First Iberian R1b-DF27 sample, probably from incoming East Bell Beakers


I had some more time to read the paper by Valdiosera et al. (2018) and its supplementary material.

One of the main issues since the publication of Olalde et al. (2018) (and its hundreds of Bell Beaker samples) was the lack of a clear Y-DNA R1b-DF27 subclades among East Bell Beaker migrants, which left us wondering when the subclade entered the Iberian Peninsula, since it could have (theoretically) happened from the Chalcolithic to the Iron Age.

My prediction was that this lineage found today widespread among the Iberian population crossed the Pyrenees quite early, during the Chalcolithic, with migrating East Bell Beakers expanding North-West Indo-European dialects, and that it spread slowly afterwards.

The first ancient sample clearly identified as of R1b-DF27 subclade is found in this paper, at the Late Bronze Age site Cueva de los Lagos. Although it is unidentified and has no radiocarbon date, the site as a whole is associated with the Cogotas culture and its Bouquique ceramic decoration.

Y-DNA and mtDNA haplogroups, from the paper. Sequencing statistics and contamination rates for newly generated sequence data.

It was found in the northern part of the Cogotas culture territory (which lies mainly between Castille and Aragon, in North-Central Spain), shows evident steppe admixture, and it has become obvious with the latest papers (including this one) that R1b-M269 lineages intruded south of the Pyrenees associated with East Bell Beaker migrations.

The Proto-Cogotas culture is associated with a Bell Beaker substrate influenced by either El Argar or Atlantic Bronze, and the specific type of ceramics found at this Cogotas culture site are probably from the mid-2nd millennium, which is too early for the Celtic expansion.

Supervised ADMIXTURE results.

Nevertheless, due to the quite likely late date of the sample (in the centuries around 1500 BC), there is still a possibility that incoming R1b-DF27 lineages were not among the early R1b-M269 lineages found in the Iberian Chalcolithic, and were associated with later migrations from Central Europe, potentially linked to the expansion of the Urnfield culture, and thus nearer to an Italo-Celtic community.

Diachronic map of migrations in Europe ca. 1250-750 BC.

In any of these scenarios, a Pre-Celtic expansion of North-West Indo-European in Iberia (possibly associated with Lusitanian) is still the best explanation for the origin and expansion of (at least some) modern Iberian R1b-DF27 lineages, including those found among the Basque-speaking population.

This implies that the ‘indigenous’ Neolithic lineages of Iberia (like I2 and G2a2) were replaced with subsequent internal gene flows and founder effects, such as those that evidently happened (probably quite recently) among Basques, even though indigenous languages show an obvious continuity.

I would say this is the last nail in the coffin for autochthonous Y-DNA continuity theories for Spain and France (i.e. for the traditional Vasconic-Uralic hypothesis), but we know that data is never enough for any die hard continuist…so let’s just say another nail in the coffin for endless autochthonous continuity theories.

EDIT (18/3/2018): Genetiker has published Y-SNP calls for both R1b samples, showing this one is R1b1a1a2a1a2a-BY15964 (see modern members of this subclade in ytree), and that the other one is R1b1a1a2a~L23.


Iberian prehistoric migrations in Genomics from Neolithic, Chalcolithic, and Bronze Age


New open access paper Four millennia of Iberian biomolecular prehistory illustrate the impact of prehistoric migrations at the far end of Eurasia, by Valdiosera, Günther, Vera-Rodríguez, et al. PNAS (2018) published ahead of print.

Abstract (emphasis mine)

Population genomic studies of ancient human remains have shown how modern-day European population structure has been shaped by a number of prehistoric migrations. The Neolithization of Europe has been associated with large-scale migrations from Anatolia, which was followed by migrations of herders from the Pontic steppe at the onset of the Bronze Age. Southwestern Europe was one of the last parts of the continent reached by these migrations, and modern-day populations from this region show intriguing similarities to the initial Neolithic migrants. Partly due to climatic conditions that are unfavorable for DNA preservation, regional studies on the Mediterranean remain challenging. Here, we present genome-wide sequence data from 13 individuals combined with stable isotope analysis from the north and south of Iberia covering a four-millennial temporal transect (7,500–3,500 BP). Early Iberian farmers and Early Central European farmers exhibit significant genetic differences, suggesting two independent fronts of the Neolithic expansion. The first Neolithic migrants that arrived in Iberia had low levels of genetic diversity, potentially reflecting a small number of individuals; this diversity gradually increased over time from mixing with local hunter-gatherers and potential population expansion. The impact of post-Neolithic migrations on Iberia was much smaller than for the rest of the continent, showing little external influence from the Neolithic to the Bronze Age. Paleodietary reconstruction shows that these populations have a remarkable degree of dietary homogeneity across space and time, suggesting a strong reliance on terrestrial food resources despite changing culture and genetic make-up.

(A) f4 statistics testing affinities of prehistoric European farmers to either early Neolithic Iberians or central Europeans, restricting these reference populations to SNP-captured individuals to avoid technical artifacts driving the affinities. The boxplots in A show the distributions of all individual f4 statistics belonging to the respective groups. The signal is not sensitive to the choice of reference populations and is not driven by hunter-gatherer–related admixture (Datasets S4 and S5). (B) Estimates of ancestry proportions in different prehistoric Europeans as well as modern southwestern Europeans. Individuals from regions of Iberia were grouped together for the analysis in A and B to increase sample sizes per group and reduce noise


We present a comprehensive biomolecular dataset spanning four millennia of prehistory across the whole Iberian Peninsula. Our results highlight the power of archaeogenomic studies focusing on specific regions and covering a temporal transect. The 4,000 y of prehistory in Iberia were shaped by major chronological changes but with little geographic substructure within the Peninsula. The subtle but clear genetic differences between early Neolithic Iberian farmers and early Neolithic central European farmers point toward two independent migrations, potentially originating from two slightly different source populations. These populations followed different routes, one along the Mediterranean coast, giving rise to early Neolithic Iberian farmers, and one via mainland Europe forming early Neolithic central European farmers. This directly links all Neolithic Iberians with the first migrants that arrived with the initial Mediterranean Neolithic wave of expansion. These Iberians mixed with local hunter-gatherers (but maintained farming/pastoral subsistence strategies, i.e., diet), leading to a recovery from the loss of genetic diversity emerging from the initial migration founder bottleneck. Only after the spread of Bell Beaker pottery did steppe-related ancestry arrive in Iberia, where it had smaller contributions to the population compared with the impact that it had in central Europe. This implies that the two prehistoric migrations causing major population turnovers in central Europe had differential effects at the southwestern edge of their distribution: The Neolithic migrations caused substantial changes in the Iberian gene pool (the introduction of agriculture by farmers) (6, 9, 11, 13, 24), whereas the impact of Bronze Age migrations (Yamnaya) was significantly smaller in Iberia than in north-central Europe (24). The post-Neolithic prehistory of Iberia is generally characterized by interactions between residents rather than by migrations from other parts of Europe, resulting in relative genetic continuity, while most other regions were subject to major genetic turnovers after the Neolithic (4, 6, 7, 9, 25, 48). Although Iberian populations represent the furthest wave of Neolithic expansion in the westernmost Mediterranean, the subsequent populations maintain a surprisingly high genetic legacy of the original pioneer farming migrants from the east compared with their central European counterparts. This counterintuitive result emphasizes the importance of in-depth diachronic studies in all parts of the continent.


Ancient genomes document multiple waves of migration in south-east Asian prehistory


Open access preprint at bioRxiv Ancient genomes document multiple waves of migration in Southeast Asian prehistory, by Lipson, Cheronet, Mallick, et al. (2018).

Abstract (emphasis mine):

Southeast Asia is home to rich human genetic and linguistic diversity, but the details of past population movements in the region are not well known. Here, we report genome-wide ancient DNA data from thirteen Southeast Asian individuals spanning from the Neolithic period through the Iron Age (4100-1700 years ago). Early agriculturalists from Man Bac in Vietnam possessed a mixture of East Asian (southern Chinese farmer) and deeply diverged eastern Eurasian (hunter-gatherer) ancestry characteristic of Austroasiatic speakers, with similar ancestry as far south as Indonesia providing evidence for an expansive initial spread of Austroasiatic languages. In a striking parallel with Europe, later sites from across the region show closer connections to present-day majority groups, reflecting a second major influx of migrants by the time of the Bronze Age.

Schematics of admixture graph results. (A) Wider phylogenetic context. (B) Details of the Austroasiatic clade. Branch lengths are not to scale, and the order of the two events on the Nicobarese lineage in (B) is not well determined (Supplementary Text).

Featured image, from the article: “Overview of samples. (A) Locations and dates of ancient individuals. Overlapping positions are shifted slightly for visibility. (B) PCA with East and Southeast Asians. We projected the ancient samples onto axes computed using the present-day populations (with the exception of Mlabri, who were projected instead due to their large population-speci c drift). Present-day colors indicate language family affiliation: green, Austroasiatic; blue, Austronesian; orange, Hmong-Mien; black, Sino-Tibetan; magenta, Tai-Kadai.”

See also:

Demographic research of Neolithic, Chalcolithic, and Bronze Age Europe


I mentioned in the Indo-European demic diffusion model the need to assess absolute and relative population growth – as well as other demographic changes – to interpret genomic data from the different European regions studied.

One article I referred to was Demographic traces of technological innovation, social change and mobility: from 1 to 8 million Europeans (6000–2000 BCE), by Johannes Müller.

Excerpts (emphasis mine):

  • The neolithization of Northern and Northwestern Europe (probably with new forms of slash-and-burn agriculture; Feeser et al. 2012; Schier 2009) was also one of the causes for the population increase observed.
  • The introduction of the plough and developing technologies (e.g. the introduction of the wheel) (cf. Mischka 2011) might also be causes of rising population figures from ca. 3500–3000 BC.
  • The establishment of subcontinental value systems, such as the Corded Ware and Bell-Beaker phenomena (Czebreszuk/Szmyt 2003; Furholt 2004), in contrast to regional identities, might have triggered different reactions in different areas, leading to fluctuating population levels.
  • The introduction of Bronze Age ideologies, including bronze as a technology, triggered the spread of Neolithic and Bronze Age societies to vast areas of Europe (e.g. Earle/Kristiansen 2010).A major population increase is observed in both the areas already settled as well as in new areas of interest.
Absolute population values in Europe and the Near East from 6500–1500 BCE (interpolation line: spline).

In our population estimations for Central Europe and Scandinavia, population increases are associated with the periods from 5500–5000 BCE (LBK) and 3500–3000 BCE (middle and late Funnelbeaker Culture), but not for the period from 2500–2000 BCE (Bell Beaker). Consequently, this would possibly indicate forms of immigration for the first two periods and a form of interregional networking (e.g. through marriage) for the latter. But as also for the first cases on the supra-regional level (which our enquiries investigated), no other area with a significant population decrease could be observed, therefore “proof” for larger population displacement is not given. For such inquiries, studies on a more regional level are probably necessary. Nevertheless, for the Bell Beaker period I would like to exclude the possibility of large population influxes at least to Central Europe and South Scandinavia as the population values show no indication of such an event (cf. Fig. 10). In consequence, supra-regional networks or population-exchanges between smaller regions might be responsible for the isotope values.

One could interpret from the graphics, including known anthropological and genomic data, that:

  • The population growth corresponding to the Corded Ware expansion from ca. 3300 into Central Europe was seen initially with the introduction of new technology, but then stalled – probably with population replacement during the A-horizon of the Corded Ware culture.
  • The Yamna expansion into South-East Europe must have included some population replacement, i.e. influx into progressively deserted areas (such as that of the Cucuteni-Trypillia culture), since it did not leave traces of population growth.
  • The impact of the expansion of East Bell Beakers from ca. 2500 BC is clear in South-East Europe, and especially in Western Europe – taking into account the whole population growth in Europe. In Central Europe and Scandinavia the overall impact of BB migration was more limited, which suggests some degree of population replacement.

Also important to interpret genomic data are the actual economic and social differences in the different periods and cultures – usually growing after the introduction of farming. A good example is the scarce data from Khvalynsk, where the sample of haplogroup R1b (most likely of subclade M269) shows – apart from a closer position in PCA to Yamna – a a high-status burial, similar to high-status individuals buried under kurgans in later Yamna graves. This man was therefore probably a founder of an elite group of patrilineally-related families, which dominated in the following Yamna culture, which explains the clear expansion of this haplogroup’s subclades from this region.

Figures from Rebellion and Inequality in Archaeology (2017), by Johannes Müller

Other interesting papers on European demographics by Johannes Müller include:

Check out also works by Marko Porčić (such as Radiocarbon test for demographic events in written and oral history) or Stephen Shennan.

EDIT (17 Feb 2018): For how variation in the effective population size governs genetic diversity, see:

Featured image, from the main article: “The distribution of agrarian regions in Europe and the Near East in relation to the supra-regions as defined in this study: Near East (NE) about 2.400.000 km2; South East Europe (SEE) about 1.087500 km2; Central Europe and South Scandinavia (CE/SSc) about 1.613.000 km2. Europe includes 10.050.000 km2 (without Iceland)”.

See also:

The significance of the Tollense Valley in Bronze Age North-East Germany


An early Bronze Age causeway in the Tollense Valley, Mecklenburg-Western Pomerania – The starting point of a violent conflict 3300 years ago?, by Jantzen et al. (BERICHT RGK 95, 2014).

Excerpt (emphasis mine):

The causeway in the Tollense Valley, built of timber, stones, turf and sand, and documented over a length of more than 100 m, represents a unique finding from northern Germany. For the first time, part of a Bronze Age network of land routes could be made visible in the southern Baltic area.

Together with the other evidence, the archaeological remains suggest the construction of elaborate trackways and, in some cases, even bridges in the Bronze Age. The Tollense Valley causeway can probably be attributed to the wish or the necessity to be able to cross the Tollense Valley regardless of weather and seasonally differing water level conditions. Its location, situated at a narrow section of the Tollense Valley, offered a prime position for the construction of a permanent crossing of the floodplain on the eastern bank. It is quite possible that a bridge was also part of this.

The complex causeway construction that was likely used and maintained for centuries suggests a significance of the crossing beyond just local. In this context, finds from the valley relating to Bronze Age metal crafts are of interest: along with the scrap metal hoard mentioned above found in the immediate area of the crossing, attention is drawn to a hoard from Golchen comprising an unusual accumulation of tools, as well as to two tin rings found in the same archaeological layer as the Bronze Age skeletal remains. These finds could indicate that metal crafts were of particular significance in the Tollense Valley and its surrounding areas. The middle section of the Tollense Valley that is the focus of attention here could have derived special significance from its role as a crossroads.

The documented pathway, which may have been the starting point of the violent conflict described above, not only contributes to the understanding of the entire findings and the reconstruction of the events in the early 13th century BCE in the Tollense Valley; its context also sheds new light on the cross-regional infrastructure of North-East Germany in the (Early) Bronze Age. Unfortunately, there currently is little further information to integrate it into the broader network of supraregional communication and traffic routes.

The region around the famous barrow of Seddin in Brandenburg is a further example for the significance of river systems for regional power and the exchange of goods. Similarly, the River Tollense could have played a role in the flow of commodities; the causeway at the Kessin 12 site offers a possible connection of the south-north water transportation route via the Tollense River to the Baltic Sea with an east-west land route linking the River Oder estuary region and the Mecklenburg Lake District.

The Lake District was of great importance from the Early Bronze Age; here independent bronze production was established early on. Diversity analyses indicate a shift of regions of innovation during the transition from the 3rd to the 2nd millennium BCE, as the southern Baltic Sea region and the region east of the river Oder clearly also became more important. Early Bronze Age imports from south-east Europe highlight the significance of the region west of the Oder estuary. The Tollense Valley likely played a role in connecting these areas. Therefore, the violent events in the Tollense Valley could also be seen as a result of its strategic significance for the power structure of North-East Germany and the regions on the southern Baltic coast during the Early Bronze Age.

Model of the Tollense Valley with the position of pathway (R. Scholz, using a digital model of the valley made by ArcTron [©]).

See also:

Genetic prehistory of the Baltic Sea region and Y-DNA: Corded Ware and R1a-Z645, Bronze Age and N1c


Open Access The genetic prehistory of the Baltic Sea region, by Mittnik et al., Nature Communications 9: 442 (2018), based on preprint The Genetic History of Northern Europe, at BioRxirv.

As you can see, it follows my predictions in terms of haplogroups, and sadly the same trend to substitute ‘Yamna’ for ‘steppe’ while keeping linguistic interpretations unchanged…

Important excerpts for the Indo-European question (emphasis mine):

Mesolithic to Neolithic

In the archaeological understanding, the transition from Mesolithic to Neolithic in the Eastern Baltic region does not coincide with a large-scale population turnover and a stark shift in economy as seen in Central and Southern Europe. Rather, it is signified by a change in networks of contacts and the use of pottery, among other material, cultural and economic changes. Our results suggest continued admixture between groups in the south of the Eastern Baltic region, who are more closely related to WHG, and northern or eastern groups, more closely related to EHG. Neolithic social networks from the Eastern Baltic to the River Volga could also explain similarities of the hunter-gatherer pottery styles, although morphologically analogous ceramics might also have developed independently due to similar functionality. The genetic evidence for a change in networks and possibly even a large-scale population movement is most pronounced in the Middle Neolithic in individuals attributed to the CCC. The distribution of this culture overlaps in the north with the Narva culture and extends further north to Finland and Karelia. Its spread in the Eastern Baltic is linked with a significant change in imported raw materials, artefacts, and the appearance of village-like settlements15.

Neolithic to Chalcolithic

We see a further population movement into the regions surrounding the Baltic Sea with the CWC in the Late Neolithic that was accompanied by the first evidence of extensive animal husbandry in the Eastern Baltic. The presence of ancestry from the Pontic-Caspian Steppe among Baltic CWC individuals without the genetic component from north-western Anatolian Neolithic farmers must be due to a direct migration of steppe pastoralists that did not pick up this ancestry in Central Europe. It suggests import of the new economy by an incoming steppe-like population independent of the agricultural societies that were already established to the south and west of the Baltic Sea. The presence of direct contacts to the steppe could lend support to a linguistic model that sees an early branching of Balto-Slavic from a Proto-Indo-European language, for which the west Eurasian steppe was proposed as a homeland. However, as farmer ancestry is found in later Eastern Baltic individuals, it is likely that considerable individual mobility and a network of contact throughout the range of the CWC facilitated its spread eastward, possibly through exogamous marriage practices. Conversely, the appearance of mitochondrial haplogroup U4 in the Central European Late Neolithic after millennia of absence could indicate female gene-flow from the Eastern Baltic, where this haplogroup was present at high frequency.

PCA and ADMIXTURE analysis reflecting Late Neolithic in Northern European prehistory. a Principal components analysis of 1012 present-day West Eurasians (grey points, modern Baltic populations in dark grey) with 294 projected published ancient and ancient North European samples introduced in this study (marked with a red outline). b Ancestral components in ancient individuals estimated by ADMIXTURE (k = 11)
Zoomed-in version of the European Late Neolithic PCA.

So, we see that no farmer ancestry is found in the Baltic (unlike in Western Yamna), that PCA of Late Neolithic is closer to Corded Ware samples from Europe (or to earlier samples from the region) and not to Yamna, as suggested at first by the Zvejnieki individual.

There obviously was exogamy – which may in fact justify the findings in PCA close to Yamna (like the Zvejnieki sample), although researchers obviate that.

Also, as expected, no R1b-M269 in the Baltic (during the Corded Ware period), most are R1a with the majority showing subclade R1a-Z645 (and others poor SNP coverage), which support the reduction in haplogroup diversity to this very subclade during the expansion of Corded Ware peoples, as I predicted it would happen.

Bronze Age

Local foraging societies were, however, not completely replaced and contributed a substantial proportion to the ancestry of Eastern Baltic individuals of the latest LN and Bronze Age. This ‘resurgence’ of hunter-gatherer ancestry in the local population through admixture between foraging and farming groups recalls the same phenomenon observed in the European Middle Neolithic and is responsible for the unique genetic signature of modern-day Eastern Baltic populations.

We suggest that the Siberian and East Asian related ancestry in Estonia, and Y-haplogroup N in north-eastern Europe, where it is widespread today, arrived there after the Bronze Age, ca. 500 calBCE, as we detect neither in our Bronze Age samples from Lithuania and Latvia. As Uralic speaking populations of the Volga-Ural region show high frequencies of haplogroup N, a connection was proposed with the spread of Uralic language speakers from the east that contributed to the male gene pool of Eastern Baltic populations and left linguistic descendants in the Finno-Ugric languages Finnish and Estonian. A potential future direction of research is the identification of the proximate population that contributed to the arrival of this eastern ancestry into Northern Europe.

I predicted that haplogroup N arrived probably to the region west of the Urals with the Sejma-Turbino phenomenon, and that it expanded quite late, probably through founder effects. A late arrival to the region leaves obviously (safe for these researchers and others working with old ideas) only the Corded Ware culture (represented by steppe admixture and mainly haplogroup R1a-Z645) as the vector of expansion of Uralic languages, which show obviously a dialectalization process and regional expansion much older than 500 BC…

It is funny to see how people keep trying to identify R1a with ‘Yamnaya’, now ‘steppe’, but always Indo-European (an ethnolinguistic term, mind you) supposedly because of the ‘Yamnaya’ (now ‘steppe’) admixture, but the only ‘mark’ of Uralic languages for the same researchers in the same paper using this very concept is nevertheless, paradoxically, haplogroup N, with an assumption explicitly based on prevalence in modern populations

This admixture vs. haplogroup question for language and culture identification in genetic papers is really gettting messed up with new data, now in a contortionist-like way…

Images and text: Content of the paper is licensed under CC-by 4.0.

See also:

More evidence on the recent arrival of haplogroup N and gradual replacement of R1a lineages in North-Eastern Europe


A new article (in Russian), Kinship Analysis of Human Remains from the Sargat Mounds, Baraba forest-steppe, Western Siberia, by Pilipenko et al. Археология, этнография и антропология Евразии Том 45 № 4 2017, downloadable at ResearchGate.


We present the results of a paleogenetic analysis of nine individuals from two Early Iron Age mounds in the Baraba forest -teppe, associated with the Sargat culture (fi ve from Pogorelka-2 mound 8, and four from Vengerovo-6 mound 1). Four systems of genetic markers were analyzed: mitochondrial DNA, the polymorphic part of the amelogenin gene, autosomal STR-loci, and those of the Y-chromosome. Complete or partial data, obtained for eight of the nine individuals, were subjected to kinship analysis. No direct relatives of the “parent-child” type were detected. However, the data indicate close paternal and maternal kinship among certain individuals. This was evidently one of the reasons why certain individuals were buried under a single mound. Paternal kinship appears to have been of greater importance. The diversity of mtDNA and Y-chromosome lineages among individuals from one and the same mound suggests that kinship was not the only motive behind burying the deceased people jointly. The presence of very similar, though not identical, variants of the Y chromosome in different burial grounds may indicate the existence of groups such as clans, consisting of paternally related males. Our conclusions need further confi rmation and detailed elaboration. Keywords: Paleogenetics, ancient DNA, kinship analysis, mitochondrial DNA, uniparental genetic markers, STR-loci, Y-chromosome, Baraba forest-steppe, Sargat culture, Early Iron Age.

From the older study of the same region (Baraba, numbered 4) “Location of ancient human groups with a high frequency of mtDNA haplogroups U5, U4 and U2e lineages. The area of Northern Eurasian anthropological formation is marked by yellow region on the map (References: 1. Bramanti et al., 2009; 2. Malmstrom et
al., 2009; 3. Krause et al., 2010; 4. this study)”

Chronological time scale of Bronze Age Cultures from the Baraba region
This is the same team that brought an ancient mtDNA study of different cultures within the Baraba steppe-forest region (from the Open Access book Population Dynamics in Prehistory and Early History).

The Baraba steppe-forest is a region between the Ob and Irtysh rivers (about 800 km from west to east), stretching over 200 km from the taiga zone in the north to the steppes in the south.

The new study brings a more recent picture of the region, from the Iron Age Sargat culture, ca. 500 BC – 500 AD, with five samples of haplogroup N and two samples of haplogroup R1a.

R1a lineages in the region probably derive from the previous expansion of Andronovo and related cultures, which had absorbed North Caspian steppe populations and their Late Indo-European culture.

N subclades prevalent in certain modern Eurasian populations are probably derived from the expansion of the Seima-Turbino phenomenon.

While samples are scarce, Y-DNA data keeps showing the same picture I have spoken about more than once:

N subclades (potentially originally speaking Proto-Yukaghir languages) gradually replacing haplogroup R1a (originally probably speaking Uralic languages), probably through successive founder effects (such as the bottlenecks found in Finland), which left their Uralic culture and ethnolinguistic identification intact.

Therefore, late Corded Ware groups of North-Eastern Europe (in the Forest Zone and the Baltic), mainly of R1a-Z645 subclades, probably never adopted Late Indo-European languages.


Review article about Ancient Genomics, by Pontus Skoglund and Iain Mathieson


A preprint article by two of the most prolific researchers in Human Ancestry is out, and they request feedback: Ancient genomics: a new view into human prehistory and evolution, by Skoglund and Mathieson (2017). Right now, it is downloadable on Dropbox.


The first decade of ancient genomics has revolutionized the study of human prehistory and evolution. We review new insights based on ancient genomic data, including greatly increased resolution of the timing and structure of the out-of-Africa event, the diversification of present-day non-African populations, and the earliest expansions of those populations into Eurasia and America. Prehistoric genomes now document patterns of population continuity and change on every inhabited continent–in particular the effect of agricultural expansions in Africa, Europe and Oceania–and record a history of natural selection that shapes present-day phenotypic diversity. Despite these advances, much remains unknown, in particular about the genomic histories of Asia–the most populous continent, and Africa–the continent that contains the most genetic diversity. Ancient genomes from these and other regions, integrated with a growing understanding of the genomic basis of human phenotypic diversity, will be in focus during the next decade of research in the field.

The paper may be highly recommended as an introduction for anyone interested in the field of Human Ancestry in general.

However, its short summary of steppe ancestry expansion (where the Corded Ware culture predominates) is still reminiscent of the infamous “Yamnaya -> Corded Ware -> Bell Beaker” model set forth by the 2015 Nature articles on the subject, and Kristiansen’s Indo-European Corded Ware theory.

Here is an excerpt (emphasis mine):

The next substantial change is closely related to ancestry that by around 5000 BP extended over a region of more than 2000 miles of the Eurasian steppe, including in individuals associated with the Yamnaya Cultural Complex in far-eastern Europe (1; 38) and with the Afanasievo culture in the central Asian Altai mountains (1). This “steppe” ancestry is itself a mixture between ancestry that is related to Mesolithic hunter-gatherers of eastern Europe and ancestry that is related to both present-day populations (38) and Mesolithic hunter-gatherers (46) from the Caucasus mountains, and also to the populations of Neolithic (11), and Copper Age (56) Iran. Steppe ancestry appeared in southeastern Europe by 6000 BP (72), northeastern Europe around 5000 BP (47) and central Europe at the time of the Corded Ware Complex around 4600 BP (1; 38). These dates are reasonably tight constraints, because in each case there is no evidence of steppe ancestry in individuals immediately preceding these dates (47; 72). Gene flow on the steppe was extensive and bidirectional, as shown by the eastward flow of Anatolian Neolithic ancestry– reaching well into central Eurasia by the time of the Andronovo culture ~3500 BP (1)–and the westward flow of East Asian ancestry–found in individuals associated with the Iron Age Scythian culture close to the Black Sea ~2500 BP (143).

Copper and Bronze Age population movements (14; 78 Martiniano, 2017 #8761; 85; 112), as well as later movements in the Iron Age and Historical period (70; 119) further distributed steppe ancestry around Europe. Present-day western European populations can be modeled as mixtures of these three ancestry components (Mesolithic hunter-gatherer, Anatolian Neolithic and Steppe) (38; 57). In eastern Europe, further shifts in ancestry are the result of additional or distinct gene flow from Anatolia throughout the Neolithic and Bronze Age in the Aegean (42; 51; 55; 72; 87), and gene flow from Siberian-related populations in Finland and the Baltic region (38). East-west gene flow also brought new ancestry–related to populations from 265 Copper Age Iran–to the Levant during the Copper and Bronze ages (39; 56).

The geographic structure of these population transformations gave rise to population structure of present-day Europe. For example Anatolian Neolithic ancestry is highest in southern European populations like Sardinians, and lowest in northern European populations (38). Steppe ancestry is at high frequency in north-central Europeans and low in the south. Isolation-by-distance may have contributed to these patterns to some extent, but the contribution must have been small. In much of Europe, extreme population discontinuity was the norm.

Featured image: from the article, “Major Holocene population movements and expansions that have been demonstrated using ancient DNA.”


Holocene rise in mobility in at least three stages: Strong link between technological change and human mobility in Western Eurasia


New interesting article at PNAS: Estimating mobility using sparse data: Application to human genetic variation, by Loog et al (2017).

Download links and supplemental information.


Migratory activity is a critical factor in shaping processes of biological and cultural change through time. We introduce a method to estimate changes in underlying migratory activity that can be applied to genetic, morphological, or cultural data and is well-suited to samples that are sparsely distributed in space and through time. By applying this method to ancient genome data, we infer a number of changes in human mobility in Western Eurasia, including higher mobility in pre- than post-Last Glacial Maximum hunter–gatherers, and oscillations in Holocene mobility with peaks centering on the Neolithic transition and the beginnings of the Bronze Age and the Late Iron Age.


Mobility is one of the most important processes shaping spatiotemporal patterns of variation in genetic, morphological, and cultural traits. However, current approaches for inferring past migration episodes in the fields of archaeology and population genetics lack either temporal resolution or formal quantification of the underlying mobility, are poorly suited to spatially and temporally sparsely sampled data, and permit only limited systematic comparison between different time periods or geographic regions. Here we present an estimator of past mobility that addresses these issues by explicitly linking trait differentiation in space and time. We demonstrate the efficacy of this estimator using spatiotemporally explicit simulations and apply it to a large set of ancient genomic data from Western Eurasia. We identify a sequence of changes in human mobility from the Late Pleistocene to the Iron Age. We find that mobility among European Holocene farmers was significantly higher than among European hunter–gatherers both pre- and postdating the Last Glacial Maximum. We also infer that this Holocene rise in mobility occurred in at least three distinct stages: the first centering on the well-known population expansion at the beginning of the Neolithic, and the second and third centering on the beginning of the Bronze Age and the late Iron Age, respectively. These findings suggest a strong link between technological change and human mobility in Holocene Western Eurasia and demonstrate the utility of this framework for exploring changes in mobility through space and time.

Featured image, from the article: Estimation of mobility through time from empirical data. (A) Relative mobility rate estimates in Western Eurasia over the last 14,000 y, using a 4,000-y sliding window (121 windows). The solid black line represents the mean α value from 10,000 date resampled iterations; the colored area represents the 95% confidence intervals of the jackknife distribution.

Iberian Peninsula: Discontinuity in mtDNA between hunter-gatherers and farmers, not so much during the Chalcolithic and EBA


A new preprint paper at BioRxiv, The maternal genetic make-up of the Iberian Peninsula between the Neolithic and the Early Bronze Age, by Szécsényi-Nagy et al. (2017).


Agriculture first reached the Iberian Peninsula around 5700 BCE. However, little is known about the genetic structure and changes of prehistoric populations in different geographic areas of Iberia. In our study, we focused on the maternal genetic makeup of the Neolithic (~ 5500-3000 BCE), Chalcolithic (~ 3000-2200 BCE) and Early Bronze Age (~ 2200-1500 BCE). We report ancient mitochondrial DNA results of 213 individuals (151 HVS-I sequences) from the northeast, central, southeast and southwest regions and thus on the largest archaeogenetic dataset from the Peninsula to date. Similar to other parts of Europe, we observe a discontinuity between hunter-gatherers and the first farmers of the Neolithic. During the subsequent periods, we detect regional continuity of Early Neolithic lineages across Iberia, however the genetic contribution of hunter-gatherers is generally higher than in other parts of Europe and varies regionally. In contrast to ancient DNA findings from Central Europe, we do not observe a major turnover in the mtDNA record of the Iberian Late Chalcolithic and Early Bronze Age, suggesting that the population history of the Iberian Peninsula is distinct in character.

Iberian mtDNA samples

Detailed conclusions of their work,

The present study, based on 213 new and 125 published mtDNA data of prehistoric Iberian individuals suggests a more complex mode of interaction between local hunter-gatherers and incoming early farmers during the Early and Middle Neolithic of the Iberian Peninsula, as compared to Central Europe. A characteristic of Iberian population dynamics is the proportion of autochthonous hunter-gatherer haplogroups, which increased in relation to the distance to the Mediterranean coast. In contrast, the early farmers in Central Europe showed comparatively little admixture of contemporaneous hunter-gatherer groups. Already during the first centuries of Neolithic transition in Iberia, we observe a mix of female DNA lineages of different origins. Earlier hunter-gatherer haplogroups were found together with a variety of new lineages, which ultimately derive from Near Eastern farming groups. On the other hand, some early Neolithic sites in northeast Iberia, especially the early group from the cave site of Els Trocs in the central Pyrenees, seem to exhibit affinities to Central European LBK communities. The diversity of female lineages in the Iberian communities continued even during the Chalcolithic, when populations became more homogeneous, indicating higher mobility and admixture across different geographic regions. Even though the sample size available for Early Bronze Age populations is still limited, especially with regards to El Argar groups, we observe no significant changes to the mitochondrial DNA pool until the end of our time transect (1500 BCE). The expansion of groups from the eastern steppe, which profoundly impacted Late Neolithic and EBA groups of Central and North Europe, cannot (yet) be seen in the contemporaneous population substrate of the Iberian Peninsula at the present level of genetic resolution. This highlights the distinct character of the Neolithic transition both in the Iberian Peninsula and elsewhere and emphasizes the need for further in depth archaeogenetic studies for reconstructing the close reciprocal relationship of genetic and cultural processes on the population level.

So it seems more and more likely that the North-West Indo-European invasion during the Copper Age (signaled by changes in Y-DNA lineages) was not, as in central Europe, accompanied by much mtDNA turnover. What that means – either a male-dominated invasion, or a longer internal evolution of invasive Y-DNA subclades – remains to bee seen, but I am still more inclined to see the former as the most likely interpretation, in spite of admixture results.


Featured images: from the article, licensed BY-NC-ND.