1a-Z280 lineages represent the majority of the native population, poor Slavic ‘peasants’ from the Carpathian Basin, forcibly acculturated by a minority of bad bad Hungarian hordes? Nope.

✅ Modern Hungarian R1a-Z280 subclades represent Ugric lineages in common with ancient R1a-Z645 Finno-Ugric populations from north-eastern Europe and the Trans-Urals? Yep (see Avars and Ugrians).

❌ Modern Hungarian R1a-Z93 lineages represent acculturated Iranian/Turkic peoples from the steppes? Not likely.

✅ Modern Hungarian R1a-Z93 lineages represent a remnant of the expansion of Corded Ware to the east, potentially more clearly associated with Samoyedic? Much more likely.

finno-ugric-haplogroup-n
Map of archaeological cultures in north-eastern Europe ca. 8th-3rd centuries BC. [The Mid-Volga Akozino group not depicted] Shaded area represents the Ananino cultural-historical society. Fading purple arrows represent likely stepped movements of subclades of haplogroup N for centuries (e.g. Siberian → Ananino → Akozino → Fennoscandia [N-VL29]; Circum-Arctic → forest-steppe [N1, N2]; etc.). Blue arrows represent eventual expansions of Uralic peoples to the north. Modified image from Vasilyev (2002).

Sooo, the theory of a “diluted” Y-DNA in Modern Hungarians from originally fully N-dominated conquerors subjugating native R1a-Z280 Slavs from the Carpathian Basin is not backed up by genetic studies? The ethnic Iranian-Turkic R1a-Z93 federation in the steppes that ended up speaking Magyar is not real?? Who would’ve thunk.

Another true story whose rejection in genetics could not be predicted, like, not at all.

Totally unexpected, too, the drift of “R1a=IE” fans with the newest genetic findings towards a Molgen-like “Yamna/R1b = Vasconic-Caucasian”, “N1c = Uralic-Altaic”, and “R1a = the origin of the white world in Mother Russia”. So much for the supposed interest in “Steppe ancestry” and fancy statistics.

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

Yekaterinovsky Cape, a link between the Samara culture and early Khvalynsk

ekaterinovsky-cape

We already had conflicting information about the elite individual from the Yekaterinovsky Cape and the materials of his grave, which seemed quite old:

For the burial of 45 in the laboratory of the University of Pennsylvania, a 14C date was obtained: PSUAMS-2880 (Sample ID 16068)> 30 kDa gelatin Russia. 12, Ekaterinovka Grave 45 14C age (BP) 6325 ± 25 δ 13C (‰) –23.6 δ15 N (‰) 14.5. The results of dating suggest chronological proximity with typologically close materials from Yasinovatsky and Nikolsky burial grounds (Telegini et al. 2001: 126). The date obtained also precedes the existing dates for the Khvalynsk culture (Morgunova 2009: 14–15), which, given the dominance of Mariupol traits of the burial rite and inventory, confirms its validity. However, the date obtained for human bones does not exclude the possibility of a “reservoir effect” when the age can increase three or more centuries (Shishlin et al. 2006: 135–140).

Now the same date is being confirmed by the latest study published on the site, by Korolev, Kochkina, and Stachenkov (2019) and it seems it is really going to be old. Abstract (in part the official one, in part newly translated for clarity):

For the first time, pottery of the Early Eneolithic burial ground Ekaterinovsky Cape is published. Ceramics were predominantly located on the sacrificial sites in the form of compact clusters of fragments. As a rule, such clusters were located above the burials, sometimes over the burials, some were sprinkled with ocher. The authors have identified more than 70 vessels, some of which have been partially reconstructed. Ceramic was made with inclusion of the crushed shell into molding mass. The rims of vessels had the thickened «collar»; the bottoms had a rounded shape. The ornament was located on the rims and the upper part of the potteries. Fully decorated vessels are rare. The vessels are ornamented with prints of comb and rope stamps, with small pits. A particularity of ceramics ornamentation is presented by the imprints of soft stamps (leather?) or traces of leather form for the making of vessels. The ornamentation, made up of «walking comb» and incised lines, was used rarely as well as the belts of pits made decoration under «collar» of a rim. Some features of the ceramics decoration under study relate it with ceramics of the Khvalynsk culture. The ceramics of Ekaterinovsky Cape burial ground is attributed by the authors to the Samara culture. The ceramic complex under study has proximity to the ceramics from Syezzhe burial ground and the ceramics of the second phase of Samara culture. The chronological position is determined by the authors as a later period than the ceramics from the Syezzhe burial ground, and earlier than the chronological position of ceramics of the Ivanovka stage of the Samara culture and the Khvalynsk culture.

ekaterinovsky-cape-pottery
Ceramics from Ekaterinovsky Cape burial ground. 1–2, 4–5, 7–11 – ceramics from aggregations; 3, 6 – ceramics from the cultural layer.

More specifically:

Based on ceramic fragments from a large vessel from a cluster of sq.m. 14, the date received was: SPb-2251–5673 ± 120 BP. The second date was obtained in fragments from the aggregation [see picture above] from the cluster of sq.m. 45–46: SPb-2252–6372 ± 100 BP. The difference in dating indicates that the process of determining the chronology of the burial ground is far from complete, although we note that the earlier date almost coincided with the date obtained from the human bone from individual 45 (Korolev, Kochkina, Stashenkov, 2018, p. 300).

Therefore, the ceramics of the burial ground Ekaterinovsky Cape possess an originality that determines the chronological position of the burial ground between the earliest materials of the burial type in Syezzhe and the Khvalynsk culture. Techno-typological features of dishes make it possible to attribute it to the Samara culture at the stage preceding the appearance of Ivanovska-Khvalynsk ceramics.

It seems that this site showed cultural influences from the upstream region near the Kama-Vyatka interfluve, too, according to Korolev, Kochkina, Stashenkov, and Khokhlov (2018):

In 2017, excavation of burial ground Ekaterinovsky Cape were continued, located in the area of the confl uence of the Bezenchuk River in the Volga River. During the new excavations, 14 burials were studied. The skeleton of the buried were in a position elongated on the back, less often – crooked on the back with knees bent at the knees. In one burial (No. 90), a special position of the skeleton was recorded. In the burial number 90 in the anatomical order, parts of the male skeleton. This gave grounds for the reconstruction of his original position in a semi-sitting position with the support of elbows on the bottom of the pit. Noteworthy inventory: on the pelvic bones on the left lay a bone spoon, near the right humerus, the pommel of a cruciform club was found. A conclusion is made about the high social status of the buried. The results of the analysis of the burial allow us to outline the closest circle of analogies in the materials of Khvalynsky I and Murzikhinsky burial grounds.

Important sites mentioned in both papers and in this text:

To sum up, it seems that the relative dates we have used until now have to be corrected: older Khvalynsk I Khvalynsk II individuals, supposedly dated ca. 5200-4000 BC (most likely after 4700 BC), and younger Yekaterinovsky individuals, supposedly of the fourth quarter of the 5th millennium (ca. 4250-4000 BC), are possibly to be considered, in fact, roughly reversed, if not chronologically, at least culturally speaking.

Interestingly, this gives a new perspective to the presence of a rare fish- or reptile-headed pommel-scepter, which would be natural in a variable period of expansion of the horse and horse-related symbolism, a cultural trait rooted in the Samara culture attested in Syezzhe before the unification of the symbol of power under the ubiquitous Khvalynsk-Suvorovo horse-headed scepters and related materials.

ekaterinovsky-cape-pommel-mace
Ekaterinovsky Cape Burial Ground. Inventory of the burial no 90: 1, 2 – stone pommel of the mace; 3, 4 – bone article.

The Khvalynsk chieftain

If the reported lineages from Yekaterinovsky Cape are within the R1b-P297 tree, but without further clades, as Yleaf comparisons may suggest, there is not much change to what we have, and R1b-M269 could actually represent a part of the local population, but also incomers from the south (e.g. the north Caspian steppe hunter-gatherers like Kairshak), the east (with hunter-gatherer pottery), or the west near the Don River (in contact with Mariupol-related cultures, as the authors inferred initially from material culture).

Just like R1a-M417 became incorporated into the Sredni Stog groups after the Novodanilovka-Suvorovo expansion, probably as incoming hunter-gatherer pottery groups from the north admixing with peoples of “Steppe ancestry”, R1b-M269 lineages might have expanded explosively only during the Repin expansion, and maybe (like R1b-L51 later) they formed just a tiny part of the clans that dominated the steppe during the Khvalynsk-Novodanilovka community.

On the other hand, the potential finding of various R1b-M269/L23 samples in Yekaterinovsky Cape (including an elite individual) would suggest now, as it was supported in the original report by Mathieson et al. (2015), that these ancient R1b lineages found in the Volga – Ural region are in fact most likely all R1b-M269 without enough coverage to obtain proper SNP calls, which would simplify the picture of Neolithic expansions (yet again). From the supplementary materials:

10122 / SVP35 (grave 12). Male (confirmed genetically), age 20-30, positioned on his back with raised knees, with 293 copper artifacts, mostly beads, amounting to 80% of the copper objects in the combined cemeteries of Khvalynsk I and II. Probably a high-status individual, his Y-chromosome haplotype, R1b1, also characterized the high-status individuals buried under kurgans in later Yamnaya graves in this region, so he could be regarded as a founder of an elite group of patrilineally related families. His MtDNA haplotype H2a1 is unique in the Samara series.

khvalynsk-cemetery
Khvalynsk cemetery and grave gifts. Grave 90 contained copper beads and rings, a harpoon, flint blades, and a bird-bone tube. Both graves (90 and 91) were partly covered by Sacrificial Deposit 4 with the bones from a horse, a sheep, and a cow. Center: grave goods from the Khvalynsk cemetery-copper rings and bracelets, polished stone mace heads, polished stone bracelet, Cardium shell ornaments, boars tusk chest ornaments, flint blades, and bifiacial projectile points. Bottom: shell-tempered pottery from the Khvalynsk cemetery. After Agapov, Vasiliev, and Pestrikova 1990; and Ryndina 1998, Figure 31. Modified from Anthony (2007).

This remarkable Khvalynsk chieftain, whose rich assemblage may correspond to the period of domination of the culture all over the Pontic-Caspian steppes, has been consistently reported as of hg. R1b-L754 in all publications, including Wang et al. (2018/2019) tentative SNP calls in the supplementary materials (obtained with Yleaf, as the infamous Narasimhan et al. 2018 samples), but has been variously reported by amateurs as within the R1b-M73, R1b-V88, or (lately) R1b-V1636 trees, which makes it unlikely that quality of the sample is allowing for a proper SNP call.

The fact that Mathieson et al. (2015) considered it a member of the R1b-M269 clans appearing later in Yamna seems on point right now, especially if samples from Yekaterinovka are all within this tree. The relevance of R1b-L23 in the expansion of Repin and Yamna is reminiscent of the influence of successful clans among Yamna offshoots, such as Bell Beakers, and among Bell Beaker offshoots during the Bronze Age all over Europe.

Taking these younger expansions as example, it seems quite likely based on cultural links that (at least part of) the main clans of Khvalynsk were of R1b-M269 lineage, stemming from a R1b-dominated Samara culture, in line with the known succeeding expansions and the expected strictly patriarcal and patrilineal society of Proto-Indo-Europeans, which would have exacerbated the usual reduction in Y-chromosome haplogroup variability that happens during population expansions, and the aversion towards foreign groups while the culture lasted.

pontic-steppe-neolithic
Cultures of the Pontic-Caspian steppes and forest-steppes and surrounding areas during the Neolithic.

The finding of R1b-L23 in Yekaterinovka, associated with the Samara culture, before or during the Khvalynsk expansion, and close to the Khvalynsk site, would make this Khvalynsk chieftain most likely a member of the M269 tree (paradoxically, the only R1b-L754 branch amateurs have not yet reported for it). Similarly, the sample of a “Samara hunter-gatherer” of Lebyazhinka, of hg. R1b-P297, could also be under this tree, just like most R1b-M269 from Yamna are downstream from R1b-L23, and most reported R1b-M269 or R1b-L23 from Bell Beakers are under R1b-L151.

On the other hand, we know of the shortcomings of attributing a haplogroup expansion to the best known rulers, such as the famous lineages previously wrongly attributed to Niall of the Nine Hostages or Genghis Khan. The known presence of R1b-V1636 up to modern Greeks would be in line with an ancient steppe expansion that we know will show up during the Neolithic, although it could also be a sign of a more recent migration from the Caucasus. The presence of a sister clade of R1b-L23, R1b-PF7562, among modern Balkan populations, may also be attributed to a pre-Yamna steppe expansion.

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

On SNP calls

I reckon that even informal reports on SNP calls, like any other analyses, should be offered in full: not only with a personal or automatic estimation of the result, but with a detailed explanation of the good, dubious, and bad calls, alternatives to that SNP estimation, and a motivated reasoning of why one branch should be preferred over others. Downloading a sample and giving an instruction using a free software tool is never enough, as it became crystal clear recently for the hilariously biased and flawed qpAdm reports on Dutch Bell Beakers as the ‘missing link’ between Corded Ware and Bell Beakers…

Another example I can recall is the report of a R1a-Z93 subclade in the R1a-M417 sample ca. 4000 BC from Alexandria, which seems rather unlikely, seeing how this subclade must have split and expanded explosively with R1a-Z645 to the east with eastern Corded Ware groups, i.e. 1,000 years later, just like Z282 lineages expanded mainly to the north-east. But then again, as with the Khvalynsk chieftain, I have only seen indirect reports of that supposed SNP (including Y26+!), so we should just stick with its officially reported R1a-M417 lineage. This upstream haplogroup was, in fact, repeated with Yleaf’s tentative estimates in Wang et al. (2019) supplementary materials…

The combination of inexperienced, biased, or simply careless design, analyses, and reports, including SNP calls and qpAdm analyses (whether in forums or publications), however well-intentioned (or not) they might be, are hindering a proper analysis of data, adding to the difficulties we already have due to the scarcity of samples, their limited coverage, and the lack of proper context.

Some people like to repeat ad nauseam that archaeology and/or linguistics are ‘not science’ whenever they don’t fit their beliefs and myths based on haplogroup and/or ancestry. But it’s becoming harder and harder to rely on certain genetic data, too, and on their infinite changing interpretations, much more than it is to rely on linguistic and archaeological research, including data, assessments, and discussions that are open for anyone to review…if one is truly interested in them.

The Pazyryk culture spoke a “Uralic-Altaic” language… because haplogroup N

Matrilineal and patrilineal genetic continuity of two iron age individuals from a Pazyryk culture burial, by Tikhonov, Gurkan, Peler, & Dyakonov, Int J Hum Genet (2019).

Relevant excerpts (emphasis mine):

Of particular interest to the current study are the archaeogenetic investigations associated with the exemplary mound 1 from the Ak-Alakha-1 site on the Ukok Plateau in the Altai Republic (Polosmak 1994a; Pilipenko et al. 2015). This typical Pazyryk “frozen grave” was dated around 2268±39 years before present (Bln-4977) (Gersdorff and Parzinger 2000). Initial anthropological findings suggested an undisturbed dual inhumation comprising “a middle-aged European- type man” and “a young European-type woman”, both of whom presumably had a high social status among the Pazyryk elite (Polosmak 1994a). In contrast, recent archaeogenetic investigations revealed somewhat contradicting results since analyses at both the amelogenin gene and Y-chromosome short tandem repeat (Y-STR) loci clearly established that both Scythians were actually males and had paternal and maternal lineages that are typically associated with eastern Eurasians (Pilipenko et al. 2015). Through the use of mitochondrial, autosomal and Y-chromosomal DNA typing systems, it was possible to not only investigate the potential relationships between the two ancient Scythians but also to gather initial phylogenetic and phylogeographic information on their paternal and maternal lineages (Pilipenko et al. 2015).

Based on the Y-STR data available, the two Ak-Alakha-1 Scythians had an in silico haplogroup assignment of N, which first appeared in southeastern Asia and then expanded in southern Siberia (Rootsi et al. 2007; Pilipenko et al. 2015).

Current study aims to investigate the geographical distributions of the ancient and contemporary matches and close genetic variants of the maternal and paternal lineages observed in the two Scythians from the exemplary Ak-Alakha-1 kurgan.

tikhonov-scythian-modern
Geographic distribution of the exact matches with the Scythian (PZ1) Y-STR (17-loci) and mtDNA (HVR1) haplotypes detailed in Tables 1a and 1b. Boundaries of the Altai Republic within the Russian Federation are shown with dashed lines, along with an approximate position of the Ak-Alakha-1 burial site, which is denoted with an ‘x’ on the map. Countries shaded in gray refer to those that have full 17-loci Y-STR and/or mtDNA HVR1 match(es) with the PZ1 haplotypes. Inset in the top and bottom left corners are the Altai and Uzbekistan maps, respectively, both scaled-up to allow better representation of the samples derived from these countries. There were no other exact matches from around parts of the globe that are not shown on the map, except for a single contemporary mtDNA haplotype from US, which presumably belonged to an ‘East Asian’ individual. Inset in the top right corner provides a scale for the number of haplotypes observed, but only up to three samples, which is valid for the entire map as well as the inset maps, irrespective of the differences in the scales of the actual map and inset maps themselves. For sample pools larger than three, the same linear scale provided on the inset in the top right corner still applies; please refer to Tables 1a and b for actual sample pool sizes. Samples are depicted on the entire map and the insets maps with circles and diamonds for the Y-STR and mtDNA haplotypes, respectively. Black and white coloring for samples depict whether the haplotype(s) are contemporary or ancient, respectively. Location of the PZ1 mtDNA and Y-STR haplotypes are shown on top of each other.

In response to aggressive Xiongnu expansion into the Altai region around the 2nd century BCE, some members of the Pazyryk culture may have started moving up North, and eventually reached the Vilyuy River at the beginning of 1st century CE. Notably, there is clear population continuity between the Uralic people such as Khants, Mansis and Nganasans, Paleo-Siberian people such as Yukaghirs and Chuvantsi, and the Pazyryk people even when considering just the two mtDNA and Y-STR haplotypes from the Ak-Alakha-1 mound 1 kurgan (Tables 1a, b, Table 2, Fig. 1). These concepts are also in agreement with the famous Yakut ethnographer Ksenofontov, who suggested that technologies associated with ferrous metallurgy were brought to the Vilyuy Valley at around 1st century CE by the first (proto)Turkic-speaking pioneers (Ksenofontov 1992). Yakut ethnogenesis per se possibly involved two major stages, the first being the proto-Turkic epoch through the arrival of Scytho-Siberian culture originating from Southern Siberia, such as that associated with the Pazyryk culture and the second being the proper Turkic epoch.

Nomadic peoples from the Central Asian steppes are East Iranian speakers whenever they are of haplogroup R1a, but “Uralic-Altaic” speakers whenever they are of haplogroup N. True story.

So they followed a haplogroup ca. 37,000 years old, in a sample dated some 2,300 years ago, whose precise subclade and ancient history is (yet) unknown, compared it to present-day populations, and the result is that they spoke “Uralic-Altaic” because haplogroup N and continuity. Sound familiar? Yep, it’s the kind of reasoning you might be reading right now about Iberian Bell Beakers, about Bell Beakers, or even about Yamna and their relationship to a Vasconic-Caucasian language, based on haplogroup R1b in modern Basques. Another true story.

Anyway, based on the multi-ethnic federations created during this time, and on the ancestral components visible in the different groups (see a post on Karasuk by Chad Rohlfsen), the Pazyryk culture’s language is unknown, and it could be, as a matter of fact (apart from the obvious East Iranian connection):

We also know that haplogroup N and Siberian ancestry expanded into cultures of Northern Eurasia precisely with the creation of the new social paradigm of chiefdoms and alliances, roughly at the same time as Scythians expanded, with the first sample of haplogroup N in Hungary appearing with Cimmerians.

finno-ugric-haplogroup-n
Map of archaeological cultures in north-eastern Europe ca. 8th-3rd centuries BC. [The Mid-Volga Akozino group not depicted] Shaded area represents the Ananino cultural-historical society. Fading purple arrows represent likely stepped movements of subclades of haplogroup N for centuries (e.g. Siberian → Ananino → Akozino → Fennoscandia [N-VL29]; Circum-Arctic → forest-steppe [N1, N2]; etc.). Blue arrows represent eventual expansions of Uralic peoples to the north. Modified image from Vasilyev (2002).

While the study of modern populations is interesting, the problem I have with the paper is the reasoning of “language of ancient haplogroups based on modern populations”, and especially with the concept of “Uralic-Altaic”, and the highly hypothetic “Proto-Turkic” nomadic steppe pastoralists before “Hunnic Turkic” (which is itself questionable), before the “real Turkic” layer (being the authors apparently Turkic themselves), and the supposed “continuity” of Eastern Uralic and Turkic groups in Asia since the Out of Africa migration. The combination of all of this in the same text is just disturbing.

If you look at it from the bright side, at least these samples were not of haplogroup R1a-Z280, or we would be talking about great Slavonic Scythians showing continuity from Russia with love, as the paper threatened to do in its introduction…

If you are enjoying the comeback of this retro 2000s comedy in 2019 (based on the classic nativist “R1a=IE”, “R1b=Basque”, and “N=Uralic” combo) it’s because you – like me – are putting yourself in this guy’s shoes every time a new episode of funny self-destruction appears:

chosen-poorly

Related

Ancient Sardinia hints at Mesolithic spread of R1b-V88, and Western EEF-related expansion of Vasconic

nuragic-sardinia-neolithic

New preprint Population history from the Neolithic to present on the Mediterranean island of Sardinia: An ancient DNA perspective, by Marcus et al. bioRxiv (2019)

Interesting excerpts (emphasis mine, edited for clarity):

On the high frequency of R1b-V88

Our genome-wide data allowed us to assign Y haplogroups for 25 ancient Sardinian individuals. More than half of them consist of R1b-V88 (n=10) or I2-M223 (n=7).

Francalacci et al. (2013) identi fied three major Sardinia-specifi c founder clades based on present-day variation within the haplogroups I2-M26, G2-L91 and R1b-V88, and here we found each of those broader haplogroups in at least one ancient Sardinian individual. Two major present-day Sardinian haplogroups, R1b-M269 and E-M215, are absent.

Compared to other Neolithic and present-day European populations, the number of identi fied R1b-V88 carriers is relatively high.

(…)ancient Sardinian mtDNA haplotypes belong almost exclusively to macro-haplogroups HV (n = 16), JT (n = 17) and U (n = 9), a composition broadly similar to other European Neolithic populations.

r1b-v88-europe
Geographic and temporal distribution of R1b-V88 Y-haplotypes in ancient European samples. We plot the geographic position of all ancient samples inferred to carry R1b-V88 equivalent markers. Dates are given as years BCE (means of calibrated 2s radio-carbon dates). Multiple V88 individuals with similar geographic positions are vertically stacked. We additionally color-code the status of the R1b-V88 subclade R1b-V2197, which is found in most present-day African R1b-V88 carriers.

On the origin of a Vasconic-like Paleosardo with the Western EEF

(…) the Neolithic (and also later) ancient Sardinian individuals sit between early Neolithic Iberian and later Copper Age Iberian populations, roughly on an axis that differentiates WHG and EEF populations and embedded in a cluster that additionally includes Neolithic British individuals. This result is also evident in terms of absolute genetic differentiation, with low pairwise FST ~ 0.005 +- 0.002 between Neolithic Sardinian individuals and Neolithic western mainland European populations. Pairwise outgroup-f3 analysis shows a very similar pattern, with the highest values of f3 (i.e. most shared drift) being with Neolithic and Copper Age Iberia, gradually dropping off for temporally and geographically distant populations.

In explicit admixture models (using qpAdm, see Methods) the southern French Neolithic individuals (France-N) are the most consistent with being a single source for Neolithic Sardinia (p ~ 0:074 to reject the model of one population being the direct source of the other); followed by other populations associated with the western Mediterranean Neolithic Cardial Ware expansion.

sardinians-ancient-eef
Principal Components Analysis based on the Human Origins dataset. A: Projection of ancient individuals’ genotypes onto principal component axes de fined by modern Western Eurasians (gray labels).

Pervasive Western Hunter-Gatherer ancestry in Iberian/French/Sardinian population

Similar to western European Neolithic and central European Late Neolithic populations, ancient Sardinian individuals are shifted towards WHG individuals in the top two PCs relative to early Neolithic Anatolians Admixture analysis using qpAdm infers that ancient Sardinian individuals harbour HG ancestry (~ 17%) that is higher than early Neolithic mainland populations (including Iberia, ~ 8%), but lower than Copper Age Iberians (~ 25%) and about the same as Southern French Middle-Neolithic individuals (~ 21%).

sardinia-modern-ancient-nuragic-pca
Principal Components Analysis based on the Human Origins dataset. B: Zoom into the region most relevant for Sardinian individuals.

Continuity from Sardinia Neolithic through the Nuragic

We found several lines of evidence supporting genetic continuity from the Sardinian Neolithic into the Bronze Age and Nuragic times. Importantly, we observed low genetic differentiation between ancient Sardinian individuals from various time periods.

A qpAdm analysis, which is based on simultaneously testing f-statistics with a number of outgroups and adjusts for correlations, cannot reject a model of Neolithic Sardinian individuals being a direct predecessor of Nuragic Sardinian individuals (…) Our qpAdm analysis further shows that the WHG ancestry proportion, in a model of admixture with Neolithic Anatolia, remains stable at ~17% throughout three ancient time-periods.

sardinians-modern-ancient-pca-admixture
Present-day genetic structure in Sardinia reanalyzed with aDNA. A: Scatter plot of the rst two principal components trained on 1577 present-day individuals with grand-parental ancestry from Sardinia. Each individual is labeled with a location if at least 3 of the 4 grandparents were born in the same geographical location (\small” three letter abbreviations); otherwise with \x” or if grand-parental ancestry is missing with \?”. We calculated median PC values for each Sardinian province (large abbreviations). We also projected each ancient Sardinian individual on to the top two PCs (gray points). B/C: We plot f-statistics that test for admixture of modern Sardinian individuals (grouped into provinces) when using Nuragic Sardinian individuals as one source population. Uncertainty ranges depict one standard error (calculated from block bootstrap). Karitiana are used in the f-statistic calculation as a proxy for ANE/Steppe ancestry (Patterson et al., 2012).

Steppe influx in Modern Sardinians

While contemporary Sardinian individuals show the highest affinity towards EEF-associated populations among all of the modern populations, they also display membership with other clusters (Fig. 5). In contrast to ancient Sardinian individuals, present-day Sardinian individuals carry a modest “Steppe-like” ancestry component (but generally less than continental present-day European populations), and an appreciable broadly “eastern Mediterranean” ancestry component (also inferred at a high fraction in other present-day Mediterranean populations, such as Sicily and Greece).

Related

Arrival of steppe ancestry with R1b-P312 in the Mediterranean: Balearic Islands, Sicily, and Iron Age Sardinia

steppe-balearic-sicily-sardinia

New preprint The Arrival of Steppe and Iranian Related Ancestry in the Islands of the Western Mediterranean by Fernandes, Mittnik, Olalde et al. bioRxiv (2019)

Interesting excerpts (emphasis in bold; modified for clarity):

Balearic Islands: The expansion of Iberian speakers

Mallorca_EBA dates to the earliest period of permanent occupation of the islands at around 2400 BCE. We parsimoniously modeled Mallorca_EBA as deriving 36.9 ± 4.2% of her ancestry from a source related to Yamnaya_Samara; (…). We next used qpAdm to identify “proximal” sources for Mallorca_EBA’s ancestry that are more closely related to this individual in space and time, and found that she can be modeled as a clade with the (small) subset of Iberian Bell Beaker culture associated individuals who carried Steppe-derived ancestry (p=0.442).

Suppl. Materials: The model used was with Bell_Beaker_Iberia_highsteppe, a group of outliers from Iberia buried in a Bell Beaker mortuary context who unlike most individuals from this context in that region had high proportions of Steppe ancestry (p=0.442).

Our estimates of Steppe ancestry in the two later Balearic Islands individuals are lower than the earlier one: 26.3 ± 5.1% for Formentera_MBA and 23.1 ± 3.6% for Menorca_LBA, but the Middle to Late Bronze Age Balearic individuals are not a clade relative to non-Balearic groups. Specifically, we find that f4(Mbuti.DG, X; Formentera_MBA, Menorca_LBA) is positive when X=Iberia_Chalcolithic (Z=2.6) or X=Sardinia_Nuragic_BA (Z=2.7). While it is tempting to interpret the latter statistic as suggesting a genetic link between peoples of the Talaiotic culture of the Balearic islands and the Nuragic culture of Sardinia, the attraction to Iberia_Chalcolithic is just as strong, and the mitochondrial haplogroup U5b1+16189+@16192 in Menorca_LBA is not observed in Sardinia_Nuragic_BA but is observed in multiple Iberia_Chalcolithic individuals. A possible explanation is that both the ancestors of Nuragic Sardinians and the ancestors of Talaiotic people from the Balearic Islands received gene flow from an unsampled Iberian Chalcolithic-related group (perhaps a mainland group affiliated to both) that did not contribute to Formentera_MBA.

This sample, like another one in El Argar, is of hg. R1b-P312. So there you are, the data that connects the Proto-Iberian expansion (replacing IE-speaking Bell Beakers) to the Iberian Chalcolithic population, signaled by the increase in Iberian Chalcolithic ancestry after the arrival of Bell Beakers, most likely connected originally to the Argaric and post-Argaric expansions during the MBA.

balearic-sicily-sardinia-pca
PCA with previously published ancient individuals (non-filled symbols), projected onto variation from present-day populations (gray squares).

Steppe in Sardinia IA: Phocaeans from Italy?

Most Sardinians buried in a Nuragic Bronze Age context possessed uniparental haplogroups found in European hunter-gatherers and early farmers, including Y-haplogroup R1b1a[xR1b1a1a] which is different from the characteristic R1b1a1a2a1a2 spread in association with the Bell Beaker complex. An exception is individual I10553 (1226-1056 calBCE) who carried Y-haplogroup J2b2a, previously observed in a Croatian Middle Bronze Age individual bearing Steppe ancestry, suggesting the possibility of genetic input from groups that arrived from the east after the spread of first farmers. This is consistent with the evidence of material culture exchange between Sardinians and mainland Mediterranean groups, although genome-wide analyses find no significant evidence of Steppe ancestry so the quantitative demographic impact was minimal.

Another interesting data, these (Mesolithic) remnant R1b-V88 lineages closely related to the Italian Peninsula, the most likely region of expansion of these lineages into Africa, in turn possibly connected to the expansion of Proto-Afroasiatic.

We detect definitive evidence of Iranian-related ancestry in an Iron Age Sardinian I10366 (391-209 calBCE) with an estimate of 11.9 ± 3.7.% Iran_Ganj_Dareh_Neolithic related ancestry, while rejecting the model with only Anatolian_Neolithic and WHG at p=0.0066 (Supplementary Table 9). The only model that we can fit for this individual using a pair of populations that are closer in time is as a mixture of Iberia_Chalcolithic (11.9 ± 3.2%) and Mycenaean (88.1 ± 3.2%) (p=0.067). This model fits even when including Nuragic Sardinians in the outgroups of the qpAdm analysis, which is consistent with the hypothesis that this individual had little if any ancestry from earlier Sardinians.

yamnaya-samara
Proportions of ancestry using a distal qpAdm framework on an individual basis (a), and based on qpWave clusters

Sicily EBA: The Lusitanian/Ligurian connection?

(…) While a previously reported Bell Beaker culture-associated individual from Sicily had no evidence of Steppe ancestry, (…) we find evidence of Steppe ancestry in the Early Bronze Age by ~2200 BCE. In distal qpAdm, the outlier Sicily_EBA11443 is parsimoniously modeled as harboring 40.2 ± 3.5% Steppe ancestry, and the outlier Sicily_EBA8561 is parsimoniously modeled as harboring 23.3 ± 3.5% Steppe ancestry. (…) The presence of Steppe ancestry in Early Bronze Age Sicily is also evident in Y chromosome analysis, which reveals that 4 of the 5 Early Bronze Age males had Steppe-associated Y-haplogroup R1b1a1a2a1a2. (Online Table 1). Two of these were Y-haplogroup R1b1a1a2a1a2a1 (Z195) which today is largely restricted to Iberia and has been hypothesized to have originated there 2500-2000 BCE. This evidence of west-to-east gene flow from Iberia is also suggested by qpAdm modeling where the only parsimonious proximate source for the Steppe ancestry we found in the main Sicily_EBA cluster is Iberians.

What’s this? An ancestral connection between Sicel Elymian and Galaico-Lusitanian or Ligurian (based on an origin in NE Iberia)? Impossible to say, especially if the languages of these early settlers were replaced later by non-Indo-European speakers from the eastern Mediterranean, and by Indo-European speakers from the mainland closely related to Proto-Italic during the LBA, but see below.

Regarding the comment on R1b-Z195, it is associated with modern Iberians, as DF27 in general, due to founder effects beyond the Pyrenees. It is a very old subclade, split directly from DF27 roughly at the same time as it split from the parent P312, i.e. it can be found anywhere in Europe, and it almost certainly accompanied the expansion of Celts from Central Europe under the subclade R1b-M167/SRY2627.

The connection is thus strong only because of the qpAdm modeling, since R1b-DF27 and subclade R1b-Z195 are certainly lineages expanded quite early, most likely with Yamna settlers in Hungary and East Bell Beakers.

In this case, if stemming from Iberia, it is most likely of subclade R1b-Z220 – or another Z195 (xM167) lineage – originally associated with the Old European substrate found in topo-hydronymy in Iberia, whose most likely remnants attested during the Iron Age were Lusitanians.

r1b-df27-z195
Left: Modern distribution of R1b-Z195 (YFull estimate 2700 BC); Right: Modern distribution of DF27. Both include later founder effects within Iberia, so the increase in the Basque country and the Crown of Aragon and the decrease in Portugal can safely be ignored. Contour maps of the derived allele frequencies of the SNPs analyzed in Solé-Morata et al. (2017).

We detect Iranian-related ancestry in Sicily by the Middle Bronze Age 1800-1500 BCE, consistent with the directional shift of these individuals toward Mycenaeans in PCA. Specifically, two of the Middle Bronze Age individuals can only be fit with models that in addition to Anatolia_Neolithic and WHG, include Iran_Ganj_Dareh_Neolithic. The most parsimonious model for Sicily_MBA3125 has 18.0 ± 3.6% Iranian-related ancestry (p=0.032 for rejecting the alternative model of Steppe rather than Iranian-related ancestry), and the most parsimonious model for Sicily_MBA has 14.9 ± 3.9% Iranian-related ancestry (p=0.037 for rejecting the alternative model).

The modern southern Italian Caucasus-related signal identified in Raveane et al. (2018) is plausibly related to the same Iranian-related spread of ancestry into Sicily that we observe in the Middle Bronze Age (and possibly the Early Bronze Age).

The non-Indo-European Sicanians and Elymians were possibly then connected to eastern Mediterranean groups before the expansion of the Sea Peoples.

For the Late Bronze Age group of individuals, qpAdm documented Steppe-related ancestry, modeling this group as 80.2 ± 1.8% Anatolia_Neolithic, 5.3 ± 1.6% WHG, and 14.5 ± 2.2% Yamnaya_Samara. Our modeling using sources more closely related in space and time also supports Sicily_LBA having Minoan-related ancestry or being derived from local preceding populations or individuals with ancestries similar to those of Sicily_EBA3123 (p=0.527), Sicily_MBA3124 (p=0.352), and Sicily_MBA3125 (p=0.095).

This increase in Steppe-related ancestry in a western site during the LBA most likely represents either an expansion from the Aegean or – maybe more likely, given the archaeological finds – a regional population similar to Sicily EBA re-emerging or rather being displaced from the eastern part of the island because of a westward movement from nearby Calabria.

Whether this population sampled spoke Indo-European or not at this time is questionable, since the Iron Age accounts show non-IE Elymians in this region.

Actually, Elymians seem to have spoken Indo-European, which fits well with the increase in steppe ancestry.

EDIT (21 MAR): Interesting about a proposed incoming Minoan-like ancestry is the potential origin of the Iran Neolithic-related ancestry that is going to appear in Central Italy during the LBA. This could then be potentially associated with Tyrsenians passing through the area, although the traditional description may be more more compatible with an arrival of Sea Peoples from the Adriatic.

Sad to read this:

This manuscript is dedicated to the memory of Sebastiano Tusa of the Soprintendenza del Mare in Palermo, who would have been an author of this study had he not tragically died in the crash of Ethiopia Airlines flight 302 on March 10.

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