A Game of Thrones in Indo-European: proto-languages in Westeros and Essos, and population genomics

I think proto-languages can be applied to basically any appropriate prehistoric setting, and especially to science fiction and fantasy settings. I often viewed the lack of interest for them as based on the idea that they are not fantastic enough, that they would render a fantastic world too realistic to allow for an adequate immersion of the reader (or viewer) into a new world.

With time, I have become more and more convinced that most authors don’t use proto-languages (or tweaked versions of them) simply because they can’t, and resort to the easier way: inventing some rules and words based on some basic ideas and sounds they feel would fit a certain culture or people, to get going. After all, world-building is about a good enough, not too detailed description, and books are about characters and settings, not worlds.

After the end of the 7th season of the Game of Thrones TV series, of which I have become a great fan, I had some season finale grief to deal with, so I thought about applying what we knew about Proto-Indo-Europeans to the fantasy world. Since all book translations deal with English names as if they were translations of the Common Tongue (e.g. Spanish “Invernalia” or “Poniente” for “Winterfel” or “Westeros”), the idea of a translation into Proto-Indo-European seemed quite interesting.

NOTE. I understand that, for some, the idea that “the original language is the best” would make them reject this. However, just take into account the millions who enjoy the books and the TV series only in their native language, and know nothing about the ‘original’ version…

Here are the text and images:

A Dance with Old Tongues

As you can see, the idea of the Common Tongue being Late Proto-Indo-European brings about a whole new (infinite) world of dialectal evolution, language contacts, and population expansions which must be established for the whole setting to work. This is what the text I began to write was about: to use languages (and related populations) of ca. 6000-1500 BC, and to avoid anachronisms and impossible language relationships:

As an added advantage, fans of role-playing games could expand their world with the use of the language correspondences and the maps. This way, instead of “Northern English” being spoken in the North, and “Spanish English” being spoken in Dorne, according to some selections that have been naturally criticized, you have ancient languages that fit with the ancient setting, and which were actually related to each other.

Equivalence of languages of the known world with coeval proto-languages. Solid red lines divide Graeco-Aryan from Northern Indo-European dialects (Tocharian is separated from North-West Indo-European by a dotted red line). See all maps.

I also began drawing a fantasy map, my first one – even though I have been member of Cartographer’s Guild for years – , which eventually helped me with my updates of maps of prehistoric migrations, and even with the use of arrows and colors for scientific publications. I drew details mainly to illustrate the text, not to offer a comprehensive translated world. Most of the work was done in the Summer of 2017, with some map changes done in 2018 with help of the maps and works of fans.

NOTE. I have reviewed it during some long travels lately, and included names of “bloodlines” (i.e. haplogroups), which I find more interesting today for people to understand bottlenecks during prehistoric migrations; I have also added a map using pie charts. If this doesn’t fit well with the whole picture, it’s because it’s a recent addition. The rest is more or less the same as one-two years ago.

I don’t have time now to correct much of what I wrote. I have forgotten most of the relevant details from the books, especially A World of Ice and Fire which I think helped me a lot with this, and I am sure that after writing A Song of Sheep and Horses (now you know the why of the book names) I would deal with some language identification and cognates differently.

I decided to publish it to liven up our Facebook page of Modern Indo-European now that the 8th season is near, so that people can participate and try to translate (translatable) names and expressions into Proto-Indo-European, to see how it would work out. You can also request access our Modern Indo-European and Proto-Indo-European groups; both are administered mainly by Fernando.

If you think this whole idea is crazy, or a huge loss of time, I agree; this is how you lose your time when you like fantasy, comic books, etc. But I am a great fan of fantasy and fiction, and I had a lot of free time back then, so I couldn’t help it…

On the other hand, if you feel that mixing fantasy (or SF) with the Proto-Indo-European question (especially population genomics) is a bad idea, I may have agreed with that two years ago, and maybe this is the reason why I hesitated to publish it then.

Hoewever, today we can read a whole new (2018 and 2019) bunch of “steppe ancestry=Indo-European” fantasies: invisible Nganasan reindeer hordes, a Fearsome Tisza River where Yamna settlers mysteriously disappear, shapeshifting Dutch CWC peoples who change haplogroups, languages dependent on cephalic types, or Yamna/Bell Beaker expanding Vasconic…So what’s the matter with some more fantasy?

Ancient Sardinia hints at Mesolithic spread of R1b-V88, and Western EEF-related expansion of Vasconic


New preprint Population history from the Neolithic to present on the Mediterranean island of Sardinia: An ancient DNA perspective, by Marcus et al. bioRxiv (2019)

Interesting excerpts (emphasis mine, edited for clarity):

On the high frequency of R1b-V88

Our genome-wide data allowed us to assign Y haplogroups for 25 ancient Sardinian individuals. More than half of them consist of R1b-V88 (n=10) or I2-M223 (n=7).

Francalacci et al. (2013) identi fied three major Sardinia-specifi c founder clades based on present-day variation within the haplogroups I2-M26, G2-L91 and R1b-V88, and here we found each of those broader haplogroups in at least one ancient Sardinian individual. Two major present-day Sardinian haplogroups, R1b-M269 and E-M215, are absent.

Compared to other Neolithic and present-day European populations, the number of identi fied R1b-V88 carriers is relatively high.

(…)ancient Sardinian mtDNA haplotypes belong almost exclusively to macro-haplogroups HV (n = 16), JT (n = 17) and U (n = 9), a composition broadly similar to other European Neolithic populations.

Geographic and temporal distribution of R1b-V88 Y-haplotypes in ancient European samples. We plot the geographic position of all ancient samples inferred to carry R1b-V88 equivalent markers. Dates are given as years BCE (means of calibrated 2s radio-carbon dates). Multiple V88 individuals with similar geographic positions are vertically stacked. We additionally color-code the status of the R1b-V88 subclade R1b-V2197, which is found in most present-day African R1b-V88 carriers.

On the origin of a Vasconic-like Paleosardo with the Western EEF

(…) the Neolithic (and also later) ancient Sardinian individuals sit between early Neolithic Iberian and later Copper Age Iberian populations, roughly on an axis that differentiates WHG and EEF populations and embedded in a cluster that additionally includes Neolithic British individuals. This result is also evident in terms of absolute genetic differentiation, with low pairwise FST ~ 0.005 +- 0.002 between Neolithic Sardinian individuals and Neolithic western mainland European populations. Pairwise outgroup-f3 analysis shows a very similar pattern, with the highest values of f3 (i.e. most shared drift) being with Neolithic and Copper Age Iberia, gradually dropping off for temporally and geographically distant populations.

In explicit admixture models (using qpAdm, see Methods) the southern French Neolithic individuals (France-N) are the most consistent with being a single source for Neolithic Sardinia (p ~ 0:074 to reject the model of one population being the direct source of the other); followed by other populations associated with the western Mediterranean Neolithic Cardial Ware expansion.

Principal Components Analysis based on the Human Origins dataset. A: Projection of ancient individuals’ genotypes onto principal component axes de fined by modern Western Eurasians (gray labels).

Pervasive Western Hunter-Gatherer ancestry in Iberian/French/Sardinian population

Similar to western European Neolithic and central European Late Neolithic populations, ancient Sardinian individuals are shifted towards WHG individuals in the top two PCs relative to early Neolithic Anatolians Admixture analysis using qpAdm infers that ancient Sardinian individuals harbour HG ancestry (~ 17%) that is higher than early Neolithic mainland populations (including Iberia, ~ 8%), but lower than Copper Age Iberians (~ 25%) and about the same as Southern French Middle-Neolithic individuals (~ 21%).

Principal Components Analysis based on the Human Origins dataset. B: Zoom into the region most relevant for Sardinian individuals.

Continuity from Sardinia Neolithic through the Nuragic

We found several lines of evidence supporting genetic continuity from the Sardinian Neolithic into the Bronze Age and Nuragic times. Importantly, we observed low genetic differentiation between ancient Sardinian individuals from various time periods.

A qpAdm analysis, which is based on simultaneously testing f-statistics with a number of outgroups and adjusts for correlations, cannot reject a model of Neolithic Sardinian individuals being a direct predecessor of Nuragic Sardinian individuals (…) Our qpAdm analysis further shows that the WHG ancestry proportion, in a model of admixture with Neolithic Anatolia, remains stable at ~17% throughout three ancient time-periods.

Present-day genetic structure in Sardinia reanalyzed with aDNA. A: Scatter plot of the rst two principal components trained on 1577 present-day individuals with grand-parental ancestry from Sardinia. Each individual is labeled with a location if at least 3 of the 4 grandparents were born in the same geographical location (\small” three letter abbreviations); otherwise with \x” or if grand-parental ancestry is missing with \?”. We calculated median PC values for each Sardinian province (large abbreviations). We also projected each ancient Sardinian individual on to the top two PCs (gray points). B/C: We plot f-statistics that test for admixture of modern Sardinian individuals (grouped into provinces) when using Nuragic Sardinian individuals as one source population. Uncertainty ranges depict one standard error (calculated from block bootstrap). Karitiana are used in the f-statistic calculation as a proxy for ANE/Steppe ancestry (Patterson et al., 2012).

Steppe influx in Modern Sardinians

While contemporary Sardinian individuals show the highest affinity towards EEF-associated populations among all of the modern populations, they also display membership with other clusters (Fig. 5). In contrast to ancient Sardinian individuals, present-day Sardinian individuals carry a modest “Steppe-like” ancestry component (but generally less than continental present-day European populations), and an appreciable broadly “eastern Mediterranean” ancestry component (also inferred at a high fraction in other present-day Mediterranean populations, such as Sicily and Greece).


Happy new year 2019…and enjoy our new books!


Sorry for the last weeks of silence, I have been rather busy lately. I am having more projects going on, and (because of that) I also wanted to finish a project I have been working on for many months already.

I have therefore decided to publish a provisional version of the text, in the hope that it will be useful in the following months, when I won’t be able to update it as often as I would like to:

EDIT (20 JAN 2019): For those of you who are more comfortable reading in your native language, I have placed some links to automatic translations by Google Translate. They might work especially well for the texts of A Game of Clans & A Clash of Chiefs.

Don’t forget to check out the maps included in the supplementary materials: I have added Y-DNA, mtDNA, and ADMIXTURE data using GIS software. The PCA graphics are also important to follow the main text.

NOTE. Right now the files are only in my server. I will try to upload them to Academia.edu and Research Gate when I have time, I have uploaded them to Academia.edu and ResearchGate, in case the websites are too slow.

I would have preferred to wait for a thorough revision of the section on archaeology and the linguistic sections on Uralic, but I doubt I will have time when the reviews come, so it was either now or maybe next December…

I say so in the introduction, but it is evident that certain aspects of the book are tentative to say the least: the farther back we go from Late Proto-Indo-European, the less clear are many aspects. Also, linguistically I am not convinced about Eurasiatic or Nostratic, although they do have a certain interest when we try to offer a comprehensive view of the past, including ethnolinguistic identities.

I cannot be an expert in everything, and these books cover a lot. I am bound to publish many corrections as new information appears and more reviews are sent. For example, just days ago (before SNP calls of Wang et al. 2018 were published) some paragraphs implied that AME might have expanded Nostratic from the Middle East. Now it does not seem so, and I changed them just before uploading the text. That’s how tentative certain routes are, and how much all of this may change. And that only if we accept a Nostratic phylum…

NOTE. Since the first book I wrote was the linguistic one, and I have spent the last months updating the archaeology + genetics part, now many of you will probably understand 1) why I am so convinced about certain language relationships and 2) how I used many posts to clarify certain ideas and receive comments. Many posts offer probably a good timeline of what I worked with, and when.

A Song of Sheep and Horses (ASoSaH) reread

Edit (23 Jan 2019):

To be able to revise and update the text properly, I decided to start a series of posts on different aspects.

This is an updated list of the posts:


I did not add this section to the books, because they are still not ready for print, but I think this is due somewhere now. It is impossible to reference all who have directly or indirectly contributed to this, so this is a list of those I feel have played an important role.

I am indebted to the following people (which does not mean that they share my views, obviously):

First and foremost, to Fernando López-Menchero, for having the patience to review with detail many parts on Indo-European linguistics, knowing that I won’t accept many of his comments anyway. The additional information he offers is invaluable, but I didn’t want to turn this into a huge linguistic encyclopaedia with unending discussions of tiny details of each reconstructed word. I think it is already too big as it is.

I would not have thought about doing this if it were not for the interest of Wekwos (Xavier Delamarre) in publishing a full book about the Indo-European demic diffusion model (in the second half of 2017, I think). It was them who suggested that I extended the content, when all I had done until then was write an essay and draw some maps in my free time between depositing the PhD thesis and defending it.

Sadly, as much as I would like to publish a book with a professional publisher, I don’t think ancient DNA lends itself for the traditional format, so my requests (mainly to have free licenses and being able to review the text at will, as new genetic papers are published) were logically not acceptable. Also, the main aim of all volumes, especially the linguistic one, is the teaching of essentials of Late Proto-Indo-European and related languages, and this objective would be thwarted by selling each volume for $50-70 and only in printed format. I prefer a wider distribution.

At first I didn’t think much of this proposal, because I do not benefit from this kind of publications in my scientific field, but with time my interest in writing a whole, comprehensive book on the subject grew to the point where it was already an ongoing project, probably by the start of 2018.

I would not have been in contact with Wekwos if it were not for user Camulogène Rix at Anthrogenica, so thanks for that and for the interest in this work.

I would not have thought of writing this either if not for the spontaneous support (with an unexpected phone call!) of a professor of the Complutense University of Madrid, Ángel Gómez Moreno, who is interested in this subject – as is his wife, a professor of Classics more closely associated to Indo-European studies, and who helped me with a search for Indo-Europeanists.

EDIT (1 JAN 2019): I remembered that Karin Bojs sent me her book after reading the demic diffusion model. I may have also thought about writing a whole book back then, but mid-2017 is probably too early for the project.

Professor Kortlandt is still to review the text, but he contributed to both previous essays in some very interesting ways, so I hope he can help me improve the parts on Uralic, and maybe alternative accounts of expansion for Balto-Slavic, depending on the time depth that he would consider warranted according to the Temematic hypothesis.

The maps are evidently (for those who are interested in genetics) in part the result of the effort of the late Jean Manco: As you can see from the maps including Y-DNA and mtDNA samples, I have benefitted from her way of organising data and publishing it. Similarly, the work of Iain McDonald in assessing the potential migration routes of R1b and R1a in Europe with the help of detailed maps was behind my idea for the first maps, and consequently behind these, too.

I should thank all people responsible for the release of free datasets to work with, including the Reich and Jena labs, the Veeramah Lab, and also researchers from the Max Planck Institute or the Mainz Palaeogenetics group, who didn’t mind to share with me datasets to work with.

Readers of this blog with interesting comments have also been essential for the improvement of the texts. You can probably see some of your many contributions there. I may not answer many comments, because I am always busy (and sometimes I just don’t have anything interesting to say), but I try to read all of them.

EDIT (1 JAN 2019) I think I should mention at least Chetan, Egg, or Robert George; but then I would leave out old europe, Sgr Ganesh, or Tileman Ehlen; and if I include them I would leave out others…

Users of other sites, like Anthrogenica, whose particular points of view and deep knowledge of some very specific aspects are sometimes very useful. In particular, user Anglesqueville helped me to fix some issues with the merging of datasets to obtain the PCAs and ADMIXTURE, and prepared some individual samples to merge them.

Even without posting anything, Google Analytics keeps sending me messages about increasing user fidelity (returning users), and stats haven’t really changed (which probably means more people are reading old posts), so thank you for that.

I hope you enjoy the books.

Happy new year!

Palaeolithic Caucasus samples reveal the most important component of West Eurasians


Preprint Paleolithic DNA from the Caucasus reveals core of West Eurasian ancestry, by Lazaridis et al. bioRxiv (2018).

Interesting excerpts:

We analyzed teeth from two individuals 63 recovered from Dzudzuana Cave, Southern Caucasus, from an archaeological layer previously dated to ~27-24kya (…). Both individuals had mitochondrial DNA sequences (U6 and N) that are consistent with deriving from lineages that are rare in the Caucasus or Europe today. The two individuals were genetically similar to each other, consistent with belonging to the same population and we thus analyze them jointly.

(…) our results prove that the European affinity of Neolithic Anatolians does not necessarily reflect any admixture into the Near East from Europe, as an Anatolian Neolithic-like population already existed in parts of the Near East by ~26kya. Furthermore, Dzudzuana shares more alleles with Villabruna-cluster groups than with other ESHG (Extended Data Fig. 5b), suggesting that this European affinity was specifically related to the Villabruna cluster, and indicating that the Villabruna affinity of PGNE populations from Anatolia and the Levant is not the result of a migration into the Near East from Europe. Rather, ancestry deeply related to the Villabruna cluster was present not only in Gravettian and Magdalenian-era Europeans but also in the populations of the Caucasus, by ~26kya. Neolithic Anatolians, while forming a clade with Dzudzuana with respect to ESHG, share more alleles with all other PGNE (Extended Data Fig. 5d), suggesting that PGNE share at least partially common descent to the exclusion of the much older samples from Dzudzuana.

