Most interesting video-conference on Luwians, Troy and the Sea Peoples

All archaeogenetics and no archaeology makes people go mad. So let's spice this a bit with this absolutely enticing video of a conference by Dr. Eberhard Zangger, which I have watched thrice already, twice tonight alone.

I love the general outline of the exposition even if I know some details, like the outline of Lower Troy are controversial. 

I also did pose the following questions as commentary to the video:

1. How can the professor be so sure that all ancient Western Anatolian nations were Luwian and not from other diverse ethnicities? How that they were the only ones in the last Sea Peoples' wave? Just the same we see some non-Greeks in the Greek side of the Trojan war, I would expect some non-Luwians in the Trojan side as well, assuming the Trojans were Luwians and not Tyrsenians or something else. 
2. What about the Phrygians who show up in Anatolia, West and East (Armenians) after the Bronze Age collapse, out of nothing (they seem to originate in an obscure Paeonian tribe, the Bryges)? Not a single mention of them: I guess they would blurr the nice "Luwian" homogeneity. 
3. What about the Greeks (Danaoi, Denesh) and their Pelasgian (Peleset, Philistine) neighbors and often allies (Achilles himself and his Myrmidons were that)? They seem also involved in that late Sea Peoples wave and there is coincidence of cultural Hellenization (and not Luwianization) of Cyprus precisely in that period of the late Sea Peoples' attacks against Syria, Egypt and whatever else. Let's not forget that the Egyptians speak of the foreign peoples making a COALITION in their "islands", and I would say that this coalition involved peoples from all the Aegean, and not just the Asian side of it (although very good point about Evans' racism and his horrible influence on Aegean studies). 

But please don't let my nit-pickiness wrong what I think is a great conference dealing with a topic that has been way too neglected and even purposely ignored. There is a lot of good stuff in the video.

By the way, this is the Wikipedia map of Luwian inscriptions (unsure of what exactly the German legend says, "early" and "late" maybe?, but it's definitely about Luwian inscriptions):

Credit: Hendrik Tammen (CC-license)

Oldest evidence for cooking in pots comes from Libya

Quickies

Kambiz Karami at Anthropology.net (a most interesting blog to follow) mentioned this week that Libyan pottery remains, maybe as old as 10,000 years ago, have provided the oldest evidence of vegetable cooking of we know of, at least in pots (see below for the disclaimer). The remains indicate cooking of vegetables and meat and are associated to pictures of people gathering plants, as well as grinding stones (hand mills) with remains of such provisions.

Ref. Nature Plants.

Oddly enough, just two days later, he contradicted himself, mentioning that the charred remains of nuts and seeds from Palestine, dated from 780,000 years ago, actually provide the oldest evidence of vegetable cooking, even if it's clearly a more primitive way of cuisine and not yet at all the "refined" stew of ancient Libyans, which won them three stars in the Paleo-Michelin Guide for Nomads and five pitchforks in Popular Mesolithic Cuisine. 

Ref. PNAS.



Update (Jan 4): origin of Libyan and West Asian pottery could be Sudan

Jm8 mentions it in the comments, referencing to Anthromadness. I would need more data to judge (the ref. paper is behind paywall) but  it does sounds as probably correct to me, being Sudan/Nubia one of the earliest Mesolithic areas in the Western half of the Old World, one that is way too often and unfairly neglected but that definitely influenced the Levant prior to the development of Neolithic proper (what should explain a lot of things both linguistic and genetic).

Neolithic DNA from Southern Anatolia

I know, I know: I'm decaying into a total procrastinator. I don't have any excuse other than I don't feel like blogging as of late: neither on anthropology nor on politics. I rather feel like learning new stuff and playing, rather than writing and I lack of the structured environment to force myself to do otherwise than what I feel like most of the time. Being of compulsive temperament only worsens things.

I also know that this is not the proper way to start an article. Yes, I know. Do I even care?

So getting to mention now some of the stuff that I have not discussed in these last months and is definitely worth posting about. First of all this key study on more easterly Anatolian early farmers than those seen so far.

Intriguingly they are notoriously similar to those sequenced farther West (see here), what seems to support the model of Anatolian origin of European Neolithic peoples, largely ancestral to modern Europeans. However even Western Anatolian early farmers show already some extra admixture with the Paleoeuropean "WHG" component relative to their Southern Anatolian precursors. So, as the authors suggest, admixture between immigrant farmers and native foragers was a gradual and continuous process beginning in Asia Minor itself.

Gülşah Merve Kılınç, Ayça Omrak, Füsun Özer et al., The Demographic Development of the First Farmers in Anatolia. Current Biology 2016. Open access → LINK [doi:10.1016/j.cub.2016.07.057]

Summary

The archaeological documentation of the development of sedentary farming societies in Anatolia is not yet mirrored by a genetic understanding of the human populations involved, in contrast to the spread of farming in Europe [ 1–3 ]. Sedentary farming communities emerged in parts of the Fertile Crescent during the tenth millennium and early ninth millennium calibrated (cal) BC and had appeared in central Anatolia by 8300 cal BC [ 4 ]. Farming spread into west Anatolia by the early seventh millennium cal BC and quasi-synchronously into Europe, although the timing and process of this movement remain unclear. Using genome sequence data that we generated from nine central Anatolian Neolithic individuals, we studied the transition period from early Aceramic (Pre-Pottery) to the later Pottery Neolithic, when farming expanded west of the Fertile Crescent. We find that genetic diversity in the earliest farmers was conspicuously low, on a par with European foraging groups. With the advent of the Pottery Neolithic, genetic variation within societies reached levels later found in early European farmers. Our results confirm that the earliest Neolithic central Anatolians belonged to the same gene pool as the first Neolithic migrants spreading into Europe. Further, genetic affinities between later Anatolian farmers and fourth to third millennium BC Chalcolithic south Europeans suggest an additional wave of Anatolian migrants, after the initial Neolithic spread but before the Yamnaya-related migrations. We propose that the earliest farming societies demographically resembled foragers and that only after regional gene flow and rising heterogeneity did the farming population expansions into Europe occur.

Autosomal DNA

Maybe the most informative graph is this one (fig. 2):

Genetic Structure and Diversity of Central Anatolian Neolithic Populations
(A) PCA on contemporary west Eurasian populations onto which a total of 85 ancient individuals are projected from this study and previous studies. See Table S1 for number of SNPs per individual. Neighboring modern populations and ancient Anatolian populations are shown encircled. Modern population names are in italics.
Etc. (not so interested here in B, C and D, legend too long, check in the original paper)
Click to expand

It is interesting that, in spite of the Anatolian origin of this ancient ancestral population, they do not tend so much to modern Anatolian Turks but rather to Levant populations like Cypriots (closest ones), Lebanese, Palestinians, etc