The expansion of Indo-Europeans in Y-chromosome haplogroups

Featuredyamnaya-corded-ware-y-dna-haplogroups

I have compiled for two years now the reported Y-DNA and mtDNA haplogroups of ancient DNA samples published, including also SNPs from analysis of the BAM files by hobbyists.

Y-DNA timeline

Here is a video with a timeline of the evolution of Indo-European speakers, according to what is known today about reconstructed languages, prehistoric cultures and ancient DNA:

yamnaya-expansion

NOTE. The video is best viewed in HD 1080p (1920×1080) with a display that allows for this or greater video quality, and a screen big enough to see haplogroup symbols, i.e. tablet or greater. The YouTube link is here. The Facebook link is here.

Based on the results of the past 5 years or so, which have been confirming this combined picture every single time, I doubt there will be much need to change it in any radical way, as only minor details remain to be clarified.

GIS maps

I wanted to publish a GIS tool of my own for everyone to have an updated reference of all data I use for my books.

The most complex GIS tools consume too many resources when used online in a client-server model, so I have to keep that to myself, but there are some ways to publish low quality outputs.

The files below include the possibility to zoom some levels to be able to see more samples, and also to check each one for more information on their ID, attributed culture and label, archaeological site, source paper, subclade (and people responsible for SNP inferences if any), etc.

Some usage notes:

  • Files are large (ca. 20 Mb), so they still take some time to load.
  • For the meaning of symbols and colors (for Y-DNA haplogroups), if there is any doubt, check the video above.
  • Pop-ups with sample information will work on desktop browsers by clicking on them, apparently not on smartphone and related tactile OS. I have changed the settings to show pop-ups on hover, so that it now works (to some extent) on tactile OS.
  • The search tool can look for specific samples according to their official ID, and works by highlighting the symbol of the selected individual (turning it into a bright blue dot), and leading the layer view to the location, but it seems to work best only with some browser and OS settings – in other browsers, you need to zoom out to see where the dot is located. The specific sample with its information could paradoxically disappear in search mode, so you might need to reload and look again for the same site that was highlighted.
  • Latitude and longitude values have been randomly modified to avoid samples overcrowding specific sites, so they are not the original ones.

Y-DNA GIS tool

There are three versions:

  1. Labels with more specific subclades (including negative SNPs), using YTree for R1b samples (whenever it conflicts with YFull).
  2. Labels with YFull nomenclature.
  3. Simbols without labels (more symbols visible per layer).

y-dna-haplogroups

mtDNA GIS tool

There are two versions:

  1. Symbols with labels.
  2. Symbols without labels.

NOTE. Because there are too many samples at the starting view, depending on the file you should zoom some levels to start seeing symbols.

mtdna-haplogroups

Static Maps

The following maps offer a timeline of Y-DNA and mtDNA evolution, divided into static periods corresponding to the Prehistoric Atlas.

Y-DNA + culture maps

The following files use the YTree or otherwise more comprehensive nomenclature than YFull. Symbols have a similar value as those from the GIS tools.

  1. Anatomically Modern Humans (PDF)
  2. Upper Palaeolithic (PDF)
  3. Epipalaeolithic (PDF)
  4. Early Mesolithic (PDF)
  5. Late Mesolithic (PDF)
  6. Neolithic and hunter-gatherer pottery (PDF)
  7. Early Eneolithic (PDF)
  8. Late Eneolithic (PDF)
  9. Early Chalcolithic (PDF)
  10. Late Chalcolithic (PDF)
  11. Early Bronze Age (PDF)
  12. Middle Bronze Age (PDF)
  13. Late Bronze Age (PDF)
  14. Early Iron Age (PDF)
  15. Late Iron Age (PDF)
  16. Antiquity (PDF)
  17. Middle Ages (PDF)

mtDNA + culture maps

Colours have been assigned randomly to each macro-haplogroup.

  1. Anatomically Modern Humans (PDF)
  2. Upper Palaeolithic (PDF)
  3. Epipalaeolithic (PDF)
  4. Early Mesolithic (PDF)
  5. Late Mesolithic (PDF)
  6. Neolithic and hunter-gatherer pottery (PDF)
  7. Early Eneolithic (PDF)
  8. Late Eneolithic (PDF)
  9. Early Chalcolithic (PDF)
  10. Late Chalcolithic (PDF)
  11. Early Bronze Age (PDF)
  12. Middle Bronze Age (PDF)
  13. Late Bronze Age (PDF)
  14. Early Iron Age (PDF)
  15. Late Iron Age (PDF)
  16. Antiquity (PDF)
  17. Middle Ages (PDF)

See also

R1b-L23-rich Bell Beaker-derived Italic peoples from the West vs. Etruscans from the East

final-bronze-age-italy

New paper (behind paywall) Ancient Rome: A genetic crossroads of Europe and the Mediterranean, by Antonio et al. Science (2019).

The paper offers a lot of interesting data concerning the Roman Empire and more recent periods, but I will focus on Italic and Etruscan origins.

NOTE. I have updated prehistoric maps with Y-DNA and mtDNA data, and also the PCA of ancient Eurasian samples by period including the recently published samples, now with added sample names to find them easily by searching the PDFs.

Apennine homeland problem

The traditional question of Italic vs. Etruscan origins from a cultural-historical view* lies in the opposition of the traditional way of life during the Bronze Age as opposed to increasingly foreign influences in the Final Bronze Age, which eventually brought about a proto-urban period in the Apennine Peninsula.

* From a modern archaeological perspective, as well as from the (unrelated) nativist view, “continuity” of ancient cultures, languages, and peoples is generally assumed, so this question is a no-brainer. Seeing how population genomics has essentially supported the cultural-historical view, dismissing the concepts of unscathed genomic or linguistic continuity, we have to assume that different cultures potentially represent different languages, and that genetic shift coupled with radical cultural changes show a strong support for linguistic change, although the later Imperial Roman period is an example of how this is not necessarily the case.

bronze-age-polada-proto-apennine
Early Bronze Age cultures ca. 2200 – 1750 BC. See full maps.

A little background to the Italic vs. Etruscan homeland problem, from Forsythe (2006) (emphasis mine):

While the material culture of the Po Valley developed in response to influences from central Europe and the Aegean, peninsular Italy during the late Bronze Age lagged somewhat behind for the most part. Inhumation continued to be the funerary practice of this region. Although agriculture doubtless remained the mainstay of human subsistence, other evidence (the occupation of mountainous sites not conducive to farming, the remains of cattle, sheep, pigs, and goats, and ceramic vessels used for boiling milk and making cheese) indicates that pastoralism was also very widespread. This suggests that transhumance was already a well-established pattern of human existence. In fact, since the material culture of central and southern Italy was relatively uniform at this time, it has been conjectured that this so-called Apennine Culture of c. 1600–1100 B.C. owed its uniformity in part to the migratory pattern characteristic of ancient Italian stockbreeding.

During the first quarter of the twelfth century B.C. the Bronze-Age civilizations of the eastern Mediterranean came to an abrupt end. The royal palaces of Pylos, Tiryns, and Mycenae in mainland Greece were destroyed by violence, and the Hittite kingdom that had ruled over Asia Minor was likewise swept away. The causes and reasons for this major catastrophe have long been debated without much scholarly consensus (see Drews 1993, 33–96). Apart from the archaeological evidence indicating the violent destruction of many sites, the only ancient accounts relating to this phenomenon come from Egypt. The most important one is a text inscribed on the temple of Medinet Habu at Thebes, which accompanies carved scenes portraying the pharaoh’s military victory over a coalition of peoples who had attempted to enter the Nile Delta by land and sea.

sea-peoples-egypt-rameses-iii

Iron metallurgy did not reach Italy until the ninth century B.C., and even then it was two or more centuries before iron displaced bronze as the most commonly used metal. Thus, archaeologists date the beginning of the Iron Age in Italy to c. 900 B.C.; and although the Italian Bronze Age is generally assigned to the period c. 1800–1100 B.C. and is subdivided into early, middle, and late phases, the 200-year interval between the late Bronze Age and early Iron Age has been labeled the Final Bronze Age.

During this period the practice of cremation spread south of the Po Valley and is attested at numerous sites throughout the peninsula. Since this cultural tradition developed into the Villanovan Culture which prevailed in Etruria and much of the Po Valley c. 900–700 B.C., modern archaeologists have devised the term “Proto-Villanovan” to describe the cremating cultures of the Italian Final Bronze Age.

The fact that some of the earliest urnfield sites of peninsular Italy are located on the coast (e.g. Pianello in Romagna and Timmari in Apulia) is interpreted by some archaeologists as an indication that cremating people had come into Italy by sea, and that their migration was part of the larger upheaval which affected the eastern Mediterranean at the end of the Bronze Age (so Hencken 1968, 78–90). On the other hand, the same data can be explained in terms of indigenous coastal settlements adopting new cultural traits as the result of commercial interaction with foreigners. In any case, by the end of the Final Bronze Age inhumation had reemerged as the dominant funerary custom of southern Italy, but cremation continued to be an integral aspect of the Villanovan Culture of northern and much of central Italy.

etruscan-world
Diffusion of the Villanovan culture (after M. Torelli, ed., Gli Etruschi, Milan, 2000, p. 45). Modified from The Etruscan World (2013), by Turfa.

There is a myriad of linguistic reasons why eastern foreign influences can be attributed to Indo-European (mainly Anatolian, including a hypothetic influence on Latino-Faliscan) or Tyrsenian – as well as many other less credible models – and there is ground in archaeology to support any of the linguistic models proposed, given the long-lasting complex interactions of Italy with other Mediterranean cultures.

NOTE. The lack of theoretical schemes including integral archaeological-linguistic cultural-historical models due to the radical reaction against the excesses of the early 20th century have paradoxically allowed anyone (from archaeologists or linguists to laymen) to posit infinite population movements often based on the simplest similarities in vase decoration, burial practices, or shared vocabulary.

However, recent studies in population genomics have simplified the picture of Bronze Age population movements, identifying radical changes related to population replacements as opposed to more subtle admixture events. As of today, (France Bell Beaker-like) Urnfield stands as the most likely vector of Celtic languages; NW Iberian Bell Beakers as the vector of Galaico-Lusitanian; NW Mediterranean Beakers as the most likely ancestors of Elymian; the Danish Late Neolithic as representative of expanding Proto-Germanic; or Central-East Bell Beakers of Proto-Balto-Slavic.

With this in mind, the most logical conclusion is to assume that Alpine Bell Beakers (close to the sampled Italian Beakers from Parma or from southern Germany) spread Italo-Venetic languages, which is deemed to have split in the early to mid-2nd millennium BC, with dialects found widespread from the Alps to Sicily by the early 1st millennium BC.

Therefore, the two main remaining models of Italian linguistic prehistory – with the information that we already had – were as follows, concerning Tyrsenian (the ancestor of Etruscan and Rhaetian):

  1. It is a remnant language of the Italian (or surrounding) Chalcolithic, which survived in some pockets isolated from the Bell Beaker influence;
  2. It was a foreign language that arrived and expanded at the same time as the turmoil that saw the emergence of the Sea Peoples.

NOTE. Read more on Italo-Venetic evolution and on the likely distribution of Old European and Tyrsenian in the Bronze Age.

Italic-venetic-etruscan-languages-map
Languages of pre-Roman Italy and nearby islands. Italo-Venetic languages surrounded with shadowed red border. I1, South Picene; I2, Umbrian; I3, Sabine; I4, Faliscan; I5, Latin; I6, Volscian and Hernican; I7, Central Italic (Marsian, Aequian, Paeligni, Marrucinian, Vestinian); I8, Oscan, Sidicini, Pre-Samnite; I9, Sicel; IE1, Venetic; IE2, North Picene; IE3, Ligurian; IE4, Elymian; IE5, Messapian; C1, Lepontic; C2, Gaulish; G1-G2-G3, Greek dialects (G1: Ionic, G2: Aeolic, G3: Doric); P1, Punic; N1, Rhaetian; N2, Etruscan; N3, Nuragic. Image modified from Davius Sanctex.

Proto-Villanovan

A Proto-Villanovan female from Martinsicuro in the Abruzzo coast (ca. 890 BC), of mtDNA hg. U5a2b, is the earliest mainland sample available showing foreign (i.e. not exclusively Anatolia_N ± WHG) ancestry:

Martinsicuro is a coastal site located on the border of Le Marche and Abruzzo on central Italy’s Adriatic coast. It is a proto-Villanovan village, situated on a hill above the Tronto river, dating to the late Bronze Age and Early Iron Age (…) finds from the site indicate an affinity with contemporaries in the Balkans, suggesting direct trade contacts and interaction across the Adriatic. In particular, the practice of decorating ceramics with bronze elements was shared between the Nin region in Croatia and Picene region of Italy, including Martinsicuro.

NOTE. These are just some of the models I have tried, most of them unsuccessfully. The standard errors that I get are too high, but I am not much interested in this sample that seems (based on its position in the PCA and the available qpAdm results) mostly unrelated to Italic and Etruscan ethnogenesis.

The sample clusters close to the Early Iron Age sample from Jazinka (ca. 780 BC), from the central Dalmatian onomastic region, on the east Adriatic coast opposite to Abruzzo, possibly related to the south-east Dalmatian (or Illyrian proper) onomastic region to the south. However, there is no clear boundary between hydrotoponymic regions for the Bronze Age, and it is quite close to the (possibly Venetic-related) Liburnian onomastic region to the north, so the accounts of Martinsicuro belonging to the Liburni in proto-historical times can probably be extrapolated to the Final Bronze Age.

NOTE. Based on feminine endings in -ona in the few available anthroponyms, Liburnian may have shared similarities with personal names of the Noricum province, which doesn’t seem to be related to the more recent (Celtic- or Germanic-related?) Noric language. On the other hand, anthroponyms are known to show the most recent hydrotoponymic layer of a region, so these personal names might be unrelated to the ancestral language behind place and river names.

dalmatian-toponymic-liburnian
Toponyms ending in -ona (after S. Čače 2007).

Villanovan

A Villanovan sample from the powerful Etruscan city-state of Veio in the Tyrrhenian coast (ca. 850 BC), to the north of Rome, shows a cluster similar to later Etruscans and some Latins. Veio features prominently in the emergence of the Etruscan society. From The Etruscan World (2013) by Turfa:

In the final phase of the Bronze Age (mid-twelfth to tenth century bc) the disposition of settlements appears to be better distributed, although they are no longer connected to the paths of the tratturi (drove roads for transhumance of flocks and herds) as they had been during the Middle Bronze Age. As evidence of the intensive exploitation of land and continuous population growth there are now known in Etruria at least 70 confirmed settlements, and several more sites with indications of at least temporary occupation. The typical town of this chronological phase generally occupies high ground or a tufa plateau of more than five hectares, isolated at the confluence of two watercourses. These small plateaus, naturally or artificially protected, are not completely built up: non-residential areas within the defenses were probably intended as collecting points for livestock or zones reserved for cultivation, land used only by certain groups, or areas designated for shelter in case of enemy attack.

Taken together, the data seem to indicate the presence of individuals or families at the head of different groups. And in the final phase of the Bronze Age, there must have begun the process that generated (at least two centuries later) a tribal society based on families and the increasingly widespread ownership of land.

In the ninth century bc the territory is divided instead into rather large districts, each belonging to a large village, divided internally into widely spaced groups of huts, and into a small number of isolated villages located in strategic positions, for which we can assume some form of dependence upon the larger settlements.

etruscan-proto-urban
Schematic reconstruction of the birth of a proto-urban center (after P. Tamburini, II Museo
territoriale del Lago di Bolsena. Vol 1. Dalle origini alperiodo etrusco, Bolsena 2007). Modified from The Etruscan World (2013), by Turfa.

Compared to the preceding period, this type of aggregation is characterized by a higher concentration of the population. To the number of villages located mostly on inaccessible plateaus, with defensive priority assigned to the needs of agriculture, are added settlements over wide plains where the population was grouped into a single hilltop location. It is a sort of synoikistic process, so, for example, at Vulci people were gathered from the district of the Fiora and Albegna Rivers, while to Veii came the communities that inhabited the region from the Tiber River to Lake Bracciano, including the Faliscan and Capenate territories. The reference to Halesos, son of Saturn, the mythical founder of Falerii in the genealogy of Morrius the king of Veii (Servius, Commentary on Aeneid 8.285) may conceal this close relationship between Veii and the Ager Faliscus (the territory of the historical Faliscans).

The great movement of population that characterizes this period is unthinkable without political organizations that were able to impose their decisions on the individual village communities: the different groups, undoubtedly each consisting of nuclei linked by bonds of kinship, located within or outside the tufa plateaus that would be the future seats of the Etruscan city-states, have cultural links between them, also attested to by the analysis of craft production, such as to imply affiliation to the same political unit and enabling us to speak of such human concentrations as “proto-urban”.

etruscan-expansion-padania
Map of Etruria Padana. Left: From 9th to 8th century BC. Right: From 6th to 4th century BC. Dipartimento di Archeologia di Bologna. Modified from The Etruscan World (2013), by Turfa.

Italic vs. Etruscan origins

Four out of five sampled Latins show Yamnaya-derived R1b-L23 lineages, including three R1b-U152 subclades, and one hg. R1b-Z2103 (in line with the variability found among East Bell Beakers), while one from Ardea shows hg. T1a-L208. A likely Volscian (i.e. Osco-Umbrian-speaking) sample from Boville Ernica also shows hg. R1b-Z2118*, an ‘archaic’ subclade within the P312 tree. These R1b-L23 subclades are also found later during the Imperial period, although in lesser proportion compared to East Mediterranean ones.

Among Etruscans, the only male sampled shows hg. J2b-CTS6190* (formed ca. 1800 BC, TMRCA ca. 1100 BC), sharing parent haplogroup J2b-Y15058 (formed ca. 2400 BC, TMRCA ca. 1900 BC) with a Croatian MBA sample from Veliki Vanik (ca. 1580 calBCE), who also clusters close to the IA sample from Jazinka.

Given the position of Latins and Etruscans in the PCA and the likely similar admixture, it is not striking that differences are subtle. From Antonio et al. (2019):

Interestingly, although Iron Age individuals were sampled from both Etruscan (n=3) and Latin (n=6) contexts, we did not detect any significant differences between the two groups with f4 statistics in the form of f4(RMPR_Etruscan, RMPR_Latin; test population, Onge), suggesting shared origins or extensive genetic exchange between them.

On the other hand, there are 3 clear outliers among 11 Iron Age individuals, and all Iron Age samples taken together form a wide Etrurian cluster, so it seems natural to test them in groups divided geographically:

Results seem inconsistent, especially for Italic peoples, due to their wide cluster. It could be argued that the samples with ‘northern’ admixture – a Latin from Palestrina Colombella (of hg. R1b-Z56) and the Volscian sample – might represent better the Italic-speaking population before the proto-urban development of Latium, especially given the reported strong Etruscan influences among the Rutuli in Ardea, which might explain the common cluster with Etruscans and the outlier with reported ‘eastern’ admixture.

etruscan-latino-faliscan-osco-umbrian-italic-languages
Languages of Central Italy at the beginning of Roman expansion. Image modified from original by Susana Freixeiro at Wikipedia.

It makes sense then to test for a group of Etruscans (adding the Villanovan sample) and another of Italic peoples, to distinguish between a hypothetic ancestral Italic ancestry from a Tyrrhenian one:

NOTE. Fine-tuning groups based on the position of samples in the PCA or the amount of this or that component, or – even worse – based on the good or bad fits relative to the tested populations risks breaking the rules of subgroup analysis, eventually obtaining completely useless results, so interpretations for the Italic cluster need to be taken with a pinch of salt (until more similar Italic samples are published). The lack of proper rules regarding what can and cannot be done with this combined archaeological – genomic research is already visible to some extent in genetic papers which use brute force qpAdm tests for all available sampled populations, instead of selecting those potentially ancestral to the studied groups.

Tabs are organized from ‘better’ to ‘worse’ fits. In this case, as a general guide to the spreadsheets, the first tabs (to the left) show better fits for Italic peoples, and as tabs progress to the right they show ‘better’ fits for Etruscans, until it reaches the ‘infeasible’ or otherwise bad models.

This is what can be inferred from the models:

1) Steppe ancestry: Italic peoples seem to show better fits for north-western Alpine sources, closest to Bell Beakers from France or South Germany; whereas Etruscans show a likely Transdanubian source, closest to late Bell Beakers from Hungary (excluding Steppe- and WHG-related outliers).

To see if Bell Beakers from the south-west could be related, I tried the same model as in Fernandes et al. (2019), selecting Iberian BBC samples with more Steppe ancestry – to simplify my task, I selected them according to their PCA position. In a second attempt, I tried adding those intermediate with Iberia_CA, and it shows decreasing p-values, suggesting that the most likely source is close to high Steppe-related Bell Beaker populations. In both cases, models seem worse than France or Germany Bell Beakers.

Since Celtic spread with France BBC-like Urnfield peoples, and Italic peoples appear to be also ancestrally connected to this ancestry, the most plausible explanation is that they share an origin close to the Danubian EBA culture, which would probably be easily detectable by selecting precise Bell Beaker groups from South Germany.

expansion-celtic-peoples
Hypothetic expansion of Celtic-speaking peoples during the La Tène period (source). Image used in Udolph (2009) because it reflects a homeland roughly coincident with the oldest Celtic hydrotoponymy.

2) Anatolia_Neolithic ancestry: different tests seem to show that fits for EEF-related ancestry get warmer the closer the source population selected is to North-West Anatolian farmers, in line with the apparent shift from the East Bell Beaker cluster toward the Anatolia Neolithic cluster in the PCA:

These analyses suggest that there was a renewed Anatolia_N-like contribution during the Bronze Age, older than these Iron Age populations, but later than the rebound of WHG ancestry found among Late Neolithic and Chalcolithic samples from Italy, Sicily, or Sardinia, reflected in their shift in the PCA towards the WHG cluster.

From a range of chronologically closer groups clustering near Anatolia_N, the source seems to be closest to Neolithic samples from the Peloponnese. The direct comparison of Greece_Peloponnese_N against Italy_CA in the analyses labelled “Strict” shows that the sampled Greece Late Neolithic individuals are closer to the source of Neolithic ancestry of Iron Age Etrurians than the Chalcolithic samples from Remedello, Etruria, or Sardinia.

NOTE. Most qpAdm analyses are done with a model similar to Ning et al. (2019), using Corded_Ware_Germany.SG as an outgroup instead of Italy_Villabruna, because I expected to test all models against Yamnaya, too, but in the end – due to the many potential models and my limited time – I only tested those with ‘better’ fits:

Using Yamnaya_Kalmykia as outgroup gives invariably ‘worse’ results, as expected from Bell Beaker-derived populations who are directly derived from Yamnaya, despite their potential admixture with local Corded Ware peoples through exogamy during their expansion in Central Europe. The differences between Italic and Etruscan peoples have to be looked for mainly in EEF-related contributions, not in Steppe-related populations.

pca-italic-etruscan-latins-villanovan
Detail of the PCA of Eurasian samples, including Italian samples from Antonio et al. (2019) with the selected clusters of Italic vs. Etruscans, as well as Bell Beaker and Balkan BA and related clusters and outliers. Also marked are Peloponnese Late Neolithic (Greece_N), Minoans, Mycenaeans and Armenian BA samples. See image with better resolution.

Etruscans and Sea Peoples

The sister clade of the Etruscan branch, J2b-PH1602 (TMRCA ca. 1100 BC), seems to have spread in different directions based on its modern distribution, and their global parent clade J2b-Y15058 (TMRCA ca. 1900 BC) was previously found in Veliki Vanik. J2b-L283 appears related to Neolithic expansions through the Mediterranean, based on its higher diversity in Sardinia, although its precise origin is unclear.

Based on the modern haplogroup distribution and on the TMRCA, it can be assumed that a community spread with hg. J2b-Z38240 from somewhere close to the Balkans coinciding with the population movements of the Final Bronze Age. Whether this haplogroup’s Middle Bronze Age area, probably close to the Adriatic, was initially Indo-European-speaking or was related to a regional survival of Etruscan-speaking communities remains unclear.

Greece Late Neolithic is probably the closest available population (from those sampled to date) geographically and chronologically to the Bronze Age North-Western Anatolian region, where the Tyrsenian language family is hypothesized to have expanded from.

We only have a few Iron Age samples from Etruria, dating from a period of complex interaction in the Mediterranean – evidenced by the relatively high proportion of outliers – so it is impossible to discard the existence of some remnant Bronze Age population closer to the Adriatic – from either the Italian (Apulia?) or the Balkan coasts – expanding with the Proto-Villanovan culture and responsible for the Greece_LN-like ancestry seen among the sampled Final Bronze / Iron Age populations from central Italy.

On the other hand, taking into account the ancestry of available Italian, Sardinian and Sicilian Neolithic, Chalcolithic and Bronze Age samples, the current genetic picture suggests an expansion of a different North-West Anatolia Neolithic-related population after the arrival of Bell Beakers from the north, hence probably through the Adriatic rather than through the Tyrrhenian coast, whether the common language group formed with Lemnian had a more distant origin in Bronze Age North-West Anatolian groups or in some isolated coastal community of the Adriatic.

NOTE. Admittedly, the ancestry of the Proto-Villanovan sample seems different from that of Etruscans, although a contribution of the most likely sources for Etruscans cannot be rejected for the Proto-Villanovan individual (see ‘reciprocal’ models of admixture here). In any case, I doubt that the main ancestry of the Proto-Villanovan from Abruzzo is directly related to the population that gave rise to Etruscans, and is more likely related to recent, intense bilateral exchanges in the Adriatic between (most likely) Indo-European-speaking populations.

violin-bow-fibula-italy-illyria-aegean-crete
The distribution of violin bow fibula from thirteenth century onward showing the movement of people between northern Italy, Illyria and the Aegean, Crete, and the parallel distribution of “foreign” darksurfaced handmade pottery (based on Kasuba 2008 : abb. 15; Lis 2009 ). Modified from Kristiansen (2018).

Northern Adriatic

This Adriatic connection could in turn be linked to wider population movements of the Final Bronze Age. Proto-Villanova represents the introduction of oriental influences coinciding with the demise of the local Terramare culture (see e.g. Cremaschi et al. 2016), whereas the Villanovan culture shows partial continuity with many Proto-Villanovan settlements where Etruscan-speaking communities later emerge. From Nicolis (2013):

Founded in the LBA, the village of Frattesina extended over around 20 hectares along the ‘Po di Adria’, a palaeochannel of the Po. It experienced its greatest development between the twelfth and eleventh centuries BC, when it had a dominant economic role thanks to an extraordinary range of artisan production (metalworking, working of bone and deer horn, glass) and major commercial influence due to trading with the Italian Peninsula and the eastern Mediterranean.

This is demonstrated by the presence of exotic objects and raw materials, such as Mycenaean pottery, amber, ivory, ostrich eggs, and glass paste. For the Mycenaean sherds found in settlements in the Verona valleys and the Po delta, analysis of pottery fabrics has shown that some of them very probably come from centres in Apulia where there were Aegean craftsmen and workers, whereas others would seem to have originated on the Greek mainland (Vagnetti 1996; Vagnetti 1998; Jones et al. 2002).

acqua-fredda-passo-del-redebus
Reconstruction of Acqua Fredda archaeological site, Passo del Redebus, where a group of 9 smelting furnaces has been discovered dating back to the Late Bronze Age (8-9th century BC). Image modified from Trentino Cultura.

In this context a particular system of relations seems to link one specific Alpine region with the social and economic structure of the groups settling between the Adige and the Po and the eastern Mediterranean trading system. In eastern Trentino, at Acquafredda, metallurgical production on a proto-industrial scale has been demonstrated between the end of the LBA and the FBA (twelfth–eleventh centuries BC) (Cierny 2008) (Fig. 38.3). These products must have supplied markets stretching beyond the local area, linked to the Luco/Laugen culture typical of the central Alpine environment. According to Pearce and De Guio (1999), such extensive production must have been destined for the supply of metal to other markets, first of all to other centres on the Po plain, where transactions for materials of Mediterranean origin also took place.

The picture of the Final Bronze Age of these regions, which seems to be coherent with the development of the cultural setting of the Early Iron Age, shows that the birth of the proto-urban Villanovan centres of Bologna in Emilia and Verucchio in Romagna, at the beginning of the Iron Age, seems to follow a line of continuity starting with the role played by Frattesina in the Final Bronze Age (Bietti Sestieri 2008).

naue-swords-final-bronze-age
Reconstruction of pan-European communication network represented by the geographical spread of archaeological objects. The network nodes represent sites that have yielded an above-average number of relevant finds. The links are direct connections between neighbouring nodes. Modified from Suchowska-Ducke (2015).

Tyrsenian

The close similarities shared by Rhaetian with the oldest Etruscan inscriptions – but not with the language of later periods, when Etruscan expanded further north – together with increased ‘foreign’ contacts in the Final Bronze Age and the ‘foreign’ ancestry of Etruscans (relative to Italian Chalcolithic and to near-by Bell Beakers) support a language split close to the Adriatic, and not long before they started using the Euboean-related Old Italic alphabet. All this is compatible with an expansion associated with the Proto-Villanovan period, possibly starting along the Po and the Adige.

From Nicolis (2013):

In this geographical context the most important morphological features are the Alps and the alluvial plain of the River Po. Since Roman times the former have always been considered a geographical limit and thus a cultural barrier. In actual fact the Alps have never really represented a barrier, but instead have played an active role in mediating between the central European and Mediterranean cultures. Some of the valleys have been used since the Mesolithic as communication routes, to establish contacts and for the exchange of materials and people over considerable distances. The discovery of Ötzi the Iceman high in the Alps in 1991 demonstrated incontrovertibly that this environment was accessible to individuals and groups from the end of the fourth millennium BC.

From the Early Neolithic period the plain of the Po Valley provided favourable conditions for the population of the area by human groups from central and eastern Europe, who found the wide flat spaces and fertile soils an ideal environment for developing agricultural techniques and animal husbandry. Lake Garda represents a very important morphological feature, benefiting among other things from a Mediterranean-type microclimate, the influence of which can already be seen in the Middle Neolithic. Situated between the plain and the mountains, the hills have always offered an alternative terrain for demographic development, equally important for the exploitation of economic and environmental resources.

As documented for previous periods, in the late and final phases of the Bronze Age the northern Adriatic coast would also seem to represent an important geographical feature, above all in terms of possible long-distance trading contacts with the Aegean and eastern Mediterranean coasts. However, the geographical and morphological characteristics and the river network in this area were very different to the way they are today, and the preferred communications routes must always have been the rivers, particularly the Po and the Adige.

etruscan-rhaetian-inscriptions
Map of inscriptions of Northern Italy. In green, Rhaetian inscriptions; in Pink, Etruscan inscriptions. Arrows show potential language movements through the Po and the Adige based on the relationship between both language. Image modified from Raetica.

Conclusion

Although it seems superfluous at this point, finding mostly Yamnaya-derived R1b-L23 lineages among speakers of another early North-West Indo-European dialect – and also the earliest to have split into its attested dialects – gives still more support to Yamnaya steppe herders as the vector of expansion of Late PIE, and their continuity up to the Iron Age also supports the strong patrilineal ties of Indo-Europeans.

This, in turn, further supports the nature of Afanasievo as the earliest separated branch from a Late Proto-Indo-European trunk, and of Khvalynsk as the Indo-Anatolian community, while a confirmation of R1b-L23 among early Greeks (speaking the earliest attested Graeco-Aryan dialect) will indirectly confirm East Yamnaya/Poltavka as the early Proto-Indo-Iranian community.

As it often happens with genetic sampling, due to many uncontrollable factors, there is a conspicuous lack of a proper regional and chronological transect of Bell Beaker and Bronze Age samples from Italy, which makes it impossible to determine the origin of each group’s ancestral components. Even though the sampled Italian Beakers don’t seem to be the best fit for Iron Age Italic-speaking peoples from Etruria, they still might have formed part of the migration waves that eventually developed the Apennine culture together with those of prevalent West-Central European Bell Beaker ancestry.

Similarly, the visible radical change from the increasingly WHG-shifted Italian farmers up to the sampled Chalcolithic individuals, including Parma Bell Beakers, to the Anatolia_N-shifted ancestry found in Iron Age Etruscans and Latins might be related to earlier population movements associated with Middle or Late Bronze Age contacts, and not necessarily to the radical social changes seen in the Final Bronze Age. The Etruscan subclade with a likely origin in the Balkans, on the other hand, suggests recent migrations from the Adriatic into Etruria.

middle-bronze-age-italy
Middle Bronze Age cultures of Italy and its surroundings ca. 1750-1250 BC. Potential source of the Greece_N-like admixture found widespread during the Iron Age. See full maps.

Until there is more data about these ancestry changes in Italy, the Balkans, and North-West Anatolia, I prefer to leave the Tyrsenian origins up in the air, so I deleted the Lemnian -> Etruscan arrow of the map of Late Bronze Age migrations, if only because an arrival through the Tyrrhenian Sea has become much less likely. An East -> West movement is still the most likely explanation for the common Tyrsenian language, culture, and ancestry, but the only Y-DNA haplogroup available seems to have an origin closer to the Adriatic.

The recent study of Sea Peoples showed – based on the previous hypothesis of the language and culture of the Philistines – that a minority of incoming elites must have imposed the language as their genetic ancestry (including haplogroups) became diluted among a majority of local peoples. Similarly, the original genetic pool of Tyrsenian speakers might have become diluted among different groups due to their more complex social organization, similar to what happened to Italic peoples during the Imperial period.

One of the most interesting aspects proven in the paper – and strongly suspected before it – is the reflection in population genomics of the change in the social system of the Italian Peninsula during the Roman expansion, and even before it during the Etruscan polity. In fact, it was not only Romans who spread and genetically influenced other European regions, but other regions – especially the more numerous Eastern Mediterranean populations – who became incorporated into a growing Etrurian community which nevertheless managed to spread its language.

In other words, Tyrsenian spread through central and northern Italy, and Latin throughout the whole Mediterranean area and mainland Europe, not (only) through population movements, but through acculturation, in a growing international system of more complex political organizations that can be inferred for most population and language expansions since the Early Iron Age. East Mediterranean populations, Scythians and other steppe peoples, East Germanic peoples, Vikings, or North-Eastern Europeans are other clear examples known to date.

Related

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 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

“Dinaric I2a” and the expansion of Common Slavs from East-Central Europe

late-iron-age-eastern-europe

A recently published abstract for an upcoming chapter about Early Slavs shows the generalized view among modern researchers that Common Slavs did not spread explosively from the east, an idea proper of 19th-century Romantic views about ancestral tribes of pure peoples showing continuity since time immemorial.

Migrations and language shifts as components of the Slavic spread, by Lindstedt and Salmela, In: Language contact and the early Slavs, Eds. Tomáš Klír, Vít Boček, Universitätsverlag Winter (2019):

The rapid spread of the Proto-Slavic language in the second half of the first millennium CE was long explained by the migration of its speakers out of their small primary habitat in all directions. Starting from the 1980s, alternative theories have been proposed that present language shift as the main scenario of the Slavic spread, emphasizing the presumed role of Slavic as the lingua franca of the Avar Khaganate. Both the migration and the language shift scenarios in their extreme forms suffer from factual and chronological inaccuracy. On the basis of some key facts about human population genetics (the relatively recent common ancestry of the East European populations), palaeoclimatology (the Late Antique Little Ice Age from 536 to around 660 CE), and historical epidemiology (the Justinianic Plague), we propose a scenario that includes a primary rapid demographic spread of the Slavs followed by population mixing and language shifts to and from Slavic in different regions of Europe. There was no single reason for the Slavic spread that would apply to all of the area that became Slavic-speaking. The northern West Slavic area, the East Slavic area, and the Avar sphere and South-Eastern Europe exhibit different kinds of spread: mainly migration to a sparsely populated area in the northwest, migration and language shift in the east, and a more complicated scenario in the southeast. The remarkable homogeneity of Slavic up to the jer shift was not attributable to a lingua-franca function in a great area, as is often surmised. It was a founder effect: Proto-Slavic was originally a small Baltic dialect with little internal variation, and it took time for the individual Slavic languages to develop in different directions.

While I would need to read the whole chapter, in principle it seems easier to agree with this summary than with Curta’s (sort of diffuse) Danubian origin of Common Slavic, based on the likely origin of the Balto-Slavic expansion with the Trzciniec and/or Lusatian culture, close to the Baltic.

A multi-ethnic Chernyakhov culture

In a sneak peek to the expected Järve et al. (2019) paper in review, there are three Chernyakhov samples (ca. calAD 350-550) with different ancestry probably corresponding to the different regions where they stem from (see image below), which supports the idea that Iron Age eastern Europe was a true melting pot where the eventual language of the different cultures depended on many different factors:

chernyakhov-samples-region
Map of the samples from Järve et al. (2019).

From the paper:

The Chernyakhiv culture was likely an ethnically heterogeneous mix based on Goths (Germanic tribes) but also including Sarmatians, Alans, Slavs, late Scythians and Dacians – the entire ancient population of the northern coast of the Black Sea.

Contacts with neighbouring regions were active, and the Chernyakhiv culture is associated with a number of historical events that took place in Europe at that time. In particular, during the Scythian or Gothic wars of the 230s and 270s, barbarians living in the territory of the Chernyakhiv culture (Goths, Ferules, Carps, Bastarns, etc.) carried out regular raids across the Danube Limes of the Roman Empire. However, from the end of the 3rd century the relations of the barbarians with the Roman Empire gained a certain stability. From the reign of Constantine I the Goths, who were part of the Chernyakhiv culture, became federates (military allies) of the Empire.

The Goths also interacted with the inhabitants of the East European forest zone. The Roman historian Jordanes described the military campaigns of the Gothic king Ermanaric against northern peoples (the ancestors of Vends, Slavs, etc., and the inhabitants of the northern Volga region).

NOTE. As it has become traditional in writings about eastern Europe, ‘Slavs’ are assumed – for no particular reason – to be part of the ‘northern peoples of the forest’ since who knows when exactly, and thus appear mentioned in this very text simultaneously as part of Chernyakhov, but also part of peoples to the north of Chernyakhov warring against them…

admixture-chernyakhov
Proportions of Eastern Hunter-Gatherer (EHG, blue), Natufian (red) and Altaian (green) ancestries in Scythian/Sarmatian groups and groups pre- and postdating them inferred using the a) qpAdm and b) ChromoPainter/NNLS method. c–e Correlation of qpAdm and CP/NNLS proportions for the three putative sources evaluated. Steppe populations predating the Scythians: Yamnaya_Ukraine [26], Yamnaya_Kalmykia [15], Ukr_BA (this study). Scythians and Sarmatians: Nomad_IA [15], Scythian_East and Sarmatian_SU [3], Hungarian Scythian, Sarmatian, Central Saka, Tian Shan Saka and Tagar [1], Scy_Ukr, ScySar_SU and Scy_Kaz (this study). Population postdating the Scythians: Chern (this study). See also Table S3.

Genetic variation

(…) the Chernyakhiv samples overlapped with modern Europeans, representing the most ‘western’ range of variation among the groups of this study.

After the end of the Scythian period in the western Eurasian Steppe, the Chernyakhiv culture samples have higher Near Eastern affinity compared to the Scythians preceding them, agreeing with the Gothic component in the multi-ethnic mix of the Chernyakhiv culture.

The higher proportion Near Eastern and (according to CP/NNLS) lower proportion of eastern ancestry in the Chernyakhiv culture samples were mirrored by f4 analyses where Chern showed lower affinity to Han (Z score –3.097) and EHG (Z score –3.643) than Ukrainian Scythian and Bronze Age samples, respectively, as well as higher Near Eastern (Levant_N and Anatolia_N) affinity than Ukrainian Scythians (Z scores 4.696 and 3.933, respectively). It is plausible to assume that this excess Near Eastern ancestry in Chern is related to European populations whose Near Eastern proportion has exceeded that in the steppe populations since the Neolithic expansion of early farmers. While the Chernyakhiv culture was likely ethnically heterogeneous, the three samples in our Chern group appear to represent its Gothic component.

chernyakhov-goths-uralic-clines
PCA obtained by projecting the ancient samples of this study together with published Scythian/Sarmatian and related samples onto a plot based on 537,802 autosomal SNPs in 1,422 modern Eurasians. To improve readability, the modern populations have been plotted as population medians (after outlier removal). Image modified from the paper, including Sredni Stog, Corded Ware/Uralic (with Srubna outliers) and Chernyakhov clusters.Notice the two new Late Yamna and Catacomb samples from Ukraine clustering with other published samples, despite being from the same region as Sredni Stog individuals.

Early Slavs of hg. I2-L621

A post in Anthrogenica shows some subclades of the varied haplogroups that are expected from medieval Poland:

KO_55, Kowalewko (100-300 AD), I1a3a1a1-Y6626
KO_45, Kowalewko (100-300 AD), I2a2a1b2a-L801
KO_22, Kowalewko (100-300 AD), G2a2b-L30
KO_57, Kowalewko (100-300 AD), G2a2b-L30

ME_7, Markowice (1000-1200 AD), I1a2a2a5-Y5384
NA_13, Niemcza, (900-1000 AD), I2a1b2-L621
NA_18, Niemcza, (900-1000 AD), J2a1a-L26

Just because of these samples among Early Slavs, and looking again more carefully at the modern distribution of I2a-L621 subclades, I think now I was wrong in assuming that I2a-L621 in early Hungarian Conquerors would mean they would appear around the Urals as a lineage integrated in Eastern Corded Ware groups. It seems rather a haplogroup with an origin in Central Europe. Whether it was part of a Baltic community that expanded south, or was incorporated during the expansions to the south is unclear. Like hg. E-V13, it doesn’t seem to have been incorporated precisely along the Danube, but closer to the north-east Carpathians.

Especially interesting is the finding of I2a-L621 among Early Slavs from Silesia, a zone of close interaction among early West Slavs. From Curta (2019):

On Common Slavs

In Poland, settlement discontinuity was postulated, to make room for the new, Prague culture introduced gradually from the southeast (from neighboring Ukraine). However, there is increasing evidence of 6th-century settlements in Lower Silesia (western Poland and the lands along the Middle Oder) that have nothing to do with the Prague culture. Nor is it clear how and when did the Prague culture spread over the entire territory of Poland.

On Great Moravia

Svatopluk’s remarkably strong position was immediately recognized by Pope John VIII, who ordered the immediate release of Methodius from his monastic prison in order to place him in 873 under Svatopluk’s protection. One year later (874), Louis the German himself was forced to recognize Svatopluk’s independence through the peace of Forchheim. By that time, the power of Svatopluk had extended into the upper Vistula Basin, over Bohemia, the lands between the Saale and the Elbe rivers, as well as the northern and northeastern parts of the Carpathian Basin.* The Czech prince Bořivoj, a member of the Přemyslid family which would unify and rule Bohemia in the following century, is believed to have been baptized in 874 by Methodius in Moravia together with his wife Ludmila (St. Wenceslas’s grandmother).

*Brather, Archäologie, p. 71. The expansion into the region of the Upper Vistula (Little Poland) results from one of St. Methodius’ prophecies, for which see the Life of Methodius 11, p. 72; Poleski, “Contacts between the Great Moravian empire and the tribes”; Poleski, “Contacts between the tribes in the basins.” Despite an early recognition of the Moravian influences on the material culture in 9th-century southern Poland and Silesia (e.g., Dostál, “Das Vordringen”), the question of Svatopluk’s expansion has triggered in the 1990s a fierce debate among Polish archaeologists. See Wachowski, “Problem”; Abłamowicz, “Górny Śląsk”; Wachowski, “Północny zasięg ekspansji”; Szydłowski, “Czy ślad”; Jaworski, “Elemente.”

On Piast Poland

Mieszko agreed to marry Oda, the daughter of the margrave of the North March, for his first wife had died in 977. The marriage signaled a change in the relations with the Empire, for Mieszko sent troops to help Otto II against the Slavic rebels of 983. He also attacked Bohemia and incorporated Silesia and Lesser Poland into the Piast realm, which prompted Bohemians to ally themselves with the Slavic rebels against whom Emperor Otto was now fighting. By 980, therefore, Mieszko was part of a broader configuration of power, and his political stature was recognized in Scandinavia as well. His daughter, Swietoslawa married first Erik Segersäll of Sweden (ca. 970–ca. 995) and then Sweyn Forkbeard of Denmark (986–1014).26 In the early 990s, together with his wife and children, Mieszko offered his state (called “civitas Schinesghe,” the state of Gniezno) to the pope as a fief, as attested by a unique document known as Dagome iudex and preserved in a late 11th-century summary. The document describes the inner boundaries of the state and peripheral provinces, as if Gniezno were a civitas (city) in Italy, with its surrounding territory. Regional centers, however, did indeed come into being shortly before AD 1000 in Lesser Poland (Cracow, Sandomierz), Pomerania (Gdańsk), and Silesia (Wrocław). Such regional centers came to be distinguished from other strongholds by virtue of the presence within their walls of some of the earliest churches built in stone. Mieszko got his own, probably missionary bishop.

In light of this recent find, which complements the Early Slav of the High Middle Ages from Sunghir (ca. AD 1100-1200), probably from the Vladimir-Suzdalian Rus’, we can assume now less speculatively that I2a-CTS10228 most likely expanded with Common Slavs, because alternative explanations for its emergence in the Carpathian Basin, among Early West Slavs, and among Early East Slavs within this short period of time requires too many unacceptable assumptions.

dinaric-i2a-distribution
Modern distribution of “Dinaric” I2a. Modified from Balanovsky et al. (2008)

Hungarian Conquerors

Knowing that R1a-Z280 was an Eastern Corded Ware lineage, found from Baltic Finns to Finno-Ugric populations of the Trans-Urals, we can probably assign expanding Magyars to at least R1a-Z280, R1a-Z93, and N1c-L392 (xB197) lineages.

From Curta (2019):

Earlier Latin sources, especially those of the first half of the 10th century, refer to Magyars as Huns or Avars. They most likely called themselves Magyars, a word indicating that the language they spoke was not Turkic, but Finno-Ugrian, related to a number of languages spoken in Western Siberia and the southern Ural region. The modern word—Hungarian—derives from the Slavic word for those people, U(n)gri, which is another indication of Ugric roots. This has encouraged the search for the origin of the Hungarian people in the lands to the east from the Ural Mountains, in western Siberia, where the Hungarian language is believed to have emerged between 1000 and 500 BC.

In looking for the Magyar primordial homeland, they draw comparisons with the assemblages found in Hungary that have been dated to the 10th century and attributed to the Magyars. Some of those comparisons had extraordinary results. For example, the excavation of the burial mound cemetery recently discovered near Lake Uelgi, in the Cheliabinsk region of Russia, has produced rosette-shaped harness mounts and silver objects ornamented with palmette and floral designs arranged in reticulated patterns, which are very similar to those of Hungary. But Uelgi is not dated to prehistory, and many finds from that site coincided in time with those found in burial assemblages in Hungary. In other words, although there can be no doubt about the relations between Uelgi and the sites in Hungary attributed to the first generations of Magyars, those relations indicate a migration directly from the Trans-Ural lands, and not gradually, with several other stops in the forest-steppe and steppe zones of Eastern Europe. In the lands west of the Ural Mountains, the Magyars are now associated with the Kushnarenkovo (6th to 8th century) and Karaiakupovo (8th to 10th century) cultures, and with such burial sites as Sterlitamak (near Ufa, Bashkortostan) and Bol’shie Tigany (near Chistopol, Tatarstan).14 However, the same problem with chronology makes it difficult to draw the model of a migration from the lands along the Middle Volga. Many parallels for the so typically Magyar sabretache plates found in Hungary are from that region. They have traditionally been dated to the 9th century, but more recent studies point to the coincidence in time between specimens found in Eastern Europe and those from Hungary.

Adding J2a and I1a samples to the Early Slavic stock, based on medieval samples from Poland – with G2a and E-V13 lineages probably shared with Goths from Wielbark/Chernyakhov, or becoming acculturated in the Carpathian Basin – one is left to wonder which of these lineages actually took part in Common Slavic migrations/acculturation events, whenever and wherever those actually happened.

I have tentatively re-assigned lineages of Hungarian conquerors according to their likely origins in a simplistic way – similar to how the paper classifies them – , now (I think) less speculatively, assuming that Early Slavs likely formed eventually part of them:

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

NOTE. The ancestral origin of lineages is meaningless for an ethnolinguistic identification. The only reasonable assumption is that all the individuals sampled formed part of the Magyar polity, shared Magyar culture, and likely spoke Hungarian, unless there is a clear reason to deny this: which I guess should include at least a clearly ‘foreign’ ancestry (showing a distant cluster compared to the group formed by all other samples), ‘foreign’ isotopic data (showing that he was born and/or raised outside of the Carpathian Basin), and particularly ‘foreign’ cultural assemblage of the burial, if one really wants to risk assuming that the individual didn’t speak Hungarian as his mother tongue.

“Dinaric” or Slavic I2a?

I don’t like the use of “Dinaric I2a”, because it is reminiscent of the use of “Iberian R1b-DF27”, or “Germanic R1b-U106”, when ancient DNA has shown that this terminology is most often wrong, and turns out to be misleading. As misleading as “Slavic R1a”. Recently, a Spanish reader wrote me emails wondering how could I possibly say that R1b-DF27 came from Central Europe, because modern distribution maps (see below) made it evident that the haplogroup expanded from Iberia…

DF27-iberia-france-m167
Contour maps of the derived allele frequencies of the SNPs analyzed in Solé-Morata et al. (2017).

The obvious answer is, these maps show modern distributions, not ancient ones. In the case of R1b-DF27, different Iberian lineages are not even related to the same expansion. At least R1b-M167/SRY2627 lineages seem to have expanded from Central Europe into Iberia much more recently than other DF27 subclades associated with Bell Beakers. What’s more, if R1b-M167/SRY2627 appear densest in north-east Spain it is not because of the impact of Celts or Iberians before the arrival of Romans, but because of the impact of medieval expansions during the Reconquista from northern kingdoms expanding south in the Middle Ages:

iberian-medieval-kingdoms-expansion-population-genomics
Genetic differentiation and the footprints of historical migrations in the Iberian Peninsula. Image modified from Bycroft et al. (2018).

Similarly, the term “Dinaric I2a”, based on the higher density in the Western Balkans, is misleading because it is probably the result of later bottlenecks. Just like the density of different R1a subclades among Modern Slavs is most likely the result of acculturation of different groups, especially to the east and north-east, where language shift is known to have happened in historical times, with the cradle of Russians in particular being a Finno-Volgaic hotspot, later expanding with hg. R1a-Z280 and N1c-L392 lineages.

Now, one may think that maybe Slavs expanded with ALL of these different lineages. Since we are talking about late Iron Age / medieval expansions, there might be confederations of different peoples expanding with a single lingua franca… But no, not really. Not likely in linguistics, not likely in archaeology, and apparently not in population genomics, either.

How many ancient peoples from the Iron Age and Early Middle Ages expanded with so many different lineages? We see bottlenecks in expansions even in recent times: say, in Visigoths under E-V13 (probably recently incorporated during their migrations); in Moors (mostly Berbers) with E-M81 and J; in medieval Iberians under different DF27 bottlenecks during the Reconquista (including huge bottlenecks among Basques); similarly, huge bottlenecks are found in Finnic expansions under N1c…How likely is it that Proto-Slavs (and Common Slavs) expanded with all those attested lineages to date among Early Slavs (E-V13, I2a-L621, R1a-M458, I1, J2a) AND also with other R1a subclades prevalent today, but almost absent in sampled Early Slavs?

To sum up, I am not so sure anymore about the possibility of simplistically assigning R1a-M458 to expanding Common Slavs. R1a-M458 may well have been the prevalent R1a subclade in Central Europe among early Balto-Slavic – and possibly also neighbouring Northern Indo-European-speaking – peoples (let’s see what subclades Tollense and Unetice samples bring), but it is more and more likely that most of the density we see in modern R1a-M458 distribution maps is actually the effect of medieval bottlenecks of West Slavs, similar to the case of Iberia.

r1a-m458-underhill-2015
Modern distribution of R1a-M458, after Underhill et al. (2015).

Related

ASoSaH Reread (II): Y-DNA haplogroups among Uralians (apart from R1a-M417)

corded-ware-yamna-ancestry

This is mainly a reread of from Book Two: A Game of Clans of the series A Song of Sheep and Horses: chapters iii.5. Early Indo-Europeans and Uralians, iv.3. Early Uralians, v.6. Late Uralians and vi.3. Disintegrating Uralians.

“Sredni Stog”

While the true source of R1a-M417 – the main haplogroup eventually associated with Corded Ware, and thus Uralic speakers – is still not known with precision, due to the lack of R1a-M198 in ancient samples, we already know that the Pontic-Caspian steppes were probably not it.

We have many samples from the north Pontic area since the Mesolithic compared to the Volga-Ural territory, and there is a clear prevalence of I2a-M223 lineages in the forest-steppe area, mixed with R1b-V88 (possibly a back-migration from south-eastern Europe).

R1a-M459 (xR1a-M198) lineages appear from the Mesolithic to the Chalcolithic scattered from the Baltic to the Caucasus, from the Dniester to Samara, in a situation similar to haplogroups Q1a-M25 and R1b-L754, which supports the idea that R1a, Q1a, and R1b expanded with ANE ancestry, possibly in different waves since the Epipalaeolithic, and formed the known ANE:EHG:WHG cline.

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

The first confirmed R1a-M417 sample comes from Alexandria, roughly coinciding with the so-called steppe hiatus. Its emergence in the area of the previous “early Sredni Stog” groups (see the mess of the traditional interpretation of the north Pontic groups as “Sredni Stog”) and its later expansion with Corded Ware supports Kristiansen’s interpretation that Corded Ware emerged from the Dnieper-Dniester corridor, although samples from the area up to ca. 4000 BC, including the few Middle Eneolithic samples available, show continuity of hg. I2a-M223 and typical Ukraine Neolithic ancestry.

NOTE. The further subclade R1a-Z93 (Y26) reported for the sample from Alexandria seems too early, given the confidence interval for its formation (ca. 3500-2500 BC); even R1a-Z645 could be too early. Like the attribution of the R1b-L754 from Khvalynsk to R1b-V1636 (after being previously classifed as of Pre-V88 and M73 subclade), it seems reasonable to take these SNP calls with a pinch of salt: especially because Yleaf (designed to look for the furthest subclade possible) does not confirm for them any subclade beyond R1a-M417 and R1b-L754, respectively.

The sudden appearance of “steppe ancestry” in the region, with the high variability shown by Ukraine_Eneolithic samples, suggests that this is due to recent admixture of incoming foreign peoples (of Ukraine Neolithic / Comb Ware ancestry) with Novodanilovka settlers.

The most likely origin of this population, taking into account the most common population movements in the area since the Neolithic, is the infiltration of (mainly) hunter-gatherers from the forest areas. That would confirm the traditional interpretation of the origin of Uralic speakers in the forest zone, although the nature of Pontic-Caspian settlers as hunter-gatherers rather than herders make this identification today fully unnecessary (see here).

EDIT (3 FEB 2019): As for the most common guesstimates for Proto-Uralic, roughly coinciding with the expansion of this late Sredni Stog community (ca. 4000 BC), you can read the recent post by J. Pystynen in Freelance Reconstruction, Probing the roots of Samoyedic.

eneolithic-ukraine-corded-ware
Late Sredni Stog admixture shows variability proper of recent admixture of forest-steppe peoples with steppe-like population. See full version here.

NOTE. Although my initial simplistic interpretation (of early 2017) of Comb Ware peoples – traditionally identified as Uralic speakers – potentially showing steppe ancestry was probably wrong, it seems that peoples from the forest zone – related to Comb Ware or neighbouring groups like Lublyn-Volhynia – reached forest-steppe areas to the south and eventually expanded steppe ancestry into east-central Europe through the Volhynian Upland to the Polish Upland, during the late Trypillian disintegration (see a full account of the complex interactions of the Final Eneolithic).

The most interesting aspect of ascertaining the origin of R1a-M417, given its prevalence among Uralic speakers, is to precisely locate the origin of contacts between Late Proto-Indo-European and Proto-Uralic. Traditionally considered as the consequence of contacts between Middle and Upper Volga regions, the most recent archaeological research and data from ancient DNA samples has made it clear that it is Corded Ware the most likely vector of expansion of Uralic languages, hence these contacts of Indo-Europeans of the Volga-Ural region with Uralians have to be looked for in neighbours of the north Pontic area.

sredni-stog-repin-contacts
Sredni Stog – Repin contacts representing Uralic – Late Indo-European contacts were probably concentrated around the Don River.

My bet – rather obvious today – is that the Don River area is the source of the earliest borrowings of Late Uralic from Late Indo-European (i.e. post-Indo-Anatolian). The borrowing of the Late PIE word for ‘horse’ is particularly interesting in this regard. Later contacts (after the loss of the initial laryngeal) may be attributed to the traditionally depicted Corded Ware – Yamna contact zone in the Dnieper-Dniester area.

NOTE. While the finding of R1a-M417 populations neighbouring R1b-L23 in the Don-Volga interfluve would be great to confirm these contacts, I don’t know if the current pace of more and more published samples will continue. The information we have right now, in my opinion, suffices to support close contacts of neighbouring Indo-Europeans and Uralians in the Pontic-Caspian area during the Late Eneolithic.

Classical Corded Ware

After some complex movements of TRB, late Trypillia and GAC peoples, Corded Ware apparently emerged in central-east Europe, under the influence of different cultures and from a population that probably (at least partially) stemmed from the north Pontic forest-steppe area.

Single Grave and central Corded Ware groups – showing some of the earliest available dates (emerging likely ca. 3000/2900 BC) – are as varied in their haplogroups as it is expected from a sink (which does not in the least resemble the Volga-Ural population):

Interesting is the presence of R1b-L754 in Obłaczkowo, potentially of R1b-V88 subclade, as previously found in two Central European individuals from Blätterhole MN (ca. 3650 and 3200 BC), and in the Iron Gates and north Pontic areas.

Haplogroups I2a and G have also been reported in early samples, all potentially related to the supposed Corded Ware central-east European homeland, likely in southern Poland, a region naturally connected to the north Pontic forest-steppe area and to the expansion of Neolithic groups.

corded-ware-haplogroups
Y-DNA samples from early Corded Ware groups and neighbouring cultures. See full version.

The true bottlenecks under haplogroup R1a-Z645 seem to have happened only during the migration of Corded Ware to the east: to the north into the Battle Axe culture, mainly under R1a-Z282, and to the south into Middle Dnieper – Fatyanovo-Balanovo – Abashevo, probably eventually under R1a-Z93.

This separation is in line with their reported TMRCA, and supports the split of Finno-Permic from an eastern Uralic group (Ugric and Samoyedic), although still in contact through the Russian forest zone to allow for the spread of Indo-Iranian loans.

This bottleneck also supports in archaeology the expansion of a sort of unifying “Corded Ware A-horizon” spreading with people (disputed by Furholt), the disintegrating Uralians, and thus a source of further loanwords shared by all surviving Uralic languages.

Confirming this ‘concentrated’ Uralic expansion to the east is the presence of R1a-M417 (xR1a-Z645) lineages among early and late Single Grave groups in the west – which essentially disappeared after the Bell Beaker expansion – , as well as the presence of these subclades in modern Central and Western Europeans. Central European groups became thus integrated in post-Bell Beaker European EBA cultures, and their Uralic dialect likely disappeared without a trace.

NOTE. The fate of R1b-L51 lineages – linked to North-West Indo-Europeans undergoing a bottleneck in the Yamna Hungary -> Bell Beaker migration to the west – is thus similar to haplogroup R1a-Z645 – linked to the expansion of Late Uralians to the east – , hence proving the traditional interpretation of the language expansions as male-driven migrations. These are two of the most interesting genetic data we have to date to confirm previous language expansions and dialectal classifications.

It will be also interesting to see if known GAC and Corded Ware I2a-Y6098 subclades formed eventually part of the ancient Uralic groups in the east, apart from lineages which will no doubt appear among asbestos ware groups and probably hunter-gatherers from north-eastern Europe (see the recent study by Tambets et al. 2018).

Corded Ware ancestry marked the expansion of Uralians

Sadly, some brilliant minds decided in 2015 that the so-called “Yamnaya ancestry” (now more appropriately called “steppe ancestry”) should be associated to ‘Indo-Europeans’. This is causing the development of various new pet theories on the go, as more and more data contradicts this interpretation.

There is a clear long-lasting cultural, populational, and natural barrier between Yamna and Corded Ware: they are derived from different ancestral populations, which show clearly different ancestry and ancestry evolution (although they did converge to some extent), as well as different Y-DNA bottlenecks; they show different cultures, including those of preceding and succeeding groups, and evolved in different ecological niches. The only true steppe pastoralists who managed to dominate over grasslands extending from the Upper Danube to the Altai were Yamna peoples and their cultural successors.

corded-ware-yamna-pca
Corded Ware admixture proper of expanding late Sredni Stog-like populations from the forest-steppe. See full version here.

NOTE. You can also read two recent posts by FrankN in the blog aDNA era, with detailed information on the Pontic-Caspian cultures and the formation of “steppe ancestry” during the Palaeolithic, Mesolithic and Neolithic: How did CHG get into Steppe_EMBA? Part 1: LGM to Early Holocene and How did CHG get into Steppe_EMBA? Part 2: The Pottery Neolithic. Unlike your typical amateur blogger on genetics using few statistical comparisons coupled with ‘archaeolinguoracial mumbo jumbo’ to reach unscientific conclusions, these are obviously carefully redacted texts which deserve to be read.

I will not enter into the discussion of “steppe ancestry” and the mythical “Siberian ancestry” for this post, though. I will just repost the opinion of Volker Heyd – an archaeologist specialized in Yamna Hungary and Bell Beakers who is working with actual geneticists – on the early conclusions based on “steppe ancestry”:

[A]rchaeologist Volker Heyd at the University of Bristol, UK, disagreed, not with the conclusion that people moved west from the steppe, but with how their genetic signatures were conflated with complex cultural expressions. Corded Ware and Yamnaya burials are more different than they are similar, and there is evidence of cultural exchange, at least, between the Russian steppe and regions west that predate Yamnaya culture, he says. None of these facts negates the conclusions of the genetics papers, but they underscore the insufficiency of the articles in addressing the questions that archaeologists are interested in, he argued. “While I have no doubt they are basically right, it is the complexity of the past that is not reflected,” Heyd wrote, before issuing a call to arms. “Instead of letting geneticists determine the agenda and set the message, we should teach them about complexity in past human actions.

Related

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

eneolithic-early-admixture-steppe-ancestry

Given my reduced free time in these months, I have decided to keep updating the text on Indo-European and Uralic migrations and/or this blog, simultaneously or alternatively, to make the most out of the time I can dedicate to this. I will add the different ‘A Song of Sheep and Horses (ASoSaH) reread’ posts to the original post announcing the books. I would be especially interested in comments and corrections to the book chapters rather than the posts, but any comments are welcome (including in the forum, where comments are more likely to stick).

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

Indo-Anatolians and Late Indo-Europeans

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

Luckily enough – for those of us who want precise answers to our previous infinite models of Indo-European language expansions (viz. GAC-associated expansion, IE-speaking Old Europe, Anatolian homeland, Iran homeland, Maykop as Proto-Anatolian, Palaeolithic Continuity Theory, Celtic in the Atlantic façade, etc.) – the situation has been more clear-cut than expected: it turns out that, especially during population expansions, acute Y-chromosome bottlenecks were very common in the past, at least until the Iron Age.

Khvalynsk and Repin-Yamna expansions were no different, and that seems quite natural in hindsight, given the strong familial ties and aversion to foreigners proper of the Late Proto-Indo-European society and culture – probably not really that different from other contemporary societies, like the neighbouring Late Proto-Uralic or Trypillian ones.

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

Y-DNA haplogroups

During the expansion of early Khvalynsk, the most likely Indo-Anatolian culture, the society of the Don-Volga area was probably made up of different lineages including R1b-V1636, R1b-M269, R1a-YP1272, Q1a-M25, and I2a-L699 (and possibly some R1b-V88?), a variability possibly greater than that of the contemporary north Pontic area, probably a sign of this region being a sink of different east and west migrations from steppe and forest areas.

During its expansion, the Khvalynsk society saw its haplogroup variability reduced, as evidenced by the succeeding expansive Repin culture:

Afanasevo, representing Pre-Tocharian (the earliest Late PIE dialect to branch off), expanded with R1b-L23 – especially R1b-Z2103 – lineages, while early Yamna expanded with R1b-L23 and I2a-L699 lineages, which suggests that these are the main haplogroups that survived the Y-DNA bottleneck undergone during the Khvalynsk expansion, and especially later during the late Repin expansion. Nevertheless, other old haplogroups might still pop up during the Repin and early Yamna period, such as the R1b-V1636 sample from Yamna in the Northern Caucasus.

It is still unclear if R1b-L23 sister clade R1b-PF7562 (formed ca. 4400 BC, TMRCA ca. 3400 BC), prevalent among modern Albanians, expanded with Yamna migrants, or if it was part of an earlier expansion of R1b-M269 into the Balkans, and represent thus Indo-Anatolian speakers who later hitchhiked the expansion of the Late PIE language from the north or west Pontic area. The early TMRCA seems to suggest an association with Repin (and therefore Yamna), rather than later movements in the Balkans.

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

‘Yamnaya’ or ‘steppe’ ancestry?

After the early years when population genetics relied mainly on modern Y-DNA haplogroups, geneticists and amateurs have been recently playing around with testing “ancestry percentages”, based on newly developed free statistical tools, which offer obviously just one among many types of data to achieve a proper interpretation of the past.

Today we have quite a lot Y-DNA haplogroups reported for ancient samples of more recent prehistoric periods, and they seem to offer (at least since the 2015 papers, but more evidently since the 2018 papers on Bell Beakers and Europeans, Corded Ware, or Fennoscandia among others) the most straightforward interpretation of all results published in population genomics research.

NOTE. The finding of a specific type of ancestry in one isolated 40,000-year-old sample from Tianyuan can offer very interesting information on potential population movements to the region. However, the identification of ethnolinguistic communities and their migrations among neighbouring groups in Neolithic or Bronze Age groups is evidently not that simple.

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

It is becoming more and more clear with each paper that the true “Yamnaya ancestry” – not the originally described one – was in fact associated with Indo-Europeans (see more on the very Yamnaya-like Yamna Hungary and early East Bell Beaker R1b samples, all of quite similar ancestry and PCA cluster before their further admixture with EEF- and CWC-like groups).

The so-called “steppe ancestry”, on the other hand, reflects the contribution of a Northern Caucasus-related ancestry to expanding Khvalynsk settlers, who spread through the steppes more than a thousand years before the expansion of Late Proto-Indo-Europeans with late Repin, and can thus be found among different groups related to the Pontic-Caspian steppes (see more on the emergence and evolution of “steppe ancestry”).

In fact, after the Yamna/Indo-European and Corded Ware/Uralic expansions, it is more likely to find “steppe ancestry” to the north and east in territories traditionally associated with Uralic languages, whereas to the south and west – i.e. in territories traditionally associated with Indo-European languages – it is more likely to find “EEF ancestry” with diminished “steppe ancestry”, among peoples patrilineally descended from Yamna settlers.

Y-DNA haplogroups, the only uniparental markers (see exceptions in mtDNA inheritance) – unlike ancestry percentages based on the comparison of a few samples and flawed study designs – do not admix, do not change, and therefore they do not lend themselves to infinite pet theories (see e.g. what David Reich has to say about R1b-P312 in Iberia directly derived from Yamna migrants in spite of their predominant EEF ancestry): their cultural continuity can only be challenged with carefully threaded linguistic, archaeological, and genetic data.

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Modern Sardinians show elevated Neolithic farmer ancestry shared with Basques

sardinia-europe-relation

New paper (behind paywall), Genomic history of the Sardinian population, by Chiang et al. Nature Genetics (2018), previously published as a preprint at bioRxiv (2016).

#EDIT (18 Sep 2018): Link to read paper for free shared by the main author.

Interesting excerpts (emphasis mine):

Our analysis of divergence times suggests the population lineage ancestral to modern-day Sardinia was effectively isolated from the mainland European populations ~140–250 generations ago, corresponding to ~4,300–7,000 years ago assuming a generation time of 30 years and a mutation rate of 1.25 × 10−8 per basepair per generation. (…) in terms of relative values, the divergence time between Northern and Southern Europeans is much more recent than either is to Sardinia, signaling the relative isolation of Sardinia from mainland Europe.

We documented fine-scale variation in the ancient population ancestry proportions across the island. The most remote and interior areas of Sardinia—the Gennargentu massif covering the central and eastern regions, including the present-day province of Ogliastra— are thought to have been the least exposed to contact with outside populations. We found that pre-Neolithic hunter-gatherer and Neolithic farmer ancestries are enriched in this region of isolation. Under the premise that Ogliastra has been more buffered from recent immigration to the island, one interpretation of the result is that the early populations of Sardinia were an admixture of the two ancestries, rather than the pre-Neolithic ancestry arriving via later migrations from the mainland. Such admixture could have occurred principally on the island or on the mainland before the hypothesized Neolithic era influx to the island. Under the alternative premise that Ogliastra is simply a highly isolated region that has differentiated within Sardinia due to genetic drift, the result would be interpreted as genetic drift leading to a structured pattern of pre-Neolithic ancestry across the island, in an overall background of high Neolithic ancestry.

sardinia-pca
PCA results of merged Sardinian whole-genome sequences and the HGDP Sardinians. See below for a map of the corresponding regions.

We found Sardinians show a signal of shared ancestry with the Basque in terms of the outgroup f3 shared-drift statistics. This is consistent with long-held arguments of a connection between the two populations, including claims of Basque-like, non-Indo-European words among Sardinian placenames. More recently, the Basque have been shown to be enriched for Neolithic farmer ancestry and Indo-European languages have been associated with steppe population expansions in the post-Neolithic Bronze Age. These results support a model in which Sardinians and the Basque may both retain a legacy of pre-Indo-European Neolithic ancestry. To be cautious, while it seems unlikely, we cannot exclude that the genetic similarity between the Basque and Sardinians is due to an unsampled pre-Neolithic population that has affinities with the Neolithic representatives analyzed here.

density-nuraghi-sardinia-genetics
Left: Geographical map of Sardinia. The provincial boundaries are given as black lines. The provinces are abbreviated as Cag (Cagliari), Cmp (Campidano), Car (Carbonia), Ori (Oristano), Sas (Sassari), Olb (Olbia-tempio), Nuo (Nuoro), and Ogl (Ogliastra). For sampled villages within Ogliastra, the names and abbreviations are indicated in the colored boxes. The color corresponds to the color used in the PCA plot (Fig. 2a). The Gennargentu region referred to in the main text is the mountainous area shown in brown that is centered in western Ogliastra and southeastern Nuoro.
Right: Density of Nuraghi in Sardinia, from Wikipedia.

While we can confirm that Sardinians principally have Neolithic ancestry on the autosomes, the high frequency of two Y-chromosome haplogroups (I2a1a1 at ~39% and R1b1a2 at ~18%) that are not typically affiliated with Neolithic ancestry is one challenge to this model. Whether these haplogroups rose in frequency due to extensive genetic drift and/or reflect sex-biased demographic processes has been an open question. Our analysis of X chromosome versus autosome diversity suggests a smaller effective size for males, which can arise due to multiple processes, including polygyny, patrilineal inheritance rules, or transmission of reproductive success. We also find that the genetic ancestry enriched in Sardinia is more prevalent on the X chromosome than the autosome, suggesting that male lineages may more rapidly trace back to the mainland. Considering that the R1b1a2 haplogroup may be associated with post-Neolithic steppe ancestry expansions in Europe, and the recent timeframe when the R1b1a2 lineages expanded in Sardinia, the patterns raise the possibility of recent male-biased steppe ancestry migration to Sardinia, as has been reported among mainland Europeans at large (though see Lazaridis and Reich and Goldberg et al.). Such a recent influx is difficult to square with the overall divergence of Sardinian populations observed here.

sardinian-admixture
Mixture proportions of the three-component ancestries among Sardinian populations. Using a method first presented in Haak et al. (Nature 522, 207–211, 2015), we computed unbiased estimates of mixture proportions without a parameterized model of relationships between the test populations and the outgroup populations based on f4 statistics. The three-component ancestries were represented by early Neolithic individuals from the LBK culture (LBK_EN), pre-Neolithic huntergatherers (Loschbour), and Bronze Age steppe pastoralists (Yamnaya). See Supplementary Table 5 for standard error estimates computed using a block jackknife.

Once again, haplogroup R1b1a2 (M269), and only R1b1a2, related to male-biased, steppe-related Indo-European migrations…just sayin’.

Interestingly, haplogroup I2a1a1 is actually found among northern Iberians during the Neolithic and Chalcolithic, and is therefore associated with Neolithic ancestry in Iberia, too, and consequently – unless there is a big surprise hidden somewhere – with the ancestry found today among Basques.

NOTE. In fact, the increase in Neolithic ancestry found in south-west Ireland with expanding Bell Beakers (likely Proto-Beakers), coupled with the finding of I2a subclades in Megalithic cultures of western Europe, would support this replacement after the Cardial and Epi-Cardial expansions, which were initially associated with G2a lineages.

I am not convinced about a survival of Palaeo-Sardo after the Bell Beaker expansion, though, since there is no clear-cut cultural divide (and posterior continuity) of pre-Beaker archaeological cultures after the arrival of Bell Beakers in the island that could be identified with the survival of Neolithic languages.

We may have to wait for ancient DNA to show a potential expansion of Neolithic ancestry from the west, maybe associated with the emergence of the Nuragic civilization (potentially linked with contemporaneous Megalithic cultures in Corsica and in the Balearic Islands, and thus with an Iberian rather than a Basque stock), although this is quite speculative at this moment in linguistic, archaeological, and genetic terms.

Nevertheless, it seems that the association of a Basque-Iberian language with the Neolithic expansion from Anatolia (see Villar’s latest book on the subject) is somehow strengthened by this paper. However, it is unclear when, how, and where expanding G2a subclades were replaced by native I2 lineages.

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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.

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