Ancient West Eurasian population structure. PCA of key ancient West Eurasians, including additional populations (shown with grey shells), in the space of outgroup f4-statistics (Methods).

Our co-modeling of Epipaleolithic Natufians and Ibero-Maurusians from Taforalt confirms that the Taforalt population was mixed, but instead of specifying gene flow from the ancestors of Natufians into the ancestors of Taforalt as originally reported, we infer gene flow in the reverse direction (into Natufians). The Neolithic population from Morocco, closely related to Taforalt is also consistent with being descended from the source of this gene flow, and appears to have no admixture from the Levantine Neolithic (Supplementary Information 166 section 3). If our model is correct, Epipaleolithic Natufians trace part of their ancestry to North Africa, consistent with morphological and archaeological studies that indicate a spread of morphological features and artifacts from North Africa into the Near East. Such a scenario would also explain the presence of Y-chromosome haplogroup E in the Natufians and Levantine farmers, a common link between the Levant and Africa.

(…) we cannot reject the hypothesis that Dzudzuana and the much later Neolithic Anatolians form a clade with respect to ESHG (P=0.286), consistent with the latter being a population largely descended from Dzudzuana-like pre-Neolithic populations whose geographical extent spanned both Anatolia and the Caucasus. Dzudzuana itself can be modeled as a 2-way mixture of Villabruna-related ancestry and a Basal Eurasian lineage.

In qpAdm modeling, a deeply divergent hunter-gatherer lineage that contributed in relatively unmixed form to the much later hunter-gatherers of the Villabruna cluster is specified as contributing to earlier hunter-gatherer groups (Gravettian Vestonice16: 35.7±11.3% and Magdalenian ElMiron: 60.6±11.3%) and to populations of the Caucasus (Dzudzuana: 199 72.5±3.7%, virtually identical to that inferred using ADMIXTUREGRAPH). In Europe, descendants of this lineage admixed with pre-existing hunter-gatherers related to Sunghir3 from Russia for the Gravettians and GoyetQ116-1 from Belgium for the Magdalenians, while in the Near East it did so with Basal Eurasians. Later Europeans prior to the arrival of agriculture were the product of re-settlement of this lineage after ~15kya in mainland Europe, while in eastern Europe they admixed with Siberian hunter-gatherers forming the WHG-ANE cline of ancestry [See PCA above]. In the Near East, the Dzudzuana-related population admixed with North African-related ancestry in the Levant and with Siberian hunter-gatherer and eastern non-African-related ancestry in Iran and the Caucasus. Thus, the highly differentiated populations at the dawn of the Neolithic were primarily descended from Villabruna Cluster and Dzudzuana-related ancestors, with varying degrees of additional input related to both North Africa and Ancient North/East Eurasia whose proximate sources may be clarified by future sampling of geographically and temporally intermediate populations.

An admixture graph model of Paleolithic West Eurasians. An automatically generated admixture graph models fits populations (worst Z-score of the difference between estimated and fitted f-statistics is 2.7) or populations (also including South_Africa_HG, worst Z-score is 3.5). This is a simplified model assuming binary admixture events and is not a unique solution (Supplementary Information section 2). Sampled populations are shown with ovals and select labeled internal nodes with rectangles.

Interesting excerpts from the supplementary materials:

From our analysis of Supplementary Information section 3, we showed that these sources are indeed complex, and only one of these (WHG, represented by Villabruna) appears to be a contributor to all the remaining sources. This should not be understood as showing that hunter-gatherers from mainland Europe migrated to the rest of West Eurasia, but rather that the fairly homogeneous post-15kya population of mainland Europe labeled WHG appear to represent a deep strain of ancestry that seems to have contributed to West Eurasians from the Gravettian era down to the Neolithic period.

Villabruna is representative of the WHG group. We also include ElMiron, the best sample from the Magdalenian era as we noticed that within the WHG group there were individuals that could not be modeled as a simple clade with Villabruna but also had some ElMiron-related ancestry. Ddudzuana is representative of the Ice Age Caucasus population, differentiated from Villabruna by Basal Eurasian ancestry. AG3 represents ANE/Upper Paleolithic Siberian ancestry, sampled from the vicinity of Lake Baikal, while Russia_Baikal_EN related to eastern Eurasians and represents a later layer of ancestry from the same region of Siberia as AG3 Finally, Mbuti are a deeply diverged African population that is used here to represent deep strains of ancestry (including Basal Eurasian) prior to the differentiation between West Eurasians and eastern non-Africans that are otherwise not accounted for by the remaining five sources. Collectively, we refer to this as ‘Basal’ or ‘Deep’ ancestry, which should be understood as referring potentially to both Basal Eurasian and African ancestry.

It has been suggested that there is an Anatolia Neolithic-related affinity in hunter-gatherers from the Iron Gates. Our analysis confirms this by showing that this population has Dzudzuana-related ancestry as do many hunter-gatherer populations from southeastern Europe, eastern Europe and Scandinavia. These populations cannot be modeled as a simple mixture of Villabruna and AG3 but require extra Dzudzuana-related ancestry even in the conservative estimates, with a positive admixture proportion inferred for several more in the speculative ones. Thus, the distinction between European hunter-gatherers and Near Eastern populations may have been gradual in pre-Neolithic times; samples from the Aegean (intermediate between those from the Balkans and Anatolia) may reveal how gradual the transition between Dzudzuana-like Neolithic Anatolians and mostly Villabruna-like hunter-gatherers was in southeastern Europe.

Modified image (cut, with important samples marked). Modeling present-day and ancient West-Eurasians. Mixture proportions computed with qpAdm (Supplementary Information section 4). The proportion of ‘Mbuti’ ancestry represents the total of ‘Deep’ ancestry from lineages that split prior to the 365 split of Ust’Ishim, Tianyuan, and West Eurasians and can include both ‘Basal Eurasian’ and other (e.g., Sub-Saharan African) ancestry. (a) ‘Conservative’ estimates. Each population 367 cannot be modeled with fewer admixture events than shown.

Villabruna: This type of ancestry differentiates between present-day Europeans and non-Europeans within West Eurasia, attaining a maximum of ~20% in the Baltic in accordance with previous observations and with the finding of a later persistence of significant hunter-gatherer ancestry in the region. Its proportion drops to ~0% throughout the Near East. Interestingly, a hint of such ancestry is also inferred in all North African populations west of Libya in the speculative proportions, consistent with an archaeogenetic inference of gene flow from Iberia to North Africa during the Late Neolithic.

ElMiron: This type of ancestry is absent in present-day West Eurasians. This may be because most of the Villabruna-related ancestry in Europeans traces to WHG populations that lacked it (since ElMiron-related ancestry is quite variable within European hunter-gatherers). However, ElMiron ancestry makes up only a minority component of all WHG populations sampled to date and WHG-related ancestry is a minority component of present-day Europeans. Thus, our failure to detect it in present day people may be simply be too little of it to detect with our methods.

Dzudzuana: Our analysis identifies Dzudzuana-related ancestry as the most important component of West Eurasians and the one that is found across West Eurasian-North African populations at ~46-88% levels. Thus, Dzudzuana-related ancestry can be viewed as the common core of the ancestry of West Eurasian-North African populations. Its distribution reaches its minima in northern Europe and appears to be complementary to that of Villabruna, being most strongly represented in North Africa, the Near East (including the Caucasus) and Mediterranean Europe. Our results here are expected from those of Supplementary Information section 3 in which we modeled ancient Near Eastern/North African populations (the principal ancestors of present-day people from the same regions) as deriving much of their ancestry from a Dzudzuana-related source. Migrations from the Near East/Caucasus associated with the spread of the Neolithic, but also the formation of steppe population introduced most of the Dzudzuana-related ancestry present in Europe, although (as we have seen above) some such ancestry was already present in some pre-agricultural hunter-gatherers in Europe.

AG3: Ancestry related to the AG3 sample from Siberia has a northern distribution, being strongly represented in both central-northern Europe and the north Caucasus.

Russia_Baikal_EN: Ancestry related to hunter-gatherers from Lake Baikal in Siberia (postdating AG3) appears to have affected primarily northeastern European populations which have been previously identified as having East Eurasian ancestry; some such ancestry is also identified for a Turkish population from Balıkesir, likely reflecting the Central Asian ancestry of Turkic speakers which has been recently confirmed directly in an Ottoman sample from Anatolia.

Some comments

So, to try and sum up:

  • Dzudzuana shares ancestry with ‘Common West Eurasian’ (CWE). the ancestor cluster of Villabruna.
  • Dzudzuana diverges from CWE because of a Basal Eurasian ancestry contribution [which supports that Basal Eurasian ancestry was a deep Middle Eastern lineage].
  • Dzudzuana is closest to Anatolia Neolithic, and close to Gravettian.
Palaeolithic migrations and clusters in Europe. See more maps.


  1. Aurignacian: First West Eurasians arrive ca. 36,000 BP, Goyet cluster expands probably with C1a2 lineages.
  2. After that, the early or ‘unmixed’ Villabruna cluster (‘hidden’ somewhere probably east of Europe, either North Eurasia or South Eurasia), lineages unknown (possibly IJ), contributes to:
    1. Gravettian (ca. 30,000 BP): Věstonice cluster expands, probably with IJ lineages.
    2. A (hidden) ‘Common West Eurasian’ population.
    3. In turn:

      • Dzudzuana ca. 26,000 BP derived from Common West Eurasian (curiously, haplogroup G seems to split in today’s subclades ca. 26,000 BP).
      • During the Gravettian (ca. 26,000 BP), an Anatolian Neolithic-like population exists already in the Near East. Both Věstonice and this Anatolian HG are close to Dzudzuana; in turn, Dzudzuana from CWE.

    4. Magdalenian (ca. 20,000 BP): El Mirón cluster expands, probably with more specific I lineages.
  3. Bølling-Allerød warming period (ca. 14,000 BP): ‘late’ Villabruna cluster or WHG (=CWE with greater affinity to Near Eastern populations) expands, probably spreading with R1b in mainland Europe and to the east (admixing with Siberian HG), creating the WHG — ANE ancestry cline, as reflected in Iron Gates HG, Baltic HG, etc.

[Here we have the possible “bidirectional gene flow between populations ancestral to Southeastern Europeans of the early Holocene and Anatolians of the late glacial or a dispersal of Southeastern Europeans into the Near East” inferred from Anatolian hunter-gatherers]

The Gravettian (30,000 to 20,000 years) is drawn in black and white; the subsequent Magdalenian (17,000 to 10,000 years) and Hamburgian (13,000-11,750 years) are in light blue and red. It is not known whether the spread of the Gravettian was a result of diffusion of people or cultures. This figure illustrates the possible monocentric origins of the Gravettian, in which the Gravettian is hypothesized to have its origin in the Middle Danube Basin, first spreading west (solid lines) and later spreading east and southeast (dashed lines). This scenario is largely based on the chronology of sites. Thus far, genome-wide data has been collected from only three of the ten< Gravettian regions indicated on the map. These regions are northern Austria (1 sample), the Czech Republic (6), southern Italy (3) and Belgium (3), indicating that they all share a genomic ancestry. However, it is unknown whether samples from the remaining regions also share a close genomic ancestry. Some skeletal remains associated with the Gravettian that could be investigated paleogenomically are from Sungir (Russia); Laghar Velho (central Portugal); Cussac Cave; Les Garennes, near Vilhonneur; and Level 2 at Abri Pataud116 (western France). Light blue and light red regions represent the approximate distributions of the Magdalenian Culture and the Hamburgian Culture (13,000-11,750 years). Figure adapted from Kozłowski. Image from Harris (2017)

The paper talks about possibilities for Common West Eurasian:

  1. Migration from mainland Europe to Near East or vice versa (not very likely);
  2. Migration from a geographically intermediate Ice Age refugium in southeast Europe, Anatolia, or the circum-Pontic region that explain post-glacial affinity of post-glacial Levantine and Anatolian populations.

It also re-states what was known:

  • EHG (ca. 8,000 BP) = between WHG — ANE (ca. 24,000 BP).
  • CHG (ca. 10,000 BP) = between EHG — Iran N.

I would say that the distinct CHG vs. Dzudzuana ancestry puts CHG probably to the south, within the Iranian Plateau, during the Gravettian, expanding probably later.

Also important, Ancestral North African probably accompanied by haplogroup E. Early expansion of North Africans into the Near East further confirms the impossibility of Afroasiatic (much younger) to be associated with these expansions, and confirms that the still unclear Green Sahara migrations are the key.


Scythians in Ukraine, Natufian and sub-Saharan ancestry in North Africa (ISBA 8, 21st Sep)


Interesting information from ISBA 8 sesions today, as seen on Twitter (see programme in PDF, and sessions from the 19th and the 20th september).

Official abstracts are listed first (emphasis mine), then reports and images and/or link to tweets. Here is the list for quick access:

Scythian population genetics and settlement patterns

Genetic continuity in the western Eurasian Steppe broken not due to Scythian dominance, but rather at the transition to the Chernyakhov culture (Ostrogoths), by Järve et al.

The long-held archaeological view sees the Early Iron Age nomadic Scythians expanding west from their Altai region homeland across the Eurasian Steppe until they reached the Ponto-Caspian region north of the Black and Caspian Seas by around 2,900 BP1. However, the migration theory has not found support from ancient DNA evidence, and it is still unclear how much of the Scythian dominance in the Eurasian Steppe was due to movements of people and how much reflected cultural diffusion and elite dominance. We present new whole-genome results of 31 ancient Western and Eastern Scythians as well as samples pre- and postdating them that allow us to set the Scythians in a temporal context by comparing the Western Scythians to samples before and after within the Ponto-Caspian region. We detect no significant contribution of the Scythians to the Early Iron Age Ponto-Caspian gene pool, inferring instead a genetic continuity in the western Eurasian Steppe that persisted from at least 4,800–4,400 cal BP to 2,700–2,100 cal BP (based on our radiocarbon dated samples), i.e. from the Yamnaya through the Scythian period.

However, the transition from the Scythian to the Chernyakhov culture between 2,100 and 1,700 cal BP does mark a shift in the Ponto-Caspian genetic landscape, with various analyses showing that Chernyakhov culture samples share more drift and derived alleles with Bronze/Iron Age and modern Europeans, while the Scythians position outside modern European variation. Our results agree well with the Ostrogothic origins of the Chernyakhov culture and support the hypothesis that the Scythian dominance was cultural rather than achieved through population replacement.

Detail of the slide with admixture of Scythian groups in Ukraine:


Interesting to read in combination with yesterday’s re-evaluation of Scythian mobility and settlement patterns in the west (showing adaptation to the different regional cultures), The Steppe was Sown – multi-isotopic research changes our understandings of Scythian diet and mobility, by Ventresca Miller et al.

Nomadic pastoralists conventionally known as the Scythians occupied the Pontic steppe during the Iron Age, c. 700-200 BC, a period of unprecedented pan-regional interaction. Popular science accounts of the Scythians promote narratives of roving bands of nomadic warriors traversing the steppe from the Altai Mountains to the Black Sea coastline. The quantity and scale of mobility in the region is usually emphasized based on the wide distribution of material culture and the characterization of Iron Age subsistence economies in the Pontic steppe and forest-steppe as mobile pastoralism. Yet, there remains a lack of systematic, direct analysis of the mobility of individuals and their animals. Here, we present a multi-isotopic analysis of humans from Iron Age Scythian sites in Ukraine. Mobility and dietary intake were documented through strontium, carbon and oxygen isotope analyses of tooth enamel. Our results provide direct evidence for mobility among populations in the steppe and forest-steppe zones, demonstrating a range of localized mobility strategies. However, we found that very few individuals came from outside of the broader vicinity of each site, often staying within a 90 km radius. Dietary intake varied at the intrasite level and was based in agro-pastoralism.

While terrestrial protein did form a portion of the diet for some individuals, there were also high levels of a 13C-enriched food source among many individuals, which has been interpreted as millet consumption. Individuals exhibiting 87Sr/86Sr ratios that fell outside the local range were more likely to have lower rates of millet consumption than those that fell within the local range. This suggests that individuals moving to the site later in life had different economic pursuits and consumed less millet. There is also strong evidence that children and infants moved at the pan-regional scale. Contrary to the popular narrative, the majority of Scythians engaged in localized mobility as part of agricultural lifeways while pan-regional movements included family groups.

North-Africans show ancestry from the ancient Near East and sub-Saharan Africa

Pleistocene North Africans show dual genetic ancestry from the ancient Near East and sub-Saharan Africa, by van de Loosdrecht et al.

