The Lusatian culture, the most likely vector of Balto-Slavic expansions

early-bronze-age-languages-europe

New archaeological paper (behind paywall) New evidence on the southeast Baltic Late Bronze Age agrarian intensification and the earliest AMS dates of Lens culinaris and Vicia faba, by Minkevičius et al. Vegetation History and Archaeobotany (2019).

Interesting excerpts (emphasis mine):

Arrival of farming in the south-east Baltic

The current state of research reveals no firm evidence of crop cultivation in the region before the LBA (Piličiauskas et al. 2017b; Grikpėdis and Motuzaitė-Matuzevičiūtė 2018). Current archaeobotanical data firmly suggest the adoption of farming during the EBA to LBA transition. (…) By comparison, in other parts of N Europe subsistence economy of CWC groups was characterized by strong emphasis on animal husbandry, however crop cultivation was also used (Kirleis 2019; Vanhanen et al. 2019). CWC sites from the Netherlands, Denmark, Sweden and Germany reveal evidence of the cultivation of H. vulgare var. nudum, T. dicoccum, Linum usitatissimum (flax) (Oudemans and Kubiak-Martens 2014; Beckerman 2015; Kubiak- Martens et al. 2015).

It is (…) striking that earliest evidence of farming in the SE Baltic only appears in the deposits dating over 4,000 years later.

The environmental conditions of the SE Baltic presented a significant barrier and numerous genetic adaptations were required before farming could successfully spread into the region (Motuzaitė-Matuzevičiūtė 2018). Adaptations through seasonality changes usually play a major role in adapting to new environments (Sherratt 1980). These include establishing genetic controls on seasonality, especially flowering times and length of growing season (Fuller and Lucas 2017). Therefore, it could be argued that farming was only firmly established in the region around the LBA after several crop species, primarily barley, became adapted to the local environment and the risk of crop failure was reduced (Motuzaitė-Matuzevičiūtė 2018). The transition to farming was further aided by the climate warming which started around 750 cal bc (Gaigalas 2004; Sillasoo et al. 2009). In such a case the fragmented evidence from earlier periods is a likely illustration of the early attempts that have failed.

south-east-baltic-agrarian-communities
Map of sites mentioned in the text: 1 Duba and Palesa Lakes, 2 Šventoji, 3 Šarnelė, 4 Iru, 5 Kvietiniai, 6 Kreiči, 7 Turlojiškė, 8 Narkūnai, 9 Luokesa 1, 10 Mūkakalns, 11 Kivutkalns, 12 Asva, 13 Kukuliškiai

Social change

The LBA agrarian intensification of the SE Baltic was most likely not an isolated case but rather a part of broader social, economic and technological developments sweeping across northern Europe.

Evidence from sites across the Baltic Sea shows that the end of the EBA (ca. 1200 bc onward, after Gustafsson 1998) was marked by intensification of agriculture and changes in landscape management. This coincides with the agricultural developments observed on the SE fringes of the Baltic Sea and provides a context for the eventual arrival of farming, followed shortly by the rapid agrarian intensification of the region. Looking just south from the study region, we see that data from northern Poland reveal a sharp increase in both scale and intensity of agricultural activities during the EBA to LBA transition. Pollen records show significant environmental changes starting around 1400/1300 bc (Wacnik 2005, 2009; Wacnik et al. 2012). These were mostly a result of development of a production economy based on plant cultivation and animal raising. Even more significant changes during this period are visible in southern Scandinavia. Pollen records from S Sweden present evidence for an opening up of the forested landscape and the creation of extensive grasslands (Berglund 1991; Gustafsson 1998). Major changes are also apparent in archaeobotanical assemblages.

In general, during the end of the EBA northern Europe underwent a massive transformation of the farming system moving towards a more intensified agriculture aimed at surplus production. However, this should not be regarded as an isolated occurrence, but rather as a radical change of the whole society which took place throughout Europe (Gustafsson 1998). Intensification of contacts across northern Europe have integrated previously isolated regions into a wider network (Kristiansen and Larsson 2005; Wehlin 2013; Earle et al. 2015). It is therefore likely that farming spread into the SE fringes of the Baltic Sea alongside other innovations including malleable technologies and developments of social structure.

bronze-age-late-baltic
Late Bronze Age cultures in the Baltic. See full map.

The presence and scale of intensifying connections is well illustrated by SE Baltic archaeological material.

Firstly, the appearance of stone ship graves has served as a basis for locating the Nordic communication zones. Construction of such graves was limited to the coastal regions of Kurzeme, Saaremaa Island and the Northern Estonian coast near Tallinn and Kaliningrad (Graudonis 1967; Okulicz 1976; Lang 2007) and is generally regarded as a foreign burial custom which was common in Gotland and along the Scandinavian coast. This is also supported by the Staldzene and Tehumardi hoards (Vasks and Vijups 2004; Sperling 2013), which contained artefacts typical of Nordic culture.

Secondly, studies of early metallurgy and its products, both imported and created in the SE Baltic, have concluded that metal consumption in the LBA had more than doubled compared to the EBA (Sidrys and Luchtanas 1999). The SE Baltic region lacks any metal artefact types exclusive to the region and metal objects are dominated by artefact types originating from Nordic and Lusatian cultures (Sidrys and Luchtanas 1999; Lang 2007; Čivilytė 2014). This indicates that even after metal crafting reached the region, the technology remained exclusively of foreign origin. Rarely identifiable negatives of clay casting moulds were also made for artefacts of Nordic influence, such as Mälar type axes or Härnevi type pins (Čivilytė 2014; Sperling 2014).

Lastly, emerging social diversification was accompanied by the establishment of the first identifiable settlement pattern. Settlement locations were strategically chosen alongside economically significant routes, primarily on the coast and near the Daugava River. Hilltop areas were prioritized over the lowlands, and excavations on these sites have often revealed several stages of enclosure construction (Graudonis 1989). This has also been explained as a reflection of intensifying communication networks between Nordic and Lusatian cultures, and the indigenous communities of the SE Baltic.

Proto-Balto-Slavic

One of the aspects of my description of Balto-Slavic I am least convinced about is my acceptance of Kortlandt’s dialectal classification into Proto-East Baltic, Proto-West Baltic, and Proto-Slavic, due to its strong reliance on his own controversial theory of late laryngeal loss.

Kortlandt’s position regarding Balto-Slavic is that it is in fact simply ‘Proto-Baltic’, a language that would stem thus from an Indo-Baltic branch, which would be originally represented by Corded Ware, and which would have split suddenly in its three dialects without any common development between branches, including some intermediate “Centum” Temematic substrate which explains everything that his model can’t…

As more genetic and archaeological data on northern Europe appears, his ideas about Balto-Slavic are becoming even less credible, fully at odds with his predicted population and cultural movements, in particular because of the evident shaping of Indo-European-speaking Europe through the expansion of the Bell Beaker culture from the Yamnaya of the Carpathian Basin, and of the shaping of Uralic-speaking Europe through the expansion of the Corded Ware culture.

bronze-age-middle-northern-europe
Middle Bronze Age cultures close to the Baltic ca. 1750-1250 BC. See full map.

The site of Turlojiškė in southern Lithuania (ca. 908-485 BC) – which Mittnik et al. (2018) classified as “Bronze Age, Trzciniec culture?” – can be more reasonably considered a settlement of incoming intensive agrarian communities under the influence of the Lusatian culture, like the Narkūnai hilltop settlement in eastern Lithuania (ca. 800–550 BC), or the enclosed hilltop settlement of Kukuliškiai in western Lithuania (ca. 887-506 BC), just 300 m east of the Baltic Sea, also referred to in the paper.

While the dates of sampled individuals include a huge span (ca. 2100-600 BC), those with confirmed radiocarbon dates are more precisely dated to the LBA-EIA transition. More specifically, the first clearly western influence is seen in a sample with an early date (ca. 1230-920 BC), while later samples and samples from Kivutkalns, in Latvia, show major genetic continuity with indigenous populations, compatible with the new chiefdom-based systems of the Baltic and the known lack of massive migrations to the region.

Contacts with western groups of the Nordic Bronze Age and Lusatian cultures intensified – based on existing archaeological and archaeobotanical evidence – in the LBA, especially from ca. 1100/1000 BC on, and Baltic languages seem to have thus little to do with the disappearing Trzciniec culture, and more with the incoming Lusatian influence.

Both facts – more simple dialectalization scheme, and more recent Indo-European expansion to the east – support the spread of Proto-Baltic into the south-east Baltic area precisely around this time, and is also compatible with an internal separation from Proto-Slavic during the expansion of the Lusatian culture.

pca-late-bronze-age-balto-slavic-finnic
Top Left:Likely Baltic, Slavic, and Balto-Finnic-speaking territories (asynchronous), overlaid over Late Bronze Age cultures. Balto-Slavic in green: West(-East?) Baltic (B1), unattested early Baltic (B2), and Slavic (S). Late Balto-Finnic (F) in cyan. In red, Tollense and Turlojiškė sampling. Dashed black line: Balto-Slavic/West Uralic hydrotoponymy border until ca. 1000 AD. Top right: PCA of groups from the Early Bronze Age to the Late Bronze Age. Marked are Iwno/Pre-Trzciniec of Gustorzyn (see below), Late Trzciniec/Iron Age samples from Turlojiškė, and in dashed line approximate extent of Tollense cluster; Y-DNA haplogroups during the Late Bronze Age (Bottom left) and during the Early Iron Age (Bottom right). Notice a majority non-R1a lineages among sampled Early Slavs. See full maps and PCAs.

Even though comparative grammar is traditionally known to be wary of resorting to language contamination or language contact, the truth is that – very much like population genomics – trying to draw a ‘pure’ phylogenetic tree for Balto-Slavic has never worked very well, and the most likely culprit is the Slavic expansion to the south-east into territories which underwent different and complex genetic and linguistic influences for centuries (see here and here).

The close interaction of Nordic BA and Lusatian cultures (and their cultural predominance over) indigenous eastern Baltic peoples from ca. 1100 BC fits (part of) the known intense lexical borrowings of Balto-Finnic from Palaeo-Germanic and from early Proto-Baltic, as well as (part of) the known Germanic–Balto-Slavic contacts, whereas the evident Balto-Finnic-like substrate of Balto-Slavic, and especially of Baltic, must stem from the acculturation of those indigenous East Baltic peoples.

The relative chronology of hydrotoponymy in the East Baltic shows that essentially all ancestral layers to the north of the Daugava must have been Uralic, while roughly south of the Daugava they seem to be mostly Indo-European. The question remains, though, when did this Indo-European layer start?

Despite the many centuries that could separate the attestation of southern place- and river-names from northern ones, Old European is also defined by linguistic traits, which would imply that the same language inferred from Western and Southern European hydrotoponymy is that found in the Baltic, hence all from North-West Indo-European-speaking Bell Beakers and derived Early European Bronze Age groups.

Interestingly, though, it is well-known that some modern Baltic toponyms can’t be easily distinguished from the Old European layers – unlike those of Iberia or the British Isles, which show some attested language change in the proto-historical and historical period – which may imply both (a) continuity of Baltic languages since the EBA, but also that (b) the Baltic naming system is a confounding factor in assessing the ancestral expansion of Old European. The latter is becoming more and more likely with each new linguistic, archaeological, and genetic paper.

up-river
Hydronyms in up-. One among many examples of scarcely attested appellatives that appear inflated in the Baltic due to modern use.

To sum up, a survival of a hypothetical late Trzciniec language in Lithuania or as part of the expanding Lusatian community is not the most economic explanation for what is seen in genetics and archaeology. On the other hand, the cluster formed by the Tollense samples (a site corresponding to the Nordic Bronze Age), the Turlojiškė outlier, and the early Slavs from Bohemia all depict an eastward expansion of Balto-Slavic languages from Central Europe, at the same time as Celtic expanded to the west with the Urnfield culture.

NOTE. For a more detailed discussion of this, see European hydrotoponymy (IV): tug of war between Balto-Slavic and West Uralic.

As I said recently, the slight increase in Corded Ware-like ancestry among Iron Age Estonians, if it were statistically relevant and representative of an incoming population – and not just the product of “usual” admixture with immediate neighbours – need not be from south-eastern Corded Ware groups, because the Akozino-Malär cultural exchange seems to have happened as an interaction in both directions, and not just as an eastward migration imagined by Carpelan and Parpola.

Archaeology and genetics could actually suggest then (at least in part) an admixture with displaced indigenous West Uralic-speaking peoples from the south-west, to the south of the Daugava River, at the same time as the Indo-European – Uralic language frontier must have shifted to its traditional location, precisely during the LBA / EIA transition around 1000 BC.

NOTE. For more on this, see the supplementary materials of Saag et al. (2019).

fortified-settlements-lba-ia
Distribution of fortified settlements (filled circles) and other hilltop sites (empty circles) of the Late Bronze Age and Pre-Roman Iron Ages in the East Baltic region. Tentative area of most intensive contacts between Baltic and Balto-Finnic communities marked with a dashed line. Image modified from (Lang 2016).

The tight relationship of the three communities also accounts for the homogeneous distribution of expanding haplogroup N1c-VL29 (possibly associated with Akozino warrior-traders) in the whole Baltic Sea area, such as those appearing in the Estonian Iron Age samples, which have no clearly defined route(s) of expansion.

It is even possible that they emerged first in the south, linked to marriage alliances of Akozino chieftains with Baltic- and Germanic-speaking chiefdoms around the Baltic Sea (see N1c in Germanic Iron Age), since the expansion of (some) N1c lineages with Gulf of Finland Finnic to the north was more clearly associated with their known bottleneck ca. 2,000 years ago.

Related

More Celts of hg. R1b, more Afanasievo ancestry, more maps

iron-age-early-celtic-expansion

Interesting recent developments:

Celts and hg. R1b

Gauls

Recent paper (behind paywall) Multi-scale archaeogenetic study of two French Iron Age communities: From internal social- to broad-scale population dynamics, by Fischer et al. J Archaeol Sci (2019).

In it, Fischer and colleagues update their previous data for the Y-DNA of Gauls from the Urville-Nacqueville necropolis, Normandy (ca. 300-100 BC), with 8 samples of hg. R, at least 5 of them R1b. They also report new data from the Gallic cemetery at Gurgy ‘Les Noisats’, Southern Paris Basin (ca. 120-80 BC), with 19 samples of hg. R, at least 13 of them R1b.

In both cases, it is likely that both communities belonged (each) to the same paternal lineages, hence the patrilocal residence rules and patrilineality described for Gallic groups, also supported by the different maternal gene pools.

The interesting data would be whether these individuals were of hg. R1b-L21, hence mainly local lineages later replaced or displaced to the west, or – a priori much more likely – of some R1b-U152 and/or R1b-DF27 subclades from Central Europe that became less and less prevalent as Celts expanded into more isolated regions south of the Pyrenees and into the British Isles. Such information is lacking in the paper, probably due to the poor coverage of the samples.

early-iron-age-europe-y-dna
Y-DNA haplogroups in Europe during the Early Iron Age. See full map.

Other Celts

As for early Celts, we already have:

Celtiberians from the Basque Country (one of hg. I2a) and likely Celtic genetic influence in north-east Iberia (all R1b), where Iberian languages spread later, showing that Celts expanded from some place in Central Europe, probably already with the Urnfield culture (ca. 1300 BC on).

Two Hallstatt samples from Bylany, Bohemia (ca. 836-780 BC), by Damgaard et al. Nature (2018), one of them of hg. R1b-U152.

mitterkirchen-grab-hu-i-8-hallstatt
Photo and diagram of burial HÜ-I/8, Mitterkirchen, Oberösterreich, Leskovar 1998.

Another Hallstatt HaC/D1 sample from Mittelkirchen, Austria (ca. 850-650/600), by Kiesslich et al. (2012), with predicted hg. G2a (see Athey’s haplogroup prediction).

One sample of early La Tène culture A from Putzenfeld am Dürrnberg, Hallein, Austria (ca 450–380 BC), by Kiesslich et al. (2012), with predicted hg. R1b (see Athey’s haplogroup prediction).

NOTE. For potential unreliability of haplogroup prediction with Whit Atheys’ haplogroup predictor, see e.g. Zhang et al. (2017).

kelten-dna-putzenfeld-duerrnberg-grab-376
Photo and diagram of Burial 376, Putzenfeld, Dürrnberg bei Hallein, Moser 2007.

Three Britons from Hinxton, South Cambridgeshire (ca. 170 BC – AD 80) from Schiffels et al. (2016), two of them of local hg. R1b-S461.

Indirectly, data of Vikings by Margaryan et al. (2019) from the British Isles and beyond show hg. R1b associated with modern British-like ancestry, also linked to early “Picts”, hence likely associated with Britons even after the Anglo-Saxon settlement. Supporting both (1) my recent prediction of hg. R1b-M167 expanding with Celts and (2) the reason for its presence among modern Scandinavians, is the finding of the first ancient sample of this subclade (VK166) among the Vikings of St John’s College Oxford, associated with the ‘St Brice’s Day Massacre’ (see Margaryan et al. 2019 supplementary materials).

The R1b-M167 sample shows 23.5% British-like ancestry, hence autosomally closer to other local samples (and related to the likely Picts from Orkney) than to some of his deceased partners at the site. Other samples with sizeable British-like ancestry include VK177 (32.6%, hg. R1b-U152), VK173 (33.3%, hg. I2a1b1a), or VK150 (25.6%, hg. I2a1b1a), while typical Germanic subclades like I1 or R1b-U106 – which may be associated with Anglo-Saxons, too – tend to show less.

late-iron-age-europe-y-dna
Y-DNA haplogroups in Europe during the Late Iron Age. See full map.

I remember some commenter asking recently what would happen to the theory of Proto-Indo-European-speaking R1b-rich Yamnaya culture if Celts expanded with hg. R1a, because there were only one hg. R1b and one (possibly) G2a from Hallstatt. As it turns out, they were mostly R1b. However, the increasingly frequent obsession of searching for specific haplogroups and ancestry during the Iron Age and the Middle Ages is weird, even as a desperate attempt, because:

  1. it is evident that the more recent the ancient DNA samples are, the more they are going to resemble modern populations of the same area, so ancient DNA would become essentially useless;
  2. cultures from the early Iron Age onward (and even earlier) were based on increasingly complex sociopolitical systems everywhere, which is reflected in haplogroup and ancestry variability, e.g. among Balts, East Germanic peoples, Slavs (of hg. E1b-V13, I2a-L621), or Tocharians.

In fact, even the finding of hg. R1b among Celts of central and western Europe during the Iron Age is rather unenlightening, because more specific subclades and information on ancestry changes are needed to reach any meaningful conclusion as to migration vs. acculturation waves of expanding Celtic languages, which spread into areas that were mostly Indo-European-speaking since the Bell Beaker expansion.

Afanasevo ancestry in Asia

Wang and colleagues continue to publish interesting analyses, now in the preprint Inland-coastal bifurcation of southern East Asians revealed by Hmong-Mien genomic history, by Xia et al. bioRxiv (2019).

Interesting excerpt (emphasis mine):

Although the Devil’s Cave ancestry is generally the predominant East Asian lineage in North Asia and adjacent areas, there is an intriguing discrepancy between the eastern [Korean, Japanese, Tungusic (except northernmost Oroqen), and Mongolic (except westernmost Kalmyk) speakers] and the western part [West Xiōngnú (~2,150 BP), Tiānshān Hun (~1,500 BP), Turkic-speaking Karakhanid (~1,000 BP) and Tuva, and Kalmyk]. Whereas the East Asian ancestry of populations in the western part has entirely belonged to the Devil’s Cave lineage till now, populations in the eastern part have received the genomic influence from an Amis-related lineage (17.4–52.1%) posterior to the presence of the Devil’s Cave population roughly in the same region (~7,600 BP)12. Analogically, archaeological record has documented the transmission of wet-rice cultivation from coastal China (Shāndōng and/or Liáoníng Peninsula) to Northeast Asia, notably the Korean Peninsula (Mumun pottery period, since ~3,500 BP) and the Japanese archipelago (Yayoi period, since ~2,900 BP)2. Especially for Japanese, the Austronesian-related linguistic influence in Japanese may indicate a potential contact between the Proto-Japonic speakers and population(s) affiliating to the coastal lineage. Thus, our results imply that a southern-East-Asian-related lineage could be arguably associated with the dispersal of wet-rice agriculture in Northeast Asia at least to some extent.

afanasevo-namazga-devils-gate-xiongnu-huns-tianshan-admixture
Spatial and temporal distribution of ancestries in East Asians. Reference populations and corresponding hypothesized ancestral populations: (1) Devil’s Cave (~7,600 BP), the northern East Asian lineage; (2) Amis, the southern East Asian lineage (= AHM + AAA + AAN); (3) Hòabìnhian (~7,900 BP), a lineage related to Andamanese and indigenous hunter-gatherer of MSEA; (4) Kolyma (~9,800 BP), “Ancient Palaeo-Siberians”; (5) Afanasievo (~4,800 BP), steppe ancestry; (6) Namazga (~5,200 BP), the lineage of Chalcolithic Central Asian. Here, we report the best-fitting results of qpAdm based on following criteria: (1) a feasible p-value (&mt; 0.05), (2) feasible proportions of all the ancestral components (mean &mt; 0 and standard error < mean), and (3) with the highest p-value if meeting previous conditions.

In this case, the study doesn’t compare Steppe_MLBA, though, so the findings of Afanasievo ancestry have to be taken with a pinch of salt. They are, however, compared to Namazga, so “Steppe ancestry” is there. Taking into account the limited amount of Yamnaya-like ancestry that could have reached the Tian Shan area with the Srubna-Andronovo horizon in the Iron Age (see here), and the amount of Yamnaya-like ancestry that appears in some of these populations, it seems unlikely that this amount of “Steppe ancestry” would emerge as based only on Steppe_MLBA, hence the most likely contacts of Turkic peoples with populations of both Afanasievo (first) and Corded Ware-derived ancestry (later) to the west of Lake Baikal.

(1) The simplification of ancestral components into A vs. B vs. C… (when many were already mixed), and (2) the simplistic selection of one OR the other in the preferred models (such as those published for Yamnaya or Corded Ware), both common strategies in population genomics pose evident problems when assessing the actual gene flow from some populations into others.

Also, it seems that when the “Steppe”-like contribution is small, both Yamnaya and Corded Ware ancestry will be good fits in admixed populations of Central Asia, due to the presence of peoples of EHG-like (viz. West Siberia HG) and/or CHG-like (viz. Namazga) ancestry in the area. Unless and until these problems are addressed, there is little that can be confidently said about the history of Yamnaya vs. Corded Ware admixture among Asian peoples.

Maps, maps, and more maps

As you have probably noticed if you follow this blog regularly, I have been experimenting with GIS software in the past month or so, trying to map haplogroups and ancestry components (see examples for Vikings, Corded Ware, and Yamnaya). My idea was to show the (pre)historical evolution of ancestry and haplogroups coupled with the atlas of prehistoric migrations, but I have to understand first what I can do with GIS statistical tools.

My latest exercise has been to map modern haplogroup distribution (now added to the main menu above) using data from the latest available reports. While there have been no great surprises – beyond the sometimes awful display of data by some papers – I think it is becoming clearer with each new publication how wrong it was for geneticists to target initially those populations considered “isolated” – hence subject to strong founder effects – to extrapolate language relationships. For example:

  • The mapping of R1b-M269, in particular basal subclades, corresponds nicely with the Indo-European expansions.
  • There is no clear relationship of R1b, not even R1b-DF27 (especially basal subclades), with Basques. There is no apparent relationship between the distribution of R1b-M269 and some mythical non-Indo-European “Old Europeans”, like Etruscans or Caucasian speakers, either.
  • Basal R1a-M417 shows an interesting distribution, as do maps of basal Z282 and Z93 subclades, despite the evident late bottlenecks and acculturation among Slavs.
  • The distribution of hg. N1a-VL29 (and other N1a-L392 subclades) is clearly dissociated from Uralic peoples, and their expansion in the whole Baltic Sea during the Iron Age doesn’t seem to be related to any specific linguistic expansion.
  • haplogroup-n1a-vl29
    Modern distribution of haplogroup N1a-VL29. See full map.
  • Even the most recent association in Post et al. (2019) with hg. N1a-Z1639 – due to the lack of relationship of Uralic with N1a-VL29 – seems like a stretch, seeing how it probably expanded from the Kola Peninsula and the East Urals, and neither the Lovozero Ware nor forest hunter-fishers of the Cis- and Trans-Urals regions were Uralic-speaking cultures.
  • The current prevalence of hg. R1b-M73 supports its likely expansion with Turkic-speaking peoples.
  • The distribution of haplogroup R1b-V88 in Africa doesn’t look like it was a mere founder effect in Chadic peoples – although they certainly underwent a bottleneck under it.
  • The distribution of R1a-M420 (xM198) and hg. R1b-M343 (possibly not fully depicted in the east) seem to be related to expansions close to the Caucasus, supporting once more their location in Eastern Europe / West Siberia during the Mesolithic.
  • The mapping of E1b-V13 and I-M170 (I haven’t yet divided it into subclades) are particularly relevant for the recent eastward expansion of early Slavic peoples.

All in all, modern haplogroup distribution might have been used to ascertain prehistoric language movements even in the 2000s. It was the obsession with (and the wrong assumptions about) the “purity” of certain populations – say, Basques or Finns – what caused many of the interpretation problems and circular reasoning we are still seeing today.

I have also updated maps of Y-chromosome haplogroups reported for ancient samples in Europe and/or West Eurasia for the Early Eneolithic, Early Chalcolithic, Late Chalcolithic, Early Bronze Age, Middle Bronze Age, Late Bronze Age, Early Iron Age, Late Iron Age, Antiquity, and Middle Ages.

Haplogroup inference

I have also tried Yleaf v.2 – which seems like an improvement over the infamous v.1 – to test some samples that hobbyists and/or geneticists have reported differently in the past. I have posted the results in this ancient DNA haplogroup page. It doesn’t mean that the inferences I obtain are the correct ones, but now you have yet another source to compare.

Not many surprises here, either:

  • M15-1 and M012, two Proto-Tocharians from Shirenzigou, are of hg. R1b-PH155, not R1b-M269.
  • I0124, the Samara HG, is of hg. R1b-P297, but uncertain for both R1b-M73 and R1b-M269.
  • I0122, the Khvalynsk chieftain, is of hg. R1b-V1636.
  • I2181, the Smyadovo outlier of poor coverage, is possibly of hg. R, and could be of hg. R1b-M269, but could also be even non-P.
  • I6561 from Alexandria is probably of hg. R1a-M417, likely R1a-Z645, maybe R1a-Z93, but can’t be known beyond that, which is more in line with the TMRCA of R1a subclades and the radiocarbon date of the sample.
  • I2181, the Yamnaya individual (supposedly Pre-R1b-L51) at Lopatino II is R1b-M269, negative for R1b-L51. Nothing beyond that.

You can ask me to try mapping more data or to test the haplogroup of more samples, provided you give me a proper link to the relevant data, they are interesting for the subject of this blog…and I have the time to do it.

Related

Yamnaya ancestry: mapping the Proto-Indo-European expansions

steppe-ancestry-expansion-europe

The latest papers from Ning et al. Cell (2019) and Anthony JIES (2019) have offered some interesting new data, supporting once more what could be inferred since 2015, and what was evident in population genomics since 2017: that Proto-Indo-Europeans expanded under R1b bottlenecks, and that the so-called “Steppe ancestry” referred to two different components, one – Yamnaya or Steppe_EMBA ancestry – expanding with Pro-Indo-Europeans, and the other one – Corded Ware or Steppe_MLBA ancestry – expanding with Uralic speakers.

The following maps are based on formal stats published in the papers and supplementary materials from 2015 until today, mainly on Wang et al. (2018 & 2019), Mathieson et al. (2018) and Olalde et al. (2018), and others like Lazaridis et al. (2016), Lazaridis et al. (2017), Mittnik et al. (2018), Lamnidis et al. (2018), Fernandes et al. (2018), Jeong et al. (2019), Olalde et al. (2019), etc.

NOTE. As in the Corded Ware ancestry maps, the selected reports in this case are centered on the prototypical Yamnaya ancestry vs. other simplified components, so everything else refers to simplistic ancestral components widespread across populations that do not necessarily share any recent connection, much less a language. In fact, most of the time they clearly didn’t. They can be interpreted as “EHG that is not part of the Yamnaya component”, or “CHG that is not part of the Yamnaya component”. They can’t be read as “expanding EHG people/language” or “expanding CHG people/language”, at least no more than maps of “Steppe ancestry” can be read as “expanding Steppe people/language”. Also, remember that I have left the default behaviour for color classification, so that the highest value (i.e. 1, or white colour) could mean anything from 10% to 100% depending on the specific ancestry and period; that’s what the legend is for… But, fere libenter homines id quod volunt credunt.

Sections:

  1. Neolithic or the formation of Early Indo-European
  2. Eneolithic or the expansion of Middle Proto-Indo-European
  3. Chalcolithic / Early Bronze Age or the expansion of Late Proto-Indo-European
  4. European Early Bronze Age and MLBA or the expansion of Late PIE dialects

1. Neolithic

Anthony (2019) agrees with the most likely explanation of the CHG component found in Yamnaya, as derived from steppe hunter-fishers close to the lower Volga basin. The ultimate origin of this specific CHG-like component that eventually formed part of the Pre-Yamnaya ancestry is not clear, though:

The hunter-fisher camps that first appeared on the lower Volga around 6200 BC could represent the migration northward of un-admixed CHG hunter-fishers from the steppe parts of the southeastern Caucasus, a speculation that awaits confirmation from aDNA.

neolithic-chg-ancestry
Natural neighbor interpolation of CHG ancestry among Neolithic populations. See full map.

The typical EHG component that formed part eventually of Pre-Yamnaya ancestry came from the Middle Volga Basin, most likely close to the Samara region, as shown by the sampled Samara hunter-gatherer (ca. 5600-5500 BC):

After 5000 BC domesticated animals appeared in these same sites in the lower Volga, and in new ones, and in grave sacrifices at Khvalynsk and Ekaterinovka. CHG genes and domesticated animals flowed north up the Volga, and EHG genes flowed south into the North Caucasus steppes, and the two components became admixed.

neolithic-ehg-ancestry
Natural neighbor interpolation of EHG ancestry among Neolithic populations. See full map.

To the west, in the Dnieper-Dniester area, WHG became the dominant ancestry after the Mesolithic, at the expense of EHG, revealing a likely mating network reaching to the north into the Baltic:

Like the Mesolithic and Neolithic populations here, the Eneolithic populations of Dnieper-Donets II type seem to have limited their mating network to the rich, strategic region they occupied, centered on the Rapids. The absence of CHG shows that they did not mate frequently if at all with the people of the Volga steppes (…)

neolithic-whg-ancestry
Natural neighbor interpolation of WHG ancestry among Neolithic populations. See full map.

North-West Anatolia Neolithic ancestry, proper of expanding Early European farmers, is found up to border of the Dniester, as Anthony (2007) had predicted.

neolithic-anatolia-farmer-ancestry
Natural neighbor interpolation of Anatolia Neolithic ancestry among Neolithic populations. See full map.

2. Eneolithic

From Anthony (2019):

After approximately 4500 BC the Khvalynsk archaeological culture united the lower and middle Volga archaeological sites into one variable archaeological culture that kept domesticated sheep, goats, and cattle (and possibly horses). In my estimation, Khvalynsk might represent the oldest phase of PIE.

(…) this middle Volga mating network extended down to the North Caucasian steppes, where at cemeteries such as Progress-2 and Vonyuchka, dated 4300 BC, the same Khvalynsk-type ancestry appeared, an admixture of CHG and EHG with no Anatolian Farmer ancestry, with steppe-derived Y-chromosome haplogroup R1b. These three individuals in the North Caucasus steppes had higher proportions of CHG, overlapping Yamnaya. Without any doubt, a CHG population that was not admixed with Anatolian Farmers mated with EHG populations in the Volga steppes and in the North Caucasus steppes before 4500 BC. We can refer to this admixture as pre-Yamnaya, because it makes the best currently known genetic ancestor for EHG/CHG R1b Yamnaya genomes.

From Wang et al (2019):

Three individuals from the sites of Progress 2 and Vonyuchka 1 in the North Caucasus piedmont steppe (‘Eneolithic steppe’), which harbour EHG and CHG related ancestry, are genetically very similar to Eneolithic individuals from Khvalynsk II and the Samara region. This extends the cline of dilution of EHG ancestry via CHG-related ancestry to sites immediately north of the Caucasus foothills

eneolithic-pre-yamnaya-ancestry
Natural neighbor interpolation of Pre-Yamnaya ancestry among Neolithic populations. See full map. This map corresponds roughly to the map of Khvalynsk-Novodanilovka expansion, and in particular to the expansion of horse-head pommel-scepters (read more about Khvalynsk, and specifically about horse symbolism)

NOTE. Unpublished samples from Ekaterinovka have been previously reported as within the R1b-L23 tree. Interestingly, although the Varna outlier is a female, the Balkan outlier from Smyadovo shows two positive SNP calls for hg. R1b-M269. However, its poor coverage makes its most conservative haplogroup prediction R-M343.

The formation of this Pre-Yamnaya ancestry sets this Volga-Caucasus Khvalynsk community apart from the rest of the EHG-like population of eastern Europe.

eneolithic-ehg-ancestry
Natural neighbor interpolation of non-Pre-Yamnaya EHG ancestry among Eneolithic populations. See full map.

Anthony (2019) seems to rely on ADMIXTURE graphics when he writes that the late Sredni Stog sample from Alexandria shows “80% Khvalynsk-type steppe ancestry (CHG&EHG)”. While this seems the most logical conclusion of what might have happened after the Suvorovo-Novodanilovka expansion through the North Pontic steppes (see my post on “Steppe ancestry” step by step), formal stats have not confirmed that.

In fact, analyses published in Wang et al. (2019) rejected that Corded Ware groups are derived from this Pre-Yamnaya ancestry, a reality that had been already hinted in Narasimhan et al. (2018), when Steppe_EMBA showed a poor fit for expanding Srubna-Andronovo populations. Hence the need to consider the whole CHG component of the North Pontic area separately:

eneolithic-chg-ancestry
Natural neighbor interpolation of non-Pre-Yamnaya CHG ancestry among Eneolithic populations. See full map. You can read more about population movements in the late Sredni Stog and closer to the Proto-Corded Ware period.

NOTE. Fits for WHG + CHG + EHG in Neolithic and Eneolithic populations are taken in part from Mathieson et al. (2019) supplementary materials (download Excel here). Unfortunately, while data on the Ukraine_Eneolithic outlier from Alexandria abounds, I don’t have specific data on the so-called ‘outlier’ from Dereivka compared to the other two analyzed together, so these maps of CHG and EHG expansion are possibly showing a lesser distribution to the west than the real one ca. 4000-3500 BC.

eneolithic-whg-ancestry
Natural neighbor interpolation of WHG ancestry among Eneolithic populations. See full map.

Anatolia Neolithic ancestry clearly spread to the east into the north Pontic area through a Middle Eneolithic mating network, most likely opened after the Khvalynsk expansion:

eneolithic-anatolia-farmer-ancestry
Natural neighbor interpolation of Anatolia Neolithic ancestry among Eneolithic populations. See full map.
eneolithic-iran-chl-ancestry
Natural neighbor interpolation of Iran Chl. ancestry among Eneolithic populations. See full map.

Regarding Y-chromosome haplogroups, Anthony (2019) insists on the evident association of Khvalynsk, Yamnaya, and the spread of Pre-Yamnaya and Yamnaya ancestry with the expansion of elite R1b-L754 (and some I2a2) individuals:

eneolithic-early-y-dna
Y-DNA haplogroups in West Eurasia during the Early Eneolithic in the Pontic-Caspian steppes. See full map, and see culture, ADMIXTURE, Y-DNA, and mtDNA maps of the Early Eneolithic and Late Eneolithic.

3. Early Bronze Age

Data from Wang et al. (2019) show that Corded Ware-derived populations do not have good fits for Eneolithic_Steppe-like ancestry, no matter the model. In other words: Corded Ware populations show not only a higher contribution of Anatolia Neolithic ancestry (ca. 20-30% compared to the ca. 2-10% of Yamnaya); they show a different EHG + CHG combination compared to the Pre-Yamnaya one.

eneolithic-steppe-best-fits
Supplementary Table 13. P values of rank=2 and admixture proportions in modelling Steppe ancestry populations as a three-way admixture of Eneolithic steppe Anatolian_Neolithic and WHG using 14 outgroups.
Left populations: Test, Eneolithic_steppe, Anatolian_Neolithic, WHG.
Right populations: Mbuti.DG, Ust_Ishim.DG, Kostenki14, MA1, Han.DG, Papuan.DG, Onge.DG, Villabruna, Vestonice16, ElMiron, Ethiopia_4500BP.SG, Karitiana.DG, Natufian, Iran_Ganj_Dareh_Neolithic.

Yamnaya Kalmykia and Afanasievo show the closest fits to the Eneolithic population of the North Caucasian steppes, rejecting thus sizeable contributions from Anatolia Neolithic and/or WHG, as shown by the SD values. Both probably show then a Pre-Yamnaya ancestry closest to the late Repin population.

wang-eneolithic-steppe-caucasus-yamnaya
Modelling results for the Steppe and Caucasus cluster. Admixture proportions based on (temporally and geographically) distal and proximal models, showing additional AF ancestry in Steppe groups and additional gene flow from the south in some of the Steppe groups as well as the Caucasus groups. See tables above. Modified from Wang et al. (2019). Within a blue square, Yamnaya-related groups; within a cyan square, Corded Ware-related groups. Green background behind best p-values. In red circle, SD of AF/WHG ancestry contribution in Afanasevo and Yamnaya Kalmykia, with ranges that almost include 0%.

EBA maps include data from Wang et al. (2018) supplementary materials, specifically unpublished Yamnaya samples from Hungary that appeared in analysis of the preprint, but which were taken out of the definitive paper. Their location among Yamnaya settlers from Hungary is speculative, although most uncovered kurgans in Hungary are concentrated in the Tisza-Danube interfluve.

eba-yamnaya-ancestry
Natural neighbor interpolation of Pre-Yamnaya ancestry among Early Bronze Age populations. See full map. This map corresponds roughly with the known expansion of late Repin/Yamnaya settlers.

The Y-chromosome bottleneck of elite males from Proto-Indo-European clans under R1b-L754 and some I2a2 subclades, already visible in the Khvalynsk sampling, became even more noticeable in the subsequent expansion of late Repin/early Yamnaya elites under R1b-L23 and I2a-L699:

chalcolithic-early-y-dna
Y-DNA haplogroups in West Eurasia during the Yamnaya expansion. See full map and maps of cultures, ADMIXTURE, Y-DNA, and mtDNA of the Early Chalcolithic and Yamnaya Hungary.

Maps of CHG, EHG, Anatolia Neolithic, and probably WHG show the expansion of these components among Corded Ware-related groups in North Eurasia, apart from other cultures close to the Caucasus:

NOTE. For maps with actual formal stats of Corded Ware ancestry from the Early Bronze Age to the modern times, you can read the post Corded Ware ancestry in North Eurasia and the Uralic expansion.

eba-chg-ancestry
Natural neighbor interpolation of non-Pre-Yamnaya CHG ancestry among Early Bronze Age populations. See full map.
eba-ehg-ancestry
Natural neighbor interpolation of non-Pre-Yamnaya EHG ancestry among Early Bronze Age populations. See full map.
eba-whg-ancestry
Natural neighbor interpolation of WHG ancestry among Early Bronze Age populations. See full map.
eba-anatolia-farmer-ancestry
Natural neighbor interpolation of Anatolia Neolithic ancestry among Early Bronze Age populations. See full map.
eba-iran-chl-ancestry
Natural neighbor interpolation of Iran Chl. ancestry among Early Bronze Age populations. See full map.

4. Middle to Late Bronze Age

The following maps show the most likely distribution of Yamnaya ancestry during the Bell Beaker-, Balkan-, and Sintashta-Potapovka-related expansions.

4.1. Bell Beakers

The amount of Yamnaya ancestry is probably overestimated among populations where Bell Beakers replaced Corded Ware. A map of Yamnaya ancestry among Bell Beakers gets trickier for the following reasons:

  • Expanding Repin peoples of Pre-Yamnaya ancestry must have had admixture through exogamy with late Sredni Stog/Proto-Corded Ware peoples during their expansion into the North Pontic area, and Sredni Stog in turn had probably some Pre-Yamnaya admixture, too (although they don’t appear in the simplistic formal stats above). This is supported by the increase of Anatolia farmer ancestry in more western Yamna samples.
  • Later, Yamnaya admixed through exogamy with Corded Ware-like populations in Central Europe during their expansion. Even samples from the Middle to Upper Danube and around the Lower Rhine will probably show increasing contributions of Steppe_MLBA, at the same time as they show an increasing proportion of EEF-related ancestry.
  • To complicate things further, the late Corded Ware Espersted family (from ca. 2500 BC or later) shows, in turn, what seems like a recent admixture with Yamnaya vanguard groups, with the sample of highest Yamnaya ancestry being the paternal uncle of other individuals (all of hg. R1a-M417), suggesting that there might have been many similar Central European mating networks from the mid-3rd millennium BC on, of (mainly) Yamnaya-like R1b elites displaying a small proportion of CW-like ancestry admixing through exogamy with Corded Ware-like peoples who already had some Yamnaya ancestry.
mlba-yamnaya-ancestry
Natural neighbor interpolation of Yamnaya ancestry among Middle to Late Bronze Age populations (Esperstedt CWC site close to BK_DE, label is hidden by BK_DE_SAN). See full map. You can see how this map correlated with the map of Late Copper Age migrations and Yamanaya into Bell Beaker expansion.

NOTE. Terms like “exogamy”, “male-driven migration”, and “sex bias”, are not only based on the Y-chromosome bottlenecks visible in the different cultural expansions since the Palaeolithic. Despite the scarce sampling available in 2017 for analysis of “Steppe ancestry”-related populations, it appeared to show already a male sex bias in Goldberg et al. (2017), and it has been confirmed for Neolithic and Copper Age population movements in Mathieson et al. (2018) – see Supplementary Table 5. The analysis of male-biased expansion of “Steppe ancestry” in CWC Esperstedt and Bell Beaker Germany is, for the reasons stated above, not very useful to distinguish their mutual influence, though.

Based on data from Olalde et al. (2019), Bell Beakers from Germany are the closest sampled ones to expanding East Bell Beakers, and those close to the Rhine – i.e. French, Dutch, and British Beakers in particular – show a clear excess “Steppe ancestry” due to their exogamy with local Corded Ware groups:

Only one 2-way model fits the ancestry in Iberia_CA_Stp with P-value>0.05: Germany_Beaker + Iberia_CA. Finding a Bell Beaker-related group as a plausible source for the introduction of steppe ancestry into Iberia is consistent with the fact that some of the individuals in the Iberia_CA_Stp group were excavated in Bell Beaker associated contexts. Models with Iberia_CA and other Bell Beaker groups such as France_Beaker (P-value=7.31E-06), Netherlands_Beaker (P-value=1.03E-03) and England_Beaker (P-value=4.86E-02) failed, probably because they have slightly higher proportions of steppe ancestry than the true source population.

olalde-iberia-chalcolithic

The exogamy with Corded Ware-like groups in the Lower Rhine Basin seems at this point undeniable, as is the origin of Bell Beakers around the Middle-Upper Danube Basin from Yamnaya Hungary.

To avoid this excess “Steppe ancestry” showing up in the maps, since Bell Beakers from Germany pack the most Yamnaya ancestry among East Bell Beakers outside Hungary (ca. 51.1% “Steppe ancestry”), I equated this maximum with BK_Scotland_Ach (which shows ca. 61.1% “Steppe ancestry”, highest among western Beakers), and applied a simple rule of three for “Steppe ancestry” in Dutch and British Beakers.

NOTE. Formal stats for “Steppe ancestry” in Bell Beaker groups are available in Olalde et al. (2018) supplementary materials (PDF). I didn’t apply this adjustment to Bk_FR groups because of the R1b Bell Beaker sample from the Champagne/Alsace region reported by Samantha Brunel that will pack more Yamnaya ancestry than any other sampled Beaker to date, hence probably driving the Yamnaya ancestry up in French samples.

The most likely outcome in the following years, when Yamnaya and Corded Ware ancestry are investigated separately, is that Yamnaya ancestry will be much lower the farther away from the Middle and Lower Danube region, similar to the case in Iberia, so the map above probably overestimates this component in most Beakers to the north of the Danube. Even the late Hungarian Beaker samples, who pack the highest Yamnaya ancestry (up to 75%) among Beakers, represent likely a back-migration of Moravian Beakers, and will probably show a contribution of Corded Ware ancestry due to the exogamy with local Moravian groups.

Despite this decreasing admixture as Bell Beakers spread westward, the explosive expansion of Yamnaya R1b male lineages (in words of David Reich) and the radical replacement of local ones – whether derived from Corded Ware or Neolithic groups – shows the true extent of the North-West Indo-European expansion in Europe:

chalcolithic-late-y-dna
Y-DNA haplogroups in West Eurasia during the Bell Beaker expansion. See full map and see maps of cultures, ADMIXTURE, Y-DNA, and mtDNA of the Late Copper Age and of the Yamnaya-Bell Beaker transition.

4.2. Palaeo-Balkan

There is scarce data on Palaeo-Balkan movements yet, although it is known that:

  1. Yamnaya ancestry appears among Mycenaeans, with the Yamnaya Bulgaria sample being its best current ancestral fit;
  2. the emergence of steppe ancestry and R1b-M269 in the eastern Mediterranean was associated with Ancient Greeks;
  3. Thracians, Albanians, and Armenians also show R1b-M269 subclades and “Steppe ancestry”.

4.3. Sintashta-Potapovka-Filatovka

Interestingly, Potapovka is the only Corded Ware derived culture that shows good fits for Yamnaya ancestry, despite having replaced Poltavka in the region under the same Corded Ware-like (Abashevo) influence as Sintashta.

This proves that there was a period of admixture in the Pre-Proto-Indo-Iranian community between CWC-like Abashevo and Yamnaya-like Catacomb-Poltavka herders in the Sintashta-Potapovka-Filatovka community, probably more easily detectable in this group because of the specific temporal and geographic sampling available.

srubnaya-yamnaya-ehg-chg-ancestry
Supplementary Table 14. P values of rank=3 and admixture proportions in modelling Steppe ancestry populations as a four-way admixture of distal sources EHG, CHG, Anatolian_Neolithic and WHG using 14 outgroups.
Left populations: Steppe cluster, EHG, CHG, WHG, Anatolian_Neolithic
Right populations: Mbuti.DG, Ust_Ishim.DG, Kostenki14, MA1, Han.DG, Papuan.DG, Onge.DG, Villabruna, Vestonice16, ElMiron, Ethiopia_4500BP.SG, Karitiana.DG, Natufian, Iran_Ganj_Dareh_Neolithic.

Srubnaya ancestry shows a best fit with non-Pre-Yamnaya ancestry, i.e. with different CHG + EHG components – possibly because the more western Potapovka (ancestral to Proto-Srubnaya Pokrovka) also showed good fits for it. Srubnaya shows poor fits for Pre-Yamnaya ancestry probably because Corded Ware-like (Abashevo) genetic influence increased during its formation.

On the other hand, more eastern Corded Ware-derived groups like Sintashta and its more direct offshoot Andronovo show poor fits with this model, too, but their fits are still better than those including Pre-Yamnaya ancestry.

mlba-ehg-ancestry
Natural neighbor interpolation of non-Pre-Yamnaya EHG ancestry among Middle to Late Bronze Age populations. See full map.
mlba-chg-ancestry
Natural neighbor interpolation of non-Pre-Yamnaya CHG ancestry among Middle to Late Bronze Age populations. See full map.
mlba-anatolia-farmer-ancestry
Natural neighbor interpolation of Anatolia Neolithic ancestry among Middle to Late Bronze Age populations. See full map.
mlba-iran-chl-ancestry
Natural neighbor interpolation of Iran Chl. ancestry among Middle to Late Bronze Age populations. See full map.

NOTE For maps with actual formal stats of Corded Ware ancestry from the Early Bronze Age to the modern times, you should read the post Corded Ware ancestry in North Eurasia and the Uralic expansion instead.

The bottleneck of Proto-Indo-Iranians under R1a-Z93 was not yet complete by the time when the Sintashta-Potapovka-Filatovka community expanded with the Srubna-Andronovo horizon:

early-bronze-age-y-dna
Y-DNA haplogroups in West Eurasia during the European Early Bronze Age. See full map and see maps of cultures, ADMIXTURE, Y-DNA, and mtDNA of the Early Bronze Age.

4.4. Afanasevo

At the end of the Afanasevo culture, at least three samples show hg. Q1a2-M25 (ca. 2900-2500 BC), which seemed to point to a resurgence of local lineages, despite continuity of the prototypical Pre-Yamnaya ancestry. On the other hand, Anthony (2019) makes this cryptic statement:

Yamnaya men were almost exclusively R1b, and pre-Yamnaya Eneolithic Volga-Caspian-Caucasus steppe men were principally R1b, with a significant Q1a minority.

Since the only available samples from the Khvalynsk community are R1b (x3), Q1a(x1), and R1a(x1), it seems strange that Anthony would talk about a “significant minority”, unless Q1a will pop up in some more individuals of those ca. 30 new to be published. Because he also mentions I2a2 as appearing in one elite burial, it seems Q1a (like R1a-M459) will not appear under elite kurgans, although it is still possible that hg. Q1a was involved in the expansion of Afanasevo to the east.

middle-bronze-age-y-dna
Y-DNA haplogroups in West Eurasia during the Middle Bronze Age. See full map and see maps of cultures, ADMIXTURE, Y-DNA, and mtDNA of the Middle Bronze Age and the Late Bronze Age.

Okunevo, which replaced Afanasevo in the Altai region, shows a majority of hg. Q1a2-M25, and at least one Q1a1-B284, but also some R1b-M269 samples proper of Afanasevo, suggesting partial genetic continuity.

NOTE. Other sampled Siberian populations clearly show a variety of Q subclades that likely expanded during the Palaeolithic, such as Baikal EBA samples from Ust’Ida and Shamanka with a majority of Q1a2-M25 (in particular Q1a2-L712), and hg. Q reported from Elunino, Sagsai, Khövsgöl, and also among peoples of the Srubna-Andronovo horizon (the Krasnoyarsk MLBA outlier), and in Karasuk. Q1a-M25 was earlier found in a Baltic hunter-gatherer, which supports a widespread distribution of Q1a2 and Q1a1 in North Eurasia during the Neolithic and Bronze Age.

From Damgaard et al. Science (2018):

(…) in contrast to the lack of identifiable admixture from Yamnaya and Afanasievo in the CentralSteppe_EMBA, there is an admixture signal of 10 to 20% Yamnaya and Afanasievo in the Okunevo_EMBA samples, consistent with evidence of western steppe influence. This signal is not seen on the X chromosome (qpAdm P value for admixture on X 0.33 compared to 0.02 for autosomes), suggesting a male-derived admixture, also consistent with the fact that 1 of 10 Okunevo_EMBA males carries a R1b1a2a2 Y chromosome related to those found in western pastoralists. In contrast, there is no evidence of western steppe admixture among the more eastern Baikal region region Bronze Age (~2200 to 1800 BCE) samples.

This Yamnaya ancestry has been also recently found to be the best fit for the Iron Age population of Shirenzigou in Xinjiang – where Tocharian languages were attested centuries later – despite the haplogroup diversity acquired during their evolution, likely through an intermediate Chemurchek culture (see a recent discussion on the elusive Proto-Tocharians).

Haplogroup diversity seems to be common in Iron Age populations all over Eurasia, most likely due to the spread of different types of sociopolitical structures where alliances played a more relevant role in the expansion of peoples. A well-known example of this is the spread of Akozino warrior-traders in the whole Baltic region under a partial N1a-VL29-bottleneck associated with the emerging chiefdom-based systems under the influence of expanding steppe nomads.

early-iron-age-y-dna
Y-DNA haplogroups in West Eurasia during the Early Iron Age. See full map and see maps of cultures, ADMIXTURE, Y-DNA, and mtDNA of the Early Iron Age and Late Iron Age.

Surprisingly, then, Proto-Tocharians from Shirenzigou pack up to 74% Yamnaya ancestry, in spite of the 2,000 years that separate them from the demise of the Afanasevo culture. They show more Yamnaya ancestry than any other population by that time, being thus a sort of Late PIE fossils not only in their archaic dialect, but also in their genetic profile:

shirenzigou-afanasievo-yamnaya-andronovo-srubna-ulchi-han

The recent intrusion of Corded Ware-like ancestry, as well as the variable admixture with Siberian and East Asian populations, both point to the known intense Old Iranian and Old/Middle Chinese contacts. The scarce Proto-Samoyedic and Proto-Turkic loans in Tocharian suggest a rather loose, probably more distant connection with East Uralic and Altaic peoples from the forest-steppe and steppe areas to the north (read more about external influences on Tocharian).

Interestingly, both R1b samples, MO12 and M15-2 – likely of Asian R1b-PH155 branch – show a best fit for Andronovo/Srubna + Hezhen/Ulchi ancestry, suggesting a likely connection with Iranians to the east of Xinjiang, who later expanded as the Wusun and Kangju. How they might have been related to Huns and Xiongnu individuals, who also show this haplogroup, is yet unknown, although Huns also show hg. R1a-Z93 (probably most R1a-Z2124) and Steppe_MLBA ancestry, earlier associated with expanding Iranian peoples of the Srubna-Andronovo horizon.

All in all, it seems that prehistoric movements explained through the lens of genetic research fit perfectly well the linguistic reconstruction of Proto-Indo-European and Proto-Uralic.

Related

Volga Basin R1b-rich Proto-Indo-Europeans of (Pre-)Yamnaya ancestry

yamnaya-expansion

New paper (behind paywall) by David Anthony, Archaeology, Genetics, and Language in the Steppes: A Comment on Bomhard, complementing in a favourable way Bomhard’s Caucasian substrate hypothesis in the current issue of the JIES.

NOTE. I have tried to access this issue for some days, but it’s just not indexed in my university library online service (ProQuest) yet. This particular paper is on Academia.edu, though, as are Bomhard’s papers on this issue in his site.

Interesting excerpts (emphasis mine):

Along the banks of the lower Volga many excavated hunting-fishing camp sites are dated 6200-4500 BC. They could be the source of CHG ancestry in the steppes. At about 6200 BC, when these camps were first established at Kair Shak III and Varfolomievka (42 and 28 on Figure 2), they hunted primarily saiga antelope around Dzhangar, south of the lower Volga, and almost exclusively onagers in the drier desert-steppes at Kair-Shak, north of the lower Volga. Farther north at the lower/middle Volga ecotone, at sites such as Varfolomievka and Oroshaemoe hunter-fishers who made pottery similar to that at Kair-Shak hunted onagers and saiga antelope in the desert-steppe, horses in the steppe, and aurochs in the riverine forests. Finally, in the Volga steppes north of Saratov and near Samara, hunter-fishers who made a different kind of pottery (Samara type) and hunted wild horses and red deer definitely were EHG. A Samara hunter-gatherer of this era buried at Lebyazhinka IV, dated 5600-5500 BC, was one of the first named examples of the EHG genetic type (Haak et al. 2015). This individual, like others from the same region, had no or very little CHG ancestry. The CHG mating network had not yet reached Samara by 5500 BC.

morgunova-eneolithic-pontic-caspian
Eneolithic settlements (1–5, 7, 10–16, 20, 22–43, 48, 50), burial grounds (6, 8–9, 17–19, 21, 47, 49) and kurgans (44–46) of the steppe Ural-Volga region: 1 Ivanovka; 2 Turganik; 3 Kuzminki; 4 Mullino; 5 Davlekanovo; 6 Sjezheye (burial ground); 7 Vilovatoe; 8 Ivanovka; 9 Krivoluchye; 10–13 LebjazhinkaI-III-IV-V; 14 Gundorovka; 15–16 Bol. Rakovka I-II; 17–18 Khvalunsk I-II; 19 Lipoviy Ovrag; 20 Alekseevka; 21 Khlopkovskiy; 22 Kuznetsovo I; 23 Ozinki II; 24 Altata; 25 Monakhov I; 26 Oroshaemoe; 27 Rezvoe; 28 Varpholomeevka; 29 Vetelki; 30 Pshenichnoe; 31 Kumuska; 32 Inyasovo; 33 Shapkino VI; 34 Russkoe Truevo I; 35 Tsaritsa I-II; 36 Kamenka I; 37 Kurpezhe-Molla; 38 Istay; 39 Isekiy; 40 Koshalak; 41 Kara-Khuduk; 42 Kair-Shak VI; 43 Kombakte; 44 Berezhnovka I-II; 45 Rovnoe; 46 Politotdelskoe; 47 burial near s. Pushkino; 48 Elshanka; 49 Novoorsk; 50 Khutor Repin. Modified from Morgunova (2014).

But before 4500 BC, CHG ancestry appeared among the EHG hunter-fishers in the middle Volga steppes from Samara to Saratov, at the same time that domesticated cattle and sheep-goats appeared. The Reich lab now has whole-genome aDNA data from more than 30 individuals from three Eneolithic cemeteries in the Volga steppes between the cities of Saratov and Samara (Khlopkov Bugor, Khvalynsk, and Ekaterinovka), all dated around the middle of the fifth millennium BC. Many dates from human bone are older, even before 5000 BC, but they are affected by strong reservoir effects, derived from a diet rich in fish, making them appear too old (Shishlina et al 2009), so the dates I use here accord with published and unpublished dates from a few dated animal bones (not fish-eaters) in graves.

Only three individuals from Khvalynsk are published, and they were first published in a report that did not mention the site in the text (Mathieson et al. 2015), so they went largely unnoticed. Nevertheless, they are crucial for understanding the evolution of the Yamnaya mating network in the steppes. They were mentioned briefly in Damgaard et al (2018) but were not graphed. They were re-analyzed and their admixture components were illustrated in a bar graph in Wang et al (2018: figure 2c), but they are not the principal focus of any published study. All of the authors who examined them agreed that these three Khvalynsk individuals, dated about 4500 BC, showed EHG ancestry admixed substantially with CHG, and not a trace of Anatolian Farmer ancestry, so the CHG was a Hotu-Cave or Kotias-Cave type of un-admixed CHG. The proportion of CHG in the Wang et al. (2018) bar graphs is about 20-30% in two individuals, substantially less CHG than in Yamnaya; but the third Khvalynsk individual had more than 50% CHG, like Yamnaya. The ca. 30 additional unpublished individuals from three middle Volga Eneolithic cemeteries, including Khvalynsk, preliminarily show the same admixed EHG/CHG ancestry in varying proportions. Most of the males belonged to Y-chromosome haplogroup R1b1a, like almost all Yamnaya males, but Khvalynsk also had some minority Y-chromosome haplogroups (R1a, Q1a, J, I2a2) that do not appear or appear only rarely (I2a2) in Yamnaya graves.

eneolithic-steppes
Pontic-Caspian steppe and neighbouring groups in the Neolithic. See full map.

Wang et al. (2018) discovered that this middle Volga mating network extended down to the North Caucasian steppes, where at cemeteries such as Progress-2 and Vonyuchka, dated 4300 BC, the same Khvalynsk-type ancestry appeared, an admixture of CHG and EHG with no Anatolian Farmer ancestry, with steppe-derived Y-chromosome haplogroup R1b. These three individuals in the North Caucasus steppes had higher proportions of CHG, overlapping Yamnaya. Without any doubt, a CHG population that was not admixed with Anatolian Farmers mated with EHG populations in the Volga steppes and in the North Caucasus steppes before 4500 BC. We can refer to this admixture as pre-Yamnaya, because it makes the best currently known genetic ancestor for EHG/CHG R1b Yamnaya genomes. The Progress-2 individuals from North Caucasus steppe graves lived not far from the pre-Maikop farmers of the Belaya valley, but they did not exchange mates, according to their DNA.

The hunter-fisher camps that first appeared on the lower Volga around 6200 BC could represent the migration northward of un-admixed CHG hunter-fishers from the steppe parts of the southeastern Caucasus, a speculation that awaits confirmation from aDNA. After 5000 BC domesticated animals appeared in these same sites in the lower Volga, and in new ones, and in grave sacrifices at Khvalynsk and Ekaterinovka. CHG genes and domesticated animals flowed north up the Volga, and EHG genes flowed south into the North Caucasus steppes, and the two components became admixed. After approximately 4500 BC the Khvalynsk archaeological culture united the lower and middle Volga archaeological sites into one variable archaeological culture that kept domesticated sheep, goats, and cattle (and possibly horses). In my estimation, Khvalynsk might represent the oldest phase of PIE.

eneolithic-early-steppes
Pontic-Caspian steppe and neighbouring groups in the Early Eneolithic. See full map.

Anatolian Farmer ancestry and Yamnaya origins

The Eneolithic Volga-North Caucasus mating network (Khvalynsk/Progress-2 type) exhibited EHG/CHG admixtures and Y-chromosome haplogroups similar to Yamnaya, but without Yamnaya’s additional Anatolian Farmer ancestry. (…)

Like the Mesolithic and Neolithic populations here, the Eneolithic populations of Dnieper-Donets II type seem to have limited their mating network to the rich, strategic region they occupied, centered on the Rapids. The absence of CHG shows that they did not mate frequently if at all with the people of the Volga steppes, a surprising but undeniable discovery. Archaeologists have seen connections in ornament types and in some details of funeral ritual between Dnieper-Donets cemeteries of the Mariupol-Nikol’skoe type and cemeteries in the middle Volga steppes such as Khvalynsk and S’yez’zhe (Vasiliev 1981:122-123). Also their cranio-facial types were judged to be similar (Bogdanov and Khokhlov 2012:212). So it it surprising that their aDNA does not indicate any genetic admixture with Khvalynsk or Progress-2. Also, neither they nor the Volga steppe Eneolithic populations showed any Anatolian Farmer ancestry. (…)

All three of the steppe-admixed exceptions were from the Varna region (Mathieson et al. 2018). One of them was the famous “golden man’ at Varna (Krause et al. 2016), Grave 43, whose steppe ancestry was the most doubtful of the three. If he had steppe ancestry, it was sufficiently distant (five+ generations before him) that he was not a statistically significant outlier, but he was displaced in the steppe direction, away from the central values of the majority of typical Anatolian Farmers at Varna and elsewhere. The other two, at Varna (grave 158, a 5-7-year-old girl) and Smyadovo (grave 29, a male 20-25 years old), were statistically significant outliers who had recent steppe ancestry (consistent with grandparents or great-grandparents) of the EHG/CHG Khvalynsk/Progress-2 type, not of the Dnieper Rapids EHG/WHG type.

(…) I believe that the Suvorovo-Cernavoda I movement into the lower Danube valley and the Balkans about 4300 BC separated early PIE-speakers (pre-Anatolian) from the steppe population that stayed behind in the steppes and that later developed into late PIE and Yamnaya.

This archaeological transition marked the breakdown of the mating barrier between steppe and Anatolian Farmer mating networks. After this 4300-4200 BC event, Anatolian Farmer ancestry began to pop up in the steppes. The currently oldest sample with Anatolian Farmer ancestry in the steppes in an individual at Aleksandriya, a Sredni Stog cemetery on the Donets in eastern Ukraine. Sredni Stog has often been discussed as a possible Yamnaya ancestor in Ukraine (Anthony 2007: 239- 254). The single published grave is dated about 4000 BC (4045– 3974 calBC/ 5215±20 BP/ PSUAMS-2832) and shows 20% Anatolian Farmer ancestry and 80% Khvalynsk-type steppe ancestry (CHG&EHG). His Y-chromosome haplogroup was R1a-Z93, similar to the later Sintashta culture and to South Asian Indo-Aryans, and he is the earliest known sample to show the genetic adaptation to lactase persistence (I3910-T). Another pre-Yamnaya grave with Anatolian Farmer ancestry was analyzed from the Dnieper valley at Dereivka, dated 3600-3400 BC (grave 73, 3634–3377 calBC/ 4725±25 BP/ UCIAMS-186349). She also had 20% Anatolian Farmer ancestry, but she showed less CHG than Aleksandriya and more Dereivka-1 ancestry, not surprising for a Dnieper valley sample, but also showing that the old fifth-millennium-type EHG/WHG Dnieper ancestry survived into the fourth millennium BC in the Dnieper valley (Mathieson et al. 2018).

late-eneolithic-repin
Pontic-Caspian steppe and neighbouring groups in the Late Eneolithic. See full map.

Probably, late PIE (Yamnaya) evolved in the same part of the steppes—the Volga-Caucasus steppes between the lower Don, the lower and middle Volga, and the North Caucasus piedmont—where early PIE evolved, and where appropriate EHG/CHG admixtures and Y-chromosome haplogroups were seen already in the Eneolithic (without Anatolian Farmer). There have always been archaeologists who argued for an origin of Yamnaya in the Volga steppes, including Gimbutas (1963), Merpert (1974), and recently Morgunova (2014), who argued that this was where Repin-type ceramics, an important early Yamnaya pottery type, first appeared in dated contexts before Yamnaya, about 3600 BC. The genetic evidence is consistent with Yamnaya EHG/CHG origins in the Volga-Caucasus steppes. Also, if contact with the Maikop culture was a fundamental cause of the innovations in transport and metallurgy that defined the Yamnaya culture, then the lower Don-North Caucasus-lower Volga steppes, closest to the North Caucasus, would be where the earliest phase is expected.

I would still guess that the Darkveti-Meshoko culture and its descendant Maikop culture established the linguistic ancestor of the Northwest Caucasian languages in approximately the region where they remained. I also accept the general consensus that the appearance of the hierarchical Maikop culture about 3600 BC had profound effects on pre-Yamnaya and early Yamnaya steppe cultures. Yamnaya metallurgy borrowed from the Maikop culture two-sided molds, tanged daggers, cast shaft hole axes with a single blade, and arsenical copper. Wheeled vehicles might have entered the steppes through Maikop, revolutionizing steppe economies and making Yamnaya pastoral nomadism possible after 3300 BC.

For those who still hoped that Proto-Indo-Europeans of Yamnaya/Afanasievo ancestry from the Don-Volga region were associated with the expansion of hg. R1a-M417, in a sort of mythical “R1-rich” Indo-European society, it seems this is going to be yet another prediction based on ancestry magic that goes wrong.

Proto-Indo-Europeans were, however, associated with other subclades beyond R1b-M269, probably (as I wrote recently) R1b-V1636, I2a-L699, Q1a-M25, and R1a-YP1272, but also interestingly some J subclade, so let’s see what surprises the new study on Khvalynsk and Yamnaya settlers from the Carpathian Basin brings…

On the bright side, it is indirectly confirmed that late Sredni Stog formed part of the neighbouring Corded Ware-like populations of ca. 20-30%+ Anatolian farmer ancestry that gave Yamnaya its share (ca. 6-10%), relative to the comparatively unmixed Khvalynsk and late Repin population (as shown by Afanasevo).

In this steppe mating network that opened up after the Khvalynsk expansion, the increasing admixture of Anatolian farmer-related ancestry in Yamnaya from east (ca. 2-10%) to west (ca. 6-15%) points to an exogamy of late Repin males in their western/south-western regions with populations around the Don River basin and beyond (and endogamy within the Yamnaya community), in an evolution relevant for language expansions and language contacts during the Late Eneolithic.

NOTE. “Mating network” is my new preferred term for “ancestry”. Also great to see scholars finally talk about “Pre-Yamnaya” ancestry, which – combined with the distinction of Yamnaya from Corded Ware ancestry – will no doubt help differentiate fine-scale population movements of steppe- and forest-steppe-related populations.

north-pontic-kvityana-dereivka-repin
Modified from Rassamakin (1999), adding red color to Repin expansion. The system of the latest Eneolithic Pointic cultures and the sites of the Zhivotilovo-Volchanskoe type: 1) Volchanskoe; 2) Zhivotilovka; 3) Vishnevatoe; 4) Koisug.

The whole issue of the JIES is centered on Caucasian influences on Early PIE as an Indo-Uralic dialect, and this language contact/substrate is useful to locate the most likely candidates for the Northeast and Northwest Caucasian and the Proto-Indo-European homelands.

On the other hand, it would also be interesting to read a discussion of how this Volga homeland of Middle PIE and Don-Volga-Ural homeland of Late PIE would be reconciled with the known continuous contacts of Uralic with Middle and Late PIE (see here) to locate the most likely Proto-Uralic homeland.

Especially because Corded Ware fully replaced all sub-Neolithic groups to the north and east of Khvalynsk/Yamnaya, like Volosovo, so no other population neighbouring Middle and Late Proto-Indo-Europeans survived into the Bronze Age…

EDIT: For those new to this blog, this information on unpublished samples from the Volga River basin is yet another confirmation of Khokhlov’s report on the R1b-L23 samples from Yekaterinovka, and its confirmation by a co-author of The unique elite Khvalynsk male from a Yekaterinovskiy Cape burial, apart from more support to the newest data placing Yekaterinovka culturally and probably chronologically between Samara and Khvalynsk.

Related

Corded Ware ancestry in North Eurasia and the Uralic expansion

uralic-clines-nganasan

Now that it has become evident that Late Repin (i.e. Yamnaya/Afanasevo) ancestry was associated with the migration of R1b-L23-rich Late Proto-Indo-Europeans from the steppe in the second half of the the 4th millennium BC, there’s still the question of how R1a-rich Uralic speakers of Corded Ware ancestry expanded , and how they spread their languages throughout North Eurasia.

Modern North Eurasians

I have been collecting information from the supplementary data of the latest papers on modern and ancient North Eurasian peoples, including Jeong et al. (2019), Saag et al. (2019), Sikora et al. (2018), or Flegontov et al. (2019), and I have tried to add up their information on ancestral components and their modern and historical distributions.

Fortunately, the current obsession with simplifying ancestry components into three or four general, atemporal groups, and the common use of the same ones across labs, make it very simple to merge data and map them.

Corded Ware ancestry

There is no doubt about the prevalent ancestry among Uralic-speaking peoples. A map isn’t needed to realize that, because ancient and modern data – like those recently summarized in Jeong et al. (2019) – prove it. But maps sure help visualize their intricate relationship better:

natural-modern-srubnaya-ancestry
Natural neighbor interpolation of Srubnaya ancestry among modern populations. See full map.
kriging-modern-srubnaya-ancestry
Kriging interpolation of Srubnaya ancestry among modern populations. See full map

Interestingly, the regions with higher Corded Ware-related ancestry are in great part coincident with (pre)historical Finno-Ugric-speaking territories:

uralic-languages-modern
Modern distribution of Uralic languages, with ancient territory (in the Common Era) labelled and delimited by a red line. For more information on the ancient territory see here.

Edit (29/7/2019): Here is the full Steppe_MLBA ancestry map, including Steppe_MLBA (vs. Indus Periphery vs. Onge) in modern South Asian populations from Narasimhan et al. (2018), apart from the ‘Srubnaya component’ in North Eurasian populations. ‘Dummy’ variables (with 0% ancestry) have been included to the south and east of the map to avoid weird interpolations of Steppe_MLBA into Africa and East Asia.

modern-steppe-mlba-ancestry2
Natural neighbor interpolation of Steppe MLBA-like ancestry among modern populations. See full map.

Anatolia Neolithic ancestry

Also interesting are the patterns of non-CWC-related ancestry, in particular the apparent wedge created by expanding East Slavs, which seems to reflect the intrusion of central(-eastern) European ancestry into Finno-Permic territory.

NOTE. Read more on Balto-Slavic hydrotoponymy, on the cradle of Russians as a Finno-Permic hotspot, and about Pre-Slavic languages in North-West Russia.

natural-modern-lbk-en-ancestry
Natural neighbor interpolation of LBK EN ancestry among modern populations. See full map.
kriging-modern-lbk-en-ancestry
Kriging interpolation of LBK EN ancestry among modern populations. See full map

WHG ancestry

The cline(s) between WHG, EHG, ANE, Nganasan, and Baikal HG are also simplified when some of them excluded, in this case EHG, represented thus in part by WHG, and in part by more eastern ancestries (see below).

modern-whg-ancestry
Natural neighbor interpolation of WHG ancestry among modern populations. See full map.
kriging-modern-whg-ancestry
Kriging interpolation of WHG ancestry among modern populations. See full map.

Arctic, Tundra or Forest-steppe?

Data on Nganasan-related vs. ANE vs. Baikal HG/Ulchi-related ancestry is difficult to map properly, because both ancestry components are usually reported as mutually exclusive, when they are in fact clearly related in an ancestral cline formed by different ancient North Eurasian populations from Siberia.

When it comes to ascertaining the origin of the multiple CWC-related clines among Uralic-speaking peoples, the question is thus how to properly distinguish the proportions of WHG-, EHG-, Nganasan-, ANE or BaikalHG-related ancestral components in North Eurasia, i.e. how did each dialectal group admix with regional groups which formed part of these clines east and west of the Urals.

The truth is, one ought to test specific ancient samples for each “Siberian” ancestry found in the different Uralic dialectal groups, but the simplistic “Siberian” label somehow gets a pass in many papers (see a recent example).

Below qpAdm results with best fits for Ulchi ancestry, Afontova Gora 3 ancestry, and Nganasan ancestry, but some populations show good fits for both and with similar proportions, so selecting one necessarily simplifies the distribution of both.

Ulchi ancestry

modern-ulchi-ancestry
Natural neighbor interpolation of Ulchi ancestry among modern populations. See full map.
kriging-modern-ulchi-ancestry
Kriging interpolation of Ulchi ancestry among modern populations. See full map.

ANE ancestry

natural-modern-ane-ancestry
Natural neighbor interpolation of ANE ancestry among modern populations. See full map.
kriging-modern-ane-ancestry
Kriging interpolation of ANE ancestry among modern populations. See full map.

Nganasan ancestry

modern-nganasan-ancestry
Natural neighbor interpolation of Nganasan ancestry among modern populations. See full map.
kriging-modern-nganasan-ancestry
Kriging interpolation of Nganasan ancestry among modern populations. See full map.

Iran Chalcolithic

A simplistic Iran Chalcolithic-related ancestry is also seen in the Altaic cline(s) which (like Corded Ware ancestry) expanded from Central Asia into Europe – apart from its historical distribution south of the Caucasus:

modern-iran-chal-ancestry
Natural neighbor interpolation of Iran Neolithic ancestry among modern populations. See full map.
kriging-modern-iran-neolithic-ancestry
Kriging interpolation of Iran Chalcolithic ancestry among modern populations. See full map.

Other models

The first question I imagine some would like to know is: what about other models? Do they show the same results? Here is the simplistic combination of ancestry components published in Damgaard et al. (2018) for the same or similar populations:

NOTE. As you can see, their selection of EHG vs. WHG vs. Nganasan vs. Natufian vs. Clovis of is of little use, but corroborate the results from other papers, and show some interesting patterns in combination with those above.

EHG

damgaard-modern-ehg-ancestry
Natural neighbor interpolation of EHG ancestry among modern populations, data from Damgaard et al. (2018). See full map.
damgaard-kriging-ehg-ancestry
Kriging interpolation of EHG ancestry among modern populations. See full map.

Natufian ancestry

damgaard-modern-natufian-ancestry
Natural neighbor interpolation of Natufian ancestry among modern populations, data from Damgaard et al. (2018). See full map.
damgaard-kriging-natufian-ancestry
Kriging interpolation of Natufian ancestry among modern populations. See full map.

WHG ancestry

damgaard-modern-whg-ancestry
Natural neighbor interpolation of WHG ancestry among modern populations, data from Damgaard et al. (2018). See full map.
damgaard-kriging-whg-ancestry
Kriging interpolation of WHG ancestry among modern populations. See full map.

Baikal HG ancestry

damgaard-modern-baikalhg-ancestry
Natural neighbor interpolation of Baikal hunter-gatherer ancestry among modern populations, data from Damgaard et al. (2018). See full map.
damgaard-kriging-baikal-hg-ancestry
Kriging interpolation of Baikal HG ancestry among modern populations. See full map.

Ancient North Eurasians

Once the modern situation is clear, relevant questions are, for example, whether EHG-, WHG-, ANE, Nganasan-, and/or Baikal HG-related meta-populations expanded or became integrated into Uralic-speaking territories.

When did these admixture/migration events happen?

How did the ancient distribution or expansion of Palaeo-Arctic, Baikalic, and/or Altaic peoples affect the current distribution of the so-called “Siberian” ancestry, and of hg. N1a, in each specific population?

NOTE. A little excursus is necessary, because the calculated repetition of a hypothetic opposition “N1a vs. R1a” doesn’t make this dichotomy real:

  1. There was not a single ethnolinguistic community represented by hg. R1a after the initial expansion of Eastern Corded Ware groups, or by hg. N1a-L392 after its initial expansion in Siberia:
  2. Different subclades became incorporated in different ways into Bronze Age and Iron Age communities, most of which without an ethnolinguistic change. For example, N1a subclades became incorporated into North Eurasian populations of different languages, reaching Uralic- and Indo-European-speaking territories of north-eastern Europe during the late Iron Age, at a time when their ancestral origin or language in Siberia was impossible to ascertain. Just like the mix found among Proto-Germanic peoples (R1b, R1a, and I1)* or among Slavic peoples (I2a, E1b, R1a)*, the mix of many Uralic groups showing specific percentages of R1a, N1a, or Q subclades* reflect more or less recent admixture or acculturation events with little impact on their languages.

*other typically northern and eastern European haplogroups are also represented in early Germanic (N1a, I2, E1b, J, G2), Slavic (I1, G2, J) and Finno-Permic (I1, R1b, J) peoples.

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

The problem with mapping the ancestry of the available sampling of ancient populations is that we lack proper temporal and regional transects. The maps that follow include cultures roughly divided into either “Bronze Age” or “Iron Age” groups, although the difference between samples may span up to 2,000 years.

NOTE. Rough estimates for more external groups (viz. Sweden Battle Axe/Gotland_A for the NW, Srubna from the North Pontic area for the SW, Arctic/Nganasan for the NE, and Baikal EBA/”Ulchi-like” for the SE) have been included to offer a wider interpolated area using data already known.

Bronze Age

Similar to modern populations, the selection of best fit “Siberian” ancestry between Baikal HG vs. Nganasan, both potentially ± ANE (AG3), is an oversimplification that needs to be addressed in future papers.

Corded Ware ancestry

bronze-age-corded-ware-ancestry
Natural neighbor interpolation of Srubnaya ancestry among Bronze Age populations. See full map.

Nganasan-like ancestry

bronze-age-nganasan-like-ancestry
Natural neighbor interpolation of Nganasan-like ancestry among Bronze Age populations. See full map.

Baikal HG ancestry

bronze-age-baikal-hg-ancestry
Natural neighbor interpolation of Baikal Hunter-Gatherer ancestry among Bronze Age populations. See full map.

Afontova Gora 3 ancestry

bronze-age-afontova-gora-ancestry
Natural neighbor interpolation of Afontova Gora 3 ancestry among Bronze Age populations. See full map.

Iron Age

Corded Ware ancestry

Interestingly, the moderate expansion of Corded Ware-related ancestry from the south during the Iron Age may be related to the expansion of hg. N1a-VL29 into the chiefdom-based system of north-eastern Europe, including Ananyino/Akozino and later expanding Akozino warrior-traders around the Baltic Sea.

NOTE. The samples from Levänluhta are centuries older than those from Estonia (and Ingria), and those from Chalmny Varre are modern ones, so this region has to be read as a south-west to north-east distribution from the Iron Age to modern times.

iron-age-corded-ware-ancestry
Natural neighbor interpolation of Srubnaya ancestry among Iron Age populations. See full map.

Baikal HG-like ancestry

The fact that this Baltic N1a-VL29 branch belongs in a group together with typically Avar N1a-B197 supports the Altaic origin of the parent group, which is possibly related to the expansion of Baikalic ancestry and Iron Age nomads:

iron-age-baikal-ancestry
Natural neighbor interpolation of Baikal HG ancestry among Iron Age populations. See full map.

Nganasan-like ancestry

The dilution of Nganasan-like ancestry in an Arctic region featuring “Siberian” ancestry and hg. N1a-L392 at least since the Bronze Age supports the integration of hg. N1a-Z1934, sister clade of Ugric N1a-Z1936, into populations west and east of the Urals with the expansion of Uralic languages to the north into the Tundra region (see here).

The integration of N1a-Z1934 lineages into Finnic-speaking peoples after their migration to the north and east, and the displacement or acculturation of Saami from their ancestral homeland, coinciding with known genetic bottlenecks among Finns, is yet another proof of this evolution:

iron-age-nganasan-ancestry
Natural neighbor interpolation of Nganasan ancestry among Iron Age populations. See full map.

WHG ancestry

Similarly, WHG ancestry doesn’t seem to be related to important population movements throughout the Bronze Age, which excludes the multiple North Eurasian populations that will be found along the clines formed by WHG, EHG, ANE, Nganasan, Baikal HG ancestry as forming part of the Uralic ethnogenesis, although they may be relevant to follow later regional movements of specific populations.

iron-age-whg-ancestry
Natural neighbor interpolation of WHG ancestry among Iron Age populations. See full map.

Conclusion

It seems natural that people used to look at maps of haplogroup distribution from the 2000s, coupled with modern language distributions, and would try to interpret them in a certain way, reaching thus the wrong conclusions whose consequences are especially visible today when ancient DNA keeps contradicting them.

In hindsight, though, assuming that Balto-Slavs expanded with Corded Ware and hg. R1a, or that Uralians expanded with “Siberian” ancestry and hg. N1a, was as absurd as looking at maps of ancestry and haplogroup distribution of ancient and modern Native Americans, trying to divide them into “Germanic” or “Iberian”…

The evolution of each specific region and cultural group of North Eurasia is far from being clear. However, the general trend speaks clearly in favour of an ancient, Bronze Age distribution of North Eurasian ancestry and haplogroups that have decreased, diluted, or become incorporated into expanding Uralians of Corded Ware ancestry, occasionally spreading with inter-regional expansions of local groups.

Given the relatively recent push of Altaic and Indo-European languages into ancestral Uralic-speaking territories, only the ancient Corded Ware expansion remains compatible with the spread of Uralic languages into their historical distribution.

Related

Iron Age Tocharians of Yamnaya ancestry from Afanasevo show hg. R1b-M269 and Q1a1

New open access Ancient Genomes Reveal Yamnaya-Related Ancestry and a Potential Source of Indo-European Speakers in Iron Age Tianshan, by Ning et al. Cell (2019).

Interesting excerpts (emphasis mine, changes for clarity):

Here, we report the first genome-wide data of 10 ancient individuals from northeastern Xinjiang. They are dated to around 2,200 years ago and were found at the Iron Age Shirenzigou site. We find them to be already genetically admixed between Eastern and Western Eurasians. We also find that the majority of the East Eurasian ancestry in the Shirenzigou individuals is related to northeastern Asian populations, while the West Eurasian ancestry is best presented by ∼20% to 80% Yamnaya-like ancestry. Our data thus suggest a Western Eurasian steppe origin for at least part of the ancient Xinjiang population. Our findings furthermore support a Yamnaya-related origin for the now extinct Tocharian languages in the Tarim Basin, in southern Xinjiang.

Haplogroups

The dominant mtDNA lineages of the Shirenzigou people are commonly found in modern and ancient West Eurasian populations, such as U4, U5, and H, while they also have East Eurasian-specific haplogroups A, D4, and G3, preliminarily documenting admixed ancestry from eastern and western Eurasia.

The admixture profile is also shown on the paternal Y chromosome side that 4 out of 6 males in Shirenzigou (Figure S2) belong to the West Eurasian-specific haplogroup R1b (n = 2) and East Eurasian-specific haplogroup Q1a (n = 2), the former is predominant in ancient Yamnaya and nearly 100% in Afanasievo, different from the Middle and Late Bronze Age Steppe groups (Steppe_MLBA) such as Andronovo, [Potapovka], Srubnaya, and Sintashta whose Y chromosomal haplogroup is mainly R1a.

tocharians-y-dna-mtdna

Autosomal

We first carried out principal component analysis (PCA) to assess the genetic affinities of the ancient individuals qualitatively by projecting them onto present-day Eurasian variation (Figure 2). We observed a distinct separation between East and West Eurasians. Our ancient Shirenzigou samples and present-day populations from Central Asia and northwestern China form a genetic cline from East to West in the first PC. The distribution of Shirenzigou samples on the cline is relatively scattered with two major clusters, one being closer to modern-day Uygurs and Kazakhs and the other being closer to recently published ancient Saka and Huns from the Tianshan in Kazakhstan (…).

We applied a formal admixture test using f3 statistics in the form of f3 (Shirenzigou; X, Y) where X and Y are worldwide populations that might be the genetic sources for the Shirenzigou individuals. We observed the most significant signals of admixture in the Shirenzigou samples when using Yamnaya_Samara or Srubnaya as the West Eurasian source and some Northern Asians or Koreans as the East Eurasian source (Table S1). We also plotted the outgroup f3 statistics in the form of f3 (Mbuti; X, Anatolia_Neolithic) and f3 (Mbuti; X, Kostenki14) to visualize the allele sharing between population X and Anatolian farmers. As shown in Figure S3, the Steppe_MLBA populations including Srubnaya, Andronovo, and Sintashta were shifted toward farming populations compared with Yamnaya groups and the Shirenzigou samples. This observation is consistent with ADMIXTURE analysis that Steppe_MLBA populations have an Anatolian and European farmer-related component that Yamnaya groups and the Shirenzigou individuals do not seem to have. The analysis consistently suggested Yamnaya-related Steppe populations were the better source in modeling the West Eurasian ancestry in Shirenzigou.

tocharians-pca-admixture
PCA and ADMIXTURE for Shirenzigou Samples. Modified from the original to include in black squares samples related to Yamnaya.

Genetic Composition of Iron Age Shirenzigou Individuals

We continued to use qpAdm to estimate the admixture proportions in the Shirenzigou samples by using different pairs of source populations, such as Yamnaya_Samara, Afanasievo, Srubnaya, Andronovo, BMAC culture (Bustan_BA and Sappali_Tepe_BA) and Tianshan_Hun as the West Eurasian source and Han, Ulchi, Hezhen, Shamanka_EN as the East Eurasian source. In all cases, Yamnaya, Afanasievo, or Tianshan_Hun always provide the best model fit for the Shirenzigou individuals, while Srubnaya, Andronovo, Bustan_BA and Sappali_Tepe_BA only work in some cases. The Yamnaya_Samara or Afanasievo-related ancestry ranges from ∼20% to 80% in different Shirenzigou individuals, consistent with the scattered distribution on the East-West cline in the PCA

ancestry-tocharians

(…) we then modeled Shirenzigou as a three-way admixture of Yamnaya_Samara, Ulchi (or Hezhen) and Han to infer the source from the East Eurasia side that contributed to Shirenzigou. We found the Ulchi or Hezhen and Han-related ancestry had a complicated and unevenly distribution in the Shirenzigou samples. The most Shirenzigou individuals derived the majority of their East Eurasian ancestry from Ulchi or Hezhen-related populations, while the following two individuals M820 and M15-2 have more Han related than Ulchi/Hezhen-related ancestry.

One important question remains, though: how and when did these Proto-Tocharian speakers migrate from the Afanasevo culture in the Altai into the Tarim Basin? The traditional answer, now more likely than ever, is through the Chemurchek culture. See e.g. A re-analysis of the Qiemu’erqieke (Shamirshak) cemeteries, Xinjiang, China, by Jia and Betts JIES (2010) 38(4).

Also, given the apparent lack of (extra farmer ancestry that characterizes) Corded Ware ancestry, if the results were already suspicious before, how likely are now the published R1a(xZ93) and/or radiocarbon dates of the Xiaohe mummies from Li et al. (2010, 2015)? Because, after all, one should have expected in such a late date a generalized admixture with neighbouring Srubna/Andronovo-like populations.

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

Volosovo hunter-gatherers started to disappear earlier than previously believed

volosovo-corded-ware

Recent paper (behind paywall) Marmot incisors and bear tooth pendants in Volosovo hunter-gatherer burials. New radiocarbon and stable isotope data from the Sakhtysh complex, Upper-Volga region, by Macānea, Nordqvist, and Kostyleva, J. Archaeol. Sci. (2019) 26:101908.

Interesting excerpts (emphasis mine):

The Sakhtysh micro-region is located in the Volga-Oka interfluve, along the headwaters of the Koyka River in the Ivanovo Region, central European Russia (Fig. 1). The area has evidence of human habitation from the Early Mesolithic to the Iron Age, and includes altogether 11 long-term and seasonal settlements (Sakhtysh I–II, IIa, III–IV, VII–XI, XIV) and four artefact scatters (sites V–VI, XII–XIII), in addition to which burials have been detected at five sites (I–II, IIa, VII, VIII) (Kostyleva and Utkin, 2010). The locations have been known since the 1930s and intensively studied since the 1960s under the leadership of D.A. Kraynov, M.G. Zhilin, E.L. Kostyleva, and A.V. Utkin.

Sakhtysh II and IIa are the most extensively studied sites of the complex, with ca. 1500m2 and around 800m2 excavated, respectively. The burial grounds at both sites are considered as fully investigated.

volosovo-sakhtysh-dates
AMS datings from the sites Sakhtysh II and IIa. Sampled contexts are given in parentheses (burial/hoard), “crust” indicates samples of charred organic
residues on pottery from cultural layer. For data, see Tables 1–2.

Sakhtysh chronology

The AMS dates do not support the previously proposed phasing of the Sakhtysh burials to early (4750–4375 BP/3600–3000 cal BCE), late (or developed; 4375–4000 BP/3000–2500 cal BCE), and final (4000–3750 BP/2500–2200 cal BCE): the early and late burials at Sakhtysh IIa do not stand out as two separate groups, and also the burials and hoards from Sakhtysh II, connected to the final phase, are temporally overlapping with these. Neither the use sequence, where the settlement and burial phases are non-overlapping and also complementary between the sites (Kostyleva and Utkin, 2010, 2014), finds support in the present material.

The AMS datings indicate that the Volosovo people started to bury their dead at Sakhtysh IIa after 3700 cal BCE; dates earlier than this may be affected by FRE or suffer from mixed contexts and poor quality of dates. The present data questions the interpretation that the Sakhtysh IIa cemetery was used without interruptions between 4800 and 4080 BP (Kostyleva and Utkin, 2010), i.e. for a millennium between 3550 and 2600 cal BCE. The AMS dates rather suggest a use period of some centuries only around the mid-4th millennium cal BCE, tentatively 3650–3400 cal BCE. This would also be more realistic considering the number of burials at the site.

volosovo-sakhtysh-sites
The core area of Volosovo culture (after Kraynov, 1987) and the sites of the Sakhtysh complex (after Kostyleva and Utkin, 2010). Eurasian map base made with Natural Earth. Illustration: K. Nordqvist.

Volosovo chronology

The absolute dating of Volosovo culture was for a long time hampered by the small number of radiocarbon dates (see Kraynov, 1987). Today,>100 datings connected with it can be found in literature (Korolev and Shalapinin, 2010; Chernykh et al., 2011; Nikitin, 2012; Mosin et al., 2014). Unfortunately, the available dates do not form solid grounds for dating the cultural phenomenon, as many of them have quality-related issues, large measurement errors, and ambiguous cultural or physical contexts. Consequently, particular datings may be connected to different cultural phases by different scholars. Finally, a large part of the newly-published datings are obtained through direct dating of potsherds (Kovaliukh and Skripkin, 2007; Zaitseva et al., 2009), and therefore, their cogency must be faced with reservation (see Van der Plicht et al., 2016; Dolbunova et al., 2017).

The datings connected with Volosovo cover a wide time range between ca. 5500 BP (4400 cal BCE) and ca. 3700 BP (2100 cal BCE). However, datings from secure contexts, with good quality (error ca. 50 years or below) and no probable FRE, place the beginning of Volosovo culture to the first half of the 4th millennium cal BCE, around 3700–3600 cal BCE. This is also supported by the roughly coeval terminal dates given for the preceding Lyalovo (Zaretskaya and Kostyleva, 2011) and Volga-Kama cultures (Lychagina, 2018), as well as the appearance of related neighbouring cultures, for example, in the Kama region (Nikitin, 2012; Lychagina, 2018), the southern forest steppe area (Korolev and Shalapinin, 2014), and north-western Russia and Finland (Nordqvist, 2018). Still, the dating of many of these cultural phases suffers from the same problems as of Volosovo.

A handful of contested datings place the end of Volosovo culture to the final centuries of the 3rd millennium cal BCE, or even later (Kostyleva and Utkin, 2010; Chernykh et al., 2011; Nikitin, 2012). On the other hand, the new AMS dates indicate that Volosovo activities at Sakhtysh II and IIa ceased before or towards the early 3rd millennium cal BCE; if this reflects the general decline of Volosovo culture must be still confirmed by more dates from Sakhtysh and elsewhere. In this context, the general cultural development must be accounted for. To what extent – if at all – the Volosovo people were present after the arrival of the Corded Ware culture-related Fatyanovo-Balanovo populations? Based on the current, albeit scant and inconclusive radiocarbon data this took place from ca. 2700 cal BCE onwards (Krenke et al., 2013).

volosovo-fatyanovo-balanovo
Corded Ware and Comb Ware hunter-gatherer-related populations in north-eastern Europe from ca. 2600 BC. See full map.

Comments

One of the interesting genetic papers in the near future will be the one that finally includes samples from Corded Ware groups in the forest zone (i.e. Fatyanovo-Balanovo and Abashevo), which will most likely confirm that they are the origin of the known genetic profile of Central and East Uralic-speaking peoples, seeing how West Uralic peoples show genetic continuity in the East Baltic area, coinciding with the Battle Axe culture.

Uralicists have come a long way from the 1990s, when the picture of Uralic before Balto-Slavic in the Baltic was already evident, and Uralians were identified with Comb Ware peoples. The linguistic data and relative chronology are still valid, despite the now outdated interpretations of absolute archaeological chronology, as happens with interpretations of Krahe or Villar about Old European.

As an example, here are some relevant excerpts from Languages in the Prehistoric Baltic Sea Region, by Kallio (2003):

NOTE. Kallio’s contribution appeared in the book Languages in Prehistoric Europe (2003), which I hold nostalgically close in my Indo-European library (now almost impossible to read fully). It is still one of my preferred books (from those made up of mostly unconnected chunks on European linguistic prehistory), because it contains Oettinger’s essential update of North-West Indo-European common vocabulary, which led us indirectly to our Modern Indo-European project from 2005 on.

In any case, the Uralic arrival in the region east of the Baltic Sea preceded the Indo-European one (…).

This theory that the ancestors of Finno-Saamic speakers arrived in the Baltic Sea region earlier than those of Balto-Slavic speakers is still rejected by some scholars (e.g. Napolskikh 1993: 41-44), who claim, for instance, that Finno-Saamic speakers would not have known salmons before they met Balts because the Finno-Saamic word for ‘salmon’ (i.e. *losi) is a borrowing from Baltic. Similarly, one could claim that English speakers would not have known salmons before they met Frenchmen because English salmon is a borrowing from French. In other words, Worter und Sachen are not necessarily borrowed hand in hand. Otherwise, it would not be so easy to explain how many Finnish names of body parts are borrowings from Baltic (e.g. hammas ‘tooth’, kaula ‘neck’, reisi ‘thigh’) and from Germanic (e.g. hartia ‘shoulder’, lantio ‘loin’, maha ‘stomach’).

A more probative argument is the fact that Balto-Slavic features in Finno-Saamic are mostly lexical ones (i.e. typical superstrate features), where Finno-Saamic features in Balto-Slavic are mostly non-lexical ones (i.e. typical substrate features). Note that there are more Balto-Slavic features in Finnic than in Saamic and more Finno-Saamic features in Baltic than in Slavic. This fact could be explained by presuming that Pre-Saamic was spoken north of the Corded Ware area and Pre-Slavic was spoken south of the Typical Pit-Comb Ware area, whereas Pre-Finnic and Pre-Baltic alone were spoken in the area, where both the Typical Pit-Comb Ware culture (ca. 4000-3600 BC) and the Corded Ware culture (ca. 3200-2300 BC) were situated. This area was most probably bilingual, until Finnic and Baltic won in the north and in the south, respectively.

As is well-known, the idea of Uralic substrate features in Balto-Slavic is not new (cf. e.g. Pokorny 1936/1968: 181-185). As recent studies (e.g. Bednarczuk 1997) have shown, their density is the most remarkable in the four Balto-Slavic languages spoken in the earlier Pit-Comb Ware area (i.e. Latvian, Lithuanian, Belorussian, Russian). On the other hand, occasional Uralisms in the other Balto-Slavic languages spoken west of the Vistula and south of the Pripyat may rather be considered adstrate features spread from the northeast.

comb-ware-uralic
Our beliefs from the 2000s. A hypothetic Uralic Comb Ware distribution before the arrival of a hypothetic North-West Indo-European-speaking Corded Ware. “Generalized distribution of the Pit-Comb Ware cultural complex (Mallory & Adams 1997: 430, Carpelan 1999: 257) and the most probable homelands of Saamic, Finnic, Mordvin, Mari, and Permic.”

The idea of Indo-European superstrate features in Finnic is not new either (cf. e.g. Posti 1953). As Jorma Koivulehto (1983) has recently shown, the earliest Indo-European loanword stratum in the westernmost Uralic branches alone can be considered Northwest Indo-European and connected with the Corded Ware culture (ca. 3200-2300 BC). Since this layer, there have been continuous contacts between Baltic and Finnic. According to Koivulehto (1990), the following stratum can be called Proto-Balt(o-Slav)ic and dated to the Late Neolithic period (ca. 2300-1500 BC). Note that this Proto-Balt(o-Slav)ic dating agrees with the established ones (cf. e.g. Shevelov 1964: 613-614, Kortlandt 1982: 181), when we remember the fact that archaeologists have also moved their datings back by centuries during the last decades.

Finally, there is also a Baltic loanword stratum which was not borrowed from the ancestral stage of Latvian, Lithuanian and/or Old Prussian but from some extinct Baltic language or dialect (Nieminen 1957). However, as these words still go back to the early Proto-Finnic stage, they can hardly be dated later than Bronze-Age ( ca. 1500-500 BC). Therefore, we may conclude that they were probably borrowed from a Baltic superstrate, which arrived in the Finnish Gulf area during the Corded Ware period and survived there until the Bronze Age, when it was no longer identical with other Baltic dialects. In any case, as later Baltic loanword strata concern southern Finnic languages alone, we may presume that this ‘North Baltic’superstrate had become extinct.

The traditional association of Uralic with Volosovo hunter-gatherers doesn’t make sense, since they neither miraculously survived for thousands of years nor mixed for hundreds of years with Corded Ware peoples, so we can now more confidently reject the recent assumption by Carpelan & Parpola that their language was adopted by incoming Fatyanovo, Balanovo and Abashevo groups, to develop into the known Uralic languages (more here). This includes one of the many models of the the Copenhagen group, who simplistically follow “Steppe ancestry” for Indo-Europeannes.

If one combines the known relative linguistic chronology with the North-West Indo-European hydrotoponymy layer, now more clearly identified as Old Europeans expanding with East Bell Beakers and derived Early Bronze Age groups, I think there is little space left for maneuvering out of the overwhelming evidence for a Uralic homeland in the forest-steppes, linked to the spread of late Sredni Stog/Corded Ware ancestry into north-eastern Europe and beyond the Urals.

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