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

Corded Ware—Uralic (II): Finno-Permic and the expansion of N-L392/Siberian ancestry

finno-ugric-samoyedic

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

I read from time to time that “we have not sampled Uralic speakers yet”, and “we are waiting to see when Uralic-speaking peoples are sampled”. Are we, though?

Proto-language homelands are based on linguistic data, such as guesstimates for dialectal evolution, loanwords and phonetic changes for language contacts, toponymy for ancient territories, etc. depending on the available information. The trace is then followed back, using available archaeological data, from the known historic speakers and territory to the appropriate potential prehistoric cultures. Only then can genetic analyses help us clarify the precise prehistoric population movements that better fit the models.

uralic-language-family
The traditional family tree of the Uralic branches. Kallio (2014)

The linguistic homeland

We thought – using linguistic guesstimates and fitting prehistoric cultures and their expansion – that Yamna was the Late Proto-Indo-European culture, so when Yamna was sampled, we had Late Proto-Indo-Europeans sampled. Simple deduction.

We thought that north-eastern Europe was a Uralic-speaking area during the Neolithic:

  • For those supporting a western continuity (and assuming CWC was Indo-European), the language was present at least since the Comb Ware culture, potentially since the Mesolithic.
  • For those supporting a late introduction into Finland, Uralic expanded the latest with Abashevo-related movements after its incorporation of Volosovo and related hunter-gatherers.

The expansion to the east must have happened through progressive infiltrations with Seima-Turbino / Andronovo-related expansions.

uralic-time-space
Some datings for the traditional proto-stages from Uralic to Finnic. Kallio (2014).

Finding the linguistic homeland going backwards can be described today as follows:

I. Proto-Fennic homeland

Based on the number of Baltic loanwords, not attested in the more eastern Uralic branches (and reaching only partially Mordvinic), the following can be said about western Finno-Permic languages (Junttila 2014):

The Volga-Kama Basin lies still too far east to be included in a list of possible contact locations. Instead, we could look for the contact area somewhere between Estonia in the west and the surroundings of Moscow in the east, a zone with evidence of Uralic settlement in the north and Baltic on the south side.

The only linguistically well-grounded version of the Stone Age continuation theory was presented by Mikko Korhonen in 1976. Its validity, however, became heavily threatened when Koivulehto 1983a-b proved the existence of a Late Proto-Indo-European or Pre-Baltic loanword layer in Saami, Finnic, and Mordvinic. Since this layer must precede the Baltic one and it was presumably acquired in the Baltic Sea region, Koivulehto posited it on the horizon of the Battle Axe period. This forces a later dating for the Baltic–Finnic contacts.

Today the Battle Axe culture is dated at 3200 to 3000 BC, a period far too remote to correspond linguistically with Proto-Baltic (Kallio 1998a).

Since the Baltic contacts began at a very initial phase of Proto-Finnic, the language must have been relatively uniform at that time. Hence, if we consider that the layer of Baltic loanwords may have spread over the Gulf of Finland at that time, we could also insist that the whole of the Proto-Finnic language did so.

migration-theory
Prehistoric Balts as the southern neighbours of Proto-Finnic speakers. 1 = The approximated area of Proto-Uralic. 2 = The approximated area of Finnic during the Iron Age. 3 = The area of ancient Baltic hydronyms. 4 = The area of Baltic languages in about 1200 AD. 5 = The problem: When did Uralic expand westwards and when did it meet Baltic? Junntila (2012).

II. Proto-Finno-Saamic homeland

The evidence of continued Palaeo-Germanic loanwords (from Pre- to Proto-Germanic stages) is certainly the most important data to locate the Finno-Saamic homeland, and from there backwards into the true Uralic homeland. Following Kallio (2017):

(…) the loanword evidence furthermore suggests that the ancestors of Finnic and Saamic had at least phonologically remained very close to Proto-Uralic as late as the Bronze Age (ca. 1700–500 BC). In particular, certain loanwords, whose Baltic and Germanic sources point to the first millennium BC, after all go back to the Finno-Saamic proto-stage, which is phonologically almost identical to the Uralic proto-stage (see especially the table in Sammallahti 1998: 198–202). This being the case, Dahl’s wave model could perhaps have some use in Uralic linguistics, too.

The presence of Pre-Germanic loanwords points rather to the centuries around the turn of the 2nd – 1st millennium BC or earlier. Proto-Germanic words must have been borrowed before the end of Germanic influence in the eastern Baltic at the beginning of the Iron Age, which sets a clear terminus ante quem ca. 800 BC.

The arrival of Bell Beaker peoples in Scandinavia ca. 2350 BC, heralding the formation of the Dagger Period, as well as the development of Pre-Germanic in common with Finnic-like populations point to the late 3rd / early 2nd millennium BC as the first time of close interaction through the Baltic region.

III. Proto-Uralic homeland

(…) the earliest Indo-European loanwords in the Uralic languages (…) show that Proto-Uralic cannot have been spoken much earlier than Proto-Indo-European dated about 3500 BC (Koivulehto 2001: 235, 257). As the same loanword evidence naturally also shows that the Uralic and Indo-European homelands were not located far from one another, the Uralic homeland can most likely be located in the Middle and Upper Volga region, right north of the Indo-European homeland*. From the beginning of the Subneolithic period about 5900 BC onwards, this region was an important innovation centre, from where several cultural waves spread to the Finnish Gulf area, such as the Sperrings Ware wave about 4900 BC, the Combed Ware wave about 3900 BC, and the Netted Ware wave about 1900 BC (Carpelan & Parpola 2001: 78–90).

The mainstream position is nowadays trying to hold together the traditional views of Corded Ware as Indo-European, and a Uralic Fennoscandia during the Bronze Age.

The following is an example of how this “Volosovo/Forest Zone hunter-gatherer theory” of Uralic origins looks like, as a ‘mixture’ of cultures and languages that benefits from the lack of genetic data for certain regions and periods (taken from Parpola 2018):

asbestos-ware
The extent of Typical Comb Ware (TCW), Asbestos- and Organic-tempered Wares (AOW) and Volosovo and Garino-Bor cultures; areas with deposits of native copper in Karelia and copperbearing sandstone in Volga-Kama-area are marked dark gray (after Zhuravlev 1977; Krajnov 1987; Nagovitsyn 1987; Chernykh 1992; Carpelan 1999; Zhul´nikov 1999). From Nordqvist et al. (2012).

The Corded Ware (or Battle Axe) culture intruded into the Eastern Baltic and coastal Finland already around 3100 BCE. The continuity hypothesis maintains that the early Proto-Finnic speakers of the coastal regions, who had come to Finland in the 4th millennium BCE with the Comb-Pitted Ware, coexisted with the Corded Ware newcomers, gradually adopting their pastoral culture and with it a number of NW-IE loanwords, but assimilating the immigrants linguistically.

The fusion of the Corded Ware and the local Comb-Pitted Ware culture resulted into the formation of the Kiukais culture (c. 2300–1500) of southwestern Finland, which around 2300 received some cultural impulses from Estonia, manifested in the appearance of the Western Textile Ceramic (which is different from the more easterly Textile Ceramic or Netted Ware, and which is first attested in Estonia c. 2700 BCE, cf. Kriiska & Tvauri 2007: 88), and supposed to have been accompanied by an influx of loanwords coming from Proto-Baltic. At the same time, the Kiukais culture is supposed to have spread the custom of burying chiefs in stone cairns to Estonia.

The coming of the Corded Ware people and their assimilation created a cultural and supposedly also a linguistic split in Finland, which the continuity hypothesis has interpreted to mean dividing Proto-Saami-Finnic unity into its two branches. Baltic Finnic, or simply Finnic, would have emerged in the coastal regions of Finland and in the northern East Baltic, while preforms of Saami would have been spoken in the inland parts of Finland.

The Nordic Bronze Age culture, correlated above with early Proto-Germanic, exerted a strong influence upon coastal Finland and Estonia 1600–700 BCE. Due to this, the Kiukais culture was transformed into the culture of Paimio ceramics (c. 1600–700 BCE), later continued by Morby ceramics (c. 700 BCE – 200 CE). The assumption is that clear cultural continuity was accompanied by linguistic continuity. Having assimilated the language of the Germanic traders and relatively few settlers of the Bronze Age, the language of coastal Finland is assumed to have reached the stage of Proto-Finnish at the beginning of the Christian era. In Estonia, the Paimio ceramics have a close counterpart in the contemporaneous Asva ceramics.

Eastern homelands?

I will not comment on Siberian or Central Asian homeland proposals, because they are obviously not mainstream, still less today when we know that Uralic was certainly in contact with Proto-Indo-European, and then with Pre- and Proto-Indo-Iranian, as supported even by the Copenhagen group in Damgaard et al. (2018).

This is what Kallio (2017) has to say about the agendas behind such proposals:

Interestingly, the only Uralicists who generally reject the Central Russian homeland are the Russian ones who prefer the Siberian homeland instead. Some Russians even advocate that the Central Russian homeland is only due to Finnish nationalism or, as one of them put it a bit more tactfully, “the political and ideological situation in Finland in the first decades of the 20th century” (Napolskikh 1995: 4).

Still, some Finns (and especially those who also belong to the “school who wants it large and wants it early”) simultaneously advocate that exactly the same Central Russian homeland is due to Finnlandisierung (Wiik 2001: 466).

Hence, for those of you willing to learn about fringe theories not related to North-Eastern Europe, you also have then the large and early version of the Uralic homeland, with Wiik’s Palaeolithic continuity of Uralic peoples spread over all of eastern and central Europe (hence EHG and R1a included):

atlantic-finnic-theory
Palaeolithic boat peoples and Finno-Ugric. Source

These fringe Finnish theories look a lot like the Corded Ware expansion… Better not go the Russian or Finnish nationalist ways? Agreed then, let’s discuss only rational proposals based on current data.

The archaeological homeland

For a detailed account of the Corded Ware expansion with Battle Axe, Fatyanovo-Balanovo, and Abashevo groups into the area, you can read my recent post on the origin of R1a-Z645.

1. Textile ceramics

During the 2nd millennium BC, textile impressions appear in pottery as a feature across a wide region, from the Baltic area through the Volga to the Urals, in communities that evolve from late Corded Ware groups without much external influence.

While it has been held that this style represents a north-west expansion from the Volga region (with the “Netted Ware” expansion), there are actually at least two original textile styles, one (earlier) in the Gulf of Finland, common in the Kiukainen pottery, which evolves into the Textile ware culture proper, and another which seems to have an origin in the Middle Volga region to the south-east.

The Netted ware culture is the one that apparently expands into inner Finland – a region not densely occupied by Corded Ware groups until then. There are, however, no clear boundaries between groups of both styles; textile impressions can be easily copied without much interaction or population movement; and the oldest textile ornamentation appeared on the Gulf of Finland. Hence the tradition of naming all as groups of Textile ceramics.

textile-ware-cultures
Maximum distribution of Textile ceramics during the Bronze Age (ca. 2000-800 BC). Asbestos-tempered ware lies to the north (and is also continued in western Fennoscandia).

The fact that different adjacent groups from the Gulf of Finland and Forest Zone share similar patterns making it very difficult to differentiate between ‘Netted Ware’ or ‘Textile Ware’ groups points to:

  • close cultural connections that are maintained through the Gulf of Finland and the Forest Zone after the evolution of late Corded Ware groups; and
  • no gross population movements in the original Battle Axe / Fatyanovo regions, except for the expansion of Netted Ware to inner Finland, Karelia, and the east, where the scattered Battle Axe finds and worsening climatic conditions suggest most CWC settlements disappeared at the end of the 3rd millennium BC and recovered only later.

NOTE. This lack of population movement – or at least significant replacement by external, non-CWC groups – is confirmed in genetic investigation by continuity of CWC-related lineages (see below).

The technology present in Textile ceramics is in clear contrast to local traditions of sub-Neolithic Lovozero and Pasvik cultures of asbestos-tempered pottery to the north and east, which point to a different tradition of knowledge and learning network – showing partial continuity with previous asbestos ware, since these territories host the main sources of asbestos. We have to assume that these cultures of northern and eastern Fennoscandia represent Palaeo-European (eventually also Palaeo-Siberian) groups clearly differentiated from the south.

The Chirkovo culture (ca. 1800-700 BC) forms on the middle Volga – at roughly the same time as Netted Ware formed to the west – from the fusion of Abashevo and Balanovo elites on Volosovo territory, and is also related (like Abashevo) to materials of the Seima-Turbino phenomenon.

Bronze Age ethnolinguistic groups

In the Gulf of Finland, Kiukainen evolves into the Paimio ceramics (in Finland) — Asva Ware (in Estonia) culture, which lasts from ca. 1600 to ca. 700 BC, probably representing an evolving Finno-Saamic community, while the Netted Ware from inner Finland (the Sarsa and Tomitsa groups) and the groups from the Forest Zone possibly represent a Volga-Finnic community.

NOTE. Nevertheless, the boundaries between Textile ceramic groups are far from clear, and inner Finland Netted Ware groups seem to follow a history different from Netted Ware groups from the Middle and Upper Volga, hence they could possibly be identified as an evolving Pre-Saamic community.

Based on language contacts, with Early Baltic – Early Finnic contacts starting during the Iron Age (ca. 500 BC onwards), this is a potential picture of the situation at the end of this period, when Germanic influence on the coast starts to fade, and Lusatian culture influence is stronger:

aikio-finnic-saamic
The linguistic situation in Lapland and the northern Baltic Sea Area in the Early Iron Age prior to the expansion of Saami languages; the locations of the language groups are schematic. The black line indicates the distribution of Saami languages in the 19th century, and the gray line their approximate maximal distribution before the expansion of Finnic. Aikio (2012)

The whole Finno-Permic community remains thus in close contact, allowing for the complicated picture that Kallio mentions as potentially showing Dahl’s wave model for Uralic languages.

Genetic data shows a uniform picture of these communities, with exclusively CWC-derived ancestry and haplogroups. So in Mittnik et al. (2018) all Baltic samples show R1a-Z645 subclades, while the recent session on Estonian populations in ISBA 8 (see programme in PDF) clearly states that:

[Of the 24 Bronze Age samples from stone-cist graves] all 18 Bronze Age males belong to R1a.

Regarding non-Uralic substrates found in Saami, supposedly absorbed during the expansion to the north (and thus representing languages spoken in northern Fennoscandia during the Bronze Age) this is what Aikio (2012) has to say:

The Saami substrate in the Finnish dialects thus reveals that also Lakeland Saami languages had a large number of vocabulary items of obscure origin. Most likely many of these words were substrate in Lakeland Saami, too, and ultimately derive from languages spoken in the region before Saami. In some cases the loan origin of these words is obvious due to their secondary Proto-Saami vowel combinations such as *ā–ë in *kāvë ‘bend; small bay’ and *šāpšë ‘whitefish’. This substrate can be called ‘Palaeo-Lakelandic’, in contrast to the ‘Palaeo-Laplandic’ substrate that is prominent in the lexicon of Lapland Saami. As the Lakeland Saami languages became extinct and only fragments of their lexicon can be reconstructed via elements preserved in Finnish place-names and dialectal vocabulary, we are not in a position to actually study the features of this Palaeo-Lakelandic substrate. Its existence, however, appears evident from the material above.

If we wanted to speculate further, based on the data we have now, it is very likely that two opposing groups will be found in the region:

A) The central Finnish group, in this hypothesis the Palaeo-Lakelandic group, made up of the descendants of the Mesolithic pioneers of the Komsa and Suomusjärvi cultures, and thus mainly Baltic HG / Scandinavian HG ancestry and haplogroups I / R1b(xM269) (see more on Scandinavian HG).

siberian-ancestry-map
Frequency map of the so-called ‘Siberian’ component. From Tambets et al. (2018).

B) Lapland and Kola were probably also inhabited by similar Mesolithic populations, until it was eventually assimilated by expanding Siberian groups (of Siberian ancestry and N1c-L392 lineages) from the east – entering the region likely through the Kola peninsula – , forming the Palaeo-Laplandic group, which was in turn later replaced by expanding Proto-Saamic groups.

Siberian ancestry appears first in Fennoscandia at Bolshoy Oleni Ostrov ca. 1520 BC, with haplogroup N1c-L392 (2 samples, BOO002 and BOO004), and with Siberian ancestry. This is their likely movement in north-eastern Europe, from Lamnidis et al (2018):

The large Siberian component in the Bolshoy individuals from the Kola Peninsula provides the earliest direct genetic evidence for an eastern migration into this region. Such contact is well documented in archaeology, with the introduction of asbestos-mixed Lovozero ceramics during the second millenium BC, and the spread of even-based arrowheads in Lapland from 1,900 BCE. Additionally, the nearest counterparts of Vardøy ceramics, appearing in the area around 1,600-1,300 BCE, can be found on the Taymyr peninsula, much further to the east. Finally, the Imiyakhtakhskaya culture from Yakutia spread to the Kola Peninsula during the same period.

saamic-lovozero-pca
PCA plot of 113 Modern Eurasian populations, with individuals from this study projected on the principal components. Uralic speakers are highlighted in light purple. Image modified from Lamnidis et al. (2018)

Obviously, these groups of asbestos-tempered ware are not connected to the Uralic expansion. From the same paper:

The fact that the Siberian genetic component is consistently shared among Uralic-speaking populations, with the exceptions of Hungarians and the non-Uralic speaking Russians, would make it tempting to equate this component with the spread of Uralic languages in the area. However, such a model may be overly simplistic. First, the presence of the Siberian component on the Kola Peninsula at ca. 4000 yBP predates most linguistic estimates of the spread of Uralic languages to the area. Second, as shown in our analyses, the admixture patterns found in historic and modern Uralic speakers are complex and in fact inconsistent with a single admixture event. Therefore, even if the Siberian genetic component partly spread alongside Uralic languages, it likely presented only an addition to populations carrying this component from earlier.

2. The Early Iron Age

The Ananino culture appears in the Vyatka-Kama area, famed for its metallurgy, with traditions similar to the North Pontic area, by this time developing Pre-Sauromatian traditions. It expanded to the north in the first half of the first millennium BC, remaining in contact with the steppes, as shown by the ‘Scythian’ nature of its material culture.

NOTE. The Ananino culture can be later followed through its zoomorphic styles into Iron Age Pjanoborskoi and Gljadenovskoi cultures, later to Ural-Siberian Middle Age cultures – Itkuska, Ust’-Poluiska, Kulaiska cultures –, which in turn can be related as prototypes of medieval Permian styles.

ananino-culture-homeland
Territory of (early and maximum) Ananino material culture. Vasilyev (2002).

At the same time as the Ananino culture begins to expand ca. 1000 BC, the Netted Ware tradition from the middle Oka expanded eastwards into the Oka-Vyatka interfluve of the middle Volga region, until then occupied by the Chirkovo culture. Eventually the Akozino or Akhmylovo group (ca. 800-300 BC) emerged from the area, showing a strong cultural influence from the Ananino culture, by that time already expanding into the Cis-Urals region.

The Akozino culture remains nevertheless linked to the western Forest Zone traditions, with long-ranging influences from as far as the Lusatian culture in Poland (in metallurgical techniques), which at this point is also closely related with cultures from Scandinavia (read more on genetics of the Tollense Valley).

malar-celts-ananino
Mälar celts and molds for casting (a) and the main distribution area (в) of Mälar-type celts of the Mälar type in the Volga-Kama region (according to Kuzminykh 1983: figure 92) and Scandinavia (according to Baudou 1960: Karte 10); Ananino celts and molds for casting (б) and the main distribution area (г) of the distribution of the celts of the Ananino type in the Volga-Kama area (according to Kuzminykh 1983: figure 9); dagger of Ananino type (д).Map from (Yushkova 2010)

Different materials from Akozino reach Fennoscandia late, at the end of the Bronze Age and beginning of the Early Iron Age, precisely when the influence of the Nordic Bronze Age culture on the Gulf of Finland was declining.

This is a period when Textile ceramic cultures in north-eastern Europe evolve into well-armed chiefdom-based groups, with each chiefdom including thousands or tens of thousands, with the main settlements being hill forts, and those in Fennoscandia starting ca. 1000-400 BC.

Mälar-type celts and Ananino-type celts appear simultaneously in Fennoscandia and the Forest Zone, with higher concentrations in south-eastern Sweden (Mälaren) and the Volga-Kama region, supporting the existence of a revived international trade network.

akozino-malar-axes-fennoscandia
Distribution of the Akozino-Mälar axes according to Sergej V. Kuz’minykh (1996: 8, Abb. 2).

The Paimio—Asva Ware culture evolves (ca. 700-200 BC) into the Morby (in Finland) — Ilmandu syle (in Estonia, Latvia, and Mälaren) culture. The old Paimio—Asva tradition continues side by side with the new one, showing a clear technical continuity with it, but with ornamentation compared to the Early Iron Age cultures of the Upper Volga area. This new south-eastern influence is seen especially in:

  • Akozino-Mälar axes (ca. 800-500 BC): introduced into the Baltic area in so great numbers – especially south-western Finland, the Åland islands, and the Mälaren area of eastern Sweden – that it is believed to be accompanied by a movement of warrior-traders of the Akozino-Akhmylovo culture, following the waterways that Vikings used more than a thousand years later. Rather than imports, they represent a copy made with local iron sources.
  • Tarand graves (ca. 500 BC – AD 400): these ‘mortuary houses’ appear in the coastal areas of northern and western Estonia and the islands, at the same time as similar graves in south-western Finland, eastern Sweden, northern Latvia and Courland. Similar burials are found in Akozino-Akhmylovo, with grave goods also from the upper and middle Volga region, while grave goods show continuity with Textile ware.

The use of asbestos increases in mainland Finnish wares with Kjelmøy Ware (ca. 700 BC – AD 300), which replaced the Lovozero Ware; and in the east in inner Finland and Karelia with the Luukonsaari and Sirnihta wares (ca. 700-500 BC – AD 200), where they replaced the previous Sarsa-Tomitsa ceramics.

The Gorodets culture appears during the Scythian period in the forest-steppe zone north and west of the Volga, shows fortified settlements, and there are documented incursions of Gorodets iron makers into the Samara valley, evidenced by deposits of their typical pottery and a bloom or iron in the region.

Iron Age ethnolinguistic groups

According to (Koryakova and Epimakhov 2007):

It is commonly accepted by archaeology, ethnography, and linguistics that the ancestors of the Permian peoples (the Udmurts, Komi-Permians, and Komi-Zyryans) left the sites of Ananyino cultural intercommunity.

NOTE. For more information on the Late Metal Ages and Early Medieval situation of Finno-Ugric languages, see e.g. South-eastern contact area of Finnic languages in the light of onomastics (Rahkonen 2013).

finno-saamic-mordvin
Yakhr-, -khra, yedr-, -dra and yer-/yar, -er(o), -or(o) names of lakes in Central and North Russia and the possible boundary of the proto-language words *jäkra/ä and *järka/ä. Rahkonen (2011)

Certain innovations shared between Proto-Fennic (identified with the Gulf of Finland) and Proto-Mordvinic (from the Gorodets culture) point to their close contact before the Proto-Fennic expansion, and thus to the identification of Gorodets as Proto-Mordvinic, hence Akozino as Volgaic (Parpola 2018):

  • the noun paradigms and the form and function of individual cases,
  • the geminate *mm (foreign to Proto-Uralic before the development of Fennic under Germanic influence) and other non-Uralic consonant clusters.
  • the change of numeral *luka ‘ten’ with *kümmen.
  • The presence of loanwords of non-Uralic origin, related to farming and trees, potentially Palaeo-European in nature (hence possibly from Siberian influence in north-eastern Europe).
ananino-textile-ware-cultures
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. Purple area show likely zones of predominant Siberian ancestry and N1c-L392 lineages. Blue areas likely zones of predominant CWC ancestry and R1a-Z645 lineages. Fading purple arrows represent likely stepped movements of haplogroup N1c-L392 for centuries (Siberian → Ananino → Akozino → Fennoscandia), found eventually in tarand graves. Blue arrows represent eventual expansions of Fennic and (partially displaced) Saamic. Modified image from Vasilyev (2002).

The introduction of a strongly hierarchical chiefdom system can quickly change the pre-existing social order and lead to a major genetic shift within generations, without a radical change in languages, as shown in Sintashta-Potapovka compared to the preceding Poltavka society (read more about Sintashta).

Fortified settlements in the region represented in part visiting warrior-traders settled through matrimonial relationships with local chiefs, eager to get access to coveted goods and become members of a distribution network that could guarantee them even military assistance. Such a system is also seen synchronously in other cultures of the region, like the Nordic Bronze Age and Lusatian cultures (Parpola 2013).

The most likely situation is that N1c subclades were incorporated from the Circum-Artic region during the Anonino (Permic) expansion to the north, later emerged during the formation of the Akozino group (Volgaic, under Anonino influence), and these subclades in turn infiltrated among the warrior traders that spread all over Fennoscandia and the eastern Baltic (mainly among Fennic, Saamic, Germanic, and Balto-Slavic peoples), during the age of hill forts, creating alliances partially based on exogamy strategies (Parpola 2013).

Over the course of these events, no language change is necessary in any of the cultures involved, since the centre of gravity is on the expanding culture incorporating new lineages:

  • first on the Middle Volga, when Ananino expands to the north, incorporatinig N1c lineages from the Circum-Artic region.
  • then with the expansion of the Akozino-Akhmylovo culture into Ananino territory, admixing with part of its population;
  • then on the Baltic region, when materials are imported from Akozino into Fennoscandia and the eastern Baltic (and vice versa), with local cultures being infiltrated by foreign (Akozino) warrior-traders and their materials;
  • and later with the different population movements that led eventually to a greater or lesser relevance of N1c in modern Finno-Permic populations.

To argue that this infiltration and later expansion of lineages changed the language in one culture in one of these events seems unlikely. To use this argument of “opposite movement of ethnic and language change” for different successive events, and only on selected regions and cultures (and not those where the greatest genetic and cultural impact is seen, like e.g. Sweden for Akozino materials) is illogical.

NOTE. Notice how I write here about “infiltration” and “lineages”, not “migration” or “populations”. To understand that, see below the next section on autosomal studies to compare Bronze Age, Iron Age, Medieval and Modern Estonians, and see how little the population of Estonia (homeland of Proto-Fennic and partially of Proto-Finno-Saamic) has changed since the Corded Ware migrations, suggesting genetic continuity and thus mostly close inter-regional and intra-regional contacts in the Forest Zone, hence a very limited impact of the absorbed N1c lineages (originally at some point incorporated from the Circum-Artic region). You can also check on the most recent assessment of R1a vs. N1c in modern Uralic populations.

Iron Age and later populations

From the session on Estonian samples on ISBA 8, by Tambets et al.:

[Of the 13 samples from the Iron Age tarand-graves] We found that the Iron Age individuals do in fact carry chrY hg N3 (…) Furthermore, based on their autosomal data, all of the studied individuals appear closer to hunter-gatherers and modern Estonians than Estonian CWC individuals do.

EDIT (16 OCT) A recent abstract with Saag as main author (Tambets second) cites 3 out of 5 sampled Iron Age individuals as having haplogroup N3.

EDIT (28 OCT): Notice also the appearance of N1a1a1a1a1a1a1-L1025 in Lithuania (ca. 300 AD), from Damgaard (Nature 2018); the N1c sample of the Krivichi Pskov Long Barrows culture (ca. 8th-10th c. AD), and N1a1a1a1a1a1a7-Y4341 among late Vikings from Sigtuna (ca. 10th-12th c. AD) in Krzewinska (2018).

estonian-pca
PCA of Estonian samples from the Bronze Age, Iron Age and Medieval times. Tambets et al. (2018, upcoming).

Looking at the plot, the genetic inflow marking the change from the Bronze Age to the Iron Age looks like an obvious expansion of nearby peoples with CWC-related ancestry, i.e. likely from the south-east, near the Middle Volga, where influence of steppe peoples is greater (hence likely Akozino) into a Proto-Fennic population already admixed (since the arrival of Corded Ware groups) with Comb Ware-like populations.

All of these groups were probably R1a-Z645 (likely R1a-Z283) since the expansion of Corded Ware peoples, with an introduction of some N1c lineages precisely during this Iron Age period. This infiltration of N1c-L392 with Akozino is obviously not directly related to Siberian cultures, given what we know about the autosomal description of Estonian samples.

Rather, N1c-L392 lineages were likely part of the incoming (Volgaic) Akozino warrior-traders, who settled among developing chiefdoms based on hill fort settlements of cultures all over the Baltic area, and began to appear thus in some of the new tarand graves associated with the Iron Age in north-eastern Europe.f

A good way to look at this is to realize that no new cluster appears compared to the data we already have from Baltic LN and BA samples from Mittnik et al. (2018), so the Estonian BA and IA clusters must be located (in a proper PCA) in the cline from Pit-Comb Ware culture through Baltic BA to Corded Ware groups:

baltic-samples
PCA and ADMIXTURE analysis reflecting three time periods in Northern European prehistory. a Principal components analysis of 1012 present-day West Eurasians (grey points, modern Baltic populations in dark grey) with 294 projected published ancient and 38 ancient North European samples introduced in this study (marked with a red outline). Population labels of modern West Eurasians are given in Supplementary Fig. 7 and a zoomed-in version of the European Late Neolithic and Bronze Age samples is provided in Supplementary Fig. 8. b Ancestral components in ancient individuals estimated by ADMIXTURE (k = 11)

This genetic continuity from Corded Ware (the most likely Proto-Uralic homeland) to the Proto-Fennic and Proto-Saamic communities in the Gulf of Finland correlates very well with the known conservatism of Finno-Saamic phonology, quite similar to Finno-Ugric, and both to Proto-Uralic (Kallio 2017): The most isolated region after the expansion of Corded Ware peoples, the Gulf of Finland, shielded against migrations for almost 1,500 years, is then the most conservative – until the arrival of Akozino influence.

NOTE. This has its parallel in the phonetic conservatism of Celtic or Italic compared to Finno-Ugric-influenced Germanic, Balto-Slavic, or Indo-Iranian.

Only later would certain regions (like Finland or Lappland) suffer Y-DNA bottlenecks and further admixture events associated with population displacements and expansions, such as the spread of Fennic peoples from their Estonian homeland (evidenced by the earlier separation of South Estonian) to the north and east:

diversification-finnic
The Finnic family tree. Kallio (2014).

The initial Proto-Fennic expansion was probably coupled with the expansion of Proto-Saami to the north, with the Kjelmøy Ware absorbing the Siberian population of Lovozero Ware, and potentially in inner Finland and Karelia with the Luukonsaari and Sirnihta wares (Carpelan and Parpola 2017).

This Proto-Saami population expansion from the mainland to the north, admixing with Lovozero-related peoples, is clearly reflected in the late Iron Age Saamic samples from Levänluhta (ca. 400-800 AD), as a shift (of 2 out of 3 samples) to Siberian-like ancestry from their original CWC_Baltic-like situation (see PCA from Lamnidis et al. 2018 above).

Also, Volgaic and Permic populations from inner Finland and the Forest Zone to the Cis-Urals and Circum-Artic regions probably incorporate Siberian ancestry and N1c-L392 lineages during these and later population movements, while the westernmost populations – Estonian, Mordvinic – remain less admixed (see PCA from Tambets et al. 2018 below).

We also have data of N1c-L392 in Nordic territory in the Middle Ages, proving its likely strong presence in the Mälaren area since the Iron Age, with the arrival of Akozino warrior traders. Similarly, it is found among Balto-Slavic groups along the eastern Baltic area. Obviously, no language change is seen in Nordic Bronze Age and Lusatian territory, and none is expected in Estonian or Finnish territory, either.

Therefore, no “N1c-L392 + Siberian ancestry” can be seen expanding Finno-Ugric dialects, but rather different infiltrations and population movements with limited effects on ancestry and Y-DNA composition, depending on the specific period and region.

estonians-hungarians-mordvinian
Selection of the PCA, with the group of Estonians, Mordovians, and Hungarians selected. See Tambets et al. (2018) for more information.

An issue never resolved

Because N1c-L392 subclades & Siberian ancestry, which appear in different proportions and with different origins among some modern Uralic peoples, do not appear in cultures supposed to host Uralic-speaking populations until the Iron Age, people keep looking into any direction to find the ‘true’ homeland of those ‘Uralic N1c peoples’? Kind of a full circular reasoning, anyone? The same is valid for R1a & steppe ancestry being followed for ‘Indo-Europeans’, or R1b-P312 & Neolithic farmer ancestry being traced for ‘Basques’, because of their distribution in modern populations.

I understand the caution of many pointing to the need to wait and see how samples after 2000 BC are like, in every single period, from the middle and upper Volga, Kama, southern Finland, and the Forest Zone between Fennoscandia and the steppe. It’s like waiting to see how people from Western Yamna and the Carpathian Basin after 3000 BC look like, to fill in what is lacking between East Yamna and Bell Beakers, and then between them and every single Late PIE dialect.

But the answer for Yamna-Bell Beaker-Poltavka peoples during the Late PIE expansion is always going to be “R1b-L23, but with R1a-Z645 nearby” (we already have a pretty good idea about that); and the answer for the Forest Zone and northern Cis- and Trans-Urals area – during the time when Uralic languages are known to have already been spoken there – is always going to be “R1a-Z645, but with haplogroup N nearby”, as is already clear from the data on the eastern Baltic region.

So, without a previously proposed model as to where those amateurs expressing concern about ‘not having enough data’ expect to find those ‘Uralic peoples’, all this waiting for the right data looks more like a waiting for N1c and Siberian ancestry to pop up somewhere in the historic Uralic-speaking area, to be able to say “There! A Uralic-speaking male!”. Not a very reasonable framework to deal with prehistoric peoples and their languages, I should think.

But, for those who want to do that, let me break the news to you already:

ananino-culture-balto-slavic
First N1c – Finno-Ugric person arrives in Estonia to teach Finno-Saamic to Balto-Slavic peoples.

And here it is, an appropriate fantasy description of the ethnolinguistic groups from the region. You are welcome:

  • During the Bronze Age, late Corded Ware groups evolve as the western Textile ware Fennic Balto-Slavic group in the Gulf of Finland; the Netted Ware Saamic Balto-Slavic group of inner Finland; the south Netted Ware / Akozino Volgaic Balto-Slavic groups of the Middle Volga; and the Anonino Permic Balto-Slavic group in the north-eastern Forest Zone; all developing still in close contact with each other, allowing for common traits to permeate dialects.
  • These Balto-Slavic groups would then incorporate west of the Urals during and after the Iron Age (ca. 800-500 BC first, and also later during their expansion to the north) limited ancestry and lineages from eastern European hunter-gatherer groups of Palaeo-European Fennic and Palaeo-Siberian Volgaic and Permic languages from the Circum-Artic region, but they adopted nevertheless the language of the newcomers in every single infiltration of N1c lineages and/or admixture with Siberian ancestry. Oh and don’t forget the Saamic peoples from central Sweden, of course, the famous N1c-L392 ‘Rurikid’ lineages expanding Saamic to the north and replacing Proto-Germanic…

The current model for those obsessed with modern Y-DNA is, therefore, that expanding Neolithic, Bronze Age and Iron Age cultures from north-eastern Europe adopted the languages of certain lineages originally from sub-Neolithic (Scandinavian and Siberian) hunter-gatherer populations of the Circum-Artic region; lineages that these cultures incorporated unevenly during their expansions. Hmmmm… Sounds like an inverse Western movie, where expanding Americans end up speaking Apache, and the eastern coast speaks Spanish until Italian migrants arrive and make everyone speak English… or something. A logic, no-nonsense approach to ethnolinguistic identification.

I kid you not, this is the kind of models we are going to see very soon. In 2018 and 2019, with ancient DNA able to confirm or reject archaeological hypotheses based on linguistic data, people will keep instead creating new pet theories to support preconceived ideas based on the Y-DNA prevalent among modern populations. That is, information available in the 2000s.

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

[Next post on the subject: Corded Ware—Uralic (III): Seima-Turbino and the Ugric and Samoyedic expansion]

See also

Related

Viking Age town shows higher genetic diversity than Neolithic and Bronze Age

sigtuna-vikings

Open access Genomic and Strontium Isotope Variation Reveal Immigration Patterns in a Viking Age Town, by Krzewińska et al., Current Biology (2018).

Interesting excerpts (emphasis mine, some references deleted for clarity):

The town of Sigtuna in eastern central Sweden was one of the pioneer urban hubs in the vast and complex communicative network of the Viking world. The town that is thought to have been royally founded was planned and organized as a formal administrative center and was an important focal point for the establishment of Christianity [19]. The material culture in Sigtuna indicates that the town had intense international contacts and hosted several cemeteries with a Christian character. Some of them may have been used by kin-based groups or by people sharing the same sociocultural background. In order to explore the character and magnitude of mobility and migration in a late Viking Age town, we generated and analyzed genomic (n = 23) and strontium isotope (n = 31) data from individuals excavated in Sigtuna.

y-dna-vikings

The mitochondrial genomes were sequenced at 1.5× to 367× coverage. Most of the individuals were assigned to haplogroups commonly found in current-day Europeans, such as H, J, and U [14, 26, 27]. All of these haplotypes are present in Scandinavia today.

The Y chromosome haplogroups were assigned in seven males. The Y haplogroups include I1a, I2a, N1a, G2a, and R1b. Two identified lineages (I2a and N1a) have not been found in modern-day Sweden or Norway [28, 29]. Haplogroups I and N are associated with eastern and central Europe, as well as Finno-Ugric groups [30]. Interestingly, I2a was previously identified in a middle Neolithic Swedish hunter-gatherer dating to ca. 3,000 years BCE [31].

In Sigtuna, the genetic diversity in the late Viking Age was greater than the genetic diversity in late Neolithic and Bronze Age cultures (Unetice and Yamnaya as examples) and modern East Asians; it was on par with Roman soldiers in England but lower than in modern-day European groups (GBR and FIN; Figure 2B). Within the town, the group excavated at church 1 has somewhat greater diversity than that at cemetery 1. Interestingly, the diversity at church 1 is nearly as high as that observed in Roman soldiers in England, which is remarkable, since the latter was considered to be an exceptionally heterogeneous group in contemporary Europe [39].

pca-vikings
A PCA plot visualising all 23 individuals from Sigtuna used in ancient DNA analyses (m – males, f – females).

Different sex-related mobility patterns for Sigtuna inhabitants have been suggested based on material culture, especially ceramics. Building on design and clay analyses, some female potters in Sigtuna are thought to have grown up in Novgorod in Rus’ [40]. Moreover, historical sources mention female mobility in connection to marriage, especially among the elite from Rus’ and West Slavonic regions [41, 42]. Male mobility is also known from historical sources, often in connection to clergymen moving to the town [43].

Interestingly, we found a number of individuals from Sigtuna to be genetically similar to the modern-day human variation of eastern Europeans, and most harbor close genetic affinities to Lithuanians (Figure 2A). The strontium isotope ratios in 28 adult individuals with assigned biological sex and strontium values obtained from teeth (23 M1 and five M2) show that 70% of the females and 44% of the males from Sigtuna were non-locals (STAR Methods). The difference in migrant ratios between females and male mobility patterns was not statistically significant (Fisher’s exact test, p = 0.254 for 28 individuals and p = 0.376 for 16 individuals). Hence, no evidence of a sex-specific mobility pattern was found.

(…) As these social groups are not mirrored by our genetic or strontium data, this suggests that the inclusion in them was not based on kinship. Therefore, it appears as if socio-cultural factors, not biological bonds, governed where people were interred (i.e., the choice of cemetery).

diversity-yamna
Average pairwise genetic diversity measured in complete Sigtuna, St. Gertrud (church 1) and cemetery 1 (the Nunnan block) compared to both ancient and modern populations ranked by time period (Yamnaya, Unetice, and GBR-Roman, Roman Age individuals from Great Britain; GBR-AS, Anglo-Saxon individuals from Great Britain; GBR-IA, Iron Age individuals from Great Britain; JPT-Modern, presentday Japanese from Tokyo; FIN-Modern, present-day Finnish; GBR-Modern, present-day British; GIHModern, present-day Gujarati Indian from Houston, Texas). Error bars show ±2 SEs.

Interesting from this paper is the higher genetic (especially Y-DNA) diversity found in more recent periods (see e.g. here) compared to Neolithic and Bronze Age cultures, which is probably the reason behind some obviously wrong interpretations, e.g. regarding links between Yamna and Corded Ware populations.

The sample 84001, a “first-generation short-distance migrant” of haplogroup N1c-L392 (N1a in the new nomenclature) brings yet more proof of how:

  • Admixture changes completely within a certain number of generations. In this case, the N1c-L392 sample clusters within the genetic variation of modern Norwegians, near to the Skane Iron Age sample, and not with its eastern origin (likely many generations before).
  • This haplogroup appeared quite late in Fennoscandia but still managed to integrate and expand into different ethnolinguistic groups; in this case, this individual was probably a Viking of Nordic language, given its genetic admixture and its non-local (but neighbouring Scandinavian) strontium values.

Related

Tracking material cultures with ancient DNA: medieval Norse walrus ivory trade, and leather shields from Zanzibar

norse-walrus-ivory-trade

Two papers have been recently published, offering another interesting use of ancient DNA analysis for Archaeology and, potentially, Linguistics.

Open access Ancient DNA reveals the chronology of walrus ivory trade from Norse Greenland, by Star, Barrett, Gondek, & Boessenkool, bioRxiv (2018).

Abstract (emphasis mine):

The search for walruses as a source of ivory -a popular material for making luxury art objects in medieval Europe- played a key role in the historic Scandinavian expansion throughout the Arctic region. Most notably, the colonization, peak and collapse of the medieval Norse colony of Greenland have all been attributed to the proto-globalization of ivory trade. Nevertheless, no studies have directly traced European ivory back to distinct populations of walrus in the Arctic. This limits our understanding of how ivory trade impacted the sustainability of northern societies and the ecology of the species they relied on. Here, we compare the mitogenomes of 27 archaeological walrus specimens from Europe and Greenland (most dated between 900 and 1400 CE) and 10 specimens from Svalbard (dated to the 18th and 19th centuries CE) to partial mitochondrial (MT) data of over 300 modern walruses. We discover two monophyletic mitochondrial clades, one of which is exclusively found in walrus populations of western Greenland and the Canadian Arctic. Investigating the chronology of these clades in our European archaeological remains, we identify a significant shift in resource use from predominantly eastern sources towards a near exclusive representation of walruses from western Greenland. These results provide empirical evidence for the economic importance of walrus for the Norse Greenland settlements and the integration of this remote, western Arctic resource into a medieval pan-European trade network.

walrus-ivory-trade-norse-greenland
(A) Population distribution, historic trade routes and sample locations of Atlantic walrus in the northern Atlantic region. The range of modern Atlantic walrus (dark grey) and putative dispersal routes (black arrows) follow (58) and (31). Eight breeding populations are recognized (58); 1 – Foxe Basin, 2 – Hudson Bay, 3 – Hudson Strait, 4, – West Greenland, 5 – North Water, 6 – East Greenland, 7 – Svalbard/Franz Josef land, 8 – Novaya Zemlya. Historic trade routes from Greenland –including the location of Norse settlements– and northern Fennoscandia/Russia (yellow) indicate possible sources from which walrus ivory was exported to Europe during the Middle Ages. The Svalbard specimens (orange) were originally from hunting stations of the 1700s and 1800s. The other Atlantic walrus specimens (red, grey) were obtained from museum collections. (B) Bayesian phylogenetic tree obtained using BEAST (84) based on 346 mitochondrial SNPs using Pacific walrus (PAC) as an outgroup. Numbers represent the different specimens as listed in Table S1, and colors match the sampling locations as in Fig. 1A. Branches with a posterior probability of one (grey circles) are indicated. (C) Distribution of RFLP and control region (CR) haplotypes of modern Atlantic walrus populations. The RFLP clade classification follows Born, Andersen et al. (2001). The distribution of a distinct ACC CR haplotype is from 306 modern specimens (see material and methods).

Determination of the geographical origin of leather shields from Zanzibar using ancient DNA tools, by Bastian, Jacot-des-Combes, Hänni, & Perrier, J Arch. Sci (2018) 19:323-333.

Abstract:

Zanzibar shields are documented in several books and preserved in many European, African and Omani museums. They are relatively small and decorated; therefore, we can assume that they served to not only to protect the hand during sword combat but also to attract the attention of the attacker. As with all shields, they are also an object of prestige and armorial bearing to identify the owner’s army corps. Within the incredible cultural and ethnic mosaic of this part of the Indian Ocean, the shield enables alliances, protection systems and allegiance to be specified and clarified.

This study is a step towards understanding the nature of the relationships between Oman and the various communities living on the western coast of the Indian Ocean based on their material culture, especially their shields. Identifying the animal species used to make the shields was crucial in establishing both the manufacturers and the consumers of these objects. DNA analyses indicated that the leather used for the studied Zanzibar shields is rhinoceros (Diceros bicornis michaeli); a subspecies historically only present on the coast of East Africa. Our results also indicate that the shields, used mainly in Oman, Zanzibar and other regions with a strong relationship with Oman power, were made in Zanzibar and the Arabian Peninsula.

zanzibar-shield-rhinoceros
Ancient distribution of Diceros bicornis michaeli (eastern black rhinoceros) from southern Sudan, Ethiopia, and Somalia through Kenya into northern-central Tanzania. Dark grey represents the presence of both species. At the tip of the arrow: Zanzibar Island. Copyright: Fabiola Bastian.

In a time when many geneticists seem to have shifted their full attention to novel statistical methods applied to a few ancient individuals, it feels good to see some of them using their research to complement traditional academic disciplines instead.

This kind of studies may help track with more detail the most obvious harbinger of potential prehistoric language change: the diffusion of material culture.

See also:

Network analysis of the Viking Age in Ireland as portrayed in Cogadh Gaedhel re Gallaibh

Ireland-kingdoms-vikings-900

Open access Network analysis of the Viking Age in Ireland as portrayed in Cogadh Gaedhel re Gallaibh, by Yose et al., Royal Society Open Science (2018).

Abstract

Cogadh Gaedhel re Gallaibh (‘The War of the Gaedhil with the Gaill’) is a medieval Irish text, telling how an army under the leadership of Brian Boru challenged Viking invaders and their allies in Ireland, culminating with the Battle of Clontarf in 1014. Brian’s victory is widely remembered for breaking Viking power in Ireland, although much modern scholarship disputes traditional perceptions. Instead of an international conflict between Irish and Viking, interpretations based on revisionist scholarship consider it a domestic feud or civil war. Counter-revisionists challenge this view and a long-standing and lively debate continues. Here, we introduce quantitative measures to the discussions. We present statistical analyses of network data embedded in the text to position its sets of interactions on a spectrum from the domestic to the international. This delivers a picture that lies between antipodal traditional and revisionist extremes; hostilities recorded in the text are mostly between Irish and Viking—but internal conflict forms a significant proportion of the negative interactions too.

Cogadh-network-vikings
The entire Cogadh network of interacting characters. Characters identified as Irish are represented by green nodes and those identified as Vikings are in blue. Other characters are in grey. Edges between pairs of Irish nodes are also coloured green while those between Viking pairs are blue. Edges linking Irish to Viking nodes are brown and the remaining edges are grey.

See also:

Migrations painted by Irish and Scottish genetic clusters, and their relationship with British and European ones

ireland-britain-cluster

Interesting and related publications, now appearing in pairs…

1. The Irish DNA Atlas: Revealing Fine-Scale Population Structure and History within Ireland, by Gilbert et al., in Scientific Reports (2017).

Abstract:

The extent of population structure within Ireland is largely unknown, as is the impact of historical migrations. Here we illustrate fine-scale genetic structure across Ireland that follows geographic boundaries and present evidence of admixture events into Ireland. Utilising the ‘Irish DNA Atlas’, a cohort (n = 194) of Irish individuals with four generations of ancestry linked to specific regions in Ireland, in combination with 2,039 individuals from the Peoples of the British Isles dataset, we show that the Irish population can be divided in 10 distinct geographically stratified genetic clusters; seven of ‘Gaelic’ Irish ancestry, and three of shared Irish-British ancestry. In addition we observe a major genetic barrier to the north of Ireland in Ulster. Using a reference of 6,760 European individuals and two ancient Irish genomes, we demonstrate high levels of North-West French-like and West Norwegian-like ancestry within Ireland. We show that that our ‘Gaelic’ Irish clusters present homogenous levels of ancient Irish ancestries. We additionally detect admixture events that provide evidence of Norse-Viking gene flow into Ireland, and reflect the Ulster Plantations. Our work informs both on Irish history, as well as the study of Mendelian and complex disease genetics involving populations of Irish ancestry.

european-ancestry-british-isles
The European ancestry profiles of 30 Irish and British clusters. (a) The total ancestry contribution summarised by majority European country of origin to each of the 30 Irish and British clusters. (b) (left) The ancestry contributions of 19 European clusters that donate at least 2.5% ancestry to any one Irish or British cluster. (right) The geographic distribution of the 19 European clusters, shown as the proportion of individuals in each European region belonging to each of the 19 European clusters. The proportion of individuals form each European region not a member of the 19 European clusters is shown in grey. Total numbers of individuals from each region are shown in white text. Not all Europeans included in the analysis were phenotyped geographically. The figure was generated in the statistical software language R46, version 3.4.1, using various packages. The map of Europe was sourced from the R software package “mapdata” (https://CRAN.R-project.org/package=mapdata).

2. New preprint on BioRxiv, Insular Celtic population structure and genomic footprints of migration, by Byrne, Martiniano et al. (2017).

Abstract:

Previous studies of the genetic landscape of Ireland have suggested homogeneity, with population substructure undetectable using single-marker methods. Here we have harnessed the haplotype-based method fineSTRUCTURE in an Irish genome-wide SNP dataset, identifying 23 discrete genetic clusters which segregate with geographical provenance. Cluster diversity is pronounced in the west of Ireland but reduced in the east where older structure has been eroded by historical migrations. Accordingly, when populations from the neighbouring island of Britain are included, a west-east cline of Celtic-British ancestry is revealed along with a particularly striking correlation between haplotypes and geography across both islands. A strong relationship is revealed between subsets of Northern Irish and Scottish populations, where discordant genetic and geographic affinities reflect major migrations in recent centuries. Additionally, Irish genetic proximity of all Scottish samples likely reflects older strata of communication across the narrowest inter-island crossing. Using GLOBETROTTER we detected Irish admixture signals from Britain and Europe and estimated dates for events consistent with the historical migrations of the Norse-Vikings, the Anglo-Normans and the British Plantations. The influence of the former is greater than previously estimated from Y chromosome haplotypes. In all, we paint a new picture of the genetic landscape of Ireland, revealing structure which should be considered in the design of studies examining rare genetic variation and its association with traits.

Here are some interesting excerpts (emphasis mine):

Population structure in Ireland

The geographical distribution of this deep subdivision of Leinster resembles pre-Norman territorial boundaries which divided Ireland into fifths (cúige), with north Leinster a kingdom of its own known as Meath (Mide) [15]. However interpreted, the firm implication of the observed clustering is that despite its previously reported homogeneity, the modern Irish population exhibits genetic structure that is subtly but detectably affected by ancestral population structure conferred by geographical distance and, possibly, ancestral social structure.

ChromoPainter PC1 demonstrated high diversity amongst clusters from the west coast, which may be attributed to longstanding residual ancient (possibly Celtic) structure in regions largely unaffected by historical migration. Alternatively, genetic clusters may also have diverged as a consequence of differential influence from outside populations. This diversity between western genetic clusters cannot be explained in terms of geographic distance alone.

In contrast to the west of Ireland, eastern individuals exhibited relative homogeneity; (…) The overall pattern of western diversity and eastern homogeneity in Ireland may be explained by increased gene flow and migration into and across the east coast of Ireland from geographically proximal regions, the closest of which is the neighbouring island of Britain.

Analysis of variance of the British admixture component in cluster groups showed a significant difference (p < 2×10-16), indicating a role for British Anglo-Saxon admixture in distinguishing clusters, and ChromoPainter PC2 was correlated with the British component (p < 2×10-16), explaining approximately 43% of the variance. PC2 therefore captures an east to west Anglo-Celtic cline in Irish ancestry. This may explain the relative eastern homogeneity observed in Ireland, which could be a result of the greater English influence in Leinster and the Pale during the period of British rule in Ireland following the Norman invasion, or simply geographic proximity of the Irish east coast to Britain. Notably, the Ulster cluster group harboured an exceptionally large proportion of the British component (Fig 1D and 1E), undoubtedly reflecting the strong influence of the Ulster Plantations in the 17th century and its residual effect on the ethnically British population that has remained.

ireland-population-structure
Fine-grained population structure in Ireland. (A) fineSTRUCTURE clustering dendrogram for 1,035 Irish individuals. Twenty-three clusters are defined, which are combined into cluster groups for clusters that are neighbouring in the dendrogram, overlapping in principal component space (B) and sampled from regions that are geographically contiguous. Details for each cluster in the dendrogram are provided in S1 Fig. (B) Principal components analysis (PCA) of haplotypic similarity, based on ChromoPainter coancestry matrix for Irish individuals. Points are coloured according to cluster groups defined in (A); the median location of each cluster group is plotted. (C) Map of Irelandshowing the sampling location for a subset of 588 individuals analysed in (A) and (B), coloured by cluster group. Points have been randomly jittered within a radius of 5 km to preserve anonymity. Precise sampling location for 44 Northern Irish individuals from the People of the British Isles dataset was unknown; these individuals are plotted geometrically in a circle. (D) “British admixture component” (ADMIXTURE estimates; k=2) for Irish cluster groups. This component has the largest contribution in ancient Anglo-Saxons and the SEE cluster. (E) Linear regression of principal component 2 (B) versus British admixture component (r2 = 0.43; p < 2×10-16). Points are coloured by cluster group. (Standard error for ADMIXTURE point estimates presented in S11 Fig.)

On the genetic structure of the British Isles

The genetic substructure observed in Ireland is consistent with long term geographic diversification of Celtic populations and the continuity shown between modern and Early Bronze Age Irish people

Clusters representing Celtic populations harbouring less Anglo-Saxon influence separate out above and below SEE on PC4. Notably, northern Irish clusters (NLU), Scottish (NISC, SSC and NSC), Cumbria (CUM) and North Wales (NWA) all separate out at a mutually similar level, representing northern Celtic populations. The southern Celtic populations Cornwall (COR), south Wales (SWA) and south Munster (SMN) also separate out on similar levels, indicating some shared haplotypic variation between geographically proximate Celtic populations across both Islands. It is notable that after the split of the ancestrally divergent Orkney, successive ChromoPainter PCs describe diversity in British populations where “Anglo-saxonization” was repelled [22]. PC3 is dominated by Welsh variation, while PC4 in turn splits North and South Wales significantly, placing south Wales adjacent to Cornwall and north Wales at the other extreme with Cumbria, all enclaves where Brittonic languages persisted.

In an interesting symmetry, many Northern Irish samples clustered strongly with southern Scottish and northern English samples, defining the Northern Irish/Cumbrian/Scottish (NICS) cluster group. More generally, by modelling Irish genomes as a linear mixture of haplotypes from British clusters, we found that Scottish and northern English samples donated more haplotypes to clusters in the north of Ireland than to the south, reflecting an overall correlation between Scottish/north English contribution and ChromoPainter PC1 position in Fig 1 (Linear regression: p < 2×10-16, r2 = 0.24).

North to south variation in Ireland and Britain are therefore not independent, reflecting major gene flow between the north of Ireland and Scotland (Fig 5) which resonates with three layers of historical contacts. First, the presence of individuals with strong Irish affinity among the third generation PoBI Scottish sample can be plausibly attributed to major economic migration from Ireland in the 19th and 20th centuries [6]. Second, the large proportion of Northern Irish who retain genomes indistinguishable from those sampled in Scotland accords with the major settlements (including the Ulster Plantation) of mainly Scottish farmers following the 16th Century Elizabethan conquest of Ireland which led to these forming the majority of the Ulster population. Third, the suspected Irish colonisation of Scotland through the Dál Riata maritime kingdom, which expanded across Ulster and the west coast of Scotland in the 6th and 7th centuries, linked to the introduction and spread of Gaelic languages [3]. Such a migratory event could work to homogenise older layers of Scottish population structure, in a similar manner as noted on the east coasts of Britain and Ireland. Earlier communications and movements across the Irish Sea are also likely, which at its narrowest point separates Ireland from Scotland by approximately 20 km.

ireland-britain-genetic-geography
Genes mirror geography in the British Isles. (A) fineSTRUCTURE clustering dendrogram for combined Irish and British data. Data principally split into Irish and British groups before subdividing into a total of 50 distinct clusters, which are combined into cluster groups for clusters that formed clades in the dendrogram, overlapped in principal component space (B) and were sampled from regions that are geographically contiguous. Names and labels follow the geographical provenance for the majority of data within the cluster group. Details for each cluster in the dendrogram are provided in S2 Fig. (B) Principal component analysis (PCA) of haplotypic similarity based on the ChromoPainter coancestry matrix, coloured by cluster group with their median locations labelled. We have chosen to present PC1 versus PC4 here as these components capture new information regarding correlation between haplotypic variation across Britain and Ireland and geography, while PC2 and PC3 (Fig 4) capture previously reported splitting for Orkney and Wales from Britain [7]. A map of Ireland and Britain is shown for comparison, coloured by sampling regions for cluster groups, the boundaries of which are defined by the Nomenclature of Territorial Units for Statistics (NUTS 2010), with some regions combined. Sampling regions are coloured by the cluster group with the majority presence in the sampling region; some sampling regions have significant minority cluster group representations as well, for example the Northern Ireland sampling region (UKN0; NUTS 2010) is majorly explained by the NICS cluster group but also has significant representation from the NLU cluster group. The PCA plot has been rotated clockwise by 5 degrees to highlight its similarity with the geographical map of the Ireland and Britain. NI, Northern Ireland; PC, principal component. Cluster groups that share names with groups from Fig 1 (NLU; SMN; CLN; CNN) have an average of 80% of their samples shared with the initial cluster groups. © EuroGeographics for the map and administrative boundaries, note some boundaries have been subsumed or modified to better reflect sampling regions.

Genomic footprints of migration into Ireland

Quite interesting is that it is haplogroups, and not admixture, that which defines the oldest migration layers into Ireland. Without evidence of paternal Y-DNA lineages we would probably not be able to ascertain the oldest migrations and languages broght by migrants, including Celtic languages:

Of all the European populations considered, ancestral influence in Irish genomes was best represented by modern Scandinavians and northern Europeans, with a significant single-date one-source admixture event overlapping the historical period of the Norse-Viking settlements in Ireland (p < 0.01; fit quality FQB > 0.985; Fig 6). (…) This suggests a contribution of historical Viking settlement to the contemporary Irish genome and contrasts with previous estimates of Viking ancestry in Ireland based on Y chromosome haplotypes, which have been very low [25]. The modern-day paucity of Norse-Viking Y chromosome haplotypes may be a consequence of drift with the small patrilineal effective population size, or could have social origins with Norse males having less influence after their military defeat and demise as an identifiable community in the 11th century, with persistence of the autosomal signal through recombination.

European admixture date estimates in northwest Ulster did not overlap the Viking age but did include the Norman period and the Plantations

The genetic legacies of the populations of Ireland and Britain are therefore extensively intertwined and, unlike admixture from northern Europe, too complex to model with GLOBETROTTER.

ireland-admixture-estimates
All-Ireland GLOBETROTTER admixture date estimates for European and British surrogate admixing populations. A summary of the date estimates and 95% confidence intervals for inferred admixture events into Ireland from European and British admixing sources is shown in (A), with ancestry proportion estimates for each historical source population for the two events and example coancestry curves shown in (B). In the coancestry curves Relative joint probability estimates the pairwise probability that two haplotype chunks separated by a given genetic distance come from the two modeled source populations respectively (ie FRA(8) and NOR-SG); if a single admixture event occurred, these curves are expected to decay exponentially at a rate corresponding to the number of generations since the event. The green fitted line describes this GLOBETROTTER fitted exponential decay for the coancestry curve. If the sources come from the same ancestral group the slope of this curve will be negative (as with FRA(8) vs FRA(8)), while a positive slope indicates that sources come from different admixing groups (as with FRA(8) vs NOR-SG). The adjacent bar plot shows the inferred genetic composition of the historical admixing sources modelled as a mixture of the sampled modern populations. A European admixture event was estimated by GLOBETROTTER corresponding to the historical record of the Viking age, with major contributions from sources similar to modern Scandinavians and northern Europeans and minor contributions from southern European-like sources. For admixture date estimates from British-like sources the influence of the Norman settlement and the Plantations could not be disentangled, with the point estimate date for admixture falling between these two eras and GLOBETROTTER unable to adequately resolve source and proportion details of admixture event (fit quality FQB< 0.985). The relative noise of the coancestry curves reflects the uncertainty of the British event. Cluster labels (for the European clustering dendrogram, see S4 Fig; for the PoBI clustering dendrogram, see S3 Fig): FRA(8), France cluster 8; NOR-SG, Norway, with significant minor representations from Sweden and Germany; SE_ENG, southeast England; N_SCOT(4) northern Scotland cluster 4.

Another study that strengthens the need to ascertain haplogroup-admixture differences between Yamna/Bell Beaker and Sredni Stog/Corded Ware.

Text and images from preprint article under a CC-BY-NC-ND 4.0 International license.

Featured image, from the article on Science Reports: The clustering of individuals with Irish and British ancestry based solely on genetics. Shown are 30 clusters identified by fineStructure from 2,103 Irish and British individuals. The dendrogram (left) shows the tree of clusters inferred by fineStructure and the map (right) shows the geographic origin of 192 Atlas Irish individuals and 1,611 British individuals from the Peoples of the British Isles (PoBI) cohort, labelled according to fineStructure cluster membership. Individuals are placed at the average latitude and longitude of either their great-grandparental (Atlas) or grandparental (PoBI) birthplaces. Great Britain is separated into England, Scotland, and Wales. The island of Ireland is split into the four Provinces; Ulster, Connacht, Leinster, and Munster. The outline of Britain was sourced from Global Administrative Areas (2012). GADM database of Global Administrative Areas, version 2.0. www.gadm.org. The outline of Ireland was sourced from Open Street Map Ireland, Copyright OpenStreetMap Contributors, (https://www.openstreetmap.ie/) – data available under the Open Database Licence. The figure was plotted in the statistical software language R46, version 3.4.1, with various packages.
Related:

My European Family: The First 54,000 years, by Karin Bojs

steppe-expansion-corded-ware

I have recently read the book My European Family: The First 54,000 years (2015), by Karin Bojs, a known Swedish scientific journalist, former science editor of the Dagens Nyheter.

my-european-family
My European Family: The First 54,000 Years
It is written in a fresh, dynamic style, and contains general introductory knowledge to Genetics, Archaeology, and their relation to language, and is written in a time of great change (2015) for the disciplines involved.

The book is informed, it shows a balanced exercise between responsible science journalism and entertaining content, and it is at times nuanced, going beyond the limits of popular science books. It is not written for scholars, although you might learn – as I did – interesting details about researchers and institutions of the anthropological disciplines involved. It contains, for example, interviews with known academics, which she uses to share details about their personalities and careers, which give – in my opinion – a much needed context to some of their publications.

Since I am clearly biased against some of the findings and research papers which are nevertheless considered mainstream in the field (like the identification of haplogroup R1a with the Proto-Indo-European expansion, or the concept of steppe admixture), I asked my wife (who knew almost nothing about genetics, or Indo-European studies) to read it and write a summary, if she liked it. She did. So much, that I have convinced her to read The Horse, the Wheel, and Language: How Bronze-Age Riders from the Eurasian Steppes Shaped the Modern World (2007), by David Anthony.

Here is her summary of the book, translated from Spanish:

The book is divided in three main parts: The Hunters, The Farmers, and The Indo-Europeans, and each has in turn chapters which introduce and break down information in an entertaining way, mixing them with recounts of her interactions and personal genealogical quest.

Part one, The Hunters, offers intriguing accounts about the direct role music had in the development of the first civilizations, the first mtDNA analyses of dogs (Savolainen), and the discovery of the author’s Saami roots. Explanations about the first DNA studies and their value for archaeological studies are clear and comprehensible for any non-specialized reader. Interviews help give a close view of investigations, like that of Frederic Plassard’s in Les Combarelles cave.

Part two, The Farmers, begins with her travel to Cyprus, and arouses the interest of the reader with her description of the circular houses, her notes on the Basque language, the new papers and theories related to DNA analyses, the theory of the decision of cats to live with humans, the first beers, and the houses built over graves. Karin Bojs analyses the subgroup H1g1 of her grandmother Hilda, and how it belonged to the first migratory wave into Central Europe. This interest in her grandmother’s origins lead her to a conference in Pilsen about the first farmers in Europe, where she knows firsthand of the results of studies by János Jakucs, and studies of nuclear DNA. Later on she interviews Guido Brandt and Joachim Burguer, with whom she talks about haplogroups U, H, and J.

The chapter on Ötzi and the South Tyrol Museum of Archaeology (Bolzano) introduces the reader to the first prehistoric individual whose DNA was analysed, belonging to haplogroup G2a4, but also revealing other information on the Iceman, such as his lactose intolerance.

Part three, dealing with the origin of Indo-Europeans, begins with the difficulties that researchers have in locating the origin of horse domestication (which probably happened in western Kazakhstan, in the Russian steppe between the rivers Volga and Don). She mentions studies by David Anthony and on the Yamna culture, and its likely role in the diffusion of Proto-Indo-European. In an interview with Mallory in Belfast, she recalls the potential interest of far-right extremists in genetic studies (and early links of the Journal of Indo-European Studies to certain ideology), as well as controversial statements of Gimbutas, and her potentially biased vision as a refugee from communist Europe. During the interview, Mallory had a copy of the latest genetic paper sent to Nature Magazine by Haak et al., not yet published, for review, but he didn’t share it.

Then haplogroups R1a and R1b are introduced as the most common in Europe. She visits the Halle State Museum of Prehistory (where the Nebra sky disk is exhibited), and later Krakow, where she interviews Slawomir Kadrow, dealing with the potential creation of the Corded Ware culture from a mix of Funnelbeaker and Globular Amphorae cultures. New studies of ancient DNA samples, published in the meantime, are showing that admixture analyses between Yamna and Corded Ware correlate in about 75%.

In the following chapters there is a broad review of all studies published to date, as well as individuals studied in different parts of Europe, stressing the importance of ships for the expansion of R1b lineages (Hjortspring boat).

The concluding chapter is dedicated to vikings, and is used to demystify them as aggressive warmongers, sketching their relevance as founders of the Russian state.

To sum up, it is a highly documented book, written in a clear style, and is capable of awakening the reader’s interest in genetic and anthropological research. The author enthusiastically looks for new publications and information from researchers, but is at the same time critic with them, showing often her own personal reactions to new discoveries, all of which offers a complex personal dynamic often shared by the reader, engaged with her first-person account the full length of the book.

Mayte Batalla (July 2017)

DISCLAIMER: The author sent me a copy of the book (a translation into Spanish), so there is a potential conflict of interest in this review. She didn’t ask for a review, though, and it was my wife who did it.