North Africa, connecting sub-Saharan Africa and Eurasia, is important for understanding human history. However, the genetic history of modern humans in this region is largely unknown before the introduction of agriculture. After the Last Glacial Maximum modern humans, associated with the Iberomaurusian culture, inhabited a wide area spanning from Morocco to Libya. The Iberomaurusian is part of the early Later Stone Age and characterized by a distinct microlithic bladelet technology, complex hunter-gathering and tooth evulsion.

Here we present genomic data from seven individuals, directly dated to ~15,000-year-ago, from Grotte des Pigeons, Taforalt in Morocco. Uni-parental marker analyses show mitochondrial haplogroup U6a for six individuals and M1b for one individual, and Y-chromosome haplogroup E-M78 (E1b1b1a1) for males. We find a strong genetic affinity of the Taforalt individuals with ancient Near Easterners, best represented by ~12,000 year old Levantine Natufians, that made the transition from complex hunter-gathering to more sedentary food production. This suggests that genetic connections between Africa and the Near East predate the introduction of agriculture in North Africa by several millennia. Notably, we do not find evidence for gene flow from Paleolithic Europeans into the ~15,000 year old North Africans as previously suggested based on archaeological similarities. Finally, the Taforalt individuals derive one third of their ancestry from sub-Saharan Africans, best approximated by a mixture of genetic components preserved in present-day West Africans (Yoruba, Mende) and Africans from Tanzania (Hadza). In contrast, modern North Africans have a much smaller sub-Saharan African component with no apparent link to Hadza. Our results provide the earliest direct evidence for genetic interactions between modern humans across Africa and Eurasia.

A detail of the cultures involved in these population movements:


So, most likely, Natufian-related ancestry – as sub-Saharan ancestry – not related to the Afroasiatic expansion.

NOTE. This now probably outdated already by the new preprint on Dzudzuana samples, from the Caucasus.

Impact of colonization in north-eastern Siberia

Exploring the genomic impact of colonization in north-eastern Siberia by Seguin-Orlando et al.

Yakutia is the coldest region in the northern hemisphere, with winter record temperatures below minus 70°C. The ability of Yakut people to adapt both culturally and biologically to extremely cold temperatures has been key to their subsistence. They are believed to descend from an ancestral population, which left its original homeland in the Lake Baykal area following the Mongol expansion between the 13th and 15th centuries AD. They originally developed a semi-nomadic lifestyle, based on horse and cattle breeding, providing transportation, primary clothing material, meat, and milk. The early colonization by Russians in the first half of the 17th century AD, and their further expansion, have massively impacted indigenous populations. It led not only to massive epidemiological outbreaks, but also to an important dietary shift increasingly relying on carbohydrate-rich resources, and a profound lifestyle transition with the gradual conversion from Shamanism to Christianity and the establishment of new marriage customs. Leveraging an exceptional archaeological collection of more than a hundred of bodies excavated by MAFSO (Mission Archéologique Française en Sibérie Orientale) over the last 15 years and naturally kept frozen by the extreme cold temperatures of Yakutia, we have started to characterize the (epi)genome of indigenous individuals who lived from the 16th to the 20th century AD. Current data include the genome sequence of approximately 50 individuals that lived prior to and after Russian contact, at a coverage from 2 to 40 fold. Combined with data from archaeology and physical anthropology, as well as microbial DNA preserved in the specimens, our unique dataset is aimed at assessing the biological consequences of the social and biological changes undergone by the Yakut people following their neolithisation by Russian colons.

Also interesting to read Balanovsky’s session, and a previous paper on the expansion of Yakuts.

Migrations in the Levant region during the Chalcolithic, also marked by distinct Y-DNA


Open access Ancient DNA from Chalcolithic Israel reveals the role of population mixture in cultural transformation, by Harney et al. Nature Communications (2018).

Interesting excerpts (emphasis mine, reference numbers deleted for clarity):


The material culture of the Late Chalcolithic period in the southern Levant contrasts qualitatively with that of earlier and later periods in the same region. The Late Chalcolithic in the Levant is characterized by increases in the density of settlements, introduction of sanctuaries, utilization of ossuaries in secondary burials, and expansion of public ritual practices as well as an efflorescence of symbolic motifs sculpted and painted on artifacts made of pottery, basalt, copper, and ivory. The period’s impressive metal artifacts, which reflect the first known use of the “lost wax” technique for casting of copper, attest to the extraordinary technical skill of the people of this period.

The distinctive cultural characteristics of the Late Chalcolithic period in the Levant (often related to the Ghassulian culture, although this term is not in practice applied to the Galilee region where this study is based) have few stylistic links to the earlier or later material cultures of the region, which has led to extensive debate about the origins of the people who made this material culture. One hypothesis is that the Chalcolithic culture in the region was spread in part by immigrants from the north (i.e., northern Mesopotamia), based on similarities in artistic designs. Others have suggested that the local populations of the Levant were entirely responsible for developing this culture, and that any similarities to material cultures to the north are due to borrowing of ideas and not to movements of people.

Previous genome-wide ancient DNA studies from the Near East have revealed that at the time when agriculture developed, populations from Anatolia, Iran, and the Levant were approximately as genetically differentiated from each other as present-day Europeans and East Asians are today. By the Bronze Age, however, expansion of different Near Eastern agriculturalist populations — Anatolian, Iranian, and Levantine — in all directions and admixture with each other substantially homogenized populations across the region, thereby contributing to the relatively low genetic differentiation that prevails today. Showed that the Levant Bronze Age population from the site of ‘Ain Ghazal, Jordan (2490–2300 BCE) could be fit statistically as a mixture of around 56% ancestry from a group related to Levantine Pre-Pottery Neolithic agriculturalists (represented by ancient DNA from Motza, Israel and ‘Ain Ghazal, Jordan; 8300–6700 BCE) and 44% related to populations of the Iranian Chalcolithic (Seh Gabi, Iran; 4680–3662 calBCE). Suggested that the Canaanite Levant Bronze Age population from the site of Sidon, Lebanon (~1700 BCE) could be modeled as a mixture of the same two groups albeit in different proportions (48% Levant Neolithic-related and 52% Iran Chalcolithic-related). However, the Neolithic and Bronze Age sites analyzed so far in the Levant are separated in time by more than three thousand years, making the study of samples that fill in this gap, such as those from Peqi’in, of critical importance.

This procedure produced genome-wide data from 22 ancient individuals from Peqi’in Cave (4500–3900 calBCE) (…)


We find that the individuals buried in Peqi’in Cave represent a relatively genetically homogenous population. This homogeneity is evident not only in the genome-wide analyses but also in the fact that most of the male individuals (nine out of ten) belong to the Y-chromosome haplogroup T, a lineage thought to have diversified in the Near East. This finding contrasts with both earlier (Neolithic and Epipaleolithic) Levantine populations, which were dominated by haplogroup E, and later Bronze Age individuals, all of whom belonged to haplogroup J.

Detailed sample background data for each of the 22 samples from which we successfully obtained ancient DNA. Additionally, background information for all samples from Peqi’in that were screened is included in Supplementary Data 1. *Indicates that Y-chromosome haplogroup call should be interpreted with caution, due to low coverage data.

Our finding that the Levant_ChL population can be well-modeled as a three-way admixture between Levant_N (57%), Anatolia_N (26%), and Iran_ChL (17%), while the Levant_BA_South can be modeled as a mixture of Levant_N (58%) and Iran_ChL (42%), but has little if any additional Anatolia_N-related ancestry, can only be explained by multiple episodes of population movement. The presence of Iran_ChL-related ancestry in both populations – but not in the earlier Levant_N – suggests a history of spread into the Levant of peoples related to Iranian agriculturalists, which must have occurred at least by the time of the Chalcolithic. The Anatolian_N component present in the Levant_ChL but not in the Levant_BA_South sample suggests that there was also a separate spread of Anatolian-related people into the region. The Levant_BA_South population may thus represent a remnant of a population that formed after an initial spread of Iran_ChL-related ancestry into the Levant that was not affected by the spread of an Anatolia_N-related population, or perhaps a reintroduction of a population without Anatolia_N-related ancestry to the region. We additionally find that the Levant_ChL population does not serve as a likely source of the Levantine-related ancestry in present-day East African populations.

These genetic results have striking correlates to material culture changes in the archaeological record. The archaeological finds at Peqi’in Cave share distinctive characteristics with other Chalcolithic sites, both to the north and south, including secondary burial in ossuaries with iconographic and geometric designs. It has been suggested that some Late Chalcolithic burial customs, artifacts and motifs may have had their origin in earlier Neolithic traditions in Anatolia and northern Mesopotamia. Some of the artistic expressions have been related to finds and ideas and to later religious concepts such as the gods Inanna and Dumuzi from these more northern regions. The knowledge and resources required to produce metallurgical artifacts in the Levant have also been hypothesized to come from the north.

Our finding of genetic discontinuity between the Chalcolithic and Early Bronze Age periods also resonates with aspects of the archeological record marked by dramatic changes in settlement patterns, large-scale abandonment of sites, many fewer items with symbolic meaning, and shifts in burial practices, including the disappearance of secondary burial in ossuaries. This supports the view that profound cultural upheaval, leading to the extinction of populations, was associated with the collapse of the Chalcolithic culture in this region.

Genetic structure of analyzed individuals. a Principal component analysis of 984 present-day West Eurasians (shown in gray) with 306 ancient samples projected onto the first two principal component axes and labeled by culture. b ADMIXTURE analysis of 984 and 306 ancient samples with K = 11
ancestral components. Only ancient samples are shown


I think the most interesting aspect of this paper is – as usual – the expansion of peoples associated with a single Y-DNA haplogroup. Given that the expansion of Semitic languages in the Middle East – like that of Anatolian languages from the north – must have happened after ca. 3100 BC, coinciding with the collapse of the Uruk period, these Chalcolithic north Levant peoples are probably not related to the posterior Semitic expansion in the region. This can be said to be supported by their lack of relationship with posterior Levantine migrations into Africa. The replacement of haplogroup E before the arrival of haplogroup J suggests still more clearly that Natufians and their main haplogroup were not related to the Afroasiatic expansions.

Distribution of Semitic languages. From Wikipedia.

On the other hand, while their ancestry points to neighbouring regional origins, their haplogroup T1a1a (probably T1a1a1b2) may be closely related to that of other Semitic peoples to the south, as found in east Africa and Arabia. This may be due either to a northern migration of these Chalcolithic Levantine peoples from southern regions in the 5th millennium BC, or maybe to a posterior migration of Semitic peoples from the Levant to the south, coupled with the expansion of this haplogroup, but associated with a distinct population. As we know, ancestry can change within certain generations of intense admixture, while Y-DNA haplogroups are not commonly admixed in prehistoric population expansions.

Without more data from ancient DNA, it is difficult to say. Haplogroup T1a1 is found in Morocco (ca. 3780-3650 calBC), which could point to a recent expansion of a Berbero-Semitic branch; but also in a sample from Balkans Neolithic ca. 5800-5400 calBCE, which could suggest an Anatolian origin of the specific subclades encountered here. In any case, a potential origin of Proto-Semitic anywhere near this wide Near Eastern region ca. 4500-3500 BC cannot be discarded, knowing that their ancestors came probably from Africa.

Distribution of haplogroup T of Y-chromosome. From Wikipedia.

Interesting from this paper is also that we are yet to find a single prehistoric population expansion not associated with a reduction of variability and expansion of Y-DNA haplogroups. It seems that the supposedly mixed Yamna community remains the only (hypothetical) example in history where expanding patrilineal clans will not share Y-DNA haplogroup…


Sahara’s rather pale-green and discontinuous Sahelo-Sudanian steppe corridor, and the R1b – Afroasiatic connection


Interesting new paper (behind paywall) Megalakes in the Sahara? A Review, by Quade et al. (2018).

Abstract (emphasis mine):

The Sahara was wetter and greener during multiple interglacial periods of the Quaternary, when some have suggested it featured very large (mega) lakes, ranging in surface area from 30,000 to 350,000 km2. In this paper, we review the physical and biological evidence for these large lakes, especially during the African Humid Period (AHP) 11–5 ka. Megalake systems from around the world provide a checklist of diagnostic features, such as multiple well-defined shoreline benches, wave-rounded beach gravels where coarse material is present, landscape smoothing by lacustrine sediment, large-scale deltaic deposits, and in places, tufas encrusting shorelines. Our survey reveals no clear evidence of these features in the Sahara, except in the Chad basin. Hydrologic modeling of the proposed megalakes requires mean annual rainfall ≥1.2 m/yr and a northward displacement of tropical rainfall belts by ≥1000 km. Such a profound displacement is not supported by other paleo-climate proxies and comprehensive climate models, challenging the existence of megalakes in the Sahara. Rather than megalakes, isolated wetlands and small lakes are more consistent with the Sahelo-Sudanian paleoenvironment that prevailed in the Sahara during the AHP. A pale-green and discontinuously wet Sahara is the likelier context for human migrations out of Africa during the late Quaternary.

The whole review is an interesting read, but here are some relevant excerpts:

Various researchers have suggested that megalakes coevally covered portions of the Sahara during the AHP and previous periods, such as paleolakes Chad, Darfur, Fezzan, Ahnet-Mouydir, and Chotts (Fig. 2, Table 2). These proposed paleolakes range in size by an order of magnitude in surface area from the Caspian Sea–scale paleo-Lake Chad at 350,000 km2 to Lake Chotts at 30,000 km2. At their maximum, megalakes would have covered ~ 10% of the central and western Sahara, similar to the coverage by megalakes Victoria, Malawi, and Tanganyika in the equatorial tropics of the African Rift today. This observation alone should raise questions of the existence of megalakes in the Sahara, and especially if they developed coevally. Megalakes, because of their significant depth and area, generate large waves that become powerful modifiers of the land surface and leave conspicuous and extensive traces in the geologic record.

ETOPO1 digital elevation model (1 arc-minute; Amante and Eakins, 2009) of proposed megalakes in the Sahara Desert during the late Quaternary. Colors denote Köppen-Geiger climate zones: blue, Aw, Af, Am (tropical); light tan, Bwk, BSh, BSk, Csa, Csb, Cwb, Cfa, Cfb (temperate); red-brown, Bwh (arid, hot desert and steppe climate). Lake area at proposed megalake high stands and present Lake Victoria are in blue, and contributing catchment areas are shown as thin solid black lines. The main tributaries of Lake Chad are denoted by blue lines (from west to east: the Komadougou-Yobe, Logone, and Chari Rivers; source: Global Runoff Data Center, Koblenz, Germany). Rainfall isohyets (50, 200, 800, 1200, and 1600) are marked in dashed gray-scale lines. Physical parameters of each basin are shown in white boxes: Abt, total basin area; AW, lake area; Vw, lake volume; and aW= AW/Abt. Black dots mark the location of the paleohydrological records from Lezine et al. (2011), also compiled in Supplementary Table S5.

Lakes, megalakes, and wetlands

Active ground-water discharge systems abound in the Sahara today, although they were much more widespread in the AHP. They range from isolated springs and wet ground in many oases scattered across the Sahara (e.g., Haynes et al., 1989) to wetlands and small lakes (Kröpelin et al., 2008). Ground water feeding these systems is dominated by fossil AHP-age and older water (e.g., Edmunds and Wright 1979; Sonntag et al., 1980), although recently recharged water (<50 yr) has been locally identified in Saharan ground water (e.g., Sultan et al., 2000; Maduapuchi et al., 2006).

Megalake Chad

In our view, Lake Chad is the only former megalake in the Sahara firmly documented by sedimentologic and geomorphic evidence. Mega-Lake Chad is thought to have covered ~ 345,000 km2, stretching for nearly 8° (10–18°N) of latitude (Ghienne et al., 2002) (Fig. 2). The presence of paleo- Lake Chad was at one point challenged, but several—and in our view very robust—lines of evidence have been presented to support its development during the AHP. These include: (1) clear paleo-shorelines at various elevations, visible on the ground (Abafoni et al., 2014) and in radar and satellite images (Schuster et al., 2005; Drake and Bristow, 2006; Bouchette et al., 2010); (2) sand spits and shoreline berms (Thiemeyer, 2000; Abafoni et al., 2014); and (3) evaporites and aquatic fauna such as fresh-water mollusks and diatoms in basin deposits (e.g., Servant, 1973; Servant and Servant, 1983). Age determinations for all but the Holocene history of mega- Lake Chad are sparse, but there is evidence for Mio-Pliocene lake (s) (Lebatard et al., 2010) and major expansion of paleo- Lake Chad during the AHP (LeBlanc et al., 2006; Schuster et al., 2005; Abafoni et al., 2014; summarized in Armitage et al., 2015) up to the basin overflow level at ~ 329m asl.

Insights from hydrologic mass balance of megalakes

Graph of mean annual rainfall (mm/yr) versus aw (area lake/area basin, AW/AL); their modeled relationship using our Sahelo-Sudanian hydrologic model for the different lake basins are shown as solid colored lines. Superimposed on this (dashed lines) are the aw values for individual megalake basins and the mean annual rainfall required to sustain them. Mean annual paleo-rainfall estimates of 200– 400 mm/yr during the AHP from fossil pollen and mollusk evidence is shown as a tan box. The intersection of this box with the solid colored lines describes the resulting aw for Saharan paleolakes on the y-axis. The low predicted values for aw suggest that very large lakes would not form under Sahelo-Sudanian conditions where sustained by purely local rainfall and runoff. (For interpretation of the references to color in this figure legend, the reader is referred to the web version of this article.)

Using these conservative conditions (i.e., erring in the direction that will support megalake formation), our hydrologic models for the two biggest central Saharan megalakes (Darfur and Fezzan) require minimum annual average rainfall amounts of ~ 1.1 m/yr to balance moisture losses from their respective basins (Supplementary Table S1). Lake Chad required a similar amount (~1 m/yr; Supplementary Table S1) during the AHP according to our calculations, but this is plausible, because even today the southern third of the Chad basin receives ≥1.2 m/yr (Fig. 2) and experiences a climate similar to Lake Victoria. A modest 5° shift in the rainfall belt would bring this moist zone northward to cover a much larger portion of the Chad basin, which spans N13° ±7°. Estimated rainfall rates for Darfur and Fezzan are slightly less than the average of ~ 1.3 m/yr for the Lake Victoria basin, because of the lower aw values, that is, smaller areas of Saharan megalakes compared with their respective drainage basins (Fig. 15).

Estimates of paleo-rainfall during the AHP

Here major contradictions develop between the model outcomes and paleo-vegetation evidence, because our Sahelo-Sudanian hydrologic model predicts wetter conditions and therefore more tropical vegetation assemblages than found around Lake Victoria today. In fact, none of the very wet rainfall scenarios required by all our model runs can be reconciled with the relatively dry conditions implied by the fossil plant and animal evidence. In short, megalakes cannot be produced in Sahelo-Sudanian conditions past or present; to form, they require a tropical or subtropical setting, and major displacements of the African monsoon or extra-desert moisture sources.

Change in mean annual precipitation over northern Africa between mid-Holocene (6 ka) and pre-industrial conditions in PMIP3 models (affiliations are provided in Supplementary Table S4). Lakes Victoria and Chad outlined in blue. (a) Ensemble mean change in mean annual precipitation and positions of the African summer (July–September) ensemble mean ITCZ during mid-Holocene (solid red line) and pre-industrial conditions (solid blue line). (b) Zonal average of change in mean annual precipitation over land (20°W–30°E) for the ensemble mean (thick black) and individual models are listed on right). The range of minimal estimated change in mean annual precipitation required to sustain steppe is shown in shaded green (Jolly et al., 1998).


If not megalakes, what size lakes, marshes, discharging springs, and flowing rivers in the Sahara were sustainable in Sahelo-Sudanian climatic conditions? For lakes and perennial rivers to be created and sustained, net rainfall in the basin has to exceed loss to evapotranspiration, evaporation, and infiltration, yielding runoff that then supplies a local lake or river. Our hydrologic models (see Supplementary Material) and empirical observations (Gash et al., 1991; Monteith, 1991) for the Sahel suggest that this limit is in the 200–300 mm/yr range, meaning that most of the Sahara during the AHP was probably too dry to support very large lakes or perennial rivers by means of local runoff. This does not preclude creation of local wetlands supplied by ground-water recharge focused from a very large recharge area or forced to the surface by hydrologic barriers such as faults, nor megalakes like Chad supplied by moisture from the subtropics and tropics outside the Sahel. But it does raise a key question concerning the size of paleolakes, if not megalakes, in the Sahara during the AHP. Our analysis suggests that Sahelo-Sudanian climate could perhaps support a paleolake approximately ≤5000 km2 in area in the Darfur basin and ≤10,000–20,000 km2 in the Fezzan basin. These are more than an order of magnitude smaller than the megalakes envisioned for these basins, but they are still sizable, and if enclosed in a single body of water, should have been large enough to generate clear shorelines (Enzel et al., 2015, 2017). On the other hand, if surface water was dispersed across a series of shallow and extensive but partly disconnected wetlands, as also implied by previous research (e.g., Pachur and Hoelzmann, 1991), then shorelines may not have developed.

One of the underdeveloped ideas of my Indo-European demic diffusion model was that R1b-V88 had migrated through South Italy to Northern Africa, and from it using the Sahara Green Corridor to the south, from where the “upside-down” view of Bender (2007) could have occurred, i.e. Afroasiatic expanding westwards within the Green Sahara, precisely at this time, and from a homeland near the Megalake Chad region (see here).

Whether or not R1b-V88 brought the ‘original’ lineage that expanded Afroasiatic languages may be contended, but after D’Atanasio et al. (2018) it seems that only two lineages, E-M2 and R1b-V88, fit the ‘star-like’ structure suggesting an appropriate haplogroup expansion and necessary regional distribution that could explain the spread of Afroasiatic languages within a reasonable time frame.

Palaeolithic migrations

This review shows that the hypothesized Green Sahara corridor full of megalakes that some proposed had fully connected Africa from west to east was actually a strip of Sahelo-Sudanian steppe spread to the north of its current distribution, including the Chad megalake, East Africa and Arabia, apart from other discontinuous local wetlands further to the north in Africa. This greenish belt would have probably allowed for the initial spread of early Afroasiatic proto-languages only through the southern part of the current Sahara Desert. This and the R1b-V88 haplogroup distribution in Central and North Africa (with a prevalence among Chadic speakers probably due to later bottlenecks), and the Near East, leaves still fewer possibilities for an expansion of Afroasiatic from anywhere else.

If my proposal turns out to be correct, this Afroasiatic-like language would be the one suggested by some in the vocabulary of Old European and North European local groups (viz. Kroonen for the Agricultural Substrate Hypothesis), and not Anatolian farmer ancestry or haplogroup G2, which would have been rather confined to Southern Europe, mainly south of the Loess line, where incoming Middle East farmers encountered the main difficulties spreading agriculture and herding, and where they eventually admixed with local hunter-gatherers.

NOTE. If related to attested languages before the Roman expansion, Tyrsenian would be a good candidate for a descendant of the language of Anatolian farmers, given the more recent expansion of Anatolian ancestry to the Tuscan region (even if already influenced by Iran farmer ancestry), which reinforces its direct connection to the Aegean.

The fiercest opposition to this R1b-V88 – Afroasiatic connection may come from:

  • Traditional Hamito-Semitic scholars, who try to look for any parent language almost invariably in or around the Near East – the typical “here it was first attested, ergo here must be the origin, too”-assumption (coupled with the cradle of civilization memes) akin to the original reasons behind Anatolian or Out-of-India hypotheses; and of course
  • autochthonous continuity theories based on modern subclades, of (mainly Semitic) peoples of haplogroup E or J, who will root for either one or the other as the Afroasiatic source no matter what. As we have seen with the R1a – Indo-European hypothesis (see here for its history), this is never the right way to look at prehistoric migrations, though.

I proposed that it was R1a-M417 the lineage marking an expansion of Indo-Uralic from the east near Lake Baikal, then obviously connected to Yukaghir and Altaic languages marked by R1a-M17, and that haplogroup R could then be the source of a hypothetic Nostratic expansion (where R2 could mark the Dravidian expansion), with upper clades being maybe responsible for Borean.

Simple Nostratic tree by Bomhard (2008)

However, recent studies have shown early expansions of R1b-297 to East Europe (Mathieson et al. 2017 & 2018), and of R1b-M73 to East Eurasia probably up to Siberia, and possibly reaching the Pacific (Jeong et al. 2018). Also, the Steppe Eneolithic and Caucasus Eneolithic clusters seen in Wang et al. (2018) would be able to explain the WHG – EHG – ANE ancestry cline seen in Mesolithic and Neolithic Eurasia without a need for westward migrations.

Dravidian is now after Narasimhan et al. (2018) and Damgaard et al. (Science 2018) more and more likely to be linked to the expansion of the Indus Valley civilization and haplogroup J, in turn strongly linked to Iranian farmer ancestry, thus giving support to an Elamo-Dravidian group stemming from Iran Neolithic.

NOTE. This Dravidian-IVC and Iran connection has been supported for years by knowledgeable bloggers and commenters alike, see e.g. one of Razib Khan’s posts on the subject. This rather early support for what is obvious today is probably behind the reactionary views by some nationalist Hindus, who probably saw in this a potential reason for a strengthened Indo-Aryan/Dravidian divide adding to the religious patchwork that is modern India.

I am not in a good position to judge Nostratic, and I don’t think Glottochronology, Swadesh lists, or any statistical methods applied to a bunch of words are of any use, here or anywhere. The work of pioneers like Illich-Svitych or Starostin, on the other hand, seem to me solid attempts to obtain a faithful reconstruction, if rather outdated today.

NOTE. I am still struggling to learn more about Uralic and Indo-Uralic; not because it is more difficult than Indo-European, but because – in comparison to PIE comparative grammar – material about them is scarce, and the few available sources are sometimes contradictory. My knowledge of Afroasiatic is limited to Semitic (Arabic and Akkadian), and the field is not much more developed here than for Uralic…

Spread of Y-haplogroup R1b(xM269) in Eurasia, according to Jeong et al. (2018).

If one wanted to support a Nostratic proto-language, though, and not being able to take into account genome-wide autosomal admixture, the only haplogroup right now which can connect the expansion of all its branches is R1b-M343:

  • R1b-L278 expanded from Asia to Europe through the Iranian Plateau, since early subclades are found in Iran and the Caucasus region, thus supporting the separation of Elamo-Dravidian and Kartvelian branches;
  • From the Danube or another European region ‘near’ the Villabruna 1 sample (of haplogroup R1b-L754):
    • R1b-V88 expanding everywhere in Europe, and especially the branch expanding to the south into Africa, may be linked to the initial Afroasiatic expansion through the Pale-Green Sahara corridor (and even a hypothetic expansion with E-M2 subclades and/or from the Middle East would also leave open the influence of V88 and previous R1b subclades from the Middle East in the emergence of the language);
    • R1b-297 subclades expanding to the east may be linked to Eurasiatic, giving rise to both Indo-Uralic (M269) and Macro- or Micro-Altaic (M73) expansions.

This is shameless, simplistic speculation, of course, but not more than the Nostratic hypothesis, and it has the main advantage of offering ‘small and late’ language expansions relative to other proposals spanning thousands (or even tens of thousands) of years more of language separation. On the other hand, that would leave Borean out of the question, unless the initial expansion of R1b subclades happened from a community close to lake Baikal (and Mal’ta) that was also at the origin of the other supposedly related Borean branches, whether linked to haplogroup R or to any other…

NOTE. If Afroasiatic and Indo-Uralic (or Eurasiatic) are not genetically related, my previous simplistic model, R1b-Afroasiatic vs. R1a-Eurasiatic, may still be supported, with R1a-M17 potentially marking the latest meaningful westward population expansion from which EHG ancestry might have developed (see here). Without detailed works on Nostratic comparative grammar and dialectalization, and especially without a lot more Palaeolithic and Mesolithic samples, all this will remain highly speculative, like proposals of the 2000s about Y-DNA-haplogroup – language relationships.


Ancient genomes from North Africa evidence Neolithic migrations to the Maghreb

BioRxiv preprint now published (behind paywall) Ancient genomes from North Africa evidence prehistoric migrations to the Maghreb from both the Levant and Europe, by Fregel et al., PNAS (2018).

NOTE. I think one of the important changes in this version compared to the preprint is the addition of the recent Iberomaurusian samples.

Abstract (emphasis mine):

The extent to which prehistoric migrations of farmers influenced the genetic pool of western North Africans remains unclear. Archaeological evidence suggests that the Neolithization process may have happened through the adoption of innovations by local Epipaleolithic communities or by demic diffusion from the Eastern Mediterranean shores or Iberia. Here, we present an analysis of individuals’ genome sequences from Early and Late Neolithic sites in Morocco and from Early Neolithic individuals from southern Iberia. We show that Early Neolithic Moroccans (∼5,000 BCE) are similar to Later Stone Age individuals from the same region and possess an endemic element retained in present-day Maghrebi populations, confirming a long-term genetic continuity in the region. This scenario is consistent with Early Neolithic traditions in North Africa deriving from Epipaleolithic communities that adopted certain agricultural techniques from neighboring populations. Among Eurasian ancient populations, Early Neolithic Moroccans are distantly related to Levantine Natufian hunter-gatherers (∼9,000 BCE) and Pre-Pottery Neolithic farmers (∼6,500 BCE). Late Neolithic (∼3,000 BCE) Moroccans, in contrast, share an Iberian component, supporting theories of trans-Gibraltar gene flow and indicating that Neolithization of North Africa involved both the movement of ideas and people. Lastly, the southern Iberian Early Neolithic samples share the same genetic composition as the Cardial Mediterranean Neolithic culture that reached Iberia ∼5,500 BCE. The cultural and genetic similarities between Iberian and North African Neolithic traditions further reinforce the model of an Iberian migration into the Maghreb.

Ancestry inference in ancient samples from North Africa and the Iberian Peninsula. PCA analysis using the Human Origins panel (European, Middle Eastern, and North African populations) and LASER projection of aDNA samples.

Relevant excerpts:

FST and outgroup-f3 distances indicate a high similarity between IAM and Taforalt. As observed for IAM, most Taforalt sample ancestry derives from Epipaleolithic populations from the Levant. However, van de Loosdrecht et al. (17) also reported that one third of Taforalt ancestry was of sub-Saharan African origin. To confirm whether IAM individuals show a sub-Saharan African component, we calculated f4(chimpanzee, African population; Natufian, IAM) in such a way that a positive result for f4 would indicate that IAM is composed both of Levantine and African ancestries. Consistent with the results observed for Taforalt, f4 values are significantly positive for West African populations, with the highest value observed for Gambian and Mandenka (Fig. 3 and SI Appendix, Supplementary Note 10). Together, these results indicate the presence of the same ancestral components in ∼15,000-y old and ∼7,000-y-old populations from Morocco, strongly suggesting a temporal continuity between Later Stone Age and Early Neolithic populations in the Maghreb. However, it is important to take into account that the number of ancient genomes available for comparison is still low and future sampling can provide further refinement in the evolutionary history of North Africa.

Genetic analyses have revealed that the population history of modern North Africans is quite complex (11). Based on our aDNA analysis, we identify an Early Neolithic Moroccan component that is (i) restricted to North Africa in present-day populations (11); (ii) the sole ancestry in IAM samples; and (iii) similar to the one observed in Later Stone Age samples from Morocco (17). We conclude that this component, distantly related to that of Epipaleolithic communities from the Levant, represents the autochthonous Maghrebi ancestry associated with Berber populations. Our data suggests that human populations were isolated in the Maghreb since Upper Paleolithic times. Our hypothesis is in agreement with archaeological research pointing to the first stage of the Neolithic expansion in Morocco as the result of a local population that adopted some technological innovations, such as pottery production or farming, from neighboring areas.

By 3,000 BCE, a continuity in the Neolithic spread brought Mediterranean-like ancestry to the Maghreb, most likely from Iberia. Other archaeological remains, such as African elephant ivory and ostrich eggs found in Iberian sites, confirm the existence of contacts and exchange networks through both sides of the Gibraltar strait at this time. Our analyses strongly support that at least some of the European ancestry observed today in North Africa is related to prehistoric migrations, and local Berber populations were already admixed with Europeans before the Roman conquest. Furthermore, additional European/ Iberian ancestry could have reached the Maghreb after KEB people; this scenario is supported by the presence of Iberian-like Bell-Beaker pottery in more recent stratigraphic layers of IAM and KEB caves. Future paleogenomic efforts in North Africa will further disentangle the complex history of migrations that forged the ancestry of the admixed populations we observe today.

Ancestry inference in ancient samples from North Africa and the Iberian Peninsula. (B) ADMIXTURE analysis using the Human Origins dataset (European, Middle Eastern, and North African populations) for modern and ancient samples (K = 8). (D) Detail of ADMIXTURE analysis using the Human Origins dataset (European, Middle Eastern, North African, and sub-Saharan African populations) for modern and ancient samples, including Taforalt.

Also, from the main author’s Twitter account:

I just realized that the paragraph with information on data availability is missing! Sequence data in the European Nucleotide Archive (PRJEB22699). Consensus mtDNA sequences are available at the National Center of Biotechnology Information (Accession Numbers MF991431-MF991448).

I find it hard to believe that this genetic continuity from Upper Palaeolithic to Late Neolithic could be representative of an autochthonous development of Afroasiatic. An important population movement – likely more than one – must be found in ancient DNA influencing North-Central and North-East Africa, probably during the time of the Green Sahara corridor.

See here: