Optimal Migration Routes of Initial Upper Palaeolithic Populations to Eurasia


Ecological Niche and Least-Cost Path Analyses to Estimate Optimal Migration Routes of Initial Upper Palaeolithic Populations to Eurasia, by Kondo et al. (2018), from The Middle and Upper Paleolithic Archeology of the Levant and Beyond, Replacement of Neanderthals by Modern Humans Series. Chapter downloadable at Academia.edu.


This paper presents a computer-based method to estimate optimal migration routes of early human population groups by a combination of ecological niche analysis and least-cost path analysis. In the proposed method, niche probability is predicted by MaxEnt, an ecological niche model based on the maximum entropy theory. Location of known archaeological sites and environmental factors derived from palaeoterrain and palaeoclimate models, are input to the model to calculate the niche probability at each spatial pixel and weights of the environmental factors. The inverse of probability score is then used as an index of relative dispersal rate to accumulate the travel cost from a given origin. Based on this cumulative cost surface, least-cost paths from the origin to given destinations are visualised. This method was applied to the Initial Upper Palaeolithic population group (probably of modern humans) in Eurasia. The model identified three migration routes from the Levant to (1) Central Europe via Anatolia and Eastern Europe, (2) the Russian steppe via Caucasus Mountains, and (3) the Altai region via the southern coastal Iran and Afghanistan.

Cumulative cost to the southernmost IUP site (Wadi Aghir) using the inverse of the niche probability of the recovery experiment (corresponding to a warm/humid phase) as friction value

Check out also the chapter The Middle to Upper Paleolithic Transition in the Zagros: The Appearance and Evolution of the Baradostian, by Sonia Shidrang, from the same book. Also downloadable at Academia.edu.

Featured image from the chapter: “Niche probability for the IUP lithic industry predicted by MaxEnt using the palaeoclimate model from the recovery experiment (corresponding to a warm/humid phase).”

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The Russian school and the Yamna cultural-historical community, with emphasis on the north-west Pontic region


I have already talked about the Russian school of thought and their position regarding a Mesolithic origin of Proto-Indo-European in Northern Europe (see below related posts).

Since their archaeologists (Ukrainian, Russian, and Kazakh) are the nearest to potential Indo-Uralic origins, I have also recommended to follow some renown researchers closely.

Recently Leo S. Klejn referred to the position of Svetlana Ivanova. I found a recent summary of her model for genetic finds in an article appeared in Генофонд.рф: Степное население в Центральной Европе эпохи ранней бронзы, или путешествие туда и обратно

Aspects I agree with

– There is a Yamna cultural-historical community (i.e. with a potential ethnolinguistic unity). Although many different inner groups can be distinguished (based on cultural, social, anthropological differences), one cannot divide the culture in distinct cultures.

– There is an older Neolithic cultural-historical community of the Pontic-Caspian steppe (i.e. with a potential ethnolinguistic unity), marked in genetics by CHG ancestry in the region. It is impressive that they supported that before the Mathieson et al. third revision (september 2017) with their finding of “Yamnaya” component in Ukraine Eneolithic. Chapeau for archaeology, again.

Late Eneolithic North-West Pontic zone (after Brujako and Samojlova (eds.), 2013)

Aspects I am neutral about, but are potentially quite relevant for the future

Repin is a culture different from Yamna.

– The Budzhak culture is likely the heir of Repin, which is compatible with its expansion westward. According to Klejn and Anthony (Usatovo), this region was connected to (and might have influenced) the Corded Ware culture. Therefore – that is my contribution, not theirs – a hidden community of R1a-M417 subclades (that a lot of people are eager to find) might have stemmed from there.

NOTE: In my opinion, as I have said before, the Budzhak/Usatovo-Corded Ware connection is too late to be meaningful for Genomic finds (taking into account the earliest Corded Ware samples, their steppe ancestry, and the TMRCA for R1a-M417 subclades) – it is after all a late group appeared during or after the Yamna expansion along the Danube ca. 3000 BC or later, as supported by most archaeologists today, see e.g. Rassamakin. The most likely model is that R1a-M417 subclades with steppe ancestry expanded from groups near Eneolithic Ukraine, whether from steppe, forest-steppe, or forest zone. Also, in my opinion (supporting Anthony’s original interpretation of Repin, as closely connected to Yamna) it is more likely that the (until now) ‘hidden’ western Yamna community hosting R1b-L51 subclades that later evolved into East Bell Beaker is in fact represented by Repin…

Globular Amphorae and Corded Ware cultures are related, because GAC is actually not a uniform culture, but a ‘complex’ (i.e., in the same sense that Bell Beaker was not a culture, but a complex, as genetics has shown), and CWC is also a complex of cultures, as supported by Furholt. If that is true, the sampling of certain peoples classified as from Globular Amphorae – which some have rushed to cite as the end of the GAC-CWC connection might not be the last word, and Kristiansen’s model of long-lasting GAC-CWC connection may still be open.

Yamna culture and its surroundings (after Brujako and Samojlova (eds.), 2013)

Aspects I disagree with

– There is no migration, but long-lasting contacts that show up in genetics, since the Mesolithic-Neolithic transition. In my opinion, the expansion of admixture + haplogroup reduction and expansion depict clear migratory movements (although, obviously, without cultural identification no speculative ethnolinguistic grouping can be proposed). That much is obvious from Genomics, and if we are not going to accept the most basic findings as proof in favour of certain anthropological models, then Archaeology will not benefit at all from genetic studies.

– Because this is the Russian school of thought, when they talk about Proto-Indo-European they refer to a homeland dating to the Mesolithic-Neolithic transition, and therefore cultural-historical communities after that (and long-lasting contacts) refer to potential continuations of Proto-Indo-European. Following common language guesstimates, though, there is no reason to date the split of Anatolian from a common Indo-Hittite before ca- 4500 BC, since there is no reason to date a Late Proto-Indo-European beyond 4000-3000 BC, apart from controversial glottochronological studies…

A call for collaboration

Again, please, geneticists, archaeologists, and linguists: do collaborate. Merely by talking, the so-called ‘Yamnaya ancestral component’ would have never been given that dreadful name, and maybe we could end this quest to find a Mesolithic homeland for a reconstructed language guesstimated to have been spoken millennia after that time (and maybe turn it into a quest for an older macro-language)…


Mitogenomes show ancient human migrations to and through North-East India not of males exclusively


New open article Ancient Human Migrations to and through Jammu Kashmir- India were not of Males Exclusively, by Sharma et al., Scientific Reports 8, N. 851 (2018)


Jammu and Kashmir (J&K), the Northern most State of India, has been under-represented or altogether absent in most of the phylogenetic studies carried out in literature, despite its strategic location in the Himalayan region. Nonetheless, this region may have acted as a corridor to various migrations to and from mainland India, Eurasia or northeast Asia. The belief goes that most of the migrations post-late-Pleistocene were mainly male dominated, primarily associated with population invasions, where female migration may thus have been limited. To evaluate female-centered migration patterns in the region, we sequenced 83 complete mitochondrial genomes of unrelated individuals belonging to different ethnic groups from the state. We observed a high diversity in the studied maternal lineages, identifying 19 new maternal sub-haplogroups (HGs). High maternal diversity and our phylogenetic analyses suggest that the migrations post-Pleistocene were not strictly paternal, as described in the literature. These preliminary observations highlight the need to carry out an extensive study of the endogamous populations of the region to unravel many facts and find links in the peopling of India.


To conclude, the extent of presence of variants defining novel HGs or personal variants indicate high diversity in maternal genetic component of the population of J&K. Statistical analyses indicate that maternal population in J&K have undergone expansion, along with other regions of Indian sub-continent9. However, signatures of maternal gene pool expansion in the region past LGM and early Holocene era are also seen, and this is a unique observation for the present study. These distinct signatures and maternal lineages, never reported before in India, apparently suggest that this region might have served as a corridor, yet also as a reservoir for many unreported lineages.

The overall diversity seen in the maternal gene pool of J&K suggests that the migrations to and through this region were not exclusively of males. This data has refined the existing phylogenetic tree and added to the information further diversity of mtDNA in Indian populations. Further, this preliminary study highlights the importance of the region and emphasizes that the populations of this region should be studied extensively to understand the gene pool of Indian populations. Along with the Y chromosomal and mtDNA markers, a study of autosomal markers is also warranted in these population groups. It is anticipated to help in finding some of the missing links in the evolution of modern humans and their migratory history to and from the mainland India and the Indian subcontinent, a future perspective of our study. Further, we would like to emphasize that the endogamous populations should be studied with respect to their individual evolutionary and migration histories, rather than pooling these together as one group, an underlying drawback that has plagued many of the Indian population based studies in the past, diluting individual signatures and masking stories their DNA has to tell.

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Something is very wrong with models based on the so-called ‘Yamnaya admixture’ – and archaeologists are catching up (II)

A new article by Leo S. Klejn tries to improve the Northern Mesolithic Proto-Indo-European homeland model of the Russian school of thought: The Steppe hypothesis of Indo-European origins remains to be proven, Acta Archaeologica, 88:1, 193–204.


Recent genetic studies have claimed to reveal a massive migration of the bearers of the Yamnaya culture (Pit-grave culture) to the Central and Northern Europe. This migration has supposedly lead to the formation of the Corded Ware cultures and thereby to the dispersal of Indo-European languages in Europe. The article is a summary presentation of available archaeological, linguistic, genetic and cultural data that demonstrates many discrepancies in the suggested scenario for the transformations caused by the Yamnaya “invasion” some 5000 years ago.


Both teams [Reich/Anthony, and Willerslev/Kristiansen] interpreted this resemblance in the same way: as evidence of mass migration of the Yamnaya culture from the steppes into the Central and Northern Europe, resulting in the formation of the Corded Ware cultures, and these are universally recognised as Indo-European. Since earlier in this part of Europe existed a different pool of genomes, geneticists presumed that the Yamnaya migration alone had brought the Indo-European languages into Europe. It is difficult to say to what extent the pre-convictions of the involved archaeologists influenced these conclusions, or whether the results of the genetic studies attracted archaeologists with such beliefs.

Mismatch of cultural manifestations

First, we might question the idea of the Yamnaya culture as a unity rather than a loose conglomerate of cultures. Merpert (1974) divided it into nine local groups but did not recognise them as separate cultures. However, in 1975 I suggested that Nerushay (Budzhak) monuments should be recognised as a distinct culture (Klejn 1975), although still as a part of the same broader steppe community.

This was accepted by other specialists (Ivanova 2012; 2013; 2014). Generally, in the western branch of this community, a mixture of the eastern rites of interment with local, Balkan ceramics can be observed. It should be noted that hitherto all genetic samples were taken from eastern material (in the vicinity of Samara in the Volga basin and Kalmykia), while the central thesis concerns the intrusion of the western branch of this community (Budzhak culture) into Europe.

The spread of cultural-historical communities of the Yamnaya culture and the location of the Budzhak culture. GAC – Globular Amphora culture; CWC – Corded Ware culture. After Ivanova 2013.

Simultaneity of cultures

The Yamnaya culture (Chernykh & Orlovskaya 2004a; Heyd 2011; Frȋnculeasa et al. 2015) appears not to be the predecessor of the Corded Ware cultures but is contemporary with them. The Corded Ware cultures appeared also around the turn between the fourth and third millennium BC (Stöckli 2001; Furholt 2003). Their derivation from the Yamnaya seems, therefore, to be less probable. This is evidenced by the fact that the corded beakers or amphorae found in the Budzhak culture are not the prototypes of the corded beakers or amphorae found in more northern territories, but seem instead to be an outcome of contemporaneous contacts (Ivanova 2014; Klejn 2017c).

Discrepancies across the haplogroups

Even more remarkable is the variation in the distribution of types of Y chromosome. In the Yamnaya population, R1b is not just a single occurrence (there are about seven known occurrences) while in the Corded Ware population a different clade of R1b is found and R1a is predominant (several instances). Thus the postulate of unbroken succession finds no support!

Distribution of artefacts and customs of the Yamnaya culture in the area of the Corded Ware cultures. After Bátora 2006.

Paradoxical gradient

In the tables presented in the article by Reichs’ team (Haak et al. 2015) the genetic pool connecting the Yamnaya culture with the Corded Ware people is shown to be more intense in Northern Europe (Norway and Sweden) and decreases gradually from the North to the South (Fig. 6). It is weakest around the Danube, in Hungary, i. e. areas neighbouring the western branch of the Yamnaya culture! This is the reverse image to what the proposed hypothesis by the geneticists would lead us to expect. It is true that this gradient is traced back from the contemporary materials, but it was already present during the Bronze Age (Klejn 2015a).

The author also uses questionable interpretations from selected articles to advance his (as of today) untenable positions regarding a Mesolithic origin of the reconstructible Proto-Indo-European language.

1. Glottochronology, for a PIE origin:

If based on the data of glottochronology (taking into account all disputes) the period of initial dispersal is to be dated to the 7th-5th millennium BC.

2. Doubts on the origin of R1b-L51 subclades expressed in Genetic differentiation between upland and lowland populations shapes the Y-chromosomal landscape of West Asia, by Balanovsky et al. (2017), Human Genetics 136, 4. 437-450:

The currently available dataset does not contradict the hypothesis that R-GG400 marks a link between the East European steppe dwellers and West Asians, though the route and even direction of this migration is disputable. It does, however, demonstrate that present-day West European R1b chromosomes do not originate from the Yamnaya populations analyzed in (Haak et al. 2015; Mathieson et al. 2015) and raises the question of their origin. A Bronze Age origin is more likely than a Neolithic one (Balaresque et al. 2010), but further ancient DNA studies may be necessary to identify this source.

Just yesterday I read the post The retraction paradox: Once you retract, you implicitly have to defend all the many things you haven’t yet retracted, by Andrew Gelman. While – in my opinion – the post does not live up to its title, it poses an interesting question, as to how ad logicam (fallacy fallacy) is often used today in research: One author proposes something that is later demonstrated to be wrong, so everything they wrote or write can be said ipso facto to be wrong…especially if they accept that it was wrong.

This is usual with amateur geneticists (those who don’t publish, and are therefore not subjected to criticism): if anyone is wrong (whether in Archaeology or Genetics), then they are wrong in everything else. It seems to me that Klejn’s theses against recent genetic results rest on the same assumption: The Yamna -> Corded Ware migration model is wrong, ergo the Yamna homeland model is wrong.

I guess this same fallacy is what a lot of angered geneticists (whether professional or amateurs) are going to use to dismiss Klejn’s criticism, trying to focus on what he clearly does not grasp – about genomic data of Yamna peoples and their expansion – to disregard his doubts on genetic interpretations entirely.

I have warned many times about how simplistic interpretations of genetic data would cause a general mistrust in the field, and that archaeologists won’t take the discipline seriously, no matter how many articles get published in famous research tabloids like Nature or Science…

Those who dismiss this warning lightly seem to forget the fate of other recent “scientific breakthroughs” which were initially so promising that Humanities appeared to matter no more, like glottochronology for Linguistics and, to some extent, that of radiocarbon analysis for Archaeology.
EDIT: see here a recent example of discusion on discrepancies between archaeological and 14C-based chronologies, whereby ‘scientific data’ obviously needs archaeological context for a meaningful interpretation

Featured image: The direction of the supposed migration of the bearers of the Yamnaya culture into the area of the Corded Ware cultures. After Haak et al. 2015.

NOTE: I obviously don’t agree with Klejn’s main model: he criticises the Proto-Indo-European steppe homeland, and more specifically the expansion of Yamna peoples with R1b-L23 subclades, which I support. But, probably because of his “pre-convictions” (as he puts it when describing proponents of the steppe hypotheses) about the Proto-Indo-European homeland in Northern Europe during the Mesolithic, he was one of the first renown archaeologists to criticise the obvious inconsistencies in the genetic model of migrations based exclusively on the “Yamnaya ancestral component” concept, and to provoke the necessary reaction from (until then) overconfident geneticists, and he deserves credit for that.

In my opinion, the Russian school’s “Northern European Mesolithic” homeland model – as I have said before – could be based on the appearance of EHG ancestry, or maybe on the expansion of haplogroup R1b with post-Swiderian cultures, but the timeframe proposed is too early for any reconstructible parent proto-language, even for Indo-Uralic.


Modern Hungarian mtDNA more similar to ancient Europeans than to Hungarian conquerors


New preprint at BioRxiv, MITOMIX, an Algorithm to Reconstruct Population Admixture Histories Indicates Ancient European Ancestry of Modern Hungarians, by Maroti et al. (2018).

The estimated age distribution of the shared mt Hgs between Hungarians (Hun), the best hypothetical admix (mixFreq) and the populations contributing to this admix: Belgian/Dutch (BeN), Danish (Dan), Basque (Bsq), Croatian/Serbian (CrS), Baltic Late Bronze Age culture (BalBA), Bell Beaker culture (BellB), Slovakian (Slo). The numbers in parentheses indicate the contributions to the best hypothetical admix.

Abstract (emphasis mine)

By making use of the increasing number of available mitogenomes we propose a novel population genetic distance metric, named Shared Haplogroup Distance (SHD). Unlike FST, SHD is a true mathematical distance that complies with all metric axioms, which enables our new algorithm (MITOMIX) to detect population-level admixture based on SHD minimum optimization. In order to demonstrate the effectiveness of our methodology we analyzed the relation of 62 modern and 25 ancient Eurasian human populations, and compared our results with the most widely used FST calculation. We also sequenced and performed an in-depth analysis of 272 modern Hungarian mtDNA genomes to shed light on the genetic composition of modern Hungarians. MITOMIX analysis showed that in general admixture occurred between neighboring populations, but in some cases it also indicated admixture with migrating populations. SHD and MITOMIX analysis comply with known genetic data and shows that in case of closely related and/or admixing populations, SHD gives more realistic results and provides better resolution than FST. Our results suggest that the majority of modern Hungarian maternal lineages have Late Neolith/Bronze Age European origins (partially shared also with modern Danish, Belgian/Dutch and Basque populations), and a smaller fraction originates from surrounding (Serbian, Croatian, Slovakian, Romanian) populations. However only a minor genetic contribution (<3%) was identified from the IXth Hungarian Conquerors whom are deemed to have brought Hungarians to the Carpathian Basin. Our analysis shows that SHD and MITOMIX can augment previous methods by providing novel insights into past population processes.

Unrooted hierarchic cluster of modern and archaic populations based on the SHD matrix.

It is interesting to keep receiving data as to how language does not correlate well with Genomics, whether admixture or haplogroups, even though it is already known to happen in regions such as Anatolia, the Baltic, South-Eastern or Northern Europe.

Thorough anthropological models of migration or cultural diffusion are necessary for a proper interpretation of genetic data. There is no shortcut to that.

Co-occurrence of Hungarian Bronze Age mt Hgs Distribution of mt Hgs found in Hungarian Bronze Age archaic samples in the analyzed populations. The fixation dates are based on Behar et al [6].

Images made available under a CC-BY-NC-ND 4.0 International license.
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Ancient Phoenician mtDNA from Sardinia, Lebanon reflects settlement, genetic diversity, and female mobility


New article at PLOS One, Ancient mitogenomes of Phoenicians from Sardinia and Lebanon: A story of settlement, integration, and female mobility, by Matisoo-Smith et al. (2018).


The Phoenicians emerged in the Northern Levant around 1800 BCE and by the 9th century BCE had spread their culture across the Mediterranean Basin, establishing trading posts, and settlements in various European Mediterranean and North African locations. Despite their widespread influence, what is known of the Phoenicians comes from what was written about them by the Greeks and Egyptians. In this study, we investigate the extent of Phoenician integration with the Sardinian communities they settled. We present 14 new ancient mitogenome sequences from pre-Phoenician (~1800 BCE) and Phoenician (~700–400 BCE) samples from Lebanon (n = 4) and Sardinia (n = 10) and compare these with 87 new complete mitogenomes from modern Lebanese and 21 recently published pre-Phoenician ancient mitogenomes from Sardinia to investigate the population dynamics of the Phoenician (Punic) site of Monte Sirai, in southern Sardinia. Our results indicate evidence of continuity of some lineages from pre-Phoenician populations suggesting integration of indigenous Sardinians in the Monte Sirai Phoenician community. We also find evidence of the arrival of new, unique mitochondrial lineages, indicating the movement of women from sites in the Near East or North Africa to Sardinia, but also possibly from non-Mediterranean populations and the likely movement of women from Europe to Phoenician sites in Lebanon. Combined, this evidence suggests female mobility and genetic diversity in Phoenician communities, reflecting the inclusive and multicultural nature of Phoenician society.

Haplogroup assignments, dates, locations and Genbank accession details of all aDNA samples included in analyses.

Featured image, from the article: Map showing phoenician maritime expansions across the Mediterranean starting from around 800 BCE. Arrows indicate maritime movement. Blue dots indicate coastal sites and pink shaded areas indicate the extent of Phoenician settlements. https://doi.org/10.1371/journal.pone.0190169.g001

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