Bell Beakers and Mycenaeans from Yamnaya; Corded Ware from the forest steppe

eba-yamnaya-ancestry-hungary

I have recently written about the spread of Pre-Yamnaya or Yamnaya ancestry and Corded Ware-related ancestry throughout Eurasia, using exclusively analyses published by professional geneticists, and filling in the gaps and contradictory data with the most reasonable interpretations. I did so consciously, to avoid any suspicion that I was interspersing my own data or cherry picking results.

Now I’m finished recapitulating the known public data, and the only way forward is the assessment of these populations using the available datasets and free tools.

Understanding the complexities of qpAdm is fairly difficult without a proper genetic and statistical background, which I won’t pretend to have, so its tweaking to get strictly correct results would require an unending game of trial and error. I have sadly little time for this, even taking my tendency to procrastination into account… so I have used a simple model akin to those published before – in particular, the outgroup selection by Ning, Wang et al. (2019), who seem to be part of the only group interested in distinguishing Yamnaya-related from Corded Ware-related ancestry, probably the most relevant question discussed today in population genomics regarding the Proto-Indo-European and Proto-Uralic homelands.

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

I have used for all analyses below a merged dataset including the curated one of the Reich Lab, the latest on Central and South Asia by Narasimhan, Patterson et al. (2019), on Iberia by Olalde et al. (2019), and on the East Baltic by Saag et al. (2019), as well as datasets including samples from Wang et al. (2019) and Lamnidis et al. (2018). I used (and intend to use) the same merged dataset in all cases, despite its huge size, to avoid adding one more uncontrolled variable to the analyses, so that all results obtained can be compared.

I try to prepare in advance a bunch of relevant files with left pops and right pops for each model:

  1. It seems a priori more reasonable to use geographically and chronologically closer proxy populations (say, Trypillia or GAC for Steppe-related peoples) than hypothetic combinations of ancestral ones (viz. Anatolian farmer, WHG, and EHG).
  2. This also means using subgroups closer to the most likely source population, such as (Don-Volga interfluve) Yamnaya_Kalmykia rather than (Middle Volga) Yamnaya_Samara for the western expansion of late Repin/early Yamnaya, or the early Germany_Corded_Ware.SG or Czech_Corded Ware for the group closest to the Proto-Corded Ware population (see below), likely neighbouring the Upper Vistula region.
  3. I usually test two source populations for different targets, which seems like a much more efficient way of using computer resources, whenever I know what I want to test, since I need my PC back for its normal use; whenever I don’t know exactly what to test, I use three-way admixture models and look for subsets to try and improve the results.

I have probably left out some more complex models by individualizing the most relevant groups, but for the time being this would have to do. Also, no other formal stats have been used in any case, which is an evident shortcoming, ruling out an interpretation drawn directly and only from the results below.

Full qpAdm results for each batch of samples are presented in a Google Spreadsheet, with each tab (bottom of the page) showing a different combination of sources, usually in order of formally ‘best’ (first to the left) to ‘worst’ (last to the right) fits, although the order is difficult to select in highly heterogeneous target groups, as will be readily visible.

maykop-trypillia-intrusion-steppes
Disintegration, migration, and imports of the Azov–Black Sea region. First migration event (solid arrows): Gordineşti–Maikop expansion (groups: I – Bursuchensk; II – Zhyvotylivka; III – Vovchans’k; IV – Crimean; V – Lower Don; VI – pre-Kuban). Second migration event (hollow arrows): Repin expansion. After Rassamakin (1999), Demchenko (2016).

Corded Ware origins

The latest publications on the Yampil barrow complex have not improved much our understanding of the complexity of Corded Ware origins from an archaeological point of view, involving multiple cultural (hence likely population) influences. This bit is from Ivanova et al., Baltic-Pontic Studies (2015) 20:1, and most hypotheses of the paper remain unanswered (except maybe for the relevance of the Złota group):

In the light of the above outline therefore one should argue that the ‘architecture of barrows’ associated in the ‘Yampil landscape’ of the Middle Dniester Area with the Eneolithic (specifically, mainly with the TC), precedes the development of a similar phenomenon that can be observed from 2900/2800 BC in the Upper Dniester Area and drainage basin of the Upper Vistula, associated with the CWC [Goslar et al. 2015; Włodarczak 2006; 2007; 2008; Jarosz, Włodarczak 2007]. The most consuming research question therefore is whether ritual customs making use of Eneolithic (Tripolye) ‘barrow architecture’ could have penetrated northwards along the Dniester route, where GAC communities functioned. One could also ask what role the rituals played among the autochthons [Kośko 2000; Włodarczak 2008; 2014: 335; Ivanova, Toshchev 2015b].

This issue has already been discussed with a resulting tentative systemic taxonomy in the studies of Włodarczak, arguing for the Złota culture (ZC) in the Vistula region as an illustration of one of the (Małopolska) reception centres of civilization inspirations from the oldest Pontic ‘barrow culture’ circle associated with the Eneolithic and Early Bronze Age [Włodarczak 2008]. Notably, it is in the ZC that one can notice a set of cultural traits (catacomb grave construction, burial details, forms and decoration of vessels) analogous to those shared by the north-western Black Sea Coast groups of the forest-steppe Eneolithic (chiefly Zhyvotilovka-Volchansk) and the Late Tripolye circle (chiefly Usatovo-Gordinești-Horodiștea-Kasperovtsy).

gac-trypillia-usatovo-corded-ware
Globular Amphorae culture „exodus” to the Danube Delta: a – Globular Amphorae culture; b – GAC (1), Gorodsk (2), Vykhvatintsy (3) and Usatovo (4) groups of Trypillia culture; c – Coţofeni culture; d – northern border of the late phase of Baden culture;red arrows – direction of Globular Amphora culture expansion; blue arrow – direction of „reflux” of Globular Amphora culture (apud Włodarczak, 2008, with changes).

Taking into account that I6561 might be wrongly dated, we cannot include the Corded Ware-like sample of the end-5th millennium BC in the analysis of Corded Ware origins. That uncertainty in the chronology of the appearance of “Steppe ancestry” in Proto-Corded Ware peoples complicates the selection of any potential source population from the CHG cline.

Nevertheless, the lack of hg. R1a-M417 and sizeable Pre-Yamnaya-related ancestry in the sampled Pontic forest-steppe Eneolithic populations (represented exclusively by two samples from Dereivka ca. 3600-3400 BC) would leave open the interesting possibility that a similar ancestry got to the forest-steppe region between modern Poland and Ukraine during the known complex population movements of the Late Eneolithic.

It is known that Corded Ware-derived groups and Steppe Maykop show bad fits for Pre-Yamnaya/Yamnaya ancestry, and also that Steppe Maykop is a potential source of “Steppe-related ancestry” within the Eneolithic CHG mating network of the Pontic-Caspian steppes and forest-steppes. Testing Corded Ware for recent Trypillia and Maykop influences, proper of Late Trypillia and Late Maykop groups in the North Pontic area (such as Zhyvotylivka–Vovchans’k and Gordineşti) side by side with potential Pre-Yamnaya and Yamnaya sources makes thus sense:

Now, the main obvious difference between Khvalynsk-Yamnaya and Corded Ware is the long-lasting, pervasive Y-chromosome bottlenecks under R1b lineages in the former, compared to the haplogroup variability and late bottleneck under R1a-M417 in the latter, which speaks in favour – on top of everything else – of a different community of sub-Neolithic hunter-gatherers including hg. R1a-M417 hijacking the expansion of Steppe_Maykop-related ancestry around the Volhynian-Podolian Upland.

Akin to how Yamnaya patrilineal descendants hijacked regional EEF (±CWC) ancestry components mainly through exogamy, dragging them into the different expanding Bell Beaker groups (see below), but kept their Indo-European languages, these hunter-gatherers that admixed with peoples of “Steppe ancestry” were the most likely vector of expansion of Uralic languages in Eastern Europe.

corded-ware-from-trypillia-maykop
PCA of ancient Eurasian samples. Marked likely Proto-Corded Ware samples and potential origin of its PCA cluster based on qpAdm results. See full PCA and more related files.

Baltic Corded Ware

One of the most interesting aspects of the results above is the surprising heterogeneity of the different regional groups, which is also reflected in the Y-DNA variability of early Corded Ware samples.

Seeing how Baltic CWC groups, especially the early Latvia_LN sample, show particularly bad fits with the models above, it seems necessary to test how this population might have come to be. My first impression in 2017 was that they could represent early Corded Ware groups admixed with Yamnaya settlers through their interactions along the Dnieper-Dniester corridor.

However, I recently predicted that the most likely admixture leading to their ancestry and PCA cluster would involve a Corded Ware-like group and a group related to sub-Neolithic cultures of eastern Europe, whose best proxy to date are EHG-like Khvalynsk samples (i.e. excluding the outlier with Pre-Yamnaya ancestry, I0434):

corded-ware-pca-sub-neolithic-europe
Detail of the PCA of the Corded Ware expansion. See full PCA and more related files.

Late Corded Ware + Yamnaya vanguard

Relevant are also the mixtures of Corded Ware from Esperstedt, and particularly those of the sample I0104, which I have repeated many times in this blog I suspected to be influenced by vanguard Yamnaya settlers:

The infeasible models of CWC + Yamnaya_Kalmykia ± Hungary_Baden (see below for Bell Beakers) and the potential cluster formed with other samples from the Baltic suggest that it could represent a more complex set of mixtures with sub-Neolithic populations. On the other hand, its location in Germany, late date (ca. 2500 BC or later), and position in the PCA, together with the good fits obtained for Germany_Beaker as a source, suggest that the increase in Steppe-related ancestry + EEF makes it impossible for the model (as I set it) to directly include Yamnaya_Kalmykia, despite this excess Steppe-related ancestry actually coming from Yamnaya vanguard groups.

I think it is very likely that the future publication of EEF-admixed Yamnaya_Hungary samples (or maybe even Yamnaya vanguard samples) will improve the fits of this model.

These results confirm at least the need to distrust the common interpretation of mixtures including late Corded Ware samples from Esperstedt (giving rise to the “up to 75% Yamnaya ancestry of CWC” in the 2015 papers) as representative of the Corded Ware culture as a whole, and to keep always in mind that an admixture of European BA groups including Corded Ware Esperstedt as a source also includes East BBC-like ancestry, unless proven otherwise.

yamnaya-vanguard-corded-ware-chalcolithic-early
Yamnaya vanguard groups in Corded Ware territory before the expansion of Bell Beakers (ca. 2500 BC). See full map.

Bell Beaker expansion

A hotly (re)debated topic in the past 6 months or so, and for all the wrong reasons, is the origin of the Bell Beaker folk. Archaeology, linguistics, and different Y-chromosome bottlenecks clearly indicate that Bell Beakers were at the origin of the North-West Indo-European expansion in Europe, while the survival of Corded Ware-related groups in north-eastern Europe is clearly related to the expansion of Uralic languages.

NOTE. For the interesting case of Proto-Indo-Iranians expanding with Corded Ware-like ancestry, see more on the formation of Sintashta-Potapovka-Filatovka from East Uralic-speaking Abashevo and Pre-Proto-Indo-Iranian-speaking Poltavka herders. See also more on R1a in Indo-Iranians and on the social complexity of Sintashta.

Nevertheless, every single discarded theory out there seems to keep coming back to life from time to time, and a new wave of interest in “Bell Beaker from the Single Grave culture” somehow got revived in the process, too, because this obsession – unlike the “Bell Beakers from Iberia Chalcolithic” – is apparently acceptable in certain circles, for some reason.

We know that Iberian Beakers, British Beakers, or Sicilian EBA – representing the most likely closest source population of speakers of Proto-Galaico-Lusitanian, Pre-Celtic Indo-European, and Proto-Elymian, respectively – have already been successfully tested for a direct origin among Western European Beakers in Olalde et al. (2018), Olalde et al. (2019), and Fernandes et al. (2019).

This success in ascertaining a closer Beaker source is probably due to the physical isolation of the specific groups (related to Germany_Beaker, Netherlands_Beaker, and NE_Mediterranean_Beaker samples, respectively) after their migration into regions dominated by peoples without Steppe-related ancestry. Furthermore, Celtic-speaking populations expanding with Urnfield south of the Pyrenees also show a good fit with a source close to France_Beaker.

So I decided to test sampled Bell Beaker populations, to see if it could shed light to the most likely source population of individual Beaker groups and the direction of migration within Central Europe, i.e. roughly eastwards or westwards. As it was to be expected for closely related populations (see the relevant discussion here), an attempt to offer a simplistic analysis of direction based on formal stats does not make any sense, because most of the alternative hypotheses cannot be rejected:

Not only because of the similar values obtained, but because it is absurd to take p-values as a measure of anything, especially when most of these conflicting groups with slightly ‘better’ or ‘worse’ p-values represent multiple different mixtures of the type (Yamnaya + EEF) + (Corded Ware + EEF ± Yamnaya), impossible to distinguish without selecting proper, direct ancestral populations…

A further example of how explosive the Bell Beaker expansion was into different territories, and of their extensive local admixture, is shown by the unsuccessful attempt by Olalde et al. (2018) to obtain an origin of the EEF source for all Beaker groups (excluding Iberian Beakers):

bell-beaker-local-population-iberia
Investigating the genetic makeup of Beaker-complex-associated individuals. Testing different populations as a source for the Neolithic ancestry component in Beaker-complex-associated individuals. The table shows P values (* indicates values > 0.05) for the fit of the model: ‘Steppe_EBA + Neolithic/Copper Age’ source population.
burials-yamnaya-hungary
Map of attested Yamnaya pit-grave burials in the Hungarian plains; superimposed in shades of blue are common areas covered by floods before the extensive controls imposed in the 19th century; in orange, cumulative thickness of sand, unfavourable loamy sand layer. Marked are settlements/findings of Boleráz (ca. 3500 BC on), Baden (until ca. 2800 BC), Kostolac (precise dates unknown), and Yamna kurgans (from ca. 3100/3000 BC on).

Now, there is a simpler way to understand what kind of Steppe-related ancestry is proper of Bell Beakers. I tested two simple models for some Beaker groups: Yamnaya + Hungary Baden vs. Corded Ware + GAC Poland. After all, the Bell Beaker folk should prefer a source more closely related to either Yamnaya Hungary or Central European Corded Ware:

Interestingly, models including Yamnaya + Baden show good fits for the most important groups related to North-West Indo-Europeans, including Bell Beakers from Germany, the Netherlands, Italy, and Poland, representing the most likely closest source populations of speakers of Pre-Proto-Celtic, Pre-Proto-Germanic, Proto-Italo-Venetic, and Pre-Proto-Balto-Slavic, respectively.

The admixed Yamnaya samples from Hungary that will hopefully be published soon by the Jena Lab will most likely further improve these fits, especially in combination with intermediate Chalcolithic populations of the Middle and Upper Danube and its tributaries, to a point where there will be an absolute chronological and geographical genomic trail from the fully Yamnaya-like Yamnaya settlers from Hungary to all North-West Indo-European-speaking groups of the Early Bronze Age.

The only difference between groups will be the gradual admixture events of their source Beaker group with local populations on their expansion paths, including peoples of mainly EEF, CWC+EEF, or CWC+EEF+Yamnaya related ancestry. There is ample evidence beyond ancestry models to support this, in particular continued Y-DNA bottlenecks under typical Yamnaya paternal lineages, mainly represented by R1b-L51 subclades.

east-bell-beaker-group-expansion
Distribution of the Bell Beaker East Group, with its regional provinces, as of c. 2400 cal BC (after Heyd et al. 2004, modified). See full maps.

European Early Bronze Age

European EBA groups that might show conflicting results due to multiple admixture events with Corded Ware-related populations are the Únětice culture and the Nordic Late Neolithic.

The results for Únětice groups seem to be in line with what is expected of a Central European EBA population derived from Bell Beakers admixed with surrounding poulations of East Bell Beaker and/or late (Epi-)Corded Ware descent.

Potential models of mixture for Nordic Late Neolithic samples – despite the bad fits due to the lack of direct ancestral CWC and BBC groups from Denmark – seem to be impossible to justify as derived exclusively from Single Grave or (even less) from Battle Axe peoples, supporting immigration waves of Bell Beakers from the south and further admixture events with local groups through maritime domination.

PCA of ancient European samples. Marked are Bronze Age clusters. See full PCAs.

Balkans Bronze Age

The potential origin of the typical Corded Ware Steppe-related ancestry in the social upheaval and population movements of the Dnieper-Dniester forest-steppe corridor during the 4th millennium BC raises the question: how much do Balkan Bronze Age groups owe their ancestry to a population different than the spread of Pre-Yamnaya-like Suvorovo-Novodanilovka chieftains? Furthermore, which Bronze Age groups seem to be more likely derived exclusively from Pre-Yamnaya groups, and which are more likely to be derived from a mixture of Yamnaya and Pre-Yamnaya? Do the formal stats obtained correspond to the expected results for each group?

Since the expansion of hg. I2a-L699 (TMRCA ca. 5500 BC) need not be associated with Yamnaya, some of these values – together with the assessment of each individual archaeological culture – may question their origin in a Yamnaya-related expansion rather than in a Khvalynsk-related one.

NOTE. These are the last ones I was able to test yesterday, and I have not thought these models through, so feel free to propose other source and target groups. In particular, complex movements through the North Pontic area during the Late Eneolithic would suggest that there might have been different Steppe-ancestry-related vs. EEF-related interactions in the north-west and west Pontic area before and during the expansion of Yamnaya.

Mycenaeans

One of the key Indo-European populations that should be derived from Yamnaya to confirm the Steppe hypothesis, together with North-West Indo-Europeans, are Proto-Greeks, who will in turn improve our understanding of the preceding Palaeo-Balkan community. Unfortunately, we only have Mycenaean samples from the Aegean, with slight contributions of Steppe-related ancestry.

Still, analyses with potential source populations for this Steppe ancestry show that the Yamnaya outlier from Bulgaria is a good fit:

The comparison of all results makes it quite evident the why of the good fits from (Srubnaya-related) Bulgaria_MLBA I2163 or of Sintashta_MLBA relative to the only a priori reasonable Yamnaya and Catacomb sources: it is not about some hypothetical shared ancestor in Graeco-Aryan-speaking East Yamnaya– or even Catacomb-Poltavka-related groups, because all available Yamnaya-related peoples are almost indistinguishable from each other (at least with the sampling available today). These results reflect a sizeable contribution of similar EEF-related populations from around the Carpathians in both Steppe-related groups: Corded Ware and Yamnaya settlers from the Balkans.

mycenaeans-minyan-ware-greece-minoan
Cultural groups in and around the Balkans during the Early Bronze Age. See full maps.

qpAdm magic

In hobby ancestry magic, as in magic in general, it is not about getting dubious results out of thin air: misdirection is the key. A magician needs to draw the audience attention to ‘remarkable’ ancestry percentages coupled with ‘great’ (?) p-values that purportedly “prove” what the audience expects to see, distracting everyone from the true interesting aspects, like statistical design, the data used (and its shortcomings), other opposing models, a comparison of values, a proper interpretation…you name it.

I reckon – based on the examples above – that the following problems lie at the core of bad uses of qpAdm:

  1. In the formal aspect, the poor understanding of what p-values and other formal stats obtained actually mean, and – more importantly – what they don’t mean. The simplistic trend to accept results of a few analyses at face value is necessarily wrong, in so far as there is often no proper reasoning of what is being assessed and how, and there is never a previous opinion about what could be expected if the alternative hypotheses were true.
  2. In the interpretation aspect, the poor judgement of accompanying any results with simplistic, superficial, irrelevant, and often plainly wrong archaeological or linguistic data selected a posteriori; the inclusion of some racial or sociopolitical overtones in the mixture to set a propitious mood in the target audience; and a sort of ritualistic theatrics with the main theme of ‘winning’, that is best completed with ad hominems.

If you get rid of all this, the most reasonable interpretation of the output of a model proposed and tested should be similar to Nick Patterson’s words in his explanation of qpWave and qpAdm use:

Here we see that, at least in this analysis there are reasonable models with CordedWareNeolithic is a mix of either WHG or LBKNeolithic and YamnayaEBA. (…) The point of this note is not to give a serious phylogenetic analysis but the results here certainly support a major Steppe contribution to the Corded Ware population, which is entirely concordant with the archaeology [?].

Very far, as you can see, from the childish “Eureka! I proved the source!”-kind of thinking common among hobbyists.

The Mycenaean case is an illustrative example: if the Yamnaya outlier from Bulgaria were not available, and if one were not careful when designing and assessing those mixture models, the interpretation would range from erroneous (viz. a Graeco-Aryan substrate, as I initially thought) to impossible (say, inventing migration waves of Sintashta or Srubnaya peoples into Crete). The models presented above show that a contribution of Yamnaya to Mycenaeans couldn’t be rejected, and this alone should have been enough to accept Yamnaya as the most likely source population of “Steppe ancestry” in Proto-Greeks, pending intermediate samples from the Balkans. In other words, one could actually find that ‘the best’ p-values for source populations of Mycenaeans is a combination of modern Poles + Turks, despite the impracticality of such a model…

I haven’t been able to reproduce results which supposedly showed that Corded Ware is more likely to be derived from (Pre-)Yamnaya than other source population, or that Corded Ware is better suited as the ancestral population of Bell Beakers. The analyses above show values in line with what has been published in recent scientific papers, and what should be expected based on linguistics and archaeology. So I’ll go out on a limb here and say that it’s only through a careful selection of outgroups and samples tested, and of as few compared models as possible, that you could eventually get this kind of results and interpretation, if at all.

Whether that kind of special care for outgroups and samples is about (a) an acceptable fine-tuning of the analyses, (b) a simplistic selection dragged from the first papers published and applied indiscriminately to all models, or (c) cherry picking analyses until results fit the expected outcome, is a question that will become mostly irrelevant when future publications continue to support an origin of the expansion of ancient Indo-European languages in Khvalynsk- and Yamnaya-related migrations.

Feel free to suggest (reasonable) modifications to correct some of these models in the comments. Also, be sure to check out other values such as proportions, SD or SNPs of the different results that I might have not taken into account when assessing ‘good’ or ‘bad’ fits.

Related

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

Vikings, Vikings, Vikings! “eastern” ancestry in the whole Baltic Iron Age

vikings-middle-age

Open access Population genomics of the Viking world, by Margaryan et al. bioRxiv (2019), with a huge new sampling from the Viking Age.

Interesting excerpts (emphasis mine, modified for clarity):

To understand the genetic structure and influence of the Viking expansion, we sequenced the genomes of 442 ancient humans from across Europe and Greenland ranging from the Bronze Age (c. 2400 BC) to the early Modern period (c. 1600 CE), with particular emphasis on the Viking Age. We find that the period preceding the Viking Age was accompanied by foreign gene flow into Scandinavia from the south and east: spreading from Denmark and eastern Sweden to the rest of Scandinavia. Despite the close linguistic similarities of modern Scandinavian languages, we observe genetic structure within Scandinavia, suggesting that regional population differences were already present 1,000 years ago.

Maps illustrating the following texts have been made based on data from this and other papers:

  • Maps showing ancestry include only data from this preprint (which also includes some samples from Sigtuna).
  • Maps showing haplogroup density include Vikings from other publications, such as those from Sigtuna in Krzewinska et al. (2018), and from Iceland in Ebenesersdóttir et al. (2018).
  • Maps showing haplogroups of ancient DNA samples based on their age include data from all published papers, but with slightly modified locations to avoid overcrowding (randomized distance approx. ± 0.1 long. and lat.).

middle-ages-europe-y-dna
Y-DNA haplogroups in Europe during the Viking expansions (full map). See other maps from the Middle Ages.

We find that the transition from the BA to the IA is accompanied by a reduction in Neolithic farmer ancestry, with a corresponding increase in both Steppe-like ancestry and hunter-gatherer ancestry. While most groups show a slight recovery of farmer ancestry during the VA, there is considerable variation in ancestry across Scandinavia. In particular, we observe a wide range of ancestry compositions among individuals from Sweden, with some groups in southern Sweden showing some of the highest farmer ancestry proportions (40% or more in individuals from Malmö, Kärda or Öland).

Ancestry proportions in Norway and Denmark on the other hand appear more uniform. Finally we detect an influx of low levels of “eastern” ancestry starting in the early VA, mostly constrained among groups from eastern and central Sweden as well as some Norwegian groups. Testing of putative source groups for this “eastern” ancestry revealed differing patterns among the Viking Age target groups, with contributions of either East Asian- or Caucasus-related ancestry.

saami-ancestry-vikings
Ancestry proportions of four-way models including additional putative source groups for target groups for which three-way fit was rejected (p ≤ 0.01);

Overall, our findings suggest that the genetic makeup of VA Scandinavia derives from mixtures of three earlier sources: Mesolithic hunter-gatherers, Neolithic farmers, and Bronze Age pastoralists. Intriguingly, our results also indicate ongoing gene flow from the south and east into Iron Age Scandinavia. Thus, these observations are consistent with archaeological claims of wide-ranging demographic turmoil in the aftermath of the Roman Empire with consequences for the Scandinavian populations during the late Iron Age.

Genetic structure within Viking-Age Scandinavia

We find that VA Scandinavians on average cluster into three groups according to their geographic origin, shifted towards their respective present-day counterparts in Denmark, Sweden and Norway. Closer inspection of the distributions for the different groups reveals additional complexity in their genetic structure.

vikings-danish-ancestry
Natural neighbor interpolation of “Danish ancestry” among Vikings.

We find that the ‘Norwegian’ cluster includes Norwegian IA individuals, who are distinct from both Swedish and Danish IA individuals which cluster together with the majority of central and eastern Swedish VA individuals. Many individuals from southwestern Sweden (e.g. Skara) cluster with Danish present-day individuals from the eastern islands (Funen, Zealand), skewing towards the ‘Swedish’ cluster with respect to early and more western Danish VA individuals (Jutland).

Some individuals have strong affinity with Eastern Europeans, particularly those from the island of Gotland in eastern Sweden. The latter likely reflects individuals with Baltic ancestry, as clustering with Baltic BA individuals is evident in the IBS-UMAP analysis and through f4-statistics.

vikings-norwegian-ancestry
Natural neighbor interpolation of “Norwegian ancestry” among Vikings.

For more on this influx of “eastern” ancestry see my previous posts (including Viking samples from Sigtuna) on Genetic and linguistic continuity in the East Baltic, and on the Pre-Proto-Germanic homeland based on hydrotoponymy.

Baltic ancestry in Gotland

Genetic clustering using IBS-UMAP suggested genetic affinities of some Viking Age individuals with Bronze Age individuals from the Baltic. To further test these, we quantified excess allele sharing of Viking Age individuals with Baltic BA compared to early Viking Age individuals from Salme using f4 statistics. We find that many individuals from the island of Gotland share a significant excess of alleles with Baltic BA, consistent with other evidence of this site being a trading post with contacts across the Baltic Sea.

vikings-finnish-ancestry
Natural neighbor interpolation of “Finnish ancestry” among Vikings.

The earliest N1a-VL29 sample available comes from Iron Age Gotland (VK579) ca. AD 200-400 (see Iron Age Y-DNA maps), which also proves its presence in the western Baltic before the Viking expansion. The distribution of N1a-VL29 and R1a-Z280 (compared to R1a in general) among Vikings also supports a likely expansion of both lineages in succeeding waves from the east with Akozino warrior-traders, at the same time as they expanded into the Gulf of Finland.

vikings-y-dna-haplogroup-r1a-z280-over-r1a
Density of haplogroup R1a-Z280 (samples in pink) overlaid over other R1a samples (in green, with R1a-Z284 in cyan) among Vikings.

Vikings in Estonia

(…) only one Viking raiding or diplomatic expedition has left direct archaeological traces, at Salme in Estonia, where 41 Swedish Vikings who died violently were buried in two boats accompanied by high-status weaponry. Importantly, the Salme boat-burial predates the first textually documented raid (in Lindisfarne in 793) by nearly half a century. Comparing the genomes of 34 individuals from the Salme burial using kinship analyses, we find that these elite warriors included four brothers buried side by side and a 3rd degree relative of one of the four brothers. In addition, members of the Salme group had very similar ancestry profiles, in comparison to the profiles of other Viking burials. This suggests that this raid was conducted by genetically homogeneous people of high status, including close kin. Isotope analyses indicate that the crew descended from the Mälaren area in Eastern Sweden thus confirming that the Baltic-Mid-Swedish interaction took place early in the VA.

vikings-swedish-ancestry
Natural neighbor interpolation of “Swedish ancestry” among Vikings.

Viking samples from Estonia show thus ancient Swedes from the Mälaren area, which proves once again that hg. N1a-VL29 (especially subclade N1a-L550) and tiny proportions of so-called “Siberian ancestry” expanded during the Early Iron Age into the whole Baltic Sea area, not only into Estonia, and evidently not spreading with Balto-Finnic languages (since the language influence is in the opposite direction, east-west, Germanic > Finno-Samic, during the Bronze Age).

N1a-VL29 lineages spread again later eastwards with Varangians, from Sweden into north-eastern Europe, most likely including the ancestors of the Rurikid dynasty. Unsurprisingly, the arrival of Vikings with Swedish ancestry into the East Baltic and their dispersal through the forest zone didn’t cause a language shift of Balto-Finnic, Mordvinic, or East Slavic speakers to Old Norse, either…

NOTE. For N1a-Y4339 – N1a-L550 subclade of Swedish origin – as main haplogroup of modern descendants of Rurikid princes, see Volkov & Seslavin (2019) – full text in comments below. Data from ancient samples show varied paternal lineages even among early rulers traditionally linked to Rurik’s line, which explains some of the discrepancies found among modern descendants:

  • A sample from Chernihiv (VK542) potentially belonging to Gleb Svyatoslavich, the 11th century prince of Tmutarakan/Novgorod, belongs to hg. I2a-Y3120 (a subclade of early Slavic I2a-CTS10228) and has 71% “Modern Polish” ancestry (see below).
  • Izyaslav Ingvarevych, the 13th century prince of Dorogobuzh, Principality of Volhynia/Galicia, is probably behind a sample from Lutsk (VK541), and belongs to hg. R1a-L1029 (a subclade of R1a-M458), showing ca. 95% of “Modern Polish” ancestry.
  • Yaroslav Osmomysl, the 12th century Prince of Halych (now in Western Ukraine), was probably of hg. E1b-V13, yet another clearly early Slavic haplogroup.

vikings-y-dna-haplogroup-n1a
Density of haplogroup N1a-VL29, N1a-L550 (samples in pink, most not visible) among Vikings. Samples of hg. R1b in blue, hg. R1a in green, hg. I in orange.

Finnish ancestry

Firstly, modern Finnish individuals are not like ancient Finnish individuals, modern individuals have ancestry of a population not in the reference; most likely Steppe/Russian ancestry, as Chinese are in the reference and do not share this direction. Ancient Swedes and Norwegians are more extreme than modern individuals in PC2 and 4. Ancient UK individuals were more extreme than Modern UK individuals in PC3 and 4. Ancient Danish individuals look rather similar to modern individuals from all over Scandinavia. By using a supervised ancient panel, we have removed recent drift from the signal, which would have affected modern Scandinavians and Finnish populations especially. This is in general a desirable feature but it is important to check that it has not affected inference.

ancient-modern-finns-steppe
PCA of the ancient and modern samples using the ancient palette, showing different PCs. Modern individuals are grey and the K=7 ancient panel surrogate populations are shown in strong colors, whilst the remaining M-K=7 ancient populations are shown in faded colors.

The story for Modern-vs-ancient Finnish ancestry is consistent, with ancient Finns looking much less extreme than the moderns. Conversely, ancient Norwegians look like less-drifted modern Norwegians; the Danish admixture seen through the use of ancient DNA is hard to detect because of the extreme drift within Norway that has occurred since the admixture event. PC4 vs PC5 is the most important plot for the ancient DNA story: Sweden and the UK (along with Poland, Italy and to an extent also Norway) are visibly extremes of a distribution the same “genes-mirror-geography” that was seen in the Ancient-palette analysis. PC1 vs PC2 tells the same story – and stronger, since this is a high variance-explained PC – for the UK, Poland and Italy.

Uniform manifold approximation and projection (UMAP) analysis of the VA and other ancient samples.

Evidence for Pictish Genomes

The four ancient genomes of Orkney individuals with little Scandinavian ancestry may be the first ones of Pictish people published to date. Yet a similar (>80% “UK ancestry) individual was found in Ireland (VK545) and five in Scandinavia, implying that Pictish populations were integrated into Scandinavian culture by the Viking Age.

Our interpretation for the Orkney samples can be summarised as follows. Firstly, they represent “native British” ancestry, rather than an unusual type of Scandinavian ancestry. Secondly, that this “British” ancestry was found in Britain before the Anglo-Saxon migrations. Finally, that in Orkney, these individuals would have descended from Pictish populations.

vikings-british-ancestry
Natural neighbor interpolation of “British ancestry” among Vikings.

(…) ‘UK’ represents a group from which modern British and Irish people all receive an ancestry component. This information together implies that within the sampling frame of our data, they are proxying the ‘Briton’ component in UK ancestry; that is, a pre-Roman genetic component present across the UK. Given they were found in Orkney, this makes it very likely that they were descended from a Pictish population.

Modern genetic variation within the UK sees variation between ‘native Briton’ populations Wales, Scotland, Cornwall and Ireland as large compared to that within the more ‘Anglo-Saxon’ English. This is despite subsequent gene flow into those populations from English-like populations. We have not attempted to disentangle modern genetic drift from historically distinct populations. Roman-era period people in England, Wales, Ireland and Scotland may not have been genetically close to these Orkney individuals, but our results show that they have a shared genetic component as they represent the same direction of variation.

Density of haplogroup R1b-L21 (samples in red), overlaid over all samples of hg. R1b among Vikings (R1b-U106 in green, other R1b-L151 in deep red). To these samples one may add the one from Janakkala in south-western Finland (AD ca. 1300), of hg. R1b-L21, possibly related to these population movements.

For more on Gaelic ancestry and lineages likely representing slaves among early Icelanders, see Ebenesersdóttir et al. (2018).

Y-DNA

As in the case of mitochondrial DNA, the overall distribution profile of the Y chromosomal haplogroups in the Viking Age samples was similar to that of the modern North European populations. The most frequently encountered male lineages were the haplogroups I1, R1b and R1a.

Haplogroup I (I1, I2)

The distribution of I1 in southern Scandinavia, including a sample from Sealand (VK532) ca. AD 100 (see Iron Age Y-DNA maps) proves that it had become integrated into the West Germanic population already before their expansions, something that we already suspected thanks to the sampling of Germanic tribes.

vikings-y-dna-haplogroup-i
Density of haplogroup I (samples in orange) among Vikings. Samples of hg. R1b in blue, hg. R1a in green, N1a in pink.
vikings-y-dna-haplogroup-i1-over-i
Density of haplogroup I1 (samples in red) overlaid over all samples of hg. I among Vikings.

Haplogroup R1b (M269, U106, P312)

Especially interesting is the finding of R1b-L151 widely distributed in the historical Nordic Bronze Age region, which is in line with the estimated TMRCA for R1b-P312 subclades found in Scandinavia, despite the known bottleneck among Germanic peoples under U106. Particularly telling in this regard is the finding of rare haplogroups R1b-DF19, R1b-L238, or R1b-S1194. All of that points to the impact of Bell Beaker-derived peoples during the Dagger period, when Pre-Proto-Germanic expanded into Scandinavia.

Also interesting is the finding of hg. R1b-P297 in Troms, Norway (VK531) ca. 2400 BC. R1b-P297 subclades might have expanded to the north through Finland with post-Swiderian Mesolithic groups (read more about Scandinavian hunter-gatherers), and the ancestry of this sample points to that origin.

However, it is also known that ancestry might change within a few generations of admixture, and that the transformation brought about by Bell Beakers with the Dagger Period probably reached Troms, so this could also be a R1b-M269 subclade. In fact, the few available data from this sample show that it comes from the natural harbour Skarsvågen at the NW end of the island Senja, and that its archaeologist thought it was from the Viking period or slightly earlier, based on the grave form. From Prescott (2017):

In 1995, Prescott and Walderhaug tentatively argued that a dramatic transformation took place in Norway around the Late Neolithic (2350 BCE), and that the swift nature of this transition was tied to the initial Indo-Europeanization of southern and coastal Norway, at least to Trøndelag and perhaps as far north as Troms. (…)

The Bell Beaker/early Late Neolithic, however, represents a source and beginning of these institution and practices, exhibits continuity to the following metal age periods and integrated most of Northern Europe’s Nordic region into a set of interaction fields. This happened around 2400 BCE, at the MNB to LN transition.

NOTE. This particular sample is not included in the maps of Viking haplogroups.

vikings-y-dna-haplogroup-r1b
Density of haplogroup R1b (samples in blue) among Vikings. Samples of hg. I in orange, hg. R1a in green, N1a in pink.
vikings-y-dna-haplogroup-r1b-U106-over-r1b
Density of haplogroup R1b-U106 (samples in green) overlaid over all samples of hg. R1b (other R1b-L23 samples in red) among Vikings.
vikings-y-dna-haplogroup-r1b-P312-over-r1b
Density of R1b-L151 (xR1b-U106) (samples in deep red) overlaid over all samples of hg. R1b (R1b-U106 in green, other R1b-M269 in blue) among Vikings.

Haplogroup R1a (M417, Z284)

The distribution of hg. R1a-M417, in combination with data on West Germanic peoples, shows that it was mostly limited to Scandinavia, similar to the distribution of I1. In fact, taking into account the distribution of R1a-Z284 in particular, it seems even more isolated, which is compatible with the limited impact of Corded Ware in Denmark or the Northern European Plain, and the likely origin of R1a-Z284 in the expansion with Battle Axe from the Gulf of Finland. The distribution of R1a-Z280 (see map above) is particularly telling, with a distribution around the Baltic Sea mostly coincident with that of N1a.

vikings-y-dna-haplogroup-r1a
Density of haplogroup R1a (samples in green) among Vikings. Samples of hg. R1b in blue, of hg. I in orange, N1a in pink.
vikings-y-dna-haplogroup-r1a-z284-over-r1a
Density of haplogroup R1a-Z284 (samples in cyan) overlaid over all samples of hg. R1a (in green, with R1a-Z280 in pink) among Vikings.

Other haplogroups

Among the ancient samples, two individuals were derived haplogroups were identified as E1b1b1-M35.1, which are frequently encountered in modern southern Europe, Middle East and North Africa. Interestingly, the individuals carrying these haplogroups had much less Scandinavian ancestry compared to the most samples inferred from haplotype based analysis. A similar pattern was also observed for less frequent haplogroups in our ancient dataset, such as G (n=3), J (n=3) and T (n=2), indicating a possible non-Scandinavian male genetic component in the Viking Age Northern Europe. Interestingly, individuals carrying these haplogroups were from the later Viking Age (10th century and younger), which might indicate some male gene influx into the Viking population during the Viking period.

vikings-italian-ancestry
Natural neighbor interpolation of “Italian ancestry” among Vikings.

As the paper says, the small sample size of rare haplogroups cannot distinguish if these differences are statistically relevant. Nevertheless, both E1b samples have substantial Modern Polish-like ancestry: one sample from Gotland (VK474), of hg. E1b-L791, has ca. 99% “Polish” ancestry, while the other one from Denmark (VK362), of hg. E1b-V13, has ca. 35% “Polish”, ca. 35% “Italian”, as well as some “Danish” (14%) and minor “British” and “Finnish” ancestry.

Given the E1b-V13 samples of likely Central-East European origin among Lombards, Visigoths, and especially among Early Slavs, and the distribution of “Polish” ancestry among Viking samples, VK362 is probably a close description of the typical ancestry of early Slavs. The peak of Modern Polish-like ancestry around the Upper Pripyat during the (late) Viking Age suggests that Poles (like East Slavs) have probably mixed since the 10th century with more eastern peoples close to north-eastern Europeans, derived from ancient Finno-Ugrians:

vikings-polish-ancestry
Natural neighbor interpolation of “Polish ancestry” among Vikings.

Similarly, the finding of R1a-M458 among Vikings in Funen, Denmark (VK139), in Lutsk, Poland (VK541), and in Kurevanikha, Russia (VK160), apart from the early Slav from Usedom, may attest to the origin of the spread of this haplogroup in the western Baltic after the Bell Beaker expansion, once integrated in both Germanic and Balto-Slavic populations, as well as intermediate Bronze Age peoples that were eventually absorbed by their expansions. This contradicts, again, my simplistic initial assessment of R1a-M458 expansion as linked exclusively (or even mainly) to Balto-Slavs.

antiquity-europe-y-dna
Y-DNA haplogroups in Europe during Antiquity (full map). See other maps of cultures and ancient DNA from Antiquity.

Related

Pre-Germanic and Pre-Balto-Finnic shared vocabulary from Pitted Ware seal hunters

corded-ware-pitted-ware

I said I would write a post about topo-hydronymy in Europe and Iberia based on the most recent research, but it seems we can still enjoy some more discussions about the famous Vasconic Beakers, by people longing for days of yore. I don’t want to spoil that fun with actual linguistic data (which I already summarized) so let’s review in the meantime one of the main Uralic-Indo-European interaction zones: Scandinavia.

Seal hunting

One of the many eye-catching interpretations – and one of the few interesting ones – that could be found in the relatively recent article Talking Neolithic: Linguistic and Archaeological Perspectives on How Indo-European Was Implemented in Southern Scandinavia, by Iversen & Kroonen AJA (2017) was this:

The borrowing of lexical items from hunter-gatherers into Germanic refers to the potential adoption of Proto-Germanic *selhaz “seal” (Old Norse selr, Old English seolh, Old High German selah) as well as Early Proto-Balto-Finnic *šülkeš “seal” (Finnish hylje, Estonian hüljes) from the marine-oriented Sub-Neolithic Pitted Ware culture.

kroonen-iversen
Modified from Kristiansen et al. (2017), with red circle around the hypothesized interaction of Germanic with hunter-gatherers. “Schematic representation of how different Indo-European branches have absorbed words (circles) from a lost Neolithic language or language group (dark fill) in the reconstructed European linguistic setting of the third millennium BC, possibly involving one or more hunter gatherer languages (light fill) (after Kroonen & Iversen 2017)”.

This is what Kroonen thought about this word in his Etymological Dictionary of Proto-Germanic (2006):

Gmc. *selha– m. ‘seal’ – ON selr m. ‘id.’, Far. selur m. ‘id.’, OSw. siæl m. ‘id.’, Sw. själ c. ‘id.’, OE seolh m. ‘id.’, E seal, OS selah m. ‘id.’, EDu. seel, seel-hont m. ‘id.’, Du. zee-hond c. ‘id.’, OHG selah m. ‘id.’, MHG sele m. ‘id.’ (GM).

A Germanic word with no certain IE etymology. The link with Lith. selė́ti ‘to crawl’ (Torp 1909: 436) is erroneous, as this verb corresponds to PGm. *stelan- (q.v.). The *h may nevertheless correspond to the PIE animal suffix *-ko-, for which see *elha{n)- ‘elk’ and *baruga- ‘boar’.

Focusing on this substrate etymon, coupled with archaeology and ancient DNA, in the recent SAA 84th Annual Meeting (Abstracts in PDF):

Kroonen, Guus (Leiden University) and Rune Iversen

[196] The Linguistic Legacy of the Pitted Ware Culture

The Scandinavian hunter-, fisher- and gatherer-based Pitted Ware culture is chronologically situated in the Neolithic. However, it challenges our traditional view on cultural and social evolution by representing a return to an otherwise abandoned hunter-gatherer lifestyle. In general, the Pitted Ware culture must be seen as an offshoot of the “Sub-Neolithic” societies inhabiting wide parts of northern and northeastern Europe in the fourth and third millennium B.C.E.

Isotopic and aDNA studies have shown that people of the east Swedish Pitted Ware culture, both dietarily and genetically were distinct from the early farmers in this region, the Funnel Beaker culture. Isotopic data shows a marked predominance of seal in the diet, which has given the Pitted Ware people the nickname “Inuit of the Baltic”.

As regards language, it is to be expected that people practicing a Pitted Ware lifestyle spoke a non-Indo-European language. In fact, there is some linguistic evidence that can support this claim. It is conceivable that both the Germanic and Finnish word for “seal” were ultimately borrowed from a language spoken in a Pitted Ware context. Once more, the linguistic evidence turns out to offer important information complementary to that of archaeology and archaeo-genetics.

prehistoric-seal-hunters
Stone Age Seal Hunters, by Måns Sjöberg.

Apparently, the idea of non-IE substrate languages in contact with Germanic in Scandinavia is fashionable for the Copenhagen group, probably due to their particular interpretation of the recent genetic papers, hence the multiple Germanic-Fennic connections to be reviewed through this new prism. While the ulterior motive of this proposal may be to try and connect yet again Germanic with CWC Denmark, I would argue that the effect is actually the opposite.

An early borrowing via Uralic

The word has always been considered a more likely loan from one language to the other, and – because of the quite popular idea of Uralic native to Fennoscandia – it was often seen as a likely borrowing of Germanic from Balto-Finnic. In any possible case, the borrowing in either direction must be quite early, for obvious reasons:

  • If the borrowing had been via late Palaeo-Germanic, the ending in *-xa– would have been reflected in Balto-Finnic, hence an early Palaeo-Germanic to Pre-Balto-Finnic stage would be necessary.
  • If the borrowing had been via late Balto-Finnic, the initial sibilant would be already aspirated, being adopted as *-x– in Palaeo-Germanic, while the ending in *-k– would have remained as such if it was adopted after Grimm’s law ceased to be active.
  • Similarly, a borrowing from a common, non-Indo-European & non-Uralic source would require that it happened during the early stages of both proto-languages to have undergone their respective phonetic changes, and both borrowings chronologically close to each other, to assume a similar vocalism and consonantism of the ultimate source.
wiik-indo-european-uralic-substrate
The idea of seal-hunting Uralic substrate of Pitted Ware is not new. Image modified from The Uralic and Finno-Ugric Phonetic Substratum, by Kalevi Wiik, Linguistica Uralica (1997).

Furthermore, regarding the most likely way of expansion of this loanword, due to the different vowels and sibilants present in Uralic but not in Indo-European:

  • A direct loan from Pre-Germanic **selkos – which shows a regular thematic declension – to Pre-Balto-Finnic *šülkeš doesn’t seem to be a reasonable assumption.
  • NOTE. A Germanic borrowing from alternative Gmc. genitive *silxis could only work in a Pre-Germanic to Pre-Balto-Finnic model, hence only if the Gmc. form can be reconstructed for an earlier stage. Even then, for the same reason stated above, the opposite could be more reasonably argued, i.e. that this form is the original one adopted in Germanic: Pre-PBF *šülkeš > Pre-Gmc. *silkis, reinterpreted as an -o- stem in its declension.

  • If we reconstruct an older Pre-Finno-Samic (i.e. with Finno-Permic-like vocalism) **šëlkëš, a borrowing into Pre-Germanic **selkos would work. Even though no Saami derivative exists to confirm such a possibility, this would be supported by the known common evolution of Finno-Samic dialects in close contact with Pre-Germanic.
  • Admittedly, even accepting the existence of a Finno-Samic stem, a potential substrate word could not be discarded. In fact, while **šëlkë- could perfectly be a Uralic root, the ending in *-š can’t be easily interpreted. Therefore, a third, non-Indo-European & non-Uralic source is a plausible explanation.

NOTE. Arguably, Proto-Finno-Samic could have adopted Gmc. *kh or *x exceptionally as PFS *k. However, early Palaeo-Germanic borrowings in Finno-Samic show a consistent regular consonant change as described above. For more on this, see Finno-Samic borrowings.

This likely Uralic first nature of the loanword is important for the discussion below.

Pitted Ware culture

pitted-ware-pyheensilta-ware-culture
Middle Neolithic A period. Distribution of Pyheensilta Ware, Funnel Beaker Culture in Sweden, and Pitted Ware Culture in northern Europe during the Middle Neolithic A period, c. 3300–2800 cal BC. Find locations with numbers demarcate sites where cereal grains have been found and later AMS radiocarbon dated. Figure was created by SV using QGIS 3.4. (https://www.qgis.org/) and Natural Earth data (https://www.naturalearthdata.com/). Image from Vanhanen et al. (2019).

About the Pitted Ware culture, this is what the recent paper by Vanhanen et al. (2019), from the University of Finland (including Volker Heyd) had to say:

The origins of the PWC are controversial. In one likely scenario, Comb Ceramic and Mesolithic hunter-gatherers first interacted with FBC during the last centuries of the EN and became specialized maritime hunter-gatherers. The PWC pushed south and westwards during the Middle Neolithic (MN), c. 3300–2300 BC, along the northern Baltic shoreline and adjacent islands, eventually reaching as far west as Denmark and southern Norway. Around 2800 BC, after the FBC ceased to exist, the Corded Ware Culture (CWC) migrated into the PWC area. The end date for the PWC and CWC is approximately 2300 BC, when the material culture was replaced by the Late Neolithic (LN) culture<. Spanning nearly a millennium virtually unchanged, the PWC maintained a coherent society and a successful economic model. PWC people lived in marine-oriented settlements, commonly dwelled in huts and produced relatively large amounts of ceramic vessels. This speaks to the partly sedentary nature of their habitation, at least for their base camps. These specialist hunter-gatherers obtained the great majority of their subsistence from maritime sources, such as seal, fish, and sea birds. Considering the amount of bones, sealing was of paramount importance, causing these peoples to be labelled ‘hard-core sealers’ or even the ‘Inuit of the Baltic’.

The Middle Neolithic Pitted Ware culture is dated ca. 3500–2300 BC, so we would be seeing here Pre-Germanic and Pre-Balto-Finnic peoples arriving near the Pitted Ware culture. That would leave us with one of both languages expanding with Corded Ware peoples, and the other with Bell Beakers. Since Battle Axe-derived cultures around the Gulf of Finland are associated with Balto-Finnic groups, and Bell Beakers arriving ca. 2400 started the Dagger Period, commonly associated with the Pre-Germanic community, I think the connection of each group with their language is self-evident.

pitted-ware-cored-ware-culture
Middle Neolithic B period. Distribution of Corded Ware Culture and Pitted Ware Culture in northern Europe during the Middle Neolithic B period, c. 2800–2300 cal BC. Find locations with numbers demarcate sites where cereal grains have been found and later AMS radiocarbon dated. Figure was created by SV using QGIS 3.4. (https://www.qgis.org/) and Natural Earth data (https://www.naturalearthdata.com/). Modified from Vanhanen et al. (2019).

NOTE. You can read some interesting information about prehistoric and recent seal hunting in the Baltic in the blog post “Själen” – Seal Hunting in the Northern Baltic Sea.

Germanic-Fennic phonetic evolution

The common Germanic – Balto-Finnic phonetic evolution, especially Verner’s law in Palaeo-Germanic and qualitative gradation in Proto-Balto-Finnic, has been variably interpreted as:

  • Uralic in Scandinavia influenced by Germanic (Verner’s law source of the gradation), by Koivulehto and Vennemann (1996).
  • Germanic over a Uralic substratum in Scandinavia, by Wiik (1997).
  • Both Germanic and Balto-Finnic influenced by a third language, an “extinct non-Uralic source” spoken in Fennoscandia before the arrival of Uralic and Indo-European, by Kallio (2001); maybe the same substrate proposed to have influenced the accent shift in Germanic similar to Uralic.
  • Balto-Finnic speakers adopting Pre-Germanic in Scandinavia, in contact with Balto-Finnic speakers retaining their language, by Schrijver in Language Contact and the Origins of the Germanic Languages (2014)– although first suggested by him in the 1990s.

NOTE. There are other (some much older) proposals of a Uralic substrate in Scandinavia, but I think those above summarize the most common positions tenable today.

If you add all linguistic, archaeological, and now genetic connections, it is really strange to keep arguing for so many surprisingly fitting common substrates and/or contact languages for both. Especially because the Pre-Germanic community – if originally from southern Scandinavia and not further south (see e.g. Kortlandt’s theory) – was marked by the Dagger Period, as accepted by most archaeologists (including Kristiansen), and we know that Bell Beakers – who triggered the Dagger period – might have arrived a little late to the Pitted Ware disintegration in most seal-hunting areas of southern Scandinavia.

bell-beaker-density
Density analysis based (Bell Beaker per km2) on the distribution of Bell Beaker per region (ca. 2700-2200 BC). Combination of different levels of b-spline interpolation. Exaltation of the values through square root usage. Modified from Michael Bilger (2018).

In other words, how many common substrate languages can we propose for Germanic (and Balto-Finnic)? Just from Kroonen we have already the Semitic-like TRB, and the seal-hunting Pitted Ware culture. Apparently, the culprit of the common phonetic evolution must be some (other?) culture that both Pre-Germanic and Pre-Balto-Finnic assimilated (or with which both were in contact) in Fennoscandia.

NOTE. I believe no data supports the attribution of those Germanic borrowings to the TRB culture, especially if one assumes they belong to an Afroasiatic branch, as did Kroonen. His initial assumption about an expansion of R1b-M269 associated with the Neolithic from Anatolia, and thus with Afroasiatic, must today be rejected. Much more likely is the incorporation of most of these loanwords during the expansion of North-West Indo-Europeans from Yamna Hungary.

How many “common” substrates from different regions and cultures is too much? Arguably, it’s not a question of quantity (because the overall probability remains the same), but a question of quality of arguments.

In my opinion, both a) the marked seal-hunting subsistence economy of the Pitted Ware culture and b) the difficult reconstruction of a fitting ‘natural’ PIE or PU stem warrant this proposal of a third source, just like the European agricultural substrate of North-West Indo-European and Palaeo-Balkan languages, as well as the Asian agricultural substrate of Indo-Iranian are the most logical interpretation of words not found in other IE dialects. The only problem in this case is the lack of other Scandinavian substrate words to compare its typology against.

scandinavia-neolithic-flint-daggers
Close contacts in Fennoscandia. The distribution of Scandinavian flint daggers (A) in the east and south Baltic region and possible trends of “down the line” trade (B). Good size and quality flint zone in the south-west Baltic region is hatched (C). According to: Wojciechowski 1976; Olausson 1983, fig. 1; Madsen 1993, 126; Libera 2001; Kriiska & Tvauri 2002, 86. Image modified from Piličiauskas (2010).

Common Scandinavian substratum

The theory of a Pitted Ware borrowing is therefore quite convincing from a cultural point of view, at the same time as it fits the linguistic data. However, one reason why I dislike the interpretation of a dual origin is that our knowledge of Uralic languages is fairly limited, whereas that of Indo-European branches and hence Proto-Indo-European is huge. To put it otherwise: if a common word appears in both, and it is most likely (culturally and linguistically) not Indo-European, it certainly means that it was borrowed in Germanic. What are the a priori chances of it coming directly from a third substrate language for both dialects, instead of coming directly from Pre-Balto-Finnic?

From Schrijver (2014):

What did happen, apparently, is that Finnic speakers had enough access to the way in which Germanic speakers pronounced Balto-Finnic in order to model their own pronunciation of Balto-Finnic on it. In other words, Balto-Finns conversed with bilingual speakers of Germanic and Balto-Finnic whose pronunciation of both was essentially Germanic. But access to the Germanic language itself was not sufficient to allow Balto-Finns to become bilingual themselves, either because social segregation prevented this or because contact with Germanic was severed before widespread bilingualism set in. This limited access to Germanic would allow us to understand why Balto-Finnic did not go the way of the vernacular languages that came in contact with Latin in the Roman Empire, where access to Latin was open to almost everybody and massive language shift in favour of Latin ensued.

NOTE. For a more detailed discussion, you can read the whole chapter dedicated to this question. I summarized it in Pre-Germanic born out of a Proto-Finnic substrate in Scandinavia.

On the other hand, about the ad hoc interpretation by Kallio (2001) of hypothetic third languages strongly influencing in the same way both the Palaeo-Germanic- and Balto-Finnic-speaking communities, Schrijver (2014) comments:

The idea that perhaps both languages moved towards a lost third language, whose speakers may have been assimilated to both Balto-Finnic and Germanic, provides a fuller explanation but suffers from the drawback that it shifts the full burden of the explanation to a mysterious ‘language X’ that is called upon only in order to explain the developments in Proto-Germanic and Balto-Finnic. That comes dangerously close to circular reasoning.

early-bronze-age-nordic-dagger-period
Early Bronze Age cultures of Northern Europe (roughly ca. 2200-1750). Dagger period representing the expansion of BBC-derived groups from southern Scandinavia.

NOTE. The proposal of some kind of “SHG/EHG-based Fennoscandian substrate” seems funny to me, for two reasons: firstly, there is usually no talk about which culture spread that common language, how it survived, how it was in contact with both groups and until when, etc. (see below for possibilities); secondly, apparently the evident survival of West European EEF communities driven by at least two cultural groups – El Argar and the poorly known groups from the Atlantic façade north of the Pyrenees – is, for the same people proposing this simplistic SHG/EHG idea, somehow not fitting for the prehistory of Proto-Iberian and Proto-Aquitanian, respectively…

The same argument that one could use against the direct borrowing of both dialects from Pitted Ware, but much more strongly, can be thus wielded against a common, centuries-long phonetic evolution of both Balto-Finnic and Germanic caused by close interactions with (and/or substrate influence of) some third language. Which unitary culture and when exactly could that have happened around the Baltic Sea?

  • Was it Pitted Ware the mysterious substrate language? Seems rather unlikely, due to the early demise of the Pitted Ware culture in contrast to the long-lasting common influence seen in both dialects.
  • Was it Pitted Ware in southern Scandinavia, but Comb Ware in the Gulf of Finland? Is there a direct genetic connection between both cultures? And how likely is a common phonology of an ancestral Comb Ware-like substrate language surviving separately in Finland and Sweden? Even accepting these assumptions, we would be stuck again in the Indo-European Beakers vs. Uralic Battle Axe model.
  • Was it a succession of cultures, from some Scandinavian culture that was replaced by some incoming ethnolinguistic group, then influencing the other? This non-IE, non-Uralic substrate would then need to be proposed, given the chronological and archaeological constraints, as an effect of Pitted Ware over Pre-Finno-Baltic spoken by Battle Axe peoples in Scandinavia, then replaced by Pre-Germanic peoples arriving later with Bell Beakers. A reverse direction and later chronology (say, Germanic replaced by Balto-Finnic from Netted Ware arriving from the Volga) wouldn’t work as well.
  • Was it Asbestos Ware as a late Comb Ware group influencing both? How likely is such a continued influence in Southern Scandinavia and the Gulf of Finland? Even if we accepted this influence that miraculously didn’t affect Samic (most likely located between the Balto-Finnic-speaking Gulf of Finland and northern Fennoscandian Asbestos Ware groups), it would necessarily mean that Germanic and Balto-Finnic were spoken neighbouring exactly the same Asbestos Ware groups in Scandinavia. That is, essentially, that the BBC-derived Dagger Period represented Pre-Germanic, while Battle Axe-derived groups around the Gulf of Finland were Balto-Finnic.

Mixing linguistics with archaeology (now complemented with genetics) also risks circular reasoning. But, how else can someone propose a third substrate language for a phonetic change, necessarily represented by Fennoscandian groups potentially separated by thousands of years? In this age of population genomics we can’t simply talk about theoretical models anymore: we must refer to Fennoscandian cultures and populations in a very specific time frame, as Kronen & Iversen do in their proposal. Not only is such a third unknown language usually a weak explanation for a common development of two unrelated languages; in this case it finds no support whatsoever.

Seals and the Arctic

Another interesting aspect about this Fennic-Germanic comparandum is its relevance to the Uralic homeland problem.

uralic-languages-modern
Current distribution of Uralic languages. Nenets and Saami are among the best positioned to retain the ‘original’ Uralic seal-hunting vocabulary.

Since the publication of Mittnik et al. (2018), Lamnidis et al. (2018), and Sikora et al. (2018), the new normal is apparently to consider Corded Ware Finland as Germanic-speaking, the Gulf of Finland as Balto-Slavic-speaking, while the Kola peninsula and whichever Palaeo-Arctic peoples preceded Nganasans and Nenets as ancient Uralians. Uh-huh, OK.

But, if prehistoric Arctic peoples practiced specialized seal-hunting economies, and Uralians were one among such populations – supposedly one widespread from the Barents Sea to the Lapteve Sea…how come no common Uralic word for ‘seal’ exists? In other words, why would these True™ Uralic peoples expanding from the Arctic need to borrow a word for ‘seal’ from neighbouring populations in every single seal-hunting region they are attested?

grey-seal-distribution
Historical distribution of grey seals, an important part of the diet around the Baltic Sea. Image modified from Wikimedia to include Skagerrak and Kattegat regions.

About Saami, which some have recklessly proposed to be derived from Bronze Age N1c-L392 samples from the Kola Peninsula (against the good judgment of the authors of the paper), this is what we know from their word for ‘seal’, from Grünthal (2004):

Ter Saami vīrre ‘seal; wolf’ displays two meanings that refer to clearly different animals. Neither of them is borrowed from the source language because the word descends from Russian zver’ ‘animal’ (T.I.Itkonen 1958: 756). Another word, Skolt Saami näúdd ‘seal, wolf’, has been similarly used in the two meanings. The evidence of North Saami návdi ‘wolf; creature, fur animal; beast’ (Sammallahti 1989: 305; Lagercrantz (1939: 518) presents the alternative meanings in the opposite order; E. Itkonen (1969: 148) lists the meanings ‘wildes Tier; Raubtier (bes. Wolf); Pelztier’) suggesting that ‘wolf’ is the primary sense and ‘seal’ is a metaphorical extension of it. More precisely, it is an example of a mythic metaphor (cf. Siikala 1992). According to the old folk belief, seal was a wolf and the Skolt Saamis preferred not to eat its meat (T.I.Itkonen 1958: 906). Before that the metonymic meaning ‘wolf’ rose from the less specified meanings, and originally návdi is a Scandinavian or Finnic loan word in Saamic, cf. Old Norse naut ‘vieh, rind’, Icelandic and Norwegian naut, Swedish nöt < Germanic *nauta ‘property’ (Hellquist 1980: 721, T.I.Itkonen 1958: 275, Lagercrantz 1939: 518, de Vries 1961: 406; E. Itkonen (1969: 148) considers Finnic, cf. Finnish nauta ‘bovine’ (< Germanic) as a possible alternative source for the Saamic word).

NOTE. Possibly comparable, for the mythic metaphor proper of Scandinavian folk belief, are Germanic derivatives built as ‘seal-hound’ and/or ‘sea-hound’.

sea-distribution-arctic
Seals formed a great part of the diet for Palaeo-Arctic populations. Boundaries of regions used to predict sea ice, superimposed over the distributions of the five ringed seal subspecies. Image modified from Kelly et al. (2010).

About Nenets (quite close to the Naganasans of pure “Siberian ancestry”), here is what Edward Vajda, an expert in Palaeo-Siberian languages, has to say:

Nenets techniques for hunting the animals of the Arctic Ocean seem to have been borrowed from the first Arctic aborigines. Thus, the Nenets word for seal is nyak, the Eskimo word is nesak. Also, the Nenets word for a one-piece Arctic clothing is lu; the Korak word on the Kamchatka peninsula for clothing is l’ku. All of these groups may have borrowed the words from some original circumpolar aborigines. More probably, the first settlers of Arctic Europe were cousins of the present-day Eskimo, Chukchi and other residents of the far northeast region of Asia. Nenets folklore also speaks of the aborigines living in ice dugouts (igloos).

On the other hand, Proto-Uralic shows a Chalcolithic steppe-like culture, with common words for metal and metalworking, for agriculture, and for domesticated animals, most likely including cattle. They were close to Indo-Europeans since at least before the Tocharian split, and probably earlier than that (even if one does not accept the Indo-Uralic phylum). And there were clearly strong contacts of Finno-Ugric with Indo-Iranian, and especially of Finno-Samic with Germanic.

uralic-cline
Uralic clines from Corded Ware groups to the east. A clear reason for the lack of common seal-hunting vocabulary. Modified from Tambets et al. (2018). Principal component analysis (PCA) and genetic distances of Uralic-speaking populations. a PCA (PC1 vs PC2) of the Uralic-speaking populations. You can see another PCA including ancient samples.

Some among my readers may now be thinking about these totally believable proposals of prehistoric cultures around Lake Baikal representing the True™ Uralic homeland; because haplogroup N1c, and because some 0.5% more “Devil’s Gate Cave ancestry” in Estonians than in Lithuanians; despite the fact that 1) the so-called “Siberian ancestry” formed an ancestral cline with EHG in North Eurasia, that 2) N1c-L392 lineages seem to appear among many Asian peoples of different languages, and that 3) recent prehistoric N1c-L392 lines expanded clearly with Micro-Altaic languages.

Like, who would have hunted seals in Lake Baikal, right? The problem is, seals represented one of their main game, essential for their subsistence economy. From Novokonova et al. (2015):

One of the key reasons for the density of human settlement in the Baikal region compared to adjacent areas of Siberia is that the lake and its nearby rivers offer an abundance of aquatic food resources, including several endemic species, with perhaps the most well known being the Baikal seal. This freshwater seal is only found in Lake Baikal and portions of its tributaries. It shares lifecycle and behavioral patterns with other small northern ice-adapted seals, and is genetically and morphologically most closely related to the ringed seal (Pusa hispida). The nerpa can grow up to 1.8 m long and weigh as much as 130 kg, with the males tending to be slightly larger than the females.

Zooarchaeological analyses of the 16,000 Baikal seal remains from this well-dated site clearly show that sealing began here at least 9000 calendar years ago. The use of these animals at Sagan-Zaba appears to have peaked in the Middle Holocene, when foragers used the site as a spring hunting and processing location for yearling and juvenile seals taken on the lake ice. After 4800 years ago, seal use declined at the site, while the relative importance of ungulate hunting and fishing increased. Pastoralists began occupying Sagan-Zaba at some point during the Late Holocene, and these groups too utilized the lake’s seals. Domesticated animals are increasingly common after about 2000 years ago, a pattern seen elsewhere in the region, but spring and some summer hunting of seals was still occurring. This use of seals by prehistoric herders mirrors patterns of seal use among the region’s historic and modern groups.

Bronze Age movements in Fennoscandia

Regarding the shrinkage and expansion of different farming economic strategies in Scandinavia since the Neolithic, with potential relevance for population movements and thus ethnolinguistic change – either from Balto-Finnic peoples migrating back from eastern Sweden, or Germanic peoples moving to eastern Finland – from Vanhanen et al. (2019):

Cultivated plants at CWC sites in Finland were not discovered in the current investigation (Supplementary Results) or earlier studies. In Finland, the keeping of domestic animals is indicated by the evidence of dairy lipids and mineralized goat hairs. Charred remains and impressions of cultivated plants have been discovered at CWC sites in Estonia and east-central Sweden (Fig. 3: 12). In the eastern Baltic region, the earliest bones of domestic animals and a shift in subsistence occurred with the CWC. Whether CWC produced the cereals and other agricultural products found at PWC sites is difficult to estimate because only small amounts of plant remains have ever been discovered at CWC sites. The CWC seemingly reached east-central Sweden from regions further to the east, where there is evidence of animal husbandry, but only very few signs of plant cultivation.

For the Late Neolithic (LN), cereal grains have been found north of Mälaren and along the Norrland coast. In mainland Finland, the first cereal grains occur during the LN or Bronze Age, c. 1900–1250 cal BC. The earliest bones of sheep/goat from mainland Finland are earlier, dating back to 2200–1950 cal BC. Finds of Scandinavian bronze artefacts indicate an influx from east-central Sweden, which might well be a source area for these agricultural innovations. A similar development is found in the eastern Baltic region, where the earliest directly radiocarbon-dated cereals originate from the Bronze Age, 1392–1123 cal BC (2 sigma). Thus, agriculture was evident during the Bronze Age in the eastern Baltic, but at least animal keeping and probably crop cultivation were present earlier during the CWC phase.

It has been known for a while already that the only options left for the expansion of Finno-Saami into Fennoscandia are either Battle Axe (continued in Textile Ceramics) or Netted Ware (as proposed e.g. by Parpola), based, among other data, on language contacts, language estimates, cultural evolution, and population genomics. Data like this one on seal-hunting vocabulary also support the most likely option, which entails the identification of Corded Ware as the vector of expansion of Uralic languages.

NOTE. Also interesting in this regard is the lack of Slavic words for ‘seal’ – borrowed, in Russian from Samic, and in other Slavic dialects from Russian, Latin, or other languages -, and the coinage of a new term in East Baltic. Rather odd for an “autochthonous” Proto-Baltic (supposedly in contact with Pitted Ware, Germanic, and Balto-Finnic, then), and for a Proto-Slavic stemming from the Baltic. Quite appropriate, though, for a Proto-East Baltic arriving in the Baltic with Trzciniec and for a Proto-Slavic community evolving further south.

So, what new episode in this renewed 2000s R1b/R1a/N1c soap opera is it going to be, when eastern Fennoscandia shows Corded Ware-derived peoples of “steppe ancestry” (and mainly R1a-Z645 lineages) continue during the Bronze Age? Will the resurge and/or infiltration of I2 – maybe even N1c – lineages among Corded Ware-derived cultures of north-eastern Europe support or challenge this model, and why? Make your bet below.

Related

How the genocidal Yamnaya men loved to switch cultures

yamnaya-expansion-bell-beaker

After some really interesting fantasy full of arrows, it seems Kristiansen & friends are coming back to their most original idea from 2015, now in New Scientist’s recent clickbait Story of most murderous people of all time revealed in ancient DNA (2019):

Teams led by David Reich at Harvard Medical School and Eske Willerslev at the University of Copenhagen in Denmark announced, independently, that occupants of Corded Ware graves in Germany could trace about three-quarters of their genetic ancestry to the Yamnaya. It seemed that Corded Ware people weren’t simply copying the Yamnaya; to a large degree they actually were Yamnayan in origin.

If you think you have seen that movie, it’s because you have. They are at it again, Corded Ware from Yamna, and more “steppe ancestry” = “more Indo-European. It seems we haven’t learnt anything about “Steppe ancestry” since 2015. But there’s more:

Genocidal peoples who “switch cultures”

Burial practices shifted dramatically, a warrior class appeared, and there seems to have been a sharp upsurge in lethal violence. “I’ve become increasingly convinced there must have been a kind of genocide,” says Kristian Kristiansen at the University of Gothenburg, Sweden.

The collaboration revealed that the origin and initial spread of Bell Beaker culture had little to do – at least genetically – with the expansion of the Yamnaya or Corded Ware people into central Europe. “It started in It is in that region that the earliest Bell Beaker objects – including arrowheads, copper daggers and distinctive Bell-shaped pots – have been found, in archaeological sites carbon-dated to 4700 years ago. Then, Bell Beaker culture began to spread east, although the people more or less stayed put. By about 4600 years ago, it reached the most westerly Corded Ware people around where the Netherlands now lies. For reasons still unclear, the Corded Ware people fully embraced it. “They simply take on part of the Bell Beaker package and become Beaker people,” says Kristiansen.

The fact that the genetic analysis showed the Britons then all-but disappeared within a couple of generations might be significant. It suggests the capacity for violence that emerged when the Yamnaya lived on the Eurasia steppe remained even as these people moved into Europe, switched identity from Yamnaya to Corded Ware, and then switched again from Corded Ware to Bell Beaker.

Notice what Kristiansen did there? Yamnaya men “switched identities” into Corded Ware, then “switched identities” into Bell Beakers…So, the most aggresive peoples who have ever existed, exterminating all other Europeans, were actually not so violent when embracing wholly different cultures whose main connection is that they built kurgans (yes, Gimbutas lives on).

NOTE. By the way, just so we are clear, only Indo-Europeans are “genocidal”. Not like Neolithic farmers, or Palaeolithic or Mesolithic populations, or more recent Bronze Age or Iron Age peoples, who also replaced Y-DNA from many regions…

yamnaya-corded-ware-bell-beaker

In fact, there is much stronger evidence that these Yamnaya Beakers were ruthless. By about 4500 years ago, they had pushed westwards into the Iberian Peninsula, where the Bell Beaker culture originated a few centuries earlier. Within a few generations, about 40 per cent of the DNA of people in the region could be traced back to the incoming Yamnaya Beakers, according to research by a large team including Reich that was published this month. More strikingly, the ancient DNA analysis reveals that essentially all the men have Y chromosomes characteristic of the Yamnaya, suggesting only Yamnaya men had children.

“The collision of these two populations was not a friendly one, not an equal one, but one where the males from outside were displacing local males and did so almost completely,” Reich told New Scientist Live in September. This supports Kristiansen’s view of the Yamnaya and their descendants as an almost unimaginably violent people. Indeed, he is about to publish a paper in which he argues that they were responsible for the genocide of Neolithic Europe’s men. “It’s the only way to explain that no male Neolithic lines survived,” he says.

So these unimaginably violent Yamnaya men had children exclusively with their Y chromosomes…but not Dutch Single Grave peoples. These great great steppe-like northerners switched culture, cephalic index…and Y-chromosome from R1a (and others) to R1b-L151 to expand Italo-Celtic From The West™.

It’s hilarious how (exactly like their latest funny episode of PIE from south of the Caucasus) this new visionary idea copied by Copenhagen from amateur friends (or was it the other way around?) had been already rejected before this article came out, in Olalde et al. (2019), and that “Corded Ware=Indo-European” fans have become a parody of themselves.

What’s not to love about 2019 with all this back-and-forth hopping between old and new pet theories?

NOTE. I would complain (again) that the obsessive idea of the Danes is that Denmark CWC is (surprise!) the Pre-Germanic community, so it has nothing to do with “steppe ancestry = Indo-European” (or even with “Corded Ware = Indo-European”, for that matter), but then again you have Koch still arguing for Celtic from the West, Kortlandt still arguing for Balto-Slavic from the east, and – no doubt worst of all – “R1a=IE / R1b=Vasconic / N1c=Uralic” ethnonationalists arguing for whatever is necessary right now, in spite of genetic research.

So prepare for the next episode in the nativist and haplogroup fetishist comedy, now with western and eastern Europeans hand in hand: Samara -> Khvalynsk -> Yamnaya -> Bell Beaker spoke Vasconic-Tyrsenian, because R1b. Wait for it…

Vanguard Yamnaya groups

On a serious note, interesting comment by Heyd in the article:

A striking example of this distinction is a discovery made near the town of Valencina de la Concepción in southern Spain. Archaeologists working there found a Yamnaya-like kurgan, below which was the body of a man buried with a dagger and Yamnaya-like sandals, and decorated with red pigment just as Yamnaya dead were. But the burial is 4875 years old and genetic information suggests Yamnaya-related people didn’t reach that far west until perhaps 4500 years ago. “Genetically, I’m pretty sure this burial has nothing to do with the Yamnaya or the Corded Ware,” says Heyd. “But culturally – identity-wise – there is an aspect that can be clearly linked with them.” It would appear that the ideology, lifestyle and death rituals of the Yamnaya could sometimes run far ahead of the migrants.

NOTE. I have been trying to find which kurgan is this, reviewing this text on the archaeological site, but didn’t find anything beyond occasional ochre and votive sandals, which are usual. Does some reader know which one is it?

yamna-expansion-bell-beakers
Yamna expansion and succeeding East Bell Beaker expansion, without color on Bell Beaker territories. Notice vanguard Yamna groups in blue where East Bell Beakers later emerge. See original image with Bell Beaker territories.

Notice how, if you add all those vanguard Yamna findings of Central and Western Europe, including this one from southern Spain, you begin to get a good idea of the territories occupied by East Bell Beakers expanding later. More or less like vanguard Abashevo and Sintashta finds in the Zeravshan valley heralded the steppe-related Srubna-Andronovo expansions in Turan…

It doesn’t seem like Proto-Beaker and Yamna just “crossed paths” at some precise time around the Lower Danube, and Yamna men “switched cultures”. It seems that many Yamna vanguard groups, probably still in long-distance contact with Yamna settlers from the Carpathian Basin, were already settled in different European regions in the first half of the 3rd millennium BC, before the explosive expansion of East Bell Beakers ca. 2500 BC. As Heyd says, there are potentially many Yamna settlements along the Middle and Lower Danube and tributaries not yet found, connecting the Carpathian Basin to Western and Northern Europe.

These vanguard groups would have more easily transformed their weakened eastern Yamna connections with the fashionable Proto-Beaker package expanding from the west (and surrounding all of these loosely connected settlements), just like the Yamna materials from Seville probably represent a close cultural contact of Chalcolithic Iberia with a Yamna settlement (the closest known site with Yamna traits is near Alsace, where high Yamna ancestry is probably going to be found in a Bell Beaker R1b-L151 individual).

This does not mean that there wasn’t a secondary full-scale migration from the Carpathian Basin and nearby settlements, just like Corded Ware shows a secondary (A-horizon?) migration to the east with R1a-Z645. It just means that there was a complex picture of contacts between Yamna and European Chalcolithic groups before the expansion of Bell Beakers. Doesn’t seem genocidal enough for a popular movie, tho.

Related

Phylogeny of leprosy, relevant for prehistoric Eurasian contacts

leprosy-medieval-europe

Some interesting studies were published at roughly the same time as Damgaard et al. (Nature 2018 and Science 2018), and that’s probably why they got little attention (at least by me).

Monica H. Green (also in Academia.edu), specialized in History of Medicine, summed up their relevance in Twitter quite well (her text is edited here for clarity):

I’ve been disappointed that three recent exceptional studies of one of the world’s most historically important diseases, leprosy, have gotten so little notice from the science communication. It will take me a few hours to lay out their significance. But I think it’s important to do so.

So, here are the new studies on historical distribution and evolutionary development of Mycobacterium leprae, one of two organisms that causes leprosy (fourth study dropped yesterday!).

  1. Phylogenomics and antimicrobial resistance of the leprosy bacillus Mycobacterium leprae, by Benjak et al., Nature Communications (2018) 9:352.
  2. Abstract:

    Leprosy is a chronic human disease caused by the yet-uncultured pathogen Mycobacterium leprae. Although readily curable with multidrug therapy (MDT), over 200,000 new cases are still reported annually. Here, we obtain M. leprae genome sequences from DNA extracted directly from patients’ skin biopsies using a customized protocol. Comparative and phylogenetic analysis of 154 genomes from 25 countries provides insight into evolution and antimicrobial resistance, uncovering lineages and phylogeographic trends, with the most ancestral strains linked to the Far East. In addition to known MDT-resistance mutations, we detect other mutations associated with antibiotic resistance, and retrace a potential stepwise emergence of extensive drug resistance in the pre-MDT era. Some of the previously undescribed mutations occur in genes that are apparently subject to positive selection, and two of these (ribD, fadD9) are restricted to drug-resistant strains. Finally, nonsense mutations in the nth excision repair gene are associated with greater sequence diversity and drug resistance.

  3. Ancient DNA study reveals HLA susceptibility locus for leprosy in medieval Europeans, by Krause-Kyora et al., Nature Communications (2018) 9:1569
  4. NOTE. I referred to this study in this blog.

  5. Ancient genomes reveal a high diversity of Mycobacterium leprae in medieval Europe, by Schuenemann et al., PLOS Pathogens (2018)
  6. Abstract:

    Studying ancient DNA allows us to retrace the evolutionary history of human pathogens, such as Mycobacterium leprae, the main causative agent of leprosy. Leprosy is one of the oldest recorded and most stigmatizing diseases in human history. The disease was prevalent in Europe until the 16th century and is still endemic in many countries with over 200,000 new cases reported annually. Previous worldwide studies on modern and European medieval M. leprae genomes revealed that they cluster into several distinct branches of which two were present in medieval Northwestern Europe. In this study, we analyzed 10 new medieval M. leprae genomes including the so far oldest M. leprae genome from one of the earliest known cases of leprosy in the United Kingdom—a skeleton from the Great Chesterford cemetery with a calibrated age of 415–545 C.E. This dataset provides a genetic time transect of M. leprae diversity in Europe over the past 1500 years. We find M. leprae strains from four distinct branches to be present in the Early Medieval Period, and strains from three different branches were detected within a single cemetery from the High Medieval Period. Altogether these findings suggest a higher genetic diversity of M. leprae strains in medieval Europe at various time points than previously assumed. The resulting more complex picture of the past phylogeography of leprosy in Europe impacts current phylogeographical models of M. leprae dissemination. It suggests alternative models for the past spread of leprosy such as a wide spread prevalence of strains from different branches in Eurasia already in Antiquity or maybe even an origin in Western Eurasia. Furthermore, these results highlight how studying ancient M. leprae strains improves understanding the history of leprosy worldwide.

  7. The genome sequence of a SNP type 3K strain of Mycobacterium leprae isolated from a seventh‐century Hungarian case of lepromatous leprosy, by Mendum et al., International Journal of Osteoarchaeology (2018).
  8. Abstract:

    We report on a Mycobacterium leprae genome isolated from the remains of an individual with lepromatous leprosy that were excavated from a seventh‐century Hungarian cemetery. We determined that the genome was from a single nucleotide polymorphism (SNP) type 3K0 M. leprae strain, a lineage that diverged early from other M. leprae lineages. This is one of the earliest 3K0 M. leprae genomes to be sequenced to date. A number of novel SNPs as well as SNPs characteristic of the 3K0 lineage were confirmed by conventional polymerase chain reaction and Sanger sequencing. Recovery of accompanying human DNA from the burial was poor, particularly when compared with that of the pathogen. Modern 3K0 M. leprae strains have only been isolated from East Asia and the Pacific, and so these findings require new scenarios to describe the origins and routes of dissemination of leprosy during antiquity that have resulted in the modern phylogeographical distribution of M. leprae.

A fifth study can be added to the list, which, though not as extensive, is significant because it validates findings of others: Mycobacterium leprae genomes from naturally infected nonhuman primates, by Honap et al. PLOS Neglected Tropical Diseases (2018).

Abstract:

Leprosy is caused by the bacterial pathogens Mycobacterium leprae and Mycobacterium lepromatosis. Apart from humans, animals such as nine-banded armadillos in the Americas and red squirrels in the British Isles are naturally infected with M. leprae. Natural leprosy has also been reported in certain nonhuman primates, but it is not known whether these occurrences are due to incidental infections by human M. leprae strains or by M. leprae strains specific to nonhuman primates. In this study, complete M. leprae genomes from three naturally infected nonhuman primates (a chimpanzee from Sierra Leone, a sooty mangabey from West Africa, and a cynomolgus macaque from The Philippines) were sequenced. Phylogenetic analyses showed that the cynomolgus macaque M. leprae strain is most closely related to a human M. leprae strain from New Caledonia, whereas the chimpanzee and sooty mangabey M. leprae strains belong to a human M. leprae lineage commonly found in West Africa. Additionally, samples from ring-tailed lemurs from the Bezà Mahafaly Special Reserve, Madagascar, and chimpanzees from Ngogo, Kibale National Park, Uganda, were screened using quantitative PCR assays, to assess the prevalence of M. leprae in wild nonhuman primates. However, these samples did not show evidence of M. leprae infection. Overall, this study adds genomic data for nonhuman primate M. leprae strains to the existing M. leprae literature and finds that this pathogen can be transmitted from humans to nonhuman primates as well as between nonhuman primate species. While the prevalence of natural leprosy in nonhuman primates is likely low, nevertheless, future studies should continue to explore the prevalence of leprosy-causing pathogens in the wild.

These five studies are doing whole-genome sequencing on either modern isolates of M. leprae, or genomic fragments retrieved from buried remains (aDNA). The main objective of all the studies is to understand the diversity of M. leprae, both in terms of its history and in terms of its present-day distribution. (Benjak et al. 2018 are especially concerned to study possible reasons for variance in multiple drug resistance).

The following comments are concerned only to discuss leprosy’s history.

So, let’s start with a common claim of the science communication pieces on Schuenemann et al. 2018, which was published last week. A common formula: “New Study Suggests Leprosy Began To Spread From Europe To The World“. Is it plausible that Europe was where leprosy originated as human disease?

The answer, actually, is no. There’s two reasons for this, one having to do with chronology, the other with geography.

For chronology, these studies cumulatively suggest we are looking at a bottleneck. The current Time to Most Recent Common Ancestor (TMRCA) suggested for the divergence of M. leprae from its closest known “cousin,” M. lepromatosis (which also causes leprosy in humans) is estimated to be ca. 13.9 million years. There were no humans around 13.9M ya. So we cannot have been M. leprae‘s original host. All studies being discussed here agree on a consensus phylogeny, which puts the origin of all known strains of M. leprae at about 4-5K ya. So when we talk about the “origin” of M. leprae, we only talking about those lineages formed after this bottleneck.

Next we have to look at geography. Let’s start with this statement from the most recent study, Mendum et al. 2018, which is discussing a genome sequenced from an individual in Hungary from the 7th c. CE: “Modern 3K0 M. leprae strains have only been isolated from East Asia and the Pacific and so these findings require new scenarios to describe the origins and routes of dissemination of leprosy during antiquity that have resulted in the modern phylogeographical distribution of M. leprae.”

Okay, so stop and consider the implications of this. We have someone from 7th c. Hungary with leprosy. The strain of M. leprae that he has is not most closely related to strains sequenced earlier in Denmark or Sweden or England (see Schuenemann et al. 2018, refs. 9, 20, & 21). Rather, the strain he has (3K0) is most closely related to modern strains currently documented on the Pacific Rim, a very, very long way from Hungary. Here are the summary reflections of Mendum et al. 2018:

The global distribution of 3K0 and 3K1 strains is today restricted to regions of the Western Pacific such as Japan (except Okinawa), Korea, China, The Philippines, New Caledonia and Indonesia amongst others (Kai et al, 2013; Avanzi et al, 2015; Monot el al, 2009; Weng et al, 2013; Honap et al, 2018). This could indicate that the 3K lineage originated in Northern or Eastern Asia. The presence of two type 3K cases (KD271 and 222) in early medieval Hungary would then suggest a route of dissemination from Asia to central Europe, perhaps via trade links or migrations. This would be consistent with what is known of the origins of the Pannonian Avars, who are believed to have reached the Hungarian plain from the Eurasian steppe in the late 6th to early 9th centuries (Curta, 2006). The other possibility is that Europe was a centre of dissemination of the ancestral 3K0 and related strains, some of which later became less common or even absent from Europe but persisted in East Asia and the Pacific. Determining the likelihood of each of these scenarios will require more sampling and characterisation of both ancient and modern strains.

Two things bear stressing:

  1. The lineage in which the Hungarian sample has been placed, Lineage 0, has now been documented in historical remains from Denmark, too. (Schuenemann et al. 2018) So whatever transmission routes are postulated to connect the Pacific Rim to Hungary, we will also need to postulate routes to connect the Pacific Rim to Denmark.
  2. Schuenemann et al. 2018 document four of the five known M. leprae lineages in medieval western Europe. (Mendum et al. 2018 now declare the existence of 6 lineages; see tree.)
leprosy-phylogenetic-tree
Phylogenetic relationships between selected modern (regular text) and ancient (bold text) M. leprae strains. The phylogeny was inferred by the Maximum Likelihood method of MEGA7 (Kumar et al, 2016) and the Tamura-3-Parameter model. The tree with the highest log likelihood value is shown. Bootstrap percentages from 1000 replicates are shown next to the branches. The scale indicates the number of substitutions per site. All positions with less than 90% site coverage were eliminated. M. lepromatosis was used as an outgroup (not shown). CM1 and Br15-1 are derived from a cynomolgus macaque and a red squirrel respectively.

Now, remember that we also need to keep chronology in mind: Lineage 0 is thought to have diverged from the common ancestor of Lineages 1-4 at least 3.5K ya. (Here’s the phylogenetic tree from Benjak et al. 2018, which I have marked with time divisions for emphasis.)

phylogeny-mleprae
Modified image by Monica H. Green. Phylogeny of M. leprae. Bayesian phylogenetic tree of 146 genomes of M. leprae calculated with BEAST 2.4.4. Hypermutated samples with mutations in the nth gene were excluded from the analysis. The tree is drawn to scale, with branch lengths representing years of age. Samples were binned according to geographic origin as given in the legend. Posterior probabilities for each node are shown in gray. Location probabilities of nodes were inferred by the Discrete Phylogeny model

So what we need to explain is how a strain (Lineage 0, or 3K0 as Mendum et al. 2018 call it) can be found all the way from Denmark to New Caledonia. An “Out of Europe” narrative isn’t really helpful, any more than the earlier “Out of Africa” narrative worked.

Given the extreme amount of suffering leprosy has caused, and continues to cause around the world, and given the extraordinary investigative power that paleogenetics has now developed, it’s really time that we did a better job pulling these global narratives together.

If you have Twitter, be sure to retweet this thread!

NOTE. Another (probably also interesting) article was published recently, Digging up the plague: A diachronic comparison of aDNA confirmed plague burials and associated burial customs in Germany, by Gutsmiedl-Schümann, Praehistorische Zeitschrift (2018) 92:2, but sadly my university does not have access to it.

Abstract:

Plague outbreaks in the past are mainly known from written sources; in particular, the Justinianic Plague of the Early Middle Ages and the Black Death of the Late Middle Ages have been described in vivid detail. Yet prior to the introduction of aDNA analysis, it was often quite difficult to associate burials with plague beyond doubt – especially in areas where written evidence of the plague is scarce. As analysis of ancient DNA now allows the detection of plague victims in the archaeological record, new ways are being developed for combining archaeological, historical and ancient DNA research. In this paper we would like to present and compare known examples of plague graves from the Early Middle Ages, the Late Middle Ages and the Thirty Years’ War in Germany that have also been confirmed by ancient DNA analyses. We would like to argue for a differentiated view of the burial customs, especially when more than one plague victim shared a grave, and would like to show possible conclusions, drawn from the aDNA-confirmed plague burials, that can indicate the different strategies adopted by ancient societies to deal with catastrophic events like a pandemic disease.

Related:

Ancient DNA study reveals HLA susceptibility locus for leprosy in medieval Europeans

danemark-medieval

Open access Ancient DNA study reveals HLA susceptibility locus for leprosy in medieval Europeans, by Krause-Kyora et al., Nature Communications (2018)

Abstract:

Leprosy, a chronic infectious disease caused by Mycobacterium leprae (M. leprae), was very common in Europe till the 16th century. Here, we perform an ancient DNA study on medieval skeletons from Denmark that show lesions specific for lepromatous leprosy (LL). First, we test the remains for M. leprae DNA to confirm the infection status of the individuals and to assess the bacterial diversity. We assemble 10 complete M. leprae genomes that all differ from each other. Second, we evaluate whether the human leukocyte antigen allele DRB1*15:01, a strong LL susceptibility factor in modern populations, also predisposed medieval Europeans to the disease. The comparison of genotype data from 69 M. leprae DNA-positive LL cases with those from contemporary and medieval controls reveals a statistically significant association in both instances. In addition, we observe that DRB1*15:01 co-occurs with DQB1*06:02 on a haplotype that is a strong risk factor for inflammatory diseases today.

danes-leprosy-positive
Relationship of 53 medieval leprosy-positive Danes to contemporary Europeans. Principal component analysis plot for 53 medieval St. Jørgen individuals in relation to European population samples from the 1000 Genomes project. (CEU, Northern Europeans from Utah; GBR, British in England and Scotland; IBS, Iberian population in Spain; TSI, Tuscans in Italy; FIN, Finnish in Finland)

The study shows mtDNA haplogroups comparable to those of northern Europeans today, and findings in general indicate no major genome-wide changes in the Danish population structure in the past 1000 years.

The paper may be of interest for earlier migrations:

rs3135388-t-allele-frequency-leprosy

Discovered via Iain Mathieson:

Related: