Vikings, Vikings, Vikings! “eastern” ancestry in the whole Baltic Iron Age

vikings-middle-age

Open access Population genomics of the Viking world, by Margaryan et al. bioRxiv (2019), with a huge new sampling from the Viking Age.

Interesting excerpts (emphasis mine, modified for clarity):

To understand the genetic structure and influence of the Viking expansion, we sequenced the genomes of 442 ancient humans from across Europe and Greenland ranging from the Bronze Age (c. 2400 BC) to the early Modern period (c. 1600 CE), with particular emphasis on the Viking Age. We find that the period preceding the Viking Age was accompanied by foreign gene flow into Scandinavia from the south and east: spreading from Denmark and eastern Sweden to the rest of Scandinavia. Despite the close linguistic similarities of modern Scandinavian languages, we observe genetic structure within Scandinavia, suggesting that regional population differences were already present 1,000 years ago.

Maps illustrating the following texts have been made based on data from this and other papers:

  • Maps showing ancestry include only data from this preprint (which also includes some samples from Sigtuna).
  • Maps showing haplogroup density include Vikings from other publications, such as those from Sigtuna in Krzewinska et al. (2018), and from Iceland in Ebenesersdóttir et al. (2018).
  • Maps showing haplogroups of ancient DNA samples based on their age include data from all published papers, but with slightly modified locations to avoid overcrowding (randomized distance approx. ± 0.1 long. and lat.).

middle-ages-europe-y-dna
Y-DNA haplogroups in Europe during the Viking expansions (full map). See other maps from the Middle Ages.

We find that the transition from the BA to the IA is accompanied by a reduction in Neolithic farmer ancestry, with a corresponding increase in both Steppe-like ancestry and hunter-gatherer ancestry. While most groups show a slight recovery of farmer ancestry during the VA, there is considerable variation in ancestry across Scandinavia. In particular, we observe a wide range of ancestry compositions among individuals from Sweden, with some groups in southern Sweden showing some of the highest farmer ancestry proportions (40% or more in individuals from Malmö, Kärda or Öland).

Ancestry proportions in Norway and Denmark on the other hand appear more uniform. Finally we detect an influx of low levels of “eastern” ancestry starting in the early VA, mostly constrained among groups from eastern and central Sweden as well as some Norwegian groups. Testing of putative source groups for this “eastern” ancestry revealed differing patterns among the Viking Age target groups, with contributions of either East Asian- or Caucasus-related ancestry.

saami-ancestry-vikings
Ancestry proportions of four-way models including additional putative source groups for target groups for which three-way fit was rejected (p ≤ 0.01);

Overall, our findings suggest that the genetic makeup of VA Scandinavia derives from mixtures of three earlier sources: Mesolithic hunter-gatherers, Neolithic farmers, and Bronze Age pastoralists. Intriguingly, our results also indicate ongoing gene flow from the south and east into Iron Age Scandinavia. Thus, these observations are consistent with archaeological claims of wide-ranging demographic turmoil in the aftermath of the Roman Empire with consequences for the Scandinavian populations during the late Iron Age.

Genetic structure within Viking-Age Scandinavia

We find that VA Scandinavians on average cluster into three groups according to their geographic origin, shifted towards their respective present-day counterparts in Denmark, Sweden and Norway. Closer inspection of the distributions for the different groups reveals additional complexity in their genetic structure.

vikings-danish-ancestry
Natural neighbor interpolation of “Danish ancestry” among Vikings.

We find that the ‘Norwegian’ cluster includes Norwegian IA individuals, who are distinct from both Swedish and Danish IA individuals which cluster together with the majority of central and eastern Swedish VA individuals. Many individuals from southwestern Sweden (e.g. Skara) cluster with Danish present-day individuals from the eastern islands (Funen, Zealand), skewing towards the ‘Swedish’ cluster with respect to early and more western Danish VA individuals (Jutland).

Some individuals have strong affinity with Eastern Europeans, particularly those from the island of Gotland in eastern Sweden. The latter likely reflects individuals with Baltic ancestry, as clustering with Baltic BA individuals is evident in the IBS-UMAP analysis and through f4-statistics.

vikings-norwegian-ancestry
Natural neighbor interpolation of “Norwegian ancestry” among Vikings.

For more on this influx of “eastern” ancestry see my previous posts (including Viking samples from Sigtuna) on Genetic and linguistic continuity in the East Baltic, and on the Pre-Proto-Germanic homeland based on hydrotoponymy.

Baltic ancestry in Gotland

Genetic clustering using IBS-UMAP suggested genetic affinities of some Viking Age individuals with Bronze Age individuals from the Baltic. To further test these, we quantified excess allele sharing of Viking Age individuals with Baltic BA compared to early Viking Age individuals from Salme using f4 statistics. We find that many individuals from the island of Gotland share a significant excess of alleles with Baltic BA, consistent with other evidence of this site being a trading post with contacts across the Baltic Sea.

vikings-finnish-ancestry
Natural neighbor interpolation of “Finnish ancestry” among Vikings.

The earliest N1a-VL29 sample available comes from Iron Age Gotland (VK579) ca. AD 200-400 (see Iron Age Y-DNA maps), which also proves its presence in the western Baltic before the Viking expansion. The distribution of N1a-VL29 and R1a-Z280 (compared to R1a in general) among Vikings also supports a likely expansion of both lineages in succeeding waves from the east with Akozino warrior-traders, at the same time as they expanded into the Gulf of Finland.

vikings-y-dna-haplogroup-r1a-z280-over-r1a
Density of haplogroup R1a-Z280 (samples in pink) overlaid over other R1a samples (in green, with R1a-Z284 in cyan) among Vikings.

Vikings in Estonia

(…) only one Viking raiding or diplomatic expedition has left direct archaeological traces, at Salme in Estonia, where 41 Swedish Vikings who died violently were buried in two boats accompanied by high-status weaponry. Importantly, the Salme boat-burial predates the first textually documented raid (in Lindisfarne in 793) by nearly half a century. Comparing the genomes of 34 individuals from the Salme burial using kinship analyses, we find that these elite warriors included four brothers buried side by side and a 3rd degree relative of one of the four brothers. In addition, members of the Salme group had very similar ancestry profiles, in comparison to the profiles of other Viking burials. This suggests that this raid was conducted by genetically homogeneous people of high status, including close kin. Isotope analyses indicate that the crew descended from the Mälaren area in Eastern Sweden thus confirming that the Baltic-Mid-Swedish interaction took place early in the VA.

vikings-swedish-ancestry
Natural neighbor interpolation of “Swedish ancestry” among Vikings.

Viking samples from Estonia show thus ancient Swedes from the Mälaren area, which proves once again that hg. N1a-VL29 (especially subclade N1a-L550) and tiny proportions of so-called “Siberian ancestry” expanded during the Early Iron Age into the whole Baltic Sea area, not only into Estonia, and evidently not spreading with Balto-Finnic languages (since the language influence is in the opposite direction, east-west, Germanic > Finno-Samic, during the Bronze Age).

N1a-VL29 lineages spread again later eastwards with Varangians, from Sweden into north-eastern Europe, most likely including the ancestors of the Rurikid dynasty. Unsurprisingly, the arrival of Vikings with Swedish ancestry into the East Baltic and their dispersal through the forest zone didn’t cause a language shift of Balto-Finnic, Mordvinic, or East Slavic speakers to Old Norse, either…

NOTE. For N1a-Y4339 – N1a-L550 subclade of Swedish origin – as main haplogroup of modern descendants of Rurikid princes, see Volkov & Seslavin (2019) – full text in comments below. Data from ancient samples show varied paternal lineages even among early rulers traditionally linked to Rurik’s line, which explains some of the discrepancies found among modern descendants:

  • A sample from Chernihiv (VK542) potentially belonging to Gleb Svyatoslavich, the 11th century prince of Tmutarakan/Novgorod, belongs to hg. I2a-Y3120 (a subclade of early Slavic I2a-CTS10228) and has 71% “Modern Polish” ancestry (see below).
  • Izyaslav Ingvarevych, the 13th century prince of Dorogobuzh, Principality of Volhynia/Galicia, is probably behind a sample from Lutsk (VK541), and belongs to hg. R1a-L1029 (a subclade of R1a-M458), showing ca. 95% of “Modern Polish” ancestry.
  • Yaroslav Osmomysl, the 12th century Prince of Halych (now in Western Ukraine), was probably of hg. E1b-V13, yet another clearly early Slavic haplogroup.

vikings-y-dna-haplogroup-n1a
Density of haplogroup N1a-VL29, N1a-L550 (samples in pink, most not visible) among Vikings. Samples of hg. R1b in blue, hg. R1a in green, hg. I in orange.

Finnish ancestry

Firstly, modern Finnish individuals are not like ancient Finnish individuals, modern individuals have ancestry of a population not in the reference; most likely Steppe/Russian ancestry, as Chinese are in the reference and do not share this direction. Ancient Swedes and Norwegians are more extreme than modern individuals in PC2 and 4. Ancient UK individuals were more extreme than Modern UK individuals in PC3 and 4. Ancient Danish individuals look rather similar to modern individuals from all over Scandinavia. By using a supervised ancient panel, we have removed recent drift from the signal, which would have affected modern Scandinavians and Finnish populations especially. This is in general a desirable feature but it is important to check that it has not affected inference.

ancient-modern-finns-steppe
PCA of the ancient and modern samples using the ancient palette, showing different PCs. Modern individuals are grey and the K=7 ancient panel surrogate populations are shown in strong colors, whilst the remaining M-K=7 ancient populations are shown in faded colors.

The story for Modern-vs-ancient Finnish ancestry is consistent, with ancient Finns looking much less extreme than the moderns. Conversely, ancient Norwegians look like less-drifted modern Norwegians; the Danish admixture seen through the use of ancient DNA is hard to detect because of the extreme drift within Norway that has occurred since the admixture event. PC4 vs PC5 is the most important plot for the ancient DNA story: Sweden and the UK (along with Poland, Italy and to an extent also Norway) are visibly extremes of a distribution the same “genes-mirror-geography” that was seen in the Ancient-palette analysis. PC1 vs PC2 tells the same story – and stronger, since this is a high variance-explained PC – for the UK, Poland and Italy.

Uniform manifold approximation and projection (UMAP) analysis of the VA and other ancient samples.

Evidence for Pictish Genomes

The four ancient genomes of Orkney individuals with little Scandinavian ancestry may be the first ones of Pictish people published to date. Yet a similar (>80% “UK ancestry) individual was found in Ireland (VK545) and five in Scandinavia, implying that Pictish populations were integrated into Scandinavian culture by the Viking Age.

Our interpretation for the Orkney samples can be summarised as follows. Firstly, they represent “native British” ancestry, rather than an unusual type of Scandinavian ancestry. Secondly, that this “British” ancestry was found in Britain before the Anglo-Saxon migrations. Finally, that in Orkney, these individuals would have descended from Pictish populations.

vikings-british-ancestry
Natural neighbor interpolation of “British ancestry” among Vikings.

(…) ‘UK’ represents a group from which modern British and Irish people all receive an ancestry component. This information together implies that within the sampling frame of our data, they are proxying the ‘Briton’ component in UK ancestry; that is, a pre-Roman genetic component present across the UK. Given they were found in Orkney, this makes it very likely that they were descended from a Pictish population.

Modern genetic variation within the UK sees variation between ‘native Briton’ populations Wales, Scotland, Cornwall and Ireland as large compared to that within the more ‘Anglo-Saxon’ English. This is despite subsequent gene flow into those populations from English-like populations. We have not attempted to disentangle modern genetic drift from historically distinct populations. Roman-era period people in England, Wales, Ireland and Scotland may not have been genetically close to these Orkney individuals, but our results show that they have a shared genetic component as they represent the same direction of variation.

Density of haplogroup R1b-L21 (samples in red), overlaid over all samples of hg. R1b among Vikings (R1b-U106 in green, other R1b-L151 in deep red). To these samples one may add the one from Janakkala in south-western Finland (AD ca. 1300), of hg. R1b-L21, possibly related to these population movements.

For more on Gaelic ancestry and lineages likely representing slaves among early Icelanders, see Ebenesersdóttir et al. (2018).

Y-DNA

As in the case of mitochondrial DNA, the overall distribution profile of the Y chromosomal haplogroups in the Viking Age samples was similar to that of the modern North European populations. The most frequently encountered male lineages were the haplogroups I1, R1b and R1a.

Haplogroup I (I1, I2)

The distribution of I1 in southern Scandinavia, including a sample from Sealand (VK532) ca. AD 100 (see Iron Age Y-DNA maps) proves that it had become integrated into the West Germanic population already before their expansions, something that we already suspected thanks to the sampling of Germanic tribes.

vikings-y-dna-haplogroup-i
Density of haplogroup I (samples in orange) among Vikings. Samples of hg. R1b in blue, hg. R1a in green, N1a in pink.
vikings-y-dna-haplogroup-i1-over-i
Density of haplogroup I1 (samples in red) overlaid over all samples of hg. I among Vikings.

Haplogroup R1b (M269, U106, P312)

Especially interesting is the finding of R1b-L151 widely distributed in the historical Nordic Bronze Age region, which is in line with the estimated TMRCA for R1b-P312 subclades found in Scandinavia, despite the known bottleneck among Germanic peoples under U106. Particularly telling in this regard is the finding of rare haplogroups R1b-DF19, R1b-L238, or R1b-S1194. All of that points to the impact of Bell Beaker-derived peoples during the Dagger period, when Pre-Proto-Germanic expanded into Scandinavia.

Also interesting is the finding of hg. R1b-P297 in Troms, Norway (VK531) ca. 2400 BC. R1b-P297 subclades might have expanded to the north through Finland with post-Swiderian Mesolithic groups (read more about Scandinavian hunter-gatherers), and the ancestry of this sample points to that origin.

However, it is also known that ancestry might change within a few generations of admixture, and that the transformation brought about by Bell Beakers with the Dagger Period probably reached Troms, so this could also be a R1b-M269 subclade. In fact, the few available data from this sample show that it comes from the natural harbour Skarsvågen at the NW end of the island Senja, and that its archaeologist thought it was from the Viking period or slightly earlier, based on the grave form. From Prescott (2017):

In 1995, Prescott and Walderhaug tentatively argued that a dramatic transformation took place in Norway around the Late Neolithic (2350 BCE), and that the swift nature of this transition was tied to the initial Indo-Europeanization of southern and coastal Norway, at least to Trøndelag and perhaps as far north as Troms. (…)

The Bell Beaker/early Late Neolithic, however, represents a source and beginning of these institution and practices, exhibits continuity to the following metal age periods and integrated most of Northern Europe’s Nordic region into a set of interaction fields. This happened around 2400 BCE, at the MNB to LN transition.

NOTE. This particular sample is not included in the maps of Viking haplogroups.

vikings-y-dna-haplogroup-r1b
Density of haplogroup R1b (samples in blue) among Vikings. Samples of hg. I in orange, hg. R1a in green, N1a in pink.
vikings-y-dna-haplogroup-r1b-U106-over-r1b
Density of haplogroup R1b-U106 (samples in green) overlaid over all samples of hg. R1b (other R1b-L23 samples in red) among Vikings.
vikings-y-dna-haplogroup-r1b-P312-over-r1b
Density of R1b-L151 (xR1b-U106) (samples in deep red) overlaid over all samples of hg. R1b (R1b-U106 in green, other R1b-M269 in blue) among Vikings.

Haplogroup R1a (M417, Z284)

The distribution of hg. R1a-M417, in combination with data on West Germanic peoples, shows that it was mostly limited to Scandinavia, similar to the distribution of I1. In fact, taking into account the distribution of R1a-Z284 in particular, it seems even more isolated, which is compatible with the limited impact of Corded Ware in Denmark or the Northern European Plain, and the likely origin of R1a-Z284 in the expansion with Battle Axe from the Gulf of Finland. The distribution of R1a-Z280 (see map above) is particularly telling, with a distribution around the Baltic Sea mostly coincident with that of N1a.

vikings-y-dna-haplogroup-r1a
Density of haplogroup R1a (samples in green) among Vikings. Samples of hg. R1b in blue, of hg. I in orange, N1a in pink.
vikings-y-dna-haplogroup-r1a-z284-over-r1a
Density of haplogroup R1a-Z284 (samples in cyan) overlaid over all samples of hg. R1a (in green, with R1a-Z280 in pink) among Vikings.

Other haplogroups

Among the ancient samples, two individuals were derived haplogroups were identified as E1b1b1-M35.1, which are frequently encountered in modern southern Europe, Middle East and North Africa. Interestingly, the individuals carrying these haplogroups had much less Scandinavian ancestry compared to the most samples inferred from haplotype based analysis. A similar pattern was also observed for less frequent haplogroups in our ancient dataset, such as G (n=3), J (n=3) and T (n=2), indicating a possible non-Scandinavian male genetic component in the Viking Age Northern Europe. Interestingly, individuals carrying these haplogroups were from the later Viking Age (10th century and younger), which might indicate some male gene influx into the Viking population during the Viking period.

vikings-italian-ancestry
Natural neighbor interpolation of “Italian ancestry” among Vikings.

As the paper says, the small sample size of rare haplogroups cannot distinguish if these differences are statistically relevant. Nevertheless, both E1b samples have substantial Modern Polish-like ancestry: one sample from Gotland (VK474), of hg. E1b-L791, has ca. 99% “Polish” ancestry, while the other one from Denmark (VK362), of hg. E1b-V13, has ca. 35% “Polish”, ca. 35% “Italian”, as well as some “Danish” (14%) and minor “British” and “Finnish” ancestry.

Given the E1b-V13 samples of likely Central-East European origin among Lombards, Visigoths, and especially among Early Slavs, and the distribution of “Polish” ancestry among Viking samples, VK362 is probably a close description of the typical ancestry of early Slavs. The peak of Modern Polish-like ancestry around the Upper Pripyat during the (late) Viking Age suggests that Poles (like East Slavs) have probably mixed since the 10th century with more eastern peoples close to north-eastern Europeans, derived from ancient Finno-Ugrians:

vikings-polish-ancestry
Natural neighbor interpolation of “Polish ancestry” among Vikings.

Similarly, the finding of R1a-M458 among Vikings in Funen, Denmark (VK139), in Lutsk, Poland (VK541), and in Kurevanikha, Russia (VK160), apart from the early Slav from Usedom, may attest to the origin of the spread of this haplogroup in the western Baltic after the Bell Beaker expansion, once integrated in both Germanic and probably Balto-Slavic populations, as well as intermediate Bronze Age peoples that were eventually absorbed by their expansions. This contradicts, again, my simplistic initial assessment of R1a-M458 expansion as linked exclusively (or even mainly) to Balto-Slavs.

antiquity-europe-y-dna
Y-DNA haplogroups in Europe during Antiquity (full map). See other maps of cultures and ancient DNA from Antiquity.

Related

More Hungarian Conquerors of hg. N1c-Z1936, and the expansion of ‘Altaic-Uralic’ N1c

Open access Y-chromosomal connection between Hungarians and geographically distant populations of the Ural Mountain region and West Siberia, by Post et al. Scientific Reports (2019) 9:7786.

Hungarian Conquerors

More interesting than the study of modern populations of the paper is the following excerpt from the introduction, referring to a paper that is likely in preparation, Európai És Ázsiai Apai Genetikai Vonalak A Honfoglaló Magyar Törzsekben, by Fóthi, E., Fehér, T., Fóthi, Á. & Keyser, C., Avicenna Institute of Middle Eastern Studies (2019):

Certain chr-Y lineages from haplogroup (hg) N have been proposed to be associated with the spread of Uralic languages. So far, hg N3 has not been reported for Indo-European speaking populations in Central Europe, but it is present among Hungarians, although the proportion of hg N in the paternal gene pool of present-day Hungarians is only marginal (up to 4%) compared to other Uralic speaking populations. It has been shown earlier that one of the sub-clades of hg N – N3a4-Z1936 – could be a potential link between two Ugric speaking populations: the Hungarians and the Mansi. It is also notable that some ancient Hungarian samples from the 9th and 10th century Carpathian Basin belonged to this hg N sub-clade: Three Z1936 samples were found in the Upper-Tisza area (Karos II, Bodrogszerdahely/Streda nad Bodrogom) and two in the Middle-Tisza basin cemeteries (Nagykörű and Tiszakécske). The haplotype of the Nagykörű sample is identical with one contemporary Hungarian sample from Transylvania that tested positive for B545 marker downstream of N3a4-Z193632. Similar findings come from the maternal gene pool of historical Hungarians: the analyses of early medieval aDNA samples from Karos-Eperjesszög cemeteries revealed the presence of mtDNA hgs of East Asian provenance.

A commenter recently wrote that in a study by Fehér (probably this one) two Hungarian conquerors, from Ormenykut and Tuzser, will be of hg. N1c-2110. Assuming no other lineages will appear, this would leave the proportion of N1c-L392 vs. R1a-Z280/Z93 closer to the reported proportion of hg. N vs. R1a (5 vs. 2) among Sargat samples, and is thus compatible with a direct migration of Hungarians from around the Urals.

However, the sampling of Iron Age populations around the Urals is scarce, and we don’t know what other lineages these studied Magyars will have, but – based on the known variability of the published ones, and on the ca. 50-60 early Magyar males available to date in previous studies to obtain Y-chromosome haplogroups – I would say these reported N1c lineages are just a tiny proportion of what’s to come…

“Altaic-Uralic” N1c

altaic-uralic-n1c-haplogroup
Phylogenetic tree of hg N3a4. Phylogenetic tree of 33 high coverage Y-chromosomes from
haplogroup N3a4 was reconstructed with BEAST v.1.7.5 software package.

Archaeogenetic studies based on mtDNA haplotypes have shown that ancient Hungarians were relatively close to contemporary Bashkirs who are a Turkic speaking population residing in the Volga-Ural region. Another study reported excessive identical-by-descent (IBD) genomic segments shared between the Ob-Ugric speaking Khantys and Bashkirs but a moderate IBD sharing between Turkic speaking Tatars and their neighbours including Bashkirs.

Phylogenetic tree of hg N3a4 has two main sub-clades defined by markers B535 and B539 that diverged around 4.9 kya (95% confidence interval [CI] = 3.7–6.3 kya). Inner sub-clades of N3a4-B539 (defined by markers B540 and B545) split 4.2 kya (95% CI = 3.0–5.6 kya). (…) The phylogenetic tree reveals that all five Hungarian samples belong to N3a4-B539 sub-clade that they share with Ob-Ugric speaking Khanty and Mansi, and Turkic speaking Bashkirs and Tatars from the Volga-Ural region. Hungarian and Bashkir chrY lineages belong to both sub-clades of N3a4-B539.

Modern distribution of the “Ugric N1c”

To test the presence and proportions of hg N3a4 lineages in a more comprehensive sample set and with a higher phylogenetic resolution level compared to earlier studies, we analysed the genotyping data of about 5000 Eurasian individuals, including West Siberian Mansi and Khanty who are linguistically closest to Hungarians

n3a4-n1c-z1936-ugric
Map of the entire hg N3a4.

There is a clear difference in geographic distribution patterns of these two hg N3a4 sub-clades. Hg N3a4-B535 (Fig. 3b) is common mostly among Finnic (Finns, Karelians, Vepsas, Estonians) and Saami speaking populations in North eastern Europe. The highest frequency is detected in Finns (~44%) but it also reaches up to 32% in Vepsas and around 20% in Karelians, Saamis and North Russians. The latter are known to have changed their language or to be an admixed population with reported similar genetic composition to their Finnic speaking neighbors. The frequency of N3a4-B535 rapidly decreases towards south to around 5% in Estonians, being almost absent in Latvians (1%) and not found among Lithuanians. Towards east its frequency is from 1–9% among Eastern European Russians and populations of the Volga-Ural region such as Komis, Mordvins and Chuvashes (…)

n3a4-n1c-z1936-finnic-samic
Map of N3a4 subclades defined by B535.

Hg N3a4-B539, on the other hand, is prevalent among Turkic speaking Bashkirs and also found in Tatars but is entirely missing from other populations of the Volga-Ural region such as Uralic speaking Udmurts, Maris, Komis and Mordvins, and in Northeast Europe, where instead N3a4-B535 lineages are frequent. Besides Bashkirs and Tatars in Volga-Ural region, N3a4-B539 is substantially represented in West Siberia among Ugric speaking Mansis and Khantys. Among Hungarians, however, N3a4-B539 has a subtle frequency of 1–4%.

n3a4-n1c-z1936-ugric-bashkir
Map of N3a4 subclades defined by B539, with a local snapshot showing the N3a4-B539 distribution among Hungarian speakers.

The battle to appropriate N1c-L392

So, basically, the team of Kristiina Tambets is arguing that N1c-VL29 expanded Finnic to the East Baltic (hence from a common Finno-Mordvinic dialect splitting ca. 600 BC on?) because, you know, apparently the agreed separation of known Uralic dialects from ca. 2000 BC, and their Bronze Age presence around the Baltic, is not valid when you follow haplogroups instead of languages or archaeology.

But now this other group of Tambets (co-author of this paper) considers that hg. N1c-Z1936 – which is probably behind the N1c-L392 samples from Lovozero Ware in the Kola Peninsula – represent either the True Uralic-speaking Palaeo-Arctic peoples, or else merely Ugric-speaking peoples which happened to expand to Fennoscandia but left no trace of their language…

To accept this identification you only have to NOT ask why:

  • N1c is first found in ancient cultures close to Lake Baikal.
  • N1c-L392 appears in ancient East Asian populations speaking completely different languages, with Altaic and Uralic being just some among many Palaeo-Siberian populations where the haplogroup will pop up.
  • Turkic populations like Bashkirs and Tatars (who expanded to the Volga through the southern Urals before the expansion of Hungarians) show a shared distribution of the B539 haplotype with Hungarians.
  • The phylogenetic tree and areas of N1c-L392 expansions don’t make any sense in light of the known linguistic and cultural expansions of Uralic-speaking peoples.

In fact, the Hungarian research group of Neparáczki – publishing the recent paper on Hungarian Conquerors – was apparently looking for a connection with Turkic peoples to support some traditional Turanian myths, and they found it in some scattered R1a-Z93 samples which supposedly connect Hungarian Conquerors to Huns (?), instead of looking for this closer link through N1c-Z1936 (especially haplotype B539)…

Also, is it me or are there two opposed trends with completely different interpretations among researchers publishing papers about hg. N1c: one systematically arguing for Altaic origins, and another for Uralic ones?

If somebody sees some complex reasoning behind the discussions of all these recent papers, beyond the simplest “let’s follow N for Uralic/Altaic”, feel free to comment below. Just so I can understand what I might be doing wrong in assessing Neolithic and Bronze Age migrations in linguistics and archaeology with help of ancient haplogroups coupled with ancestral components, but these researchers are doing right by playing with obsessive ideas born out of the 2000s coupled with phylogenetic trees and maps of modern haplogroup distributions…

This is probably going to be this blog’s most used image in 2019:

horse-meme-steppe-ancestry

Related

Baltic Finns in the Bronze Age, of hg. R1a-Z283 and Corded Ware ancestry

estonian-bronze-age-dna

Open access The Arrival of Siberian Ancestry Connecting the Eastern Baltic to Uralic Speakers further East, by Saag et al. Current Biology (2019).

Interesting excerpts:

In this study, we present new genomic data from Estonian Late Bronze Age stone-cist graves (1200–400 BC) (EstBA) and Pre-Roman Iron Age tarand cemeteries (800/500 BC–50 AD) (EstIA). The cultural background of stone-cist graves indicates strong connections both to the west and the east [20, 21]. The Iron Age (IA) tarands have been proposed to mirror “houses of the dead” found among Uralic peoples of the Volga-Kama region [22].

(…) The 33 individuals included 15 from EstBA, 6 from EstIA, 5 from Pre-Roman to Roman Iron Age Ingria (500 BC–450 AD) (IngIA), and 7 from Middle Age Estonia (1200–1600 AD) (EstMA) and yielded endogenous DNA ∼4%–88%, average genomic coverages ∼0.017–0.734×, and contamination estimates <4% (Table S1). We analyzed the data in the context of modern and other ancient individuals, including from Neolithic Estonia [13].

estonian-y-dna-bronze-iron-age
Archaeological Information, Genetic Sex, mtDNA and Y Chromosome Haplogroups, and Average Coverage of the Individuals of This Study. Modified from the paper to mark distinct Y-DNA haplogroups in the LBA and IA.

We identified chrY hgs for 30 male individuals (Tables 1 and S2; STAR Methods). All 16 successfully haplogrouped EstBA males belonged to hg R1a, showing no change from the CWC period, when this was also the only chrY lineage detected in the Eastern Baltic [11, 13, 30, 31]. Three EstIA and two IngIA individuals also belonged to hg R1a, but three EstIA males belonged to hg N3a, the earliest so far observed in the Eastern Baltic. Three EstMA individuals belonged to hg N3a, two to hg R1a, and one to hg J2b. ChrY lineages found in the Baltic Sea region before the CWC belong to hgs I, R1b, R1a5, and Q [10, 11, 12, 13, 17, 32]. Thus, it appears that these lineages were substantially replaced in the Eastern Baltic by hg R1a [10, 11, 12, 13], most likely through steppe migrations from the east [30, 31]. (…) Our results enable us to conclude that, although the expansion time for R1a1 and N3a3′5 in Eastern Europe is similar [25], hg N3a likely reached Estonia or at least became comparably frequent to modern Estonia [1] only during the BA-IA transition.

A clear shift toward West Eurasian hunter-gatherers is visible between European LN and BA (including Baltic CWC) and EstBA individuals, the latter clustering together with Latvian and Lithuanian BA individuals [11]. EstIA, IngIA, and EstMA individuals project between BA individuals and modern Estonians, partially overlapping with both.

(…) EstBA individuals are clearly distinguishable from Estonian CWC individuals as the former have more of the blue component most frequent in WHGs and less of the brown and yellow components maximized in Caucasus hunter-gatherers and modern Khanty, respectively. The individuals of EstBA, EstIA, IngIA, EstMA, and modern Estonia are quite similar to each other on average, indicating that the relatively high proportion of WHG ancestry in modern Eastern Baltic populations compared to other present-day Europeans [15] traces back to the BA.

estonian-pca-published
Detail of the PCA, modified from the paper to label populations. Estonian Bronze Age and Iron Age samples cluster close to Early Corded Ware from the Baltic.. Principal-component analysis results of modern West Eurasians with ancient individuals projected onto the first two components (PC1 and PC2). BA, Bronze Age; EF, early farmers; HG, hunter-gatherers; IA, Iron Age; IMA, Iron/Middle Ages; LN, Late Neolithic; LNBA, Late Neolithic/Bronze Age; MA, Middle Ages

When comparing Estonian CWC and EstBA using autosomal outgroup f3 and Patterson’s D statistics (Table S3), the latter is more similar to other Baltic BA populations, to Baltic IA and Middle Age (MA) populations, and also to populations similar to WHGs and Scandinavian hunter-gatherers (SHGs), but not to Estonian CCC (Figures 2A and S2A; Data S1). The increase in WHG or SHG ancestry could be connected to western influences seen in material culture [20, 21] and facilitated by a decline in local population after the CCC-CWC period [20]. A slight trend of bigger similarity of Estonian CWC to forest or steppe zone populations and of EstBA to European early farmer populations can also be seen.

(…) When comparing to modern populations, Estonian CWC is slightly more similar to Caucasus individuals but EstBA to Baltic populations and Finnic speakers (Figure 2B; Data S1). Outgroup f3 and D statistics do not reveal apparent differences when comparing EstBA to EstIA, EstIA to IngIA, and EstIA to EstMA (Data S1).

estonian-ba-ia-ancestry
qpAdm results. Error bars indicate one SE. Central MN, Central European Middle Neolithic; EstBA, Estonian Bronze Age; EstIA, Estonian Iron Age; IngIA, Ingrian Iron Age; EstMA, Estonian Middle Ages; WHG, western hunter-gatherers.

These results highlight how uniparental and autosomal data can lead to different demographic inferences—the genetic change between CWC and BA not seen in uniparental lineages is clear in autosomal data and the appearance of chrY hg N in the IA is not matched by a clear shift in autosomal profiles.

EstBA individuals have no Nganasan-related ancestry and EstIA, IngIA, and EstMA individuals on average have 2% or 4% (Figure 3; Data S1). The differentiation remains when using BA or IA Fennoscandian populations [26] instead of Nganasans (Data S1). Notably, the proportion of Nganasan-related ancestry varies between 0% and 12% among sampled EstIA, IngIA, and EstMA individuals (Data S1), which may suggest its relatively recent admixture into the target population. Moreover, two individuals from Kunda (0LS10 and V10) have the highest proportions of Nganasan ancestry among EstIA (6% and 8%), one of them has chrY hg N3a, and isotopic analysis suggests neither individual being born in Kunda [34].

About these two males from Tarand-graves, ‘foreign’ to Kunda:

0LS10: Male from tarand III (burial 9; TÜ 1325: L777), age 17–25 years [34]. He had a fragment of a sheep/goat bone and ceramics as grave goods. This burial has two radiocarbon dates: 2430 ± 35 BP (Poz-10801; 760–400 cal BC) and 2530 ± 41 BP (UBA-26114; 800–530 cal BC) [34]. According to the isotopic analysis, the person was not born in the vicinity of Kunda; his place of birth is still unknown (but south-western Finland and Sweden are excluded) [34]. Sampled tooth r P1.

V10: Male from tarand XI (burial 24; TÜ 1325: L1925), age 25–35 years [34], date 2484 ± 40 BP (UBA-26115; 790–430 cal BC) [34]. He had a few potsherds near the skull. Likewise, this person was not locally born [34]. Sampled tooth l P1.

estonia-bronze-iron-age-steppe-siberian
Autosomal Analyses’ Results for Gyvakarai1 as the closest available Corded Ware source for Balto-Finnic populations.

The paper shows thus:

  • Major continuity of ancestry from Corded Ware to modern Estonians, with only slight changes in different periods. In fact, one of the best fits for the Late Bronze Age ancestry is Gyvakarai1, one of the Corded Ware “outliers” described as “closer to Yamna”, which I already said may be closer to Sredni Stog/EHG populations instead. Another interesting take is that the change from Bronze Age to Iron Age corresponds to an increase in Baltic Corded Ware-related ancestry, rather than being driven by Siberian ancestry.
  • pca-mittnik-gyvakarai
    File modified by me from Mittnik et al. (2018) to include the approximate position of the most common ancestral components, and an identification of potential outliers. Zoomed-in version of the European Late Neolithic and Bronze Age samples. “Principal components analysis of 1012 present-day West Eurasians (grey points, modern Baltic populations in dark grey) with 294 projected published ancient and 38 ancient North European samples introduced in this study (marked with a red outline). From Mittnik et al. (2018).
  • A Volosovo-related migration of hg. N1c with Netted Ware into the area seems to be discarded, based on the full replacement of paternal lines and continuity of R1a-Z283. It is only during the Tarand-grave period when a system of chiefdoms (spread from Ananyino/Akozino) brings haplogroup N1c to the Gulf of Finland. During the Iron Age, the proportion of paternal lineages is still clearly in favour of R1a (50% in the coast, 100% in Ostrobothnia), which indicates a gradual replacement led by elites, likely because of the incorporation of Akozino warrior-traders spreading all over the Baltic, bringing the described shared Mordvinic traits in Fennic.
  • 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).
  • The arrival of Akozino warrior-traders (bringing N1c and R1a lineages) was probably linked to this minimal “Nganasan-like” ancestry of some samples in the transition to the Iron Age. This arrival is supported by samples 0LS10 (the earliest hg. N1c) and V10 (of hg. R1a), both dated to ca. 800-400 BC, with V10 showing the highest “Nganasan-like” ancestry with 4.8%, both of them neighbouring samples showing 0%. This variable admixture among local and foreign paternal lineages might support the described social system of family alliances with intermarriages. In fact, a medieval sample, 0LS03_1 (hg. R1a) also shows a recent “Nganasan-like” ancestry, which probably points to the integration of different Arctic-related ancestry components among Modern Estonians, in this case related to Finnish expansions and thus integration of Levänluhta-related ancestry, as per the supplementary data.
  • NOTE. Such minimal proportions of “Nganasan-like” ancestry evidence the process of admixture of Volga Finns in Akozino territory through their close interactions with Permians of Ananyino, who in turn acquired this Palaeo-Arctic admixture most likely during the expansion of the linguistic community to hunter-gatherer territories, to the north of the Cis-Urals. This process of stepped infiltration and expansion without language change is not dissimilar to the one seen among Indo-Iranians and Balto-Slavs of hg. R1b, or Vasconic speakers of hg. I2a, although in the case of Baltic Finns of hg. R1a the process of infiltration and expansion of hg. N1c is much less dramatic, with no radical replacement anywhere before the huge bottlenecks observable in Finns.

  • The expansion of haplogroup N1c among Finnic populations, as we are going to see in samples from the Middle Ages such as Luistari, is the consequence of late founder effects after huge bottlenecks expected based on the analysis of modern populations. The expansion of N1c-VL29 is different in origin from that of N1c-Z1936 among Samic (later integrated into Finnish populations), most likely from the east and originally associated with Lovozero Ware.
haplogroup_n3a3
Frequency-Distribution Maps of Individual Subclade N3a3 / N1a1a1a1a1a-CTS2929/VL29, probably initially with Akozino warrior-traders. Map from Ilumäe et al. (2016).

In spite of all this, the conclusion of the paper is (surprise!) that Siberian ancestry and hg. N heralded the arrival of Finnic to the Gulf of Finland in the Iron Age… However, this conclusion is supposedly* supported, not by their previous papers, but by a recent phylogenetic study by Honkola et al. (2013), which doesn’t actually argue for such a late ‘arrival’: it argues for the split of Balto-Finnic around 1500 BC.

NOTE. I say ‘supposedly’ because Kristiina Tambets, for example, has been following the link of Uralic with haplogroup N since the 2000s, so this is not some conclusion they just happened to misread from some random paper they Googled. In those initial assessments, she argued that the “ancient homeland” of the Tat C mutation suggested that Finno-Ugrians were in Fennoscandia before Indo-Europeans. Apparently, since haplogroup N appears later and from the east, it is now more important to follow this haplogroup than what is established in archaeology and linguistics.

Even in the referred paper, this split is considered an in situ development, since the phylogenetic study takes the information – among others – 1) from Parpola and Carpelan, who consider Netted Ware, a culture derived from Fatyanovo/Abashevo and Volosovo, as the culprit of the Finno-Ugric expansion; and 2) from Kallio (2006), who clearly states that Proto-Balto-Finnic (like Proto-Finno-Samic) was spoken around the Gulf of Finland during the Bronze Age. Both of them set the terminus ante quem of the language presence in the Baltic ca. 1900 BC.

Anyways, as a consequence of geneticists keeping these untenable pre-ancient DNA haplogroup-based arguments today, I expect to see this “Finnic” language expansion also described for the Western Baltic, Scandinavia or northern Europe, when this same proportion of hg. N1c and “Nganasan” ancestry is observed in Iron Age samples around the Baltic Sea. The nativist trends that this domination of “Finns” all over Northern Europe 2,500 years ago will create will be even more fun to read than the current ones…

EDIT (10 May 2019) How I see the reaction of many to ancient DNA, in keeping their old theories:

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

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

The complex origin of Samoyedic-speaking populations

uralic-turkic

Open access Siberian genetic diversity reveals complex origins of the Samoyedic-speaking populations, by Karafet et al. Am J Hum Biol (2018) e23194.

Interesting excerpts (emphasis mine):

Siberian groups

Consistent with their origin, Mongolic-speaking Buryats demonstrate genetic similarity with Mongols, and Turkic-speaking Altai-Kizhi and Teleuts are drawn close to CAS groups. The Tungusic-speaking Evenks collected in central and eastern Siberia cluster together and overlap with Yukagirs. Dolgans are widely scattered in the plot, justifying their recent origin from one Evenk clan, Yakuts, and Russian peasants in the 18th century (Popov, 1964). Uralic-speaking populations comprise a very wide cluster with Komi drawn to Europe, and Khants showing a closer affinity with Selkups, Tundra and Forest Nentsi. Yenisey-speaking Kets are intermingled with Selkups. Interestingly, Samoyedic-speaking Nganasans from the Taymyr Peninsula form a separate tight cluster closer to Evenks, Yukagirs, and Koryaks.

pca-siberian-uralic
Principal component analysis (PCA) using the “drop one in” technique for 27 present-day (N = 424) and 6 ancient populations (N = 20). PCA was performed on 281 093 SNPs from the intersection of our data with publicly available ancient Siberian samples

ADMIXTURE and the “Siberian component”

Among Siberians, the Komi are primarily Europeans, while Nganasans, Evenks, Yukagirs, and Koryaks are nearly 100% East Asians. At K = 4 finer scale subcontinental structure can be distinguished with the emergence of a “Siberian” component. This component is highly pronounced in the Nganasans. Outside Siberia, this component is present in Germany and in CAS at low frequency. Within ancient cultures, this component has the highest frequency in three BA Karasuk samples. It is also found in Mal’ta, ENE Afanasievo and BA Andronovo, but not in Ust’-Ishim and BA Okunevo. At K = 5, the “Siberian” component is roughly subdivided into two components with different geographic distributions. The “Nganasan” component is frequent in nearly all Siberian populations, except the Komi, Kets and Selkups. The newly derived “Selkup-Ket” component is found at high frequencies in western Siberian populations. It is observed in BA Karasuk and in Mal’ta. At K = 6, the western Siberian “Nentsi-Khant” ancestry component was developed in Forest and Tundra Nentsi, Khants. This component is also present at low levels in EUR, CAS, Tibet, and southern Siberia.

Identity-by-descent

The Dolgans share more segments with the Nganasans than within themselves (54.13 vs 41.72, Mann-Whitney test, P = .000000000001562546). The result is not surprising as the demographic data showed that the Nganasans were subjected to intense assimilation by the Dolgans in the second half of the 20th century (Goltsova, Osipova, Zhadanov, & Villems, 2005). Tundra Nentsi share more IBD with Forest Nentsi than within themselves (83.96 vs 50.3, P = .000055) possibly due to the common origin and long-term gene flow. The Ket and Selkup populations allocate significantly more IBD blocks between populations than with individuals from their own population (121.2 cM vs 85.9 cM for Kets, P = .000008, and 121.2 cM vs 114.9 cM for Selkups, P = .043).

admixture-siberian
ADMIXTURE plot. Clustering of 444 individuals from 27 present-day and 6 ancient populations (281 093 SNPs) assuming K6 to K7 clusters. Individuals are shown as vertical bars colored in ratio to their estimated ancestry within each cluster

Haplogroup N in Siberia

Although Siberia exhibits 42 haplogroups, the vast majority of Siberian Y-chromosomes belong only to 4 of the 18 major clades (N = 46.2%; C = 20.9%; Q = 14.4%; and R = 15.2%). The Y-chromosome haplogroup N is widely spread across Siberia and Eastern Europe (Ilumae et al., 2016; Karafet et al., 2002; Wong et al., 2016) and reaches its maximum frequency among Siberian populations such as Nganasans (94.1%) and Yakuts (91.9%). Within Siberia, two sister subclades N-P43 and N-L708 show different geographic distributions. N-P43 and derived haplogroups N-P63 and N- P362 (phylogenetically identical to N-B478* and N-B170, respectively) (Ilumae et al., 2016) are extremely rare in other major geographic regions. Likely originating in western Siberia, they are limited almost entirely to northwest Siberia, the Volga- Uralic regions, and the Taymyr Peninsula (ie, do not extend to eastern Siberia). Conversely, clade N-L708 is frequent in all Siberian populations except the Kets and Selkups, reaching its highest frequency in the Yakuts (91.9%).

Surprisingly, not a single sign of the proposed reindeer pastoralist horde led by Nganasans into north-eastern Europe. This is strange because “Siberian” migrants hypothetically imposed their language over Indo-Europeans quite recently, apparently after the Iron Age

Interesting comparisons among Siberian groups, though.

Related

Minimal gene flow from western pastoralists in the Bronze Age eastern steppes

jeong-steppes-mongolia

Open access paper Bronze Age population dynamics and the rise of dairy pastoralism on the eastern Eurasian steppe, by Jeong et al. PNAS (2018).

Interesting excerpts (emphasis mine):

To understand the population history and context of dairy pastoralism in the eastern Eurasian steppe, we applied genomic and proteomic analyses to individuals buried in Late Bronze Age (LBA) burial mounds associated with the Deer Stone-Khirigsuur Complex (DSKC) in northern Mongolia. To date, DSKC sites contain the clearest and most direct evidence for animal pastoralism in the Eastern steppe before ca. 1200 BCE.

Most LBA Khövsgöls are projected on top of modern Tuvinians or Altaians, who reside in neighboring regions. In comparison with other ancient individuals, they are also close to but slightly displaced from temporally earlier Neolithic and Early Bronze Age (EBA) populations from the Shamanka II cemetry (Shamanka_EN and Shamanka_EBA, respectively) from the Lake Baikal region. However, when Native Americans are added to PC calculation, we observe that LBA Khövsgöls are displaced from modern neighbors toward Native Americans along PC2, occupying a space not overlapping with any contemporary population. Such an upward shift on PC2 is also observed in the ancient Baikal populations from the Neolithic to EBA and in the Bronze Age individuals from the Altai associated with Okunevo and Karasuk cultures.

pca-eurasians-karasuk-khovsgol
Image modified from the article. Karasuk cluster in green, closely related to sample ARS026 in red. Principal Component Analysis (PCA) of selected 2,077 contemporary Eurasians belonging to 149 groups. Contemporary individuals are plotted using three-letter abbreviations for operational group IDs. Group IDs color coded by geographic region. Ancient Khövsgöl individuals and other selected ancient groups are represented on the plot by filled shapes. Ancient individuals are projected onto the PC space using the “lsqproject: YES” option in the smartpca program to minimize the impact of high genotype missing rate.

(…) two individuals fall on the PC space markedly separated from the others: ARS017 is placed close to ancient and modern northeast Asians, such as early Neolithic individuals from the Devil’s Gate archaeological site (22) and present-day Nivhs from the Russian far east, while ARS026 falls midway between the main cluster and western Eurasians.

Upper Paleolithic Siberians from nearby Afontova Gora and Mal’ta archaeological sites (AG3 and MA-1, respectively) (25, 26) have the highest extra affinity with the main cluster compared with other groups, including the eastern outlier ARS017, the early Neolithic Shamanka_EN, and present-day Nganasans and Tuvinians (Z > 6.7 SE for AG3). Main cluster Khövsgöl individuals mostly belong to Siberian mitochondrial (A, B, C, D, and G) and Y (all Q1a but one N1c1a) haplogroups.

mongolia-botai-ehg-ane-cline
The genetic affinity of the Khövsgöl clusters measured by outgroup-f3 and -f4 statistics. (A) The top 20 populations sharing the highest amount of >genetic drift with the Khövsgöl main cluster measured by f3(Mbuti; Khövsgöl, X). (B) The top 15 populations with the most extra affinity with each of the three Khövsgöl clusters in contrast to Tuvinian (for the main cluster) or to the main cluster (for the two outliers), measured by f4(Mbuti, X; Tuvinian/Khövsgöl, Khövsgöl/ARS017/ARS026). Ancient and contemporary groups are marked by squares and circles, respectively. Darker shades represent a larger f4 statistic.

Previous studies show a close genetic relationship between WSH populations and ANE ancestry, as Yamnaya and Afanasievo are modeled as a roughly equal mixture of early Holocene Iranian/ Caucasus ancestry (IRC) and Mesolithic Eastern European hunter-gatherers, the latter of which derive a large fraction of their ancestry from ANE. It is therefore important to pinpoint the source of ANE-related ancestry in the Khövsgöl gene pool: that is, whether it derives from a pre-Bronze Age ANE population (such as the one represented by AG3) or from a Bronze Age WSH population that has both ANE and IRC ancestry.

The amount of WSH contribution remains small (e.g., 6.4 ± 1.0% from Sintashta). Assuming that the early Neolithic populations of the Khövsgöl region resembled those of the nearby Baikal region, we conclude that the Khövsgöl main cluster obtained ∼11% of their ancestry from an ANE source during the Neolithic period and a much smaller contribution of WSH ancestry (4–7%) beginning in the early Bronze Age.

khovsgol-shamanka-sintashta
Admixture modeling of Altai populations and the Khövsgöl main cluster using qpAdm. For the archaeological populations, (A) Shamanka_EBA and (B and C) Khövsgöl, each colored block represents the proportion of ancestry derived from a corresponding ancestry source in the legend. Error bars show 1 SE. (A) Shamanka_EBA is modeled as a mixture of Shamanka_EN and AG3. The Khövsgöl main cluster is modeled as (B) a two-way admixture of Shamanka_EBA+Sintashta and (C) a three-way admixture Shamanka_EN+AG3+Sintashta.

Apparently, then, the first individual with substantial WSH ancestry in the Khövsgöl population (ARS026, of haplogroup R1a-Z2123), directly dated to 1130–900 BC, is consistent with the first appearance of admixed forest-steppe-related populations like Karasuk (ca. 1200-800 BC) in the Altai. Interestingly, haplogroup N1a1a-M178 pops up (with mtDNA U5a2d1) among the earlier Khövsgöl samples.

I will repeat what I wrote recently here: Samoyedic arrived in the Altai with Karasuk and hg R1a-Z645 + Steppe_MLBA-like ancestry, admixed with Altai populations, clustering thus within an Ancient Altai cline. Only later did N1a1a subclades infiltrate Samoyedic (and Ugric) populations, bringing them closer to their modern Palaeo-Siberian cline. The shared mtDNA may support an ancestral EHG-“Siberian” cline, or else a more recent Afanasevo-related origin.

east-uralic-clines
Modified image from Jeong et al. (2018), supplementary materials. The first two PCs summarizing the genetic structure within 2,077 Eurasian individuals. The two PCs generally mirror geography. PC1 separates western and eastern Eurasian populations, with many inner Eurasians in the middle. PC2 separates eastern Eurasians along the north-south cline and also separates Europeans from West Asians. Ancient individuals (color-filled shapes), including two Botai individuals, are projected onto PCs calculated from present-day individuals. Read more.

Also interesting, Q1a2 subclades and ANE ancestry making its appearance everywhere among ancestral Eurasian peoples, as Chetan recently pointed out.

Related

Corded Ware—Uralic (IV): Hg R1a and N in Finno-Ugric and Samoyedic expansions

haplogroup-uralians

This is the fourth of four posts on the Corded Ware—Uralic identification:

Let me begin this final post on the Corded Ware—Uralic connection with an assertion that should be obvious to everyone involved in ethnolinguistic identification of prehistoric populations but, for one reason or another, is usually forgotten. In the words of David Reich, in Who We Are and How We Got Here (2018):

Human history is full of dead ends, and we should not expect the people who lived in any one place in the past to be the direct ancestors of those who live there today.

Haplogroup N

Another recurrent argument – apart from “Siberian ancestry” – for the location of the Uralic homeland is “haplogroup N”. This is as serious as saying “haplogroup R1” to refer to Indo-European migrations, but let’s explore this possibility anyway:

Ancient haplogroups

We have now a better idea of how many ancient migrations (previously hypothesized to be associated with westward Uralic migrations) look like in genetic terms. From Damgaard et al. (Science 2018):

These serial changes in the Baikal populations are reflected in Y-chromosome lineages (Fig. SA; figs. S24 to S27, and tables S13 and SI4). MAI carries the R haplogroup, whereas the majority of Baikal_EN males belong to N lineages, which were widely distributed across Northern Eurasia (29), and the Baikal_LNBA males all carry Q haplogroups, as do most of the Okunevo_EMBA as well as some present-day Central Asians and Siberians.

The only N1c1 sample comes from Ust’Ida Late Neolithic, 180km to the north of Lake Baikal, which – together with the Bronze Age sample from the Kola peninsula, and the medieval sample from Ust’Ida – gives a good idea of the overall expansion of N subclades and Siberian ancestry among the Circum-Arctic peoples of Eurasia, speakers of Palaeo-Siberian languages.

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

Modern haplogroups

What we should expect from Uralic peoples expanding with haplogroup N – seeing how Yamna expands with R1b-L23, and Corded Ware expands with R1a-Z645 – is to find a common subclade spreading with Uralic populations. Let’s see if it works like that for any N-X subclade, in data from Ilumäe et al. (2016):

haplogroup_n1
Geographic-Distribution Map of hg N3 / N1c / N1a.

Within the Eurasian circum-Arctic spread zone, N3 and N2a reveal a well-structured spread pattern where individual sub-clades show very different distributions:

N1a1-M46 (or N-TAT), formed ca. 13900 BC, TMRCA 9800 BC

   N1a1a2-B187, formed ca. 9800 BC, TMRCA 1050 AD:

The sub-clade N3b-B187 is specific to southern Siberia and Mongolia, whereas N3a-L708 is spread widely in other regions of northern Eurasia.

     N1a1a1a-L708, formed ca. 6800 BC, TMRCA 5400 BC.

       N1a1a1a2-B211/Y9022, formed ca. 5400 BC, TMRCA 1900 BC:

The deepest clade within N3a is N3a1-B211, mostly present in the Volga-Uralic region and western Siberian Khanty and Mansi populations.

         N1a1a1a1a-L392/L1026), formed ca. 4400 BC, TMRCA 2800 BC:

The neighbor clade, N3a3’6-CTS6967, spreads from eastern Siberia to the eastern part of Fennoscandia and the Baltic States

haplogroup_n3a3
Frequency-Distribution Maps of Individual Subclade N3a3 / N1a1a1a1a1a-CTS2929/VL29, probably initially with Akozino warrior-traders.

           N1a1a1a1a1a-CTS2929/VL29, formed ca. 2100 BC, TMRCA 1600 BC:

In Europe, the clade N3a3-VL29 encompasses over a third of the present-day male Estonians, Latvians, and Lithuanians but is also present among Saami, Karelians, and Finns (Table S2 and Figure 3). Among the Slavic-speaking Belarusians, Ukrainians, and Russians, about three-fourths of their hg N3 Y chromosomes belong to hg N3a3.

In the post on Finno-Permic expansions, I depicted what seems to me the most likely way of infiltration of N1c-L392 lineages with Akozino warrior-traders into the western Finno-Ugric populations, with an origin around the Barents sea.

This includes the potential spread of (a minority of) N1c-B211 subclades due to contacts with Anonino on both sides of the Urals, through a northern route of forest and forest-steppe regions (equivalent to the distribution of Cherkaskul compared to Andronovo), given the spread of certain subclades in Ugric populations.

NOTE. An alternative possibility is the association of certain B211 subclades with a southern route of expansion with Pre-Scythian and Scythian populations, under whose influence the Ananino culture emerged -which would imply a very quick infiltration of certain groups of haplogroup N everywhere among Finno-Ugrics on both sides of the Urals – , and also the expansion of some subclades with Turkic-speaking peoples, who apparently expanded with alliances of different peoples. Both (Scythian and Turkic) populations expanded from East Asia, where haplogroup N (including N1c) was present since the Neolithic. I find this a worse model of expansion for upper clades, but – given the YFull estimates and the presence of this haplogroup among Turkic peoples – it is a possibility for many subclades.

           N1a1a1a1a2-Z1936, formed ca. 2800 BC, TMRCA 2400 BC:

The only notable exception from the pattern are Russians from northern regions of European Russia, where, in turn, about two-thirds of the hg N3 Y chromosomes belong to the hg N3a4-Z1936—the second west Eurasian clade. Thus, according to the frequency distribution of this clade, these Northern Russians fit better among other non-Slavic populations from northeastern Europe. N3a4 tends to increase in frequency toward the northeastern European regions but is also somewhat unexpectedly a dominant hg N3 lineage among most Turcic-speaking Volga Tatars and South-Ural Bashkirs.

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.

The expansion of N1a-Z1936 in Fennoscandia is most likely associated with the expansion of Saami into asbestos ware-related territory (like the Lovozero culture) during the Late Iron Age – and mixture with its population – , and with the later Fennic expansion to the east and north, replacing their language, as well as with Arctic and forest populations assimilated during Permic, Ugric, and Samoyedic expansions to the north.

           N1a1a1a1a4-M2019 (previously N3a2), formed ca. 4400 BC, TMRCA 1700 BC:

Sub-hg N3a2-M2118 is one of the two main bifurcating branches in the nested cladistic structure of N3a2’6-M2110. It is predominantly found in populations inhabiting present-day Yakutia (Republic of Sakha) in central Siberia and at lower frequencies in the Khanty and Mansi populations, which exhibit a distinct Y-STR pattern (Table S7) potentially intrinsic to an additional clade inside the sub-hg N3a2

The second widespread sub-clade of hg N is N2a. (…):

   N1a2b-P43 (B523/FGC10846/Y3184), formed ca. 6800 BC, TMRCA ca. 2700 BC:

The absolute majority of N2a individuals belong to the second sub-clade, N2a1-B523, which diversified about 4.7 kya (95% CI = 4.0–5.5 kya). Its distribution covers the western and southern parts of Siberia, the Taimyr Peninsula, and the Volga-Uralic region with frequencies ranging from from 10% to 30% and does not extend to eastern Siberia (…)

haplogroup_n2
Geographic-Distribution Map of hg N2a1 / N1a2b-P43

The “European” branch suggested earlier from Y-STR patterns turned out to consist of two clades

     N1a2b2a-Y3185/FGC10847, formed ca. 2200 BC, TMRCA 800 BC:

N2a1-L1419, spread mainly in the northern part of that region.

     N1a2b2b1-B528/Y24382, formed ca. 900 BC, TMRCA ca. 900 BC:

N2a1-B528, spread in the southern Volga-Uralic region.

Haplogroup R1a

We also have a good idea of the distribution of haplogroup R1a-Z645 in ancient samples. Its subclades were associated with the Corded Ware expansion, and some of them fit quite well the early expansion of Finno-Permic, Ugric, and Samoyedic peoples to the east.

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.

This is how the modern distribution of R1a among Uralians looks like, from the latest report in Tambets et al. (2018):

  • Among Fennic populations, Estonians and Karelians (ca. 1.1 million) have not suffered the greatest bottleneck of Finns (ca. 6-7 million), and show thus a greater proportion of R1a-Z280 than N1c subclades, which points to the original situation of Fennic peoples before their expansion. To trust Finnish Y-DNA to derive conclusions about the Uralic populations is as useful as relying on the Basque Y-DNA for the language spread by R1b-P312
  • Among Volga-Finnic populations, Mordovians (the closest to the original Uralic cluster, see above) show a majority of R1a lineages (27%).
  • Hungarians (ca. 13-15 million) represent the majority of Ugric (and Finno-Ugric) peoples. They are mainly R1a-Z280, also R1a-Z2123, have little N1c, and lack Siberian ancestry, and represent thus the most likely original situation of Ugric peoples in 4th century AD (read more on Avars and Hungarians).
  • Among Samoyedic peoples, the Selkup, the southernmost ones and latest to expand – that is, those not heavily admixed with Siberian populations – , also have a majority of R1a-Z2123 lineages (see also here for the original Samoyedic haplogroups to the south).

To understand the relevance of Hungarians for Ugric peoples, as well as Estonians, Karelians, and Mordovians (and northern Russians, Finno-Ugric peoples recently Russified) for Finno-Permic peoples, as opposed to the Circum-Arctic and East Siberian populations, one has to put demographics in perspective. Even a modern map can show the relevance of certain territories in the past:

population-density
Population density (people per km2) map of the world in 1994. From Wikipedia.

Summary of ancestry + haplogroups

Fennic and Samic populations seem to be clearly influenced by Palaeo-Laplandic peoples, whereas Volga-Finnic and especially Permic populations may have received gene flow from both, but essentially Palaeo-Siberian influence from the north and east.

The fact that modern Mansis and Khantys offer the highest variation in N1a subclades, and some of the highest “Siberian ancestry” among non-Nganasans, should have raised a red flag long ago. The fact that Hungarians – supposedly stemming from a source population similar to Mansis – do not offer the same amount of N subclades or Siberian ancestry (not even close), and offer instead more R1a, in common with Estonians (among Finno-Samic peoples) and Mordvins (among Volga-Finnic peoples) should have raised a still bigger red flag. The fact that Nganasans – the model for Siberian ancestry – show completely different N1a2b-P43 lineages should have been a huge genetic red line (on top of the anthropological one) to regard them as the Uralian-type population.

We know now that ethnolinguistic groups have usually expanded with massive (usually male-biased) migrations, and that neighbouring locals often ‘resurge’ later without changing the language. That is seen in Europe after the spread of Bell Beakers, with the increase of previous ancestry and lineages in Scandinavia during the formation of the Nordic ethnolinguistic community; in Central-West Europe, with the resurgence of Neolithic ancestry (and lineages) during the Bronze Age over steppe ancestry; and in Central-East Europe (with Unetice or East European Bronze Age groups like Mierzanowice, Trzciniec, or Lusatian) showing an increase in steppe ancestry (and resurge of R1a subclades); none of them represented a radical ethnolinguistic change.

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).

It is not hard to model the stepped arrival, infiltration, and/or resurge of N subclades and “Siberian ancestries”, as well as their gradual expansion in certain regions, associated with certain migrations first – such as the expansions to the Circum-Arctic region, and later the Scythian- and Turkic-related movements – , as well as limited regional developments, like the known bottleneck in Finns, or the clear late expansion of Ugric and Samoyedic languages to the north among nomadic Palaeo-Siberians due to traditions of exogamy and multilingualism. This fits quite well with the different arrival of N (N1c and xN1c) lineages to the different Uralic-speaking groups, and to the stepped appearance of “Siberian ancestry” in the different regions.

The aternative

It is evident that a lot of people were too attached to the idea of Palaeolithic R1b lineages ‘native’ to western Europe speaking Basque languages; of R1a lineages speaking Indo-European and spreading with Yamna; and N lineages ‘native’ to north-eastern Europe and speaking Uralic, and this is causing widespread weeping and gnashing of teeth (instead of the joy of discovering where one’s true patrilineal ancestors come from, and what language they spoke in each given period, which is the supposed objective of genetic genealogy…)

Since an Indo-Germanic branch (as revived now by some in the Copenhaguen group to fit Kristiansen’s theory of the 1980s with recent genetic data) does not make any sense in linguistics, the finding of R1a in Yamna would not have led where some think it would have, because North-West Indo-European would still be the main Late PIE branch in Europe. Don’t take my word for it; take James P. Mallory’s (2013).

mallory-adams-tree
The levels of Indo-European reconstruction, from Mallory & Adams (2006).

If an (unlikely) Indo-Slavonic group were posited, though, such a group would still be bound (with Indo-Iranian) to the steppes with East Yamna/Poltavka (admixing with Abashevo migrants, but retaining its language), developing Sintashta/Potapovka → Srubna/Andronovo, and R1a lineages would have equally undergone the known bottlenecks of the steppes where they replaced R1b-Z2103 – which this eastern group shares with Balkan languages, a haplogroup that links therefore together the Graeco-Aryan group.

As far as I know – and there might be many other similar pet theories out there – there have been proposals of “modern Balto-Slavic-like” populations (in an obvious circular reasoning based on modern populations) in some Scythian clusters of the Iron Age.

NOTE. I will not enter into “Balto-Slavic-like R1a” of the Late Bronze Age or earlier because no one can seriously believe at this point of development of Population Genetics that autosomal similarity predating 1,500+ years the appearance of Slavs equates to their (ethnolinguistic) ancestral population, without a clear intermediate cultural and genetic trail – something we lack today in the Slavic case even for the late Roman period…

finno-saamic-palaeo-germanic-substratum
The Finnic and Saamic separation looks shallower than it actually is. Invisible convergence can be ‘triangulated’ with the help of Germanic layers of mutual loanwords (Häkkinen 2012).

We also know of R1a-Z280 lineages in Srubna, probably expanding to the west. With that in mind, and knowing that Palaeo-Germanic was in close contact with Finno-Samic while both were already separated but still in contact, and that Palaeo-Germanic was also in contact and closely related to a ‘Temematic’ distinct from Balto-Slavic (and also that early Proto-Baltic and Proto-Slavic from the Roman Iron Age and later were in contact with western Uralic) this will be the linguistic map of the Iron Age if R1a is considered to expand Indo-European from some kind of “patron-client” relationship with west Yamna:

palaeo-germanic-italo-celtic
Eastern European language map during the Late Bronze Age / Iron Age, if R1a spread Indo-European languages and Eastern Yamna spoke Indo-Slavonic. Palaeo-Germanic (i.e. Pre- to Proto-Germanic) needs to be in contact with both the Samic Lovozero population and the Fennic west Circum-Arctic one. Italic and Celtic in contact with Pre-Germanic. Germanic in contact with Temematic. Balto-Slavic in contact with Iranian, and near Fennic to allow for later loanwords. For Germanic and Temematic, see Kortlandt (2018).

You might think I have some personal or political reason against this kind of proposals. I haven’t. We have been proposing Indo-European to be the language of the European Union for more than 10 years, so to support R1b-Italo-Celtic in the whole Western Europe, R1a-Germanic in Central and Eastern Europe, and R1a-Indo-Slavonic in the steppes (as the Danish group seems to be doing) has nothing inherently bad (or good) for me. If anything, it gives more reason to support the revival of North-West Indo-European in Europe.

My problem with this proposal is that it is obviously beholden to the notion of the uninterrupted cultural, historic and ethnic continuity in certain territories. This bias is common in historiography (von Falkenhausen 1993), but it extends even more easily into the lesser known prehistory of any territory, and now more than ever some people feel the need to corrupt (pre)history based on their own haplogroups (or the majority haplogroups of their modern countries). However, more than on philosophical grounds, my rejection is based on facts: this picture is not what the combination of linguistic, archaeological, and genetic data shows. Period.

Nevertheless, if Yamna + Corded Ware represented the “big and early expansion” of Germanic and Italo-Celtic peoples proper of the dream Nazi’s Lebensraum and Fascist’s spazio vitale proposals; Uralians were Siberian hunter-gatherers that controlled the whole eastern and northern Russia, and miraculously managed to push (ethnolinguistically) Neolithic agropastoralists to the west during and after the Iron Age, with gradual (and often minimal) genetic impact; and Balto-Slavic peoples were represented by horse riders from Pokrovka/Srubna, hiding then somewhere around the forest-steppe until after the Scythian expansion, and then spreading their language (without much genetic impact) during the early Middle Ages…so be it.

See also

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