Predictions about the genetic change from Single Grave to the Late Neolithic in Denmark

germanic-early-bronze-age

New open access paper Mapping human mobility during the third and second millennia BC in present-day Denmark by Frei et al. PLOS One (2019), from the Copenhagen group (including Allentoft, Sikora, and Kristiansen) of samples whose genomic profile will probably be published soon.

Interesting excerpts (emphasis mine):

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.

denmark-late-neolithic-bronze-age-sites
Map of present-day Denmark illustrating locations of the burial sites.
The dashed black line marks the maximum advance stage of the last glaciation (Weichselian). Drafted with public domain data from Natural Earth (https://www.naturalearthdata.com).

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.

strontium-isotope-denmark-neolithic-bronze-age
Diagram plotting results of strontium isotope ratios versus calibrated radiocarbon dates of the individuals investigated. The grey band shows the “local” baseline.

Middle Neolithic

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).

denmark-late-neolithic-bronze-age
Strontium isotope, 14C results and sex and age determinations from individuals from the 3rd and 2nd millennia BC from Denmark presented in chronological order. Modified from the paper, see full image.

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) [56]. On the basis of the presence of this ring-headed pin, Lomborg [56] 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.

strontium-isotope-denmark-late-neolithic-bronze-age
Strontium isotope results of the 88 investigated individuals including Late Bronze Age individuals investigated previously. The grey band shows the “local” baseline.

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:

  1. 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.
  2. 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.
  3. 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.

All this will support, once again, the expansion of Bell Beakers from Yamnaya settlers of Central-East Europe. That is probably what I will be reporting about the data as related to the Pre-Proto-Germanic homeland of the Northern European Plain, unless there is some big surprise, for example that R1b-U106 expanded later from Northern Germany, more clearly associated with later Barbed Wire Beakers or even Únětice movements, although I find this very unlikely at this point.

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. [3]. 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…

Related

Mitogenomes show discontinuity in Gotland’s LN – EBA transition

New paper (behind paywall) The stone cist conundrum: A multidisciplinary approach to investigate Late Neolithic/Early Bronze Age population demography on the island of Gotland, by Fraser et al. J. Archaeol. Sci. (2018) 20:324-337.

Interesting excerpts (emphasis mine):

Unfortunately, due to poor preservation, mitochondrial haplotype calls were only obtained from the EBA individuals in this study. However, some interesting findings were observed. We find two adult local individuals with unique haplogroup lineages [H1a, H1e], and two juvenile individuals with haplogroup lineages [H2a and T1a] previously found exclusively in the CWC individuals analyzed here, all four dated to BA I showing that new lineages had already been established on Gotland at this time period. Another unique haplogroup lineage [K1b] is found in the child dated to BA III at Suderkvie, indicating continued migration to the island. Additionally, the two nonlocal individuals [LN II; ans010 and BA I; hgb010], were adding to the already existing haplogroup lineages [U5b and T2b, respectively] previously found in the PWC individuals analyzed here. The T2b lineage has also been found in the TRB individuals from the dolmen (Table S4).

Our ancestral contribution modeling for the maternal lineages showed that, among the models tested, the only models with a good fit were 55/45 CWC/TRB contribution (Fig. 6), or a 3-way mixture of 55% CWC, 40% TRB, and 5% PWC (Fig. S15). All other models had poor support, including the models with 100% contribution from either group present on Gotland in the preceding period. The PWC group had the weakest fit of all models which is quite surprising as the PWC was the only preceding group that was established on the island in the beginning of the LN period. Although separated temporally by more than 500 years, the individuals from the Hägur burial seems to have been well established in the Eksta area which previously was inhabited by PWC groups. However, the new haplogroup lineages [H1a and H2a] in the Hägur burial suggest some migration to Gotland in a period succeeding the PWC. Archaeologically, it is difficult to reconcile a 55/45 CWC/TRB contribution on Gotland as the temporal range of the TRB culture ends around c. 2700 cal BCE, and presently there is little archaeological evidence of assimilation of TRB and BAC/CWC on Gotland during the latter part of the MN period (e.g. Andersson, 2016). The apparent decline of human activity on the island post TRB, and also later postPWC is intriguing. As we do not see cultural assimilation of TRB and PWC on Gotland one can only speculate as to why TRB disapears from the archaeological record.

The introduction of new female lineages and the mtDNA haplogroup variation within these stone cist burials, together with an increase of nonlocal individuals, and a dietary shift, indicates that a demographic event has happened. The LN period shows traces of activity all over the island compared to the MN period with ten TRB, and eighteen PWC sites (e.g. Bägerfeldt, 1992; Luthander, 1988; Wallin, 2010; Österholm, 1989). This pattern seems to continue into the EBA as seen by the well-established local groups identified by their Sr-signals, as well as the monumental burials.

gotland-funnel-beaker-culture
Fig. 1. Map indicating distribution of TRB-North group megalithic tombs (Blomqvist, 1989; Midgley, 2008; Sjögren, 2003; Tilley, 1999) and PWC areas (Larsson, 2009) modified from (Malmström et al., 2009). Swedish megalithic TRB burial sites included in the analyses: 1. Gökhem passage grave, Falköping, Västergötland, 2. Alvastra dolmen, Östergötland, 3. Mysinge passage grave, Resmo, Öland, 4. Ansarve dolmen, Tofta, Gotland, and 5. the Ostorf TRB burial ground, Mecklenburg-Vorpommern, Germany. From another recent paper by Fraser et al. (2017)

From the conclusions:

We find a shift in population demography compared to the preceding cultural developments on the island recorded from the Neolithic TRB, and sub-Neolithic PWC groups. We find that these burials were used by local groups that were well established in the regions where the burials were situated. These individuals also displayed a different dietary pattern than that noted for the preceding TRB and PWC groups on the island. We also detect sporadic reuse of the MN TRB dolmen in the LN period by nonlocal individuals, who also shows deviating dietary patterns to the LN/EBA individuals in the stone cist burials.

We see an increase of new mitochondrial lineages in the EBA individuals, of which some also were noted in the CWC reference dataset used in this study. Our modeling for maternal ancestry suggests a 3-way model of 55% CWC, 40% TRB, and 5% PWC. Given the broad absence of archaeological evidence for the typical BAC/CWC burials, as well as no archaeological evidence of assimilation of TRB and PWC during the latter part of the MN period on Gotland, it seems probable that the major process of admixture did not occur on the island. Instead, the data indicates an admixture process that occurred elsewhere and prior to migrating to Gotland. Thus, our results suggest later migration to the island during the LN period by people with a new economy, as well as new burial customs. A likely scenario, taking all these factors into account, is a sizable migration of people, with a ~50/50 (maternal) ancestry in TRB and CWC associated groups, possibly admixing with much smaller local groups of PWC associated individuals on the island.

gotland-cultures-timeline
A. Gotland cultural timelines (Apel et al., 2018; Fraser et al., 2018). B: Approximate dates for the Scandinavian Early Neolithic to Early Bronze Age time
divisions; EN: onset of TRB, MN A: onset of PWC, MN B: onset of BAC, LN-EBA time division from Vankilde (1996).

A quite interesting study that supports the predicted greater mobility during the Nordic Early Bronze Age, compared to earlier periods.

Obviously, the use of previous CWC and TRB mtDNA samples (dated necessarily before 2300 BC) to assert that the picture found during the EBA (ca. 1700-1100 BC) is due to the admixture of both cultures is not tenable.

It was more likely a mixture of descendant populations in Scandinavia, after the arrival of the Bell Beaker elites (ca. 2400 BC) and their admixture with the local population, bringing maritime integration and unifying trends to Scandinavia.

Related:

Bell Beaker/early Late Neolithic (NOT Corded Ware/Battle Axe) identified as forming the Pre-Germanic community in Scandinavia

copper-age-late-bell-beaker

I wrote recently about the newly created Indo-European Corded Ware Theory group, which represents today the last dying effort to sustain the outdated model of the ‘Kurgan peoples’.

Archaeology and Linguistics (like Genetics) keeps slowly but relentlessly rejecting all the Kurgan model‘s foundations, safe for the steppe origin of Indo-European expansion.

The book Language and Prehistory of the Indo-European Peoples. A Cross-Disciplinary perspective. Eds. A. Hyllested, B.N. Whitehead, Th. Olander and B. Anette. Copenhagen Studies in Indo-European. Museum Tusculanum Press, Copenhagen, has been recently published (December 2017).

In it, Christopher Prescott contributes to the history of Indo-European migrations to Scandinavia and the formation of a common Nordic language, ancestral to Proto-Germanic.

A draft of his chapter is downloadable in Academia.edu: Dramatic beginnings of Norway’s history? Archaeology and Indo-Europeanization.

Here are some excerpts from the text:

Thus archaeology can deal with the question of Indo-Europeans through material culture, and archaeology can contribute to unraveling the events leading up to the fact that Indo-European languages were spread from the Indian Ocean to the northwestern European Arctic in pre- and proto-history. 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. Although this interpretation cannot be “proven” in any positivist sense of the word (though aDNA and isotope studies have added a new layer of relevant data), in light of the last ten years of research and excavations, it is has become an increasingly reasonable hypothesis (e.g., Engedal 2002, Fari 2006, Håland and Håland 2000, Kristiansen 2004, Melheim 2006, Østmo 1996, also Kvalø 2007, Larsson 1997).

(…)

The Late Neolithic transformation gives rise to a cultural platform where most of southerly Norway is incorporated into the Nordic sphere. Interaction is no longer over borders, rather within a common cultural arena. Locally, the cultural institutions provide a base for the continued dynamic development through the Late Neolithic and Bronze Age. On a larger geographic and historical scale, incorporation into this field of interaction opens even the most peripheral parts of southern Norway to the streams of culture and events that shape Europe’s Bronze Age history, for example those originating from within Unetice, Tumulus Culture, Urnfeld and Hallstatt.

(…)

Changes in Scandinavia Norway are linked to wider transformations in Europe. Culturally, both Corded Ware Battle Axe and the Bell Beaker are important referential easterly and westerly European cultural horizons. Both these horizons affect and transform Northern Europe, so developments in Norway are not isolated affairs. Needless to say, though often regarded as Indo-European, the processes leading to and the affect of these cultural horizons is discussed for other parts of Europe as well (Mallory 1989:243ff).

Though there are reasonable arguments to assign both Corded Ware groups and bell Beaker groups Indo-European affiliations, the Corded Ware/Battle Axe horizon did not transform large parts of the Scandinavian Peninsula, nor can this horizon be identifies as the source of the practices, forms and institutions that characterize the ensuing Late Neolithic and Bronze Age. 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.

Though much is tentative and conjecture, multiple sources indicate that ideology, cosmology, myths social organization and probably language were Indo-European in the Bronze Age, and the development of the Bronze Age is rooted in the preceding Late Neolithic. Though the evidence also indicates that the initial Indo-European encounters, indeed “colliding worlds”, were probably experienced in the Middle Neolithic B, the archaeological record points to the time around transition to the Late Neolithic as the chronologically defining threshold for the entrenchment of an Indo-European platform throughout what would become the Nordic Bronze Age region in Norway. The Late Neolithic is therefore the most likely candidate for the introduction of the foundation for economic, social and ideological institutions, that is Giddens’ “deeply layered structure[s]”, that are fundamental to the development of the region’s identities, also ethnic, in the millennia to come.

early-bronze-age-europe
Diachronic map of migrations in Europe ca. 2250-1750 BC, after the Bell Beaker invasion, the most likely time of formation of a common Nordic language, ancestor of Proto-Germanic.

Mind you, not that these actual archaeological and linguistic models will deter anyone from supporting ancestry-based tentative sketches of a fictional ‘kurgan people’ that became outdated almost 60 years ago now – especially if they fit certain desires of ancestral ethnolinguistic identification with modern populations…

Related: