We present results of the largest multidisciplinary human mobility investigation to date of skeletal remains from present-day Denmark encompassing the 3rd and 2nd millennia BC. Through a multi-analytical approach based on 88 individuals from 37 different archaeological localities in which we combine strontium isotope and radiocarbon analyses together with anthropological investigations, we explore whether there are significant changes in human mobility patterns during this period. Overall, our data suggest that mobility of people seems to have been continuous throughout the 3rd and 2nd millennia BC. However, our data also indicate a clear shift in mobility patterns from around 1600 BC onwards, with a larger variation in the geographical origin of the migrants, and potentially including more distant regions. This shift occurred during a transition period at the beginning of the Nordic Bronze Age at a time when society flourished, expanded and experienced an unprecedented economic growth, suggesting that these aspects were closely related.
Strontium isotope analyses
The results of our strontium isotope analyses are presented in Table 2 and listed in chronological order according to the radiocarbon dates (in sites with multiple individuals we start with the oldest radiocarbon individual). The strontium isotope data set reveals a wide range of values from 87Sr/86Sr = 0.70871 (RISE 23, from the site of Debel) to 87Sr/86Sr = 0.71788 (RISE 20, from the site of Karlstrup). Despite the difficulties of establishing the baseline range some of the herein investigated individuals may be classified as non-locals. A few individuals have tooth enamel signatures that lie just above the upper baseline limit of 87Sr/86Sr = 0.711 and therefore, the classification of these humans as non-locals should be considered with caution. Nevertheless, the significant proportion of individuals with relatively radiogenic values suggest that about a quarter of the individuals studied herein seem to have originated from other places than from those they were buried, and hence implying a continuous degree of mobility during the 3rd and 2nd millennia BC.
From the Single Grave Culture (SGC) which is closely related to the Corded Ware Complex in central and eastern Europe and dates from c. 2800 BC to 2200 BC, we analyzed seven of the at least ten individuals who were buried at the site of Gjerrild in eastern Jutland (Fig 1). Gjerrild is a key SGC site, as to date it has provided the most substantial skeletal material pertaining to this culture from present-day Denmark. However, it is not a typical SGC grave, but a megalithic chamber of the so-called “Bøstrup type”. The SGC pottery was decorated with cord or stamp impressions and the stone battle axes were a common feature of male equipment. Such shared traits in the Corded Ware Complex probably reflected shared occupational, social and religious characteristics. Apart from one individual who yielded a Bronze Age date, five individuals date within the period that spans from c. 2600 BC to 2200 BC, hence representing the middle and late SGC phases (Table 1 and S1 File). Of the seven individuals, three males, one female, two infants and one adult (only represented by a disarticulated mandible, and dated to the Bronze Age), all but one yielded strontium isotope signatures that fall within the local baseline range. Only the female (RISE 1283) has a more radiogenic strontium isotope signature of 87Sr/86Sr = 0.7127, which is similar to that of the male from Kyndeløse and might indicate non-local provenance. One of the individuals at Gjerrild, a mature-old adult male, who yielded a local signature (RISE 432) was accompanied by a D-type arrowhead and an amber bead which lay on his right side. He showed signs of inflammation on his lower legs, in particular on the left one. He had a healed trepanation (Fig 2). Another individual (RISE 73a, 1282), an adult male, was found with a type D arrowhead in the sternum (Fig 3).
Late Neolithic I
We sampled individuals from a total of twelve different sites that date to the Late Neolithic period (2300/2250-1700 BC).
One of these sites is Hellested on Zealand (Fig 1 and S1 File), with four flat graves containing five individuals, four young males and one mature adult female. We conducted strontium isotope analyses of enamel from all five individuals, and our results point to two individuals being characterized by local strontium isotope values. One of these individuals, the female, was buried with no grave goods (RISE 53, grave B) while the other, a young male, was buried with a fragmented bone pin (RISE 56, grave F). The other three male individuals (RISE 54, 55, 57) yielded similar strontium isotopic values that lie slightly above the local baseline range. All these individuals had been buried with early flint daggers (type I and II), and one of them (RISE 57, grave A) additionally had a ring-headed pin (Ringkopfnadel) . On the basis of the presence of this ring-headed pin, Lomborg  suggested that these individuals had connections with the Únětice culture. Furthermore, three of them have radiocarbon dates that overlap (RISE 55, 56 and 57; Table 1).
Another Late Neolithic site is Juelsberg on the island of Funen (central Denmark, Fig 1 and S1 File) which is a gallery grave that contained at least 19 individuals. We conducted strontium isotope analyses of tooth enamel on 8 out of the 19 individuals and two of them, a male and female, yielded ratios that suggest a non-local origin (RISE 30 and 32). The grave goods comprise a (Lomborg) type I flint dagger but also some non-local type of artefacts. These consist of an early type of bone pin (type 7) mainly found in south-eastern Scandinavia, and a barbed and tanged flint arrowhead of the west-European Bell Beaker type suggesting western connections. The middle adult female (RISE 32) yielded an 87Sr/86Sr = 0.7121 and the mature to old adult male (RISE 30) yielded a 87Sr/86Sr = 0.7112. The different Sr isotope signatures of these individuals imply that they might have originated from different areas, albeit their radiocarbon dates are very similar.
The gallery grave of Marbjerg, Zealand (Fig 1), yielded 17 individuals (S1 File), and we conducted strontium isotope analyses of tooth enamel on 11 of them. The majority of the individuals were males, but females and children, too, were present. Anthropological investigations of the individuals from this site, males as well as females, indicate a relatively high life expectancy with respect to that typical for this period (S1 Table). Our radiocarbon dates revealed that this grave was in use for several hundred years from the Early Late Neolithic (2210–2030 cal BC, RISE 39) to the Late Neolithic /Early Nordic Bronze Age Period (1770–1620 cal BC, RISE 41). Despite the long-term use of this grave, 10 of the 11 individuals studied herein yielded a very narrow and overlapping range of strontium isotope values between 87Sr/86Sr = 0.7096–0.7101. Their values suggest not only that these individuals were local but that their food sources were derived from the same area over the course of several centuries. Only the tooth enamel sample of one individual, a middle to mature adult male (RISE 40), yielded a higher value of 87Sr/86Sr = 0.7117, which seems to suggest a non-local origin.
Predictions about these samples
Strontium isotope analyses only show potential movements during an individual’s lifetime, which is normally useless to assess relevant migrations if the sampling is not big enough (see more on investigating population movements). Still, if a sampling like this one shows many potentially non-local individuals from different parts of Denmark deviating from the baseline at a certain period, you can infer that something is happening within Denmark and in nearby regions.
Based on what we know now, I bet these are the most likely events in Denmark that marked the Nordic Late Neolithic with its Bell Beaker-related Dagger Period ca. 2400/2300 BC on:
Sudden appearance of R1b-L23 lineages (probably R1b-U106 among them), originally from the Northern European Plain, ultimately from the Danube River Basin. R1a-M417 subclades, possibly prevalent in the previous period, disappear or appear rarely, to resurge later during the Bronze Age probably mostly as hg R1a-Z284, originally from the Battle Axe culture in Sweden, together with I1 – these resurgence events might be shifted to a later phase, though, and there might be some isolated R1a cases in the Danish LN, too.
Shift of Middle Neolithic to Late Neolithic in the PCA away from the Corded Ware cluster and closer to the Bell Beaker cluster – whatever that means exactly for Danish SGC relative to Northern European Beakers, visible especially when enough samples are available.
Evident sign of new incoming ancestry ultimately from Yamnaya-related populations, compared to earlier peoples of Corded Ware ancestry. Yes, even this far north, despite heavy admixture of Yamnaya-like Bell Beakers through exogamy with Corded Ware-like populations all the way to the north from the Danube Basin.
The above predictions are more or less evident to everyone, despite the current mistrust in the Yamnaya – Bell Beaker expansion route of North-West Indo-European, due to the prevalent nativist and/or reactionary trends in hobby population genomics and among academics. My main prediction is therefore about human behaviour:
(1) Seeing how the Copenhagen group started to describe recently South Scandinavian genetic and linguistic prehistory, their conclusions are predictable. From the introduction of this paper:
The 3rd millennium BC stands out as a period of migrations in western Eurasia, as pastoral steppe populations settled in temperate Europe after 2800 BC e.g. [1, 2]. This was also a period of cultural and genetic admixture e.g. . From 1600 BC onwards, southern Scandinavia became more closely linked to the existing European metal trade networks (…)
See what they did there? No mention of the radical change that the Dagger Period brought to Scandinavia, in cultural or genetic terms (see e.g. here or here). Strange how the only thing that Kristiansen has changed since the 1980s – and only after the 2015 genetic papers – is his previous emphasis on the Dagger Period as the most relevant unifying cultural and population movement in Scandinavia, responsible for the formation of a common Nordic language, which is suddenly given as little weight as possible in all his publications, to support some imaginary continuity with the Corded Ware culture (see e.g. here or here).
(2) Only a few males from the Single Grave period are described in this sampling, and they are quite close to the arrival of Bell Beakers, so if someone is looking for closure about the “R1b from Corded Ware”, I bet there won’t be any. As with conspiracy theories of native Vasconic R1b-L51 hidden somewhere in Western Europe even after Olalde et al. (2018) and Olalde et al. (2019), the mythic native Nordic R1b-U106 of Corded Ware will remain hidden in some unsampled Corded Ware group in the minds of many, despite being already found in Bell Beaker-derived European EBA cultures of Bohemia and possibly of Hungary, too (ca. 2500-2200 BC, see SNP calls), apart from the Late Neolithic sample from Lilla Beddinge in Scania (ca. 2275-2032 BC, see SNP calls)
I hope that I am wrong, and that some scholar in the Danish group is capable of reporting the data as it is, even if it contradicts the theories of its leading archaeologist, Kristian Kristiansen. The apparent downplay of the increase in non-local origins of individuals during the Late Neolithic I period as they appear in this paper, as well as their summary of foreign migrations into Denmark which mysteriously stop with the arrival of “Steppe ancestry” ca. 2800 BC, make me think that a change in their narrative is not very likely. The cons of working with academic divos, I guess…
A reader commented recently that there is little information about Indo-Europeans from Central and East Asia in this blog. Regardless of the scarce archaeological data compared to European prehistory, I think it is premature to write anything detailed about population movements of Indo-Iranians in Asia, especially now that we are awaiting the updates of Narasimhan et al (2018).
Furthermore, there was little hope that Tocharians would be different than neighbouring Andronovo-like populations (see a recent post on my predicted varied admixture of Common Tocharians), so the history of both unrelated Late PIE languages would have had to be explained by the admixture of Afanasievo-related groups with peoples of Andronovo descent and their acculturation.
This genetic continuity of Tocharians will no doubt help us disentangle a great part the ethnolinguistic history of speakers of the Tocharian branch of Proto-Indo-European, from Pre-Proto-Tocharians of Afanasievo to Common Tocharians of the Late Bronze Age/Iron Age eastern Tian Shan.
NOTE. Tocharian’s isolation from the rest of Late PIE dialects and its early and intense language contacts have always been the key to support an early migration and physical separation of the group, hence the traditional association with Afanasievo, a late Repin/early Yamna offshoot. Even with the current incomplete archaeological and genetic picture, there is no other option left for the expansion of Tocharian.
It is not possible to use the currently available ancestry data to map the evolution of Afanasievo ancestry, lacking a proper geographical and temporal transect of Central and East Asian groups. In spite of this, Ning, Wang, et al. (2019) is a huge leap forward, discarding some archaeological models, and leaving only a few potential routes by which Tocharians may have spread southward from the Altai.
NOTE. I have updated the maps of prehistoric cultures accordingly, with colours – as always – reflecting the language/ancestry evolution of the different groups, even though the archaeological data of some groups of Xinjiang remains scarce, so their ethnolinguistic attribution – and the colours picked for them – remain tentative.
Although [the Xinjiang] route is not uniformly agreed upon (Shelach-Lavi 2009: 134–46), this western transmission has been thought to have passed through eastern Kazakhstan, especially as it is manifest in Semireiche, with Yamnaya, Afanasievo (copper) and Andronovo (tin bronze) peoples (Mei 2000: Fig. 3). From Xinjiang this knowledge has been thought to have traveled through the Gansu Corridor via the Qijia peoples (Bagley 1999) and then into territories controlled by dynastic China. The dating of this process is still a problem, as the sites and their contents in Xinjiang are consistently later than those in Gansu, suggesting that the point of contact was in Gansu and that the knowledge then spread from there westward.
1. Eneolithic Altai
The Afanasievo sites, as they are identified in Mongolia, for instance, make up an Eneolithic culture analogous to that of southern Siberia (3100/2500–2000 BCE) in the Upper Yenissei Valley that is characterized by copper tools and an economy reliant on horse, sheep and cattle breeding as well as hunting. (…) The Afanasievo is best known through study of its burials, which typically include groups of round barrows (kurgans), each up to 12 m in diameter with a stone kerb and covering a central pit grave containing multiple inhumations. In their Siberian context, burial pottery types and styles have suggested contacts with the slightly earlier Kelteminar culture of the Aral and Caspian Sea area.
The Afanasievo culture monuments, located in the northern Altai and in the Minusinsk Basin (the western Sayan), have been seen as analogous evidence for cross-Eurasian exchange. These complexes contain small collections of metal, and many of the items are made of brass, although golden, silver and iron ornaments were also identified. A mere one-fourth of these objects are tools and ornaments, while the rest consist of unshaped remains and semi-manufactured objects. Its metallurgical tradition has recently been dated by Chernykh to as early as 3100 to 2700 BCE (1992),making it more compatible chronologically with the early brass-using sites in Shaanxi mentioned above. Kovalev and Erdenebaatar have excavated barrows in Bayan-Ulgii, Mongolia, that have been carbon-dated to the first half of the third millennium BCE and associated by ceramic types and styles and burial patterns with the Afanasievo (Kovalev and Erdenebaatar 2009: 357–58). These mounded kurgans were covered with stone and housed rectangular, wooden-faced tombs that included Afanasievo-type bronze awls, plates and small “leaf-shaped” knife blades (Kovalev and Erdenebaatar 2009: Figs. 6 and 7).
They also excavated sites belonging to the more recently identified Chemurchek archaeological culture, located in the foothills of the Mongolian Altai (Kovalev 2014, 2015) (Fig. 2.6). These sites are carbon-dated to the same period as the Afanasievo burials or to c. 3100/2500–1800 BCE (six barrows in Khovd aimag and four in Bayan-Ulgo aimag). In the rectangular stone kerbed Chemurchek slab burials (Ulaaanhus sum, Bayan-ul’gi aimag and so forth), bronze items included awls; and at Khovd aimag, Bulgan sum, in addition to stone sculptures, three lead and one bronze ring were excavated (Kovalev and Erdenebaatar 2009: Figs. 2 and 3; Fig. 2.6). Although we will not know if they were produced locally until much further investigation is undertaken, these discoveries do document knowledge of various uses and types of metal objects in western and south central Mongolia. The types of metal items thus far recovered are simple tools (awls) and rings (ornamental?) not unlike those associated with Andronovo archaeological cultures as well.
This is a complex circumstance where archaeological evidence is not complete, but raises very important questions about transmission of metallurgical knowledge to and from areas in present-day China. In the 1970s some Afanasievo mounds were excavated in Central Mongolia by a Soviet–Mongolian expedition led by V. V. Volkov and E. A. Novgorodova (Novgorodova 1989: 81–85). Unfortunately, these mounds did not yield metal objects, only ceramics, but they show that the Afanasievo culture with the Eneolithic metallurgical tradition of manufacturing pure copper items had already moved east at least far as central Mongolia. In 2004, Kovalev and Erdenebaatar investigated a large Afanasievo mound, Kulala ula, in the extreme northwest of Mongolia, near the Russian border (Kovalev and Erdenebaatar 2009). There they found a copper knife and awl (Fig. 2.5). There are five C14 dates on wood, coal and human bones from this mound, which belong to the period 2890–2570 BCE. This shows that the Afanasievo culture were carriers of technology and produced artifacts in the first half of the third millennium BCE and that they also moved south along the foothills of the Mongolian Altai. Afanasievo culture in Altai and the Minusinsk basin is dated by C14 to 3600–2500 BCE (Svyatko et al. 2009; Polyakov 2010). In the north of Xinjiang in the Altai district, several typical egg-shaped vessels and two censers of Afanasievo types were found. Some of these have been obtained from the stone boxes (chambers of megalithic graves of the Chemurchek culture) (Kovalev 2011). Thus, the Afanasievo tradition of pure copper metallurgy must have spread to the northern foothills of the Tienshan Mountains no later than the mid-third millennium BCE. The links with Afanasievo and local cultures adjacent to and south of the mountains into present-day China can now be assumed.
2. Bronze Age Altai
Kovalev and Erdenebaatar (2014a) and later Tishkin, Grushin, Kovalev and Munkhbayar (2015) in Western Mongolia conducted large-scale excavations of megalithic barrows of the Chemurchek culture (dated about 2600–1800 BCE). This peculiar culture appeared in Dzungaria and the Mongolian Altai in the second quarter of the third millennium BCE and for some time existed together with the late Afanasievo culture, as evidenced by the findings of Afanasievo ceramics in Chemurchek graves, in the stone boxes. Unfortunately, in China we do not yet know of any metal object related,without doubt, to the Chemurchek culture. Kovalev, Erdenebaatar, Tishkin and Grushin found several leaden ear rings and one ring of tin bronze in three excavated Chemurchek stone boxes (Kovalev and Erdenebaatar 2014a; Tishkin et al. 2015). Such lead rings are typical for Elunino culture,which occupied the entire West Altai after 2400–2300 BCE (Tishkin et al. 2015). This culture had developed a tradition of bronze metallurgy with various dopants, primarily tin. Thus, the tradition of bronze metallurgy as early as this time could have penetrated the Mongolian Altai far to the south. In addition, in the Hadat ovoo Chemurchek stone box, Kovalev and Erdenebaatar discovered stone vessels refurbished with the help of copper “patches,” indicating the presence there of metallurgical production (Fig. 2.7) (Kovalev and Erdenebaatar 2014a). In one of the secondary
Chemurchek graves unearthed by Kovalev and Erdenebaatar in Bayan-Ulgi (2400–2220 BCE), a bronze awl was found (Kovalev and Erdenebaatar 2009). Kovalev and Erdenebaatar also discovered a new culture in the territory of Mongolia (Map 2.3), one that begins immediately after Chemurchek – Munkh-Khairkhan culture (Kovalev and Erdenebaatar 2009, 2014b). To date, about 17 mounds of this culture have been excavated in Khovd, Zavkhan, Khovsgol, Bulgan aimag of Mongolia. This culture dates from about 1800 to 1500 BCE, that is, contemporary with the Andronovo culture. Therefore, the Andronovo culture does not extend far into the territory of Mongolia. Three knives without dedicated handles or stems and five awls have been found in the Munkh-Khairkhan culture mounds (Fig. 2.8). All these products are made of tin bronze. (…) Additionally, eight Late Bronze Age burials (c. 1400–1100 BCE) were unearthed in the Bulgan sum of Khovd aimag and belong to another previously unknown culture called Baitag. And in the Gobi Altai, a new group of “Tevsh” sites dating to the Late Bronze Age were defined in Bayankhongor and South Gobi aimags (Miyamoto and Obata 2016: 42–50). From these Tevsh and Baitag sites, we see the expansion of burial goods to include beads of semiprecious stones (carnelian), bronze beads, buttons and rings and even the famous elaborate golden hair ornaments (Tevsh uul;Bogd sum;Uverkhanagia aimag) from the Baitag barrows (Kovalev and Erdenebaatar 2009: Fig. 5; Miyamoto and Obata 2016).
The major characteristics of Qiemu’erqieke Phase I include:
Burials with two orientations of approximately 20° or 345°.
Rectangular enclosures built using large stone slabs. The size of the enclosure varies from a maximum of 28 x 30 m.*to a minimum of 10.5 x 4.4 m. (Figure 8, Table 2).
*The stone enclosure located near Hayinar is the largest one at approximately 30 x 40 m. based on pacing of the site during a visit by the authors in 2008.
Almost life-sized anthropomorphic stone stelae erected along one side of the stone enclosures (Lin Yun 2008).
Single enclosures tend to contain one or more than one burial, all or some with stone cist coffins.
The cist coffin is usually constructed using five large stone slabs, four for the sides and one on top, leaving bare earth at the base (Zhang Yuzhong 2007). Sometimes the insides of the slabs have simple painted designs (Zhang Yuzhong 2005).
Primary and secondary burials occur in the same grave.
Some decapitated bodies (up to 20) may be associated with the main burial in one cist.
Bodies are commonly placed on the back or side with the legs drawn up.
Grave goods include stone and bronze arrowheads, handmade gray or brown round-bottomed ovoid jars, and small numbers of flat-bottomed jars (Fig. 7).
Clay lamps appear to occur together with roundbottomed jars.
Complex incised decoration on ceramics is common but some vessels are undecorated.
The stone vessels are distinctive for the high quality of manufacture.
Stone moulds indicate relatively sophisticated metallurgical expertise.
Artefacts made from pure copper occur.
Sheep knucklebones (astragali) imply a tradition (as in historical and modern times) of keeping knucklebones for ritual or other purposes. They also indicate the herding of domestic sheep as part of the subsistence economy.
Available evidence suggests that the date range for Qiemu’erqieke Phase I should fall from the later third into the early second millennium BC. There are several reasons to suggest that the time span is around the early second millennium BC. Lin Yun (2008) (…) maintains that the bronze artefacts found in Phase I show a greater sophistication in the level of copper alloy technology than that of the pure copper artefacts common to the Afanasievo tradition. On this basis it might be suggested that the Afanasievo could be considered to be Chalcolithic with a time span across much of the third millennium BC ( Gorsdorf et al. 2004: 86, Fig. 1). Qiemu’erqieke Phase I, however, should more properly be considered as Bronze Age.
Lin Yun also used the bronze arrowhead from burial Ml 7 to narrow down the date of Qiemu’erqieke Phase I. Two arrowheads were found in this burial, one of them leaf shaped with a single barb on the back (Fig. 7:4). A similar arrowhead, together with its casting mould, has been found at the Huoshaogou site of Siba tradition (Li Shuicheng 2005, Sun Shuyun and Han Rufen 1997), in Gansu province, northwest China, dated around 2000-1800 BC (Li Shuicheng and Shui Tao 2000) . This supports a date in the early second millennium BC for the Qiemu’erqieke arrowhead. The painted, round-bottomed jar from the Tianshanbeilu cemetery Qia Weiming, Betts and Wu Xinhua 2008: Fig. 7, bottom left) has been considered as a hybrid between the Upper Yellow River Bronze Age cultures of Siba in northwest China and the steppe tradition of Qiemu’erqieke in west Siberia (Li Shuicheng 1999). If this assumption is correct, the date of Tianshanbeilu, around 2000 BC, can be used as a reference for Qiemu’erqieke Phase I (Jia Weiming, Betts and Wu Xinhua 2008, Lin Yun 2008, Li Shuicheng 1999). Stone arrowheads found in Qiemu’erqieke Phase I also imply that the date is likely to fall within the earlier part of the Bronze Age as no such stone arrowheads have yet been found elsewhere in sites of the Bronze Age in Xinlang dated after the beginning of the second millennium BC.*
*For example Chawuhu and Xiaohe cemeteries (Xinjiang Institute of Archaeology 1999, 2003).
(…) Pottery “oil burners” (goblet-like ceramic vessels, possibly lamps) have been found in three traditions: Afanasievo (Gryaznov and Krizhevskaya 1986:21), Okunevo and Qiemu’erqieke. It is believed that this oil-burner found in Siberia and the Altai is a heritage from the Yamnaya and Catacomb
cultures (Sulimirski 1970: 225, 425; Shishlina 2008:46) in the Caspian steppe further to the west, but does not seem to exist in known Andronovo cultures. The oil-burner tends to disappear after around 2300 BC during the mid-Okunevo period. It is, however, possible that the tradition continues longer in the Qiemu’erqieke sites.
The construction of the stone enclosures also reveals a close connection between Qiemu’erqieke Phase I and the mid and late Okunevo tradition (Sokolova 2007). Slab built stone enclosures emerged in both the Okunevo and Afanasievo traditions (Gryaznov and Krizhevskaya 1986:15-23, Kovalev 2008, Sokolova 2007, Anthony 2007:310, Koryakova and Epimakhov 2007). In the early Afanasievo the enclosure is circular with no cist coffin (Anthony 2007:310, Gryaznov and Krizhevskaya 1986:20), but in the early stage of the Okunevo square stone enclosures with a single cist burial are dominant. Square or rectangular stone enclosures are a marked feature of Qiemu’erqieke Phase I, suggesting temporal relationships between Qiemu’erqieke Phase I and the Okunevo. In Okunevo chronological group II, possibly with influence from the Anfanasievo, circular stone enclosures appeared in combination with rectangular enclosures within individual cemeteries, referred to by Sokolova (2007: table 2) as hybrid examples. By Okunevo chronological group III, rectangular stone slab enclosures with multi-burials emerged again. This is the dominant form in Qiemu’erqieke Phase I. Okunevo burial traditions changed again to single cist burials in the late stage around chronological group V ( Sokol ova 2007). A specific mortuary rite of decapitated burials exists in both the Qiemu’erqieke and Okunevo traditions (Sokolova 2007, Chen Kwang-tzuu and Hiebert 1995), as does the occasional occurrence of painted designs on the interior of the slabs forming the cists ( e.g., Khavrin 1997: 70, fig. 4; 77: tab. IV.5). Based on these comparisons, the date of Qiemu’erqieke Phase I may well parallel that of the Okunevo from at least chronological group II around 2400 BC (Gorsdorf et al. 2004: fig. 1).
In addition to the pottery making tradition, the anthropomorphic stone stelae may also have earlier antecedents. In the Okunevo assemblage there are anthropomorphic stelae that are longer, thinner and more abstract than those of Qiemu’erqieke. There is no indication of such stelae in the Afanasievo tradition (Gryaznov and Krizhevskaya 1986:15-23). However, further to the west, anthropomorphic stone stelae are associated with the Kemi-Oba and Yamnya cultures around the third millennium BC (Telegin and Mallory 1994; Figure 13). Some major characteristics of these stelae such as the icons on the front face of the stelae (Telegin and Mallory 1994:8-9) also appear on stelae found in Qiemu’erqieke Phase I. Recalling the oil burners that may have been inherited from the Yamnya culture and which are found in the Afansievo, Okunevo and Qiemu’erqieke Phase I, it migh t be possible to speculate that Qiemu’erqieke Phase I has its origins even earlier than the first half of the third millennium BC. This idea has also been suggested by Kovalev ( 1999).
Despite the affinities with the Okunevo cultural tradition, Qiemu’erqieke Phase I appears to be a discrete regional variant. The ceramic assemblage shows traits unique to this cluster of sites, while the anthropomorphic stelae are also distinctive markers of this tradition.
It also offered a full summary of findings from prehistoric sites of Xinjiang related to the arrival of a cultural package from the Altai region, ultimately connected to Afanasievo. Relevant excerpts include the following (emphasis mine):
In Bronze Age Xinjiang, burials were diverse but also show some common features between different geographic sections. The main three mountains, including Kunlun Mountains, Tian Shan (mountains) and Altai Mountains, enclose the Tarim Basin, and the Dzungaria Basin, but leave the eastern part of the Tarim Basin and the south-eastern part of the Dzungaria Basin open (with easy access to the surroundings). The Hami Basin is located at the transitional area, connecting the two basins. Burials are mainly spread along the edge of the mountain ranges.
3.1. The Lop Nur region
In the Lop Nur region, the Xiaohe cemetery (2000-1450 BCE) and the Gumugou cemetery (1900-1800 BCE) had many common features shared, and so is the Keliyahe northern cemetery:
Cemeteries were located in sandy areas;
Rectangular/boat-shaped wooden coffins with monuments of wooden planks or poles;
Coffins had no bottoms;
The dead were placed lying straight on the back;
The dead were commonly buried in single graves.
The Gumugou cemetery contained six special sun-radiating-spokes burial pattern in addition to the normal burials, which were similar to the wooden coffin graves of the Xiaohe cemetery.
NOTE. For more on Xiaohe and Gumugou, see the recent post on Proto-Tocharians. See other papers on the Andronovo horizon for other Early to Middle Bronze Age cultural groups less clearly associated with the Xiaohe horizon, like Hazandu, Xintala, or the Chust culture.
An assemblage of early bronzes had been recovered from northwestern Xinjiang and the periphery of Dzungaria 准噶尔 Basin. It comprises a variety of utilitarian tools and weapons, and a small number of apparels. These artifacts bear the stamps of Andronovo Culture in form, artifact type and decorative pattern. The metallographic analysis on selected artifacts indicates that they comprise mainly of tin-bronzes that contain 2–10% of tin. Moreover, the chemical compositions of these artifacts are similar to that of the Andronovo Culture. Latter date (first half of the 1st millennium BC) artifacts of the assemblage include a small number of arsenic bronzes. In all, during the period between the mid-2nd and mid-1st millennium BC, copper and bronze artifacts coexisted in this region, albeit tin-bronze comprised the majority. The composition of alloy did not show significant change over time. Some colleagues pointed out that the Nulasai 奴拉赛 site at Nileke 尼勒克 County in the Yili 伊犁 River basin of Xinjiang was the pioneer in the use of “sulphuric ore–ice copper–copper”technology. It is also the only early smelting site in Euro-Asia that arsenic ore was added to deliberately produce an alloy
The Hami Basin-the Balikun Grassland area is located at the eastern part of Tian Shan. The area is divided in a northern basin and a southern basin by the east-west stretch of the Tian Shan. In the Hami Basin-the Balikun Grassland area, the main type of burials were earth-pit graves in the early Bronze Age, and burials of stone-pit with barrows became more common in the late Bronze Age. The Hami-Tianshan-Beilu cemetery is a representative of the earth-pit graves. The features of the Hami-Tianshan-Beilu cemetery (2000-1500 bce) here were:
Rectangular earth pit graves;
The dead were often in a hocker position lying on one side;
Commonly a single dead in one grave.
The Hami-Wubu cemetery (earlier than 1000 bce) and the Yanbulake cemetery (1200-600 bce) are representatives of another common earth-pit graves. Common features here were:
Rectangular earth pits, with two storeys and/or roofed with wooden boards;
The dead was placed in a hocker position lying on one side;
Mostly a single dead in one grave.
Later there appeared more stone-pit graves in this area, and the features can be summarized as:
Round burial mounds, commonly constructed by stones or a mix of stones and earth;
Burial mounds with a sunken top or a normal (dome) top;
The diameter of the burial mounds varied between 3 and 25.4 m (but not necessarily limited in this scope);
Circular or rectangular stone kerbs;
Rectangular stone pits, constructed by earth, or stones, or a mix of earth and stones;
Rectangular stone pits contained wooden coffins (represented by the Yiwu Baiqi’er cemetery).
In the Hami Basin, the Bronze Age cemeteries show common burial features like earth pits and hocker position of the dead. With similar pottery styles in the Hami-Tianshan-Beilu cemetery to those in the Machang and Siba cultures (Xinjiang 2011: 17), it suggests possible cultural influence or people’s migrating from the Hexi Corridor in the east.
In the Balikun Grassland, burials in an earlier time contained mostly earth-pit graves but also a small number of stone-pit graves. The pebbles were imbedded in the floors and the walls of the graves in a rectangular shape, e.g. the Balikun-Nanwan cemetery (1600-1000 bce). In a later time, there appeared huge burial mounds with a sunken top, and with the diameters of the burial mounds varying from 3 to 25.4 m, e.g. the Balikun-Dongheigou cemetery and the Balikun-Heigouliang cemetery. The Yiwu-Bai’erqi and the Yiwu-Kuola cemeteries contained either round stone burial mounds or circular stone kerbs on the ground surface. Considering the three burial elements including burial mounds, stone pits and circular kerbs, the later period cemeteries in the Balikun Grassland were actually similar to cemeteries from the southern edge of the Altai Mountain area.
The Nanwan 南湾 cemetery site at Kuisu 奎苏 Town, Balikun 巴里坤 (1600–1100 BC) also yielded an assemblage of early bronzes. The style of its early phase artifacts is similar to that of the burials distributed in the North Tianshan Route. Some sorts of cultural connection should have existed between the two.
The dates of Yanbulake 焉不拉克 Culture (1300–700 BC) are comparatively late. Its metallurgy was a continuation of the western China tradition. Artifact types include a variety of utilitarian tools, weapons and apparels.
3.3. The Turpan Basin-the middle part of Tian Shan
Turpan Basin is located at the western part of the Hami Basin, and lies at the southern edge of the eastern Tian Shan. In the Turpan Basin-the middle part of Tian Shan area, the main representative of the Bronze Age cemeteries is the Yanghai Nr.1 cemetery. The features here were:
Elliptic earth pit graves, commonly covered by round logs on the top;
Some graves contained burial beds made of round logs or reeds;
The dead were mainly placed lying straight on the back;
Mostly a single dead in one grave.
In Iron Age, the stone burials became dominant, but the stone burials varied in different regions of the Turpan Basin-the middle part of Tian Shan area. Graves containing burial mounds, stone pit, and circular stone kerbs are represented by the Shanshan-Ertanggou cemetery, the Tuokexun-Alagou cemetery, the Urumqi-Chaiwobu cemetery and the Urumqi-Yizihu-Sayi cemetery, etc. The stone funeral construction features here are similar to those contemporary cemeteries in the Hami Basin-the Balikun Grassland area.
3.4. The southern edge of the western and middle part of Tian Shan
In the southern edge of the western and middle part of Tian Shan area, the main representatives of the late Bronze Age cemeteries are the Hejing-Chawuhu Nr.4 cemetery (around 1000-500 bce), the Hejing-Xiaoshankou cemetery, the Baicheng-cemetery, etc. The main burial features of the late Bronze Age and the early Iron Age cemeteries (see Fig.12) here were:
Burial mounds, constructed by stones or a mix of stones and earth;
Irregular circular or rectangular stone kerbs;
Stone pit graves in a bell-shape or a rectangular shape;
Stone pit graves constructed by imbedding pebbles or stone slabs in walls and floors;
The dead were often placed lying on their back with bent legs;
The dead were commonly reburied a second time with multiple burials.
From the late Bronze Age to the early Iron Age in this area, the burial traditions tended to be in a more varied way. In the stone burials with stone kerbs, there is a mixture of stone pit and earth pit graves. The burial features of the Iron Age cemeteries in this section were similar to those contemporary both in the Hami Basin-the Balikun Grassland area and in the Turpan Basin-the middle part of Tian Shan area.
The Chawuhu 察吾呼 Culture (1100–500 BC) distributes on the foothills between the middle section of the Tianshan Mountain Ranges and Tarim River. Its bronze assemblage comprises a variety of weapons, utilitarian tools and small apparels. They show no apparent temporal change in form and type through the four cultural phases. In addition, bronzes bear the Chawuhu characteristics were found in Hejing 和静, Baicheng 拜城 and Luntai 轮台 (Bügür). Yet, sites distributed along the Tarim River, such as Heshuo 和硕, Kuga 库车and Aksu 阿克苏, yielded remains of a bronze culture different from that of Chawuhu. Bronzes recovered include double-eared socketed axe, arrowheads, awls, knives, needles and bracelets. Their absolute dates have been estimated to be earlier than that of Chawuhu.
A typical Bronze Age cemetery from the Pamir Plateau area is the Tashenku’ergan-Xiabandi cemetery (around 1000-500 bce). The burial features here were:
Mainly inhumations, but also a few cremations;
Burial mounds, constructed of stones;
Irregular circular or rectangular stone kerbs;
Mostly a single dead in one grave;
The dead was placed in a hocker position lying on one side.
The adoption of burial customs from the east supports the migration of Afanasievo-related peoples from the Tian Shan up to the Pamir Plateau, strongly influencing the findings of the Xiabandi cemetery, which has been dated from an early Bronze Age phase (ca. 1500-300 BC) to a late date up to ca. 600 BC.
While it is today unclear how far the Afanasievo admixture reached into the western Xinjiang, it seems that the Pamir Plateau remained culturally connected to neighbouring Andronovo-related cultures in pottery and metallurgical innovations, hence their language probably belonged – during most part of the Bronze and Iron Ages – to the Indo-Iranian branch, even though specific dialects might have changed with each new attested group.
In particular, it is possible that the early Andronovo groups related to the Xiaohe Horizon spoke Indo-Aryan or West Iranian dialects, while Saka-related groups replaced them – or an intermediate Tocharian-speaking group – with East Iranian dialects. A close interaction with West Iranian would justify the known ancient borrowings of Tocharian, although they could also be explained by contacts with Chust-related groups farther west. For more on this, see Ged Carling’s work on the different layers of Iranian loans.
In the early Bronze Age, there are distinct regional differences in the burial customs in and surrounding the Tarim Basin. At the southern edge of the Altai Mountains area, the burial customs included stone burial mounds, stone pit graves, circular or rectangular stone kerbs and stone human sculptures; the dead were placed lying straight on the back. In the Hami Basin-the Balikun Grassland area, the burial customs included earth pit graves; the dead were placed in a hocker position lying on one side. In the Turpan Basin-the middle part of Tian Shan area, the burial customs included earth pit graves; the dead were placed lying straight on the back. In the Lop Nur region, the burial customs included wooden coffins buried in sand; the dead were placed lying straight on the back.
But from the late Bronze Age to the early Iron Age, there was a common shift in burial customs from earth pit graves to stone burials in the Hami Basin-the Balikun Grassland area and in the Turpan Basin-the middle part of Tian Shan area. The main features of the stone burials include stone burial mounds, circular or rectangular stone kerbs, and the stone pit graves in the cemeteries. Similar stone burial customs commonly appeared at the southern edge of the western and middle part of Tian Shan area and the Pamir Plateau area in Iron Age. The burial features in most areas are in a mixture of both the earth pit graves and stone pit graves, especially in the Hami Basin-the Balikun Grassland area and the Turpan Basin-the middle part of Tian Shan area.
Historians of metallurgy conducted metallographic analyses on a sample of 234 metal specimens recovered from 16 localities in eastern Xinjiang. They concluded that the metallurgic industry in eastern Xinjiang could be roughly partitioned into three developmental phases. The early phase is represented by the burials distributed in the North Tianshan Route. The majority of the metal assemblage was tin-bronzes; however, copper and arsenic-bronzes maintained considerable proportions. The middle phase is represented by the burials at Yanbulake. During this phase, tin-bronze still maintained the majority; the proportion of arsenic-bronze increased, and some of them were high arsenic-bronzes. The late phase is represented by the burials at Heigouliang 黑沟梁. The composition of lead increased in the bronze alloy in the expense of arsenic. In addition, this phase witnessed the appearance of high tin-bronze that composed up to 16% of tin and the appearance of brass, that is, an alloy of copper and zinc. The bronze alloy consistently contained significant amount of impurities regardless of temporal difference. Casting and forging technologies coexisted throughout the three phases. The early bronzes (2000–500 BC) of eastern Xinjiang, in general, contained arsenic; however, the composition of arsenic was usually under 8%, but a few artifacts contained more than 20% arsenic. In all, arsenic had long been used in the alloy-forming of the early bronzes in eastern Xinjiang. Consequently, arsenic-bronzes were widely found in the prehistoric archaeology of the region. The artifact types, chemical compositions and manufacture techniques of the bronze assemblage of the burials of the North Tianshan Route are similar to those of Siba Culture, indicating that eastern Xinjiang had played a significant role in the East-West interactions.
An assemblage of early bronzes had been recovered from northwestern Xinjiang and the periphery of Dzungaria 准噶尔 Basin. It comprises a variety of utilitarian tools and weapons, and a small number of apparels. These artifacts bear the stamps of Andronovo Culture in form, artifact type and decorative pattern. The metallographic analysis on selected artifacts indicates that they comprise mainly of tin-bronzes that contain 2–10% of tin. Moreover, the chemical compositions of these artifacts are similar to that of the Andronovo Culture. Latter date (first half of the 1st millennium BC) artifacts of the assemblage include a small number of arsenic-bronzes. In all, during the period between the mid-2nd and mid-1st millennium BC, copper and bronze artifacts coexisted in this region, albeit tin-bronze comprised the majority.
Tocharians in population genomics
Prehistoric population movements between the Altai and the Tian Shan are difficult to pinpoint, not the least because of the division of these territories among three different countries and their archaeological teams, only recently (more) open to the international scholarship.
The available schematic archaeological picture, where migrations could only be roughly inferred, has been recently updated to a great extent by Ning, Wang et al. (2019), whose genetic analysis of the samples is as thorough as anyone could have asked for, with a level of detail which matches the complex genetic picture of the region by the Iron Age.
As a summary, here is what they described about the samples from Shirenzigou (ca 400-200 BC), corresponding to the Iron Age populations of the Hami Basin-the Balikun Grassland area, and closely related to the preceding Yanbulake Culture:
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.
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.
In the PCA, ADMIXTURE, outgroup f3 statistics [see Figure S4], as well as f4 statistics (Table S3), we observed the Shirenzigou individuals were closer to the present day Tungusic and Mongolic-speaking populations in northern Asia than to the populations in central and southern China, suggesting the northern populations might contribute more to the Shirenzigou individuals. Based on this, 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
It is unclear whether the Chemurchek population will show a sizeable local contribution from neighbouring groups. The fact that Okunevo shows 20% Yamnaya-related ancestry strongly supports the nature of neighbouring stone-grave-building peoples of the Altai and the northern Tian Shan as mostly Afanasievo-like, and the apparent lack of contributions of Srubna/Andronovo-like ancestry in the early Hami-Balikun stone burial builders also speaks for radical population replacement events reaching the areas south of Tian Shan, at least initially.
While ancestry cannot settle linguistic questions, it seems that nomads of the Gansu and Qinghai grasslands retained an ancestry close to Andronovo, whereas nomads of the Hami Basin-Balikun grasslands and related populations of Xinjiang remained closely related to Afanasievo. This doesn’t preclude that the ancestors of the Yuezhi became acculturated under the influence of peoples from eastern Xinjiang, but all data combined suggest an isolation of both populations – relative to other groups and to each other – and it is therefore more likely that they spoke Indo-Iranian-related languages rather than a language of the Tocharian branch.
In an interesting twist of events, despite the initially reported hg. R1b and Q, Tocharians from Shirenzigou actually show a haplogroup diversity comparable to that attested in other late Iron Age populations: a similar diversity is seen, for example, among Germanic, Baltic, and Balto-Finnic peoples of the Baltic region; among East Germanic or Scythians of the north Pontic region; or among Mediterranean peoples sampled to date. Iron Age peoples show thus a complex sociopolitical setting that overcame the previous patrilineal homogeneity of Bronze Age expansions.
M15-2 (with Han-related ancestry) is of the rare haplogroup Q1a-M120, while the samples with highest Steppe_MLBA-related ancestry are of hg. R1b-PH155, which points to their recent origin among Yuezhi, or to Hun-related populations showing an admixture related to the proto-historic nomads of the Gansu and Qinghai grasslands.
The expansion of Chemurchek-related peoples was probably associated more with hg. Q1a (dubious if it’s a Pre-ISOGG 2017 nomenclature, hence possibly Q1b), a haplogroup that might be found in Khvalynsk as a “significant minority” according to Anthony (2019), and it might also be attested in sampled individuals from Afanasievo in its late phase. This might be, therefore, a case similar to the early expansion of Indo-Europeans with R1b-V1636 lineages through the Volga – North Caucasus region, and of the later expansion with I2a-L699 lineages into the Balkans.
Haplogroup Q1a2-M25 is found in individual X3, whose Steppe ancestry is likely a combination of Afanasievo plus Andronovo-like ancestry heavily admixed with Hezhen/Ulchi-like populations, in line with the expected recent contacts with the neighbouring Xiongnu, Yuezhi, and other population movements affecting eastern Xinjiang.
Sample M4, which packs the most Afanasievo-like ancestry, is of hg. R1a-Z645, which – like sample M8R1 of hg. O – is most likely related to haplogroup resurgence events of local populations, which left the predominant Afanasievo-like admixture brought by builders of stone burials essentially intact, evidenced by the almost 100% of R1a found in the Xiaohe cemetery – and in most of the early Andronovo horizon – and among expanding Kangju and Wusun, as well as by the prevalence of hg. O among sampled East Asian populations.
A question that will only be answered with more samples is how and when the prevalent R1b-L23 and Q1b lineages among Afanasievo-related peoples began to be replaced to reach the high variability seen in Shirenzigou. Given the pastoralist nature of peoples around Tian Shan, the succeeding expansions of Proto-Tocharians, and the late isolation of different Common Tocharian groups, it is more than likely that this variability represents a late and local phenomenon within Xinjiang itself.
An early split of Pre-Proto-Tocharians with R1b-rich Afanasievo peoples migrating into Central Asia, evolving into these builders of stone burial constructions of Xinjiang, with a center of gravity during the Late Bronze Age and Iron Age in the Hami Basin-Balikun Grassland area, where the Shirenzigou nomads have been sampled.
A late, long-term admixture of R1b-rich Poltavka herders of Yamnaya ancestry with incoming R1a-rich Abashevo chiefs of Corded Ware ancestry to form the Sintashta-Potapovka-Filatovka community, representing the transition of a conservative Pre-Proto-Indo-Iranian into the heavily Uralic-influenced Proto-Indo-Iranian community (see here).
Just like the East Bell Beaker expansion from Yamnaya Hungary has confirmed that Corded Ware peoples did not partake in spreading Indo-European languages (spreading Uralic languages instead), data on the expansion of Tocharian speakers from Afanasievo to the Tian Shan was always there; population genomics is merely helping to connect the dots.
In summary, genetic research is supporting the expected linguistic expansions of the Neolithic and Bronze Age step by step, slowly but surely.
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.
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.
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.
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 hypothetic “Centum” Temematic substrate that would explain everything his model can’t…
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 the early outlier Turlojiškė1932 (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.
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 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?
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.
In summary, 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. Another, more complicated question, though, is if this expanding Proto-Baltic language accompanying agriculture represents the extinct
early Proto-Baltic dialect from which Balto-Finnic borrowed words, hence Proto-Baltic proper expanded later, or if this early Baltic branch could have been part of the Trzciniec expansion. Again, the answer in archaeological and genetic terms seems to be the former. For a more detailed discussion of this and more, 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.
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.
The recent data on ancient DNA from Iberia published by Olalde et al. (2019) was interesting for many different reasons, but I still have the impression that the authors – and consequently many readers – focused on not-so-relevant information about more recent population movements, or even highlighted the least interesting details related to historical events.
This post is thus a summary of its findings with the help of natural neighbour interpolation maps of the reported Germany_Beaker and France_Beaker ancestry for individual samples. Even though maps are not necessary, visualizing geographically the available data facilitates a direct comprehension of the most relevant information. What I considered key points of the paper are highlighted in bold, and enumerated.
NOTE. To get “more natural” maps, extrapolation for the whole Iberian Peninsula is obtained by interpolation through the use of external data from the British Isles, Central Europe, and Africa. This is obviously not ideal, but – lacking data from the corners of the Iberian Peninsula – this method gives a homogeneous look to all maps. Only data in direct line between labelled samples in each map is truly interpolated for the Iberian Peninsula, while the rest would work e.g. for a wider (and more simplistic) map of European Bronze Age ancestry components.
The Proto-Beaker package may or may not have expanded into Central Europe with typical Iberia_Chalcolithic ancestry. A priori, it seems a rather cultural diffusion of traits stemming from west Iberia roughly ca. 2800 BC.
The situation during the Chalcolithic is only relevant for the Indo-European question insofar as it shows a homogeneous Iberia_Chalcolithic-like ancestry with typical Y-chromosome (and mtDNA) haplogroups of the Iberian Neolithic dominating over the whole Peninsula until about 2500 BC. This might represent an original Basque-Iberian community.
(1) East Bell Beakers brought hg. R1b-L23 and Yamnaya ancestry to Iberia, ergo the Bell Beaker phenomenon was not a (mere) local development in Iberia, but involved the expansion of peoples tracing their ancestry to the Yamnaya culture who eventually replaced a great part of the local population.
(2) Classical Bell Beakers have their closest source population in Germany Beakers, and they reject an origin close to Rhine Beakers (i.e. Beakers from the British Isles, the Netherlands, or northern France), ergo the Single Grave culture was not the origin of the Bell Beaker culture, either (see here).
Early Bronze Age
Interestingly, the European Early Bronze Age in Iberia is still a period of adjustments before reaching the final equilibrium. Unlike the situation in the British Isles, where Bell Beakers brought about a swift population replacement, Iberia shows – like the Nordic Late Neolithic period – centuries of genomic balancing between Indo-European- and non-Indo-European-speaking peoples, as could be suggested by hydrotoponymic research alone.
This balancing is seen in terms of Germany_Beaker vs. Iberia_Chalcolithic ancestry, but also in terms of Y-chromosome haplogroups, with the most interesting late developments happening in southern Iberia, around the territory where El Argar eventually emerged in radical opposition to the Bell Beaker culture.
We obtained lower proportions of ancestry related to Germany_Beaker on the X-chromosome than on the autosomes (Table S14), although the Z-score for the differences between the estimates is 2.64, likely due to the large standard error associated to the mixture proportions in the X-chromosome.
Regarding the PCA, Iberia Bronze Age samples occupy an intermediate cluster between Iberia Chalcolithic and Bell Beakers of steppe ancestry, with Yamnaya-rich samples from the north (Asturias, Burgos) representing the likely source Old European population whose languages survived well into the Roman Iron Age:
Middle Bronze Age
During the Middle Bronze Age, the equilibrium reached earlier is reversed, with a (likely non-Indo-European-speaking) Argaric sphere of influence expanding to the west and north featuring Iberia Chalcolithic and lesser amount of Germany_Beaker ancestry, present now in the whole Peninsula, although in varying degrees.
All Iberian groups were probably already under a bottleneck of R1b-DF27 lineages, although it is likely that specific subclades differed among regions:
Late Bronze Age
The Late Bronze Age represents the arrival of the Urnfield culture, which probably expanded with Celtic-speaking peoples. A Late Bronze Age transect before their genetic impact still shows a prevalent Germany_Beaker-like Steppe ancestry, probably peaking in north/west Iberia:
(5) Galaico-Lusitanians were descendants of Iberian Beakers of Germany_Beaker ancestry and hg. R1b-M269. Autosomal data of samples I7688 and I7687, of the Final Bronze (end of the reported 1200-700 BC period for the samples), from Gruta do Medronhal (Arrifana, Coimbra, Portugal) confirms this.
In the 1940s, human bones, metallic artifacts (n=37) and non-human bones were discovered in the natural cave of Medronhal (Arrifana, Coimbra). All these findings are currently housed in the Department of Life Sciences of the University of Coimbra and are analyzed by a multidisciplinary team. The artifacts suggest a date at the beginning of the 1st millennium BC, which is confirmed by radiocarbon date of a human fibula: 890–780 cal BCE (2650±40 BP, Beta–223996). This natural cave has several rooms and corridors with two entrances. No information is available about the context of the human remains. Nowadays these remains are housed mixed and correspond to a minimum number of 11 individuals, 5 adults and 6 non-adults.
NOTE. To understand how the region around Coimbra was (Proto-)Lusitanian – and not just Old European in general – until the expansion of the Turduli Oppidani, see any recent paper on Bronze Age expansion of warrior stelae, hydrotoponymy, anthroponymy, or theonymy (see e.g. about Spear-vocabulary).
In a complex period of multiple population movements and language replacements, the temporal transect in Olalde et al. (2019) offers nevertheless relevant clues for the Pre-Roman Iron Age:
(6) The expansion of Celtic languages was associated with the spread of France_Beaker-like ancestry, most likely already with the LBA Urnfield culture, since a Tartessian and a Pre-Iberian samples (both dated ca. 700-500 BC) already show this admixture, in regions which some centuries earlier did not show it. Similarly, a BA sample from Álava ca. 910–840 BC doesn’t show it, and later Celtiberian samples from the same area (ca. 4th c. BC and later) show it, depicting a likely north-east to west/south-west routes of expansion of Celts.
(7) The distribution of Germany_Beaker ancestry peaked, by the Iron Age, among Old Europeans from west Iberia, including Galaico-Lusitanians and probably also Astures and Cantabri, in line with what was expected before genetic research:
A probably more precise picture of the Final Bronze – Early Iron Age transition is obtained by including the Final Bronze samples I2469 from El Sotillo, Álava (ca. 910-875 BC) as Celtic ancestry buffer to the west, and the sample I3315 from Menorca (ca. 904-861 BC), lacking more recent ones from intermediate regions:
In terms of Y-DNA and mtDNA haplogroups, the situation is difficult to evaluate without more samples and more reported subclades:
In the PCA, Proto-Lusitanian samples occupy an intermediate cluster between Iberian Bronze Age and Bronze Age North (see above), including the Final Bronze sample from Álava, while Celtic-speaking peoples (including Pre-Iberians and Iberians of Celtic descent from north-east Iberia) show a similar position – albeit evidently unrelated – due to their more recent admixture between Iberian Bronze Age and Urnfield/Hallstatt from Central Europe:
(8) Iberian-speaking peoples in north-east Iberia represent a recent expansion of the language from the south, possibly accompanied by an increase in Iberia_Chalcolithic/Germany_Beaker admixture from east/south-east Iberia.
(9) Modern Basques represent a recent isolation + Y-DNA bottlenecks after the Roman Iron Age population movements, probably from Aquitanians migrating south of the Pyrenees, admixing with local peoples, and later becoming isolated during the Early Middle Ages and thereafter:
[Modern Basques] overlap genetically with Iron Age populations showing substantial levels of Steppe ancestry.
Assuming that France_Beaker ancestry is associated with the Urnfield culture (spreading with Celtic-speaking peoples), Vasconic speakers were possibly represented by some population – most likely from France – whose ancestry is close to Rhine Beakers (see here).
Alternatively, a Vasconic language could have survived in some France/Iberia_Chalcolithic-like population that got isolated north of the Pyrenees close to the Atlantic Façade during the Bronze Age, and who later admixed with Celtic-speaking peoples south of the Pyrenees, such as the Vascones, to the point where their true ancestry got diluted.
In any case, the clear Celtic Steppe-like admixture of modern Basques supports for the time being their recent arrival to Aquitaine before the proto-historical period, which is in line with hydrotoponymic research.
The most interesting aspects to discuss after the publication of Olalde et al. (2019) would have been thus the nature of controversial Palaeohispanic peoples for which there is not much linguistic data, such as:
the Astures and the Cantabri, usually considered Pre-Celtic Indo-European (see here);
the Vaccaei, usually considered Celtic;
the Vettones, traditionally viewed as sharing the same language as Lusitanians due to their apparent shared hydrotoponymic, anthroponymic, and/or theonymic layers, but today mostly viewed as having undergone Celticization and helped the westward expansion of Celtic languages (and archaeologically clearly divided from Old European hostile neighbours to the west by their characteristic verracos);
the Pellendones or the Carpetani, who were once considered Pre-Celtic Indo-Europeans, too;
the nature of Tartessian as Indo-European, or maybe even as “Celtic”, as defended by Koch;
or the potential remote connection of Basque and Iberian languages in a common trunk featuring Iberian/France_Chalcolithic ancestry (also including Palaeo-Sardo).
Despite these interesting questions still open for discussion, the paper remarked something already known for a long time: that modern Basques had steppe ancestry and Y-DNA proper of the Yamnaya 5,000 years ago, and that Bell Beakers had brought this steppe ancestry and R1b-P312 lineages to Iberia. This common Basque-centric interpretation of Iberian prehistory is the consequence of a 19th-century tradition of obsessively imagining Vasconic-speaking peoples in their medieval territories extrapolated to Cro-Magnons and Atapuerca (no, really), inhabiting undisturbed for millennia a large territory encompassing the whole Iberia and France, “reduced” or “broken” only with the arrival of Celts just before the Roman conquests. A recursive idea of “linguistic autochthony” and “genetic purity” of the peoples of Iberia that has never had any scientific basis.
Similarly, this paper offered the Nth proof already in population genomics that traditional nativist claims for the origin of the Bell Beaker folk in Western Europe were wrong, both southern (nativist Iberian origin) and northern European (nativist Lower Rhine origin). Both options could be easily rejected with phylogeography since 2015, they were then rejected in Olalde et al. and Mathieson et al (2017), then again with the update of many samples in Olalde et al. (2018) and Mathieson et al (2018), and it has most clearly been rejected recently with data from Wang et al. (2018) and its Yamnaya Hungary samples. Findings from Olalde et al. (2019) are just another nail to coffins that should have been well buried by now.
Even David Anthony didn’t have any doubt in his latest model (2017) about the Carpathian Basin origin of North-West Indo-Europeans (see here), and his latest update to the Proto-Indo-European homeland question (2019) shows that he is convinced now about R1b bottlenecks and proper Pre-Yamnaya ancestry stemming from a time well before the Bell Beaker expansion. This won’t be the last setback to supporters of zombie theories: like the hypotheses of an Anatolian, Armenian, or OIT origin of the PIE homeland, other mythical ideas are so entrenched in nationalist and/or nativist tradition that many supporters will no doubt prefer them to die hard, under the most numerous and shameful rejections of endlessly remade reactionary models.
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.
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.
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:
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;
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.
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.
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.
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.
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 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.
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.
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 Proto-Indo-Europeans, and the other one – Corded Ware or Steppe_MLBA ancestry – expanding with Uralic speakers.
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.
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.
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.
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 (…)
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.
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
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.
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:
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.
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:
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:
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.
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.
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.
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:
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:
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.
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.
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:
There is scarce data on Palaeo-Balkan movements yet, although it is known that:
Yamnaya ancestry appears among Mycenaeans, with the Yamnaya Bulgaria sample being its best current ancestral fit;
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.
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.
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:
At the end of the Afanasevo culture, at least three samples show hg. Q1b (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 (potentially Q1b in the newer nomenclature) 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.
Okunevo, which replaced Afanasevo in the Altai region, shows a majority of hg. Q1b, 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 Q1b, and hg. Q reported from Elunino, Sagsai, Khövsgöl, and also among peoples of the Srubna-Andronovo horizon (the Krasnoyarsk MLBA outlier), and in Karasuk.
(…) 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.
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.
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:
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.
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:
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.
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.
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).
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.
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:
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.
Baikal HG ancestry
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-populationsexpanded 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:
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:
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.
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.
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.
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.
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:
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:
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.
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.
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.
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 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.).
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.
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.
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.
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.
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 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.
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.
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.
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.
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.
(…) ‘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.
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.
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.
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.
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:
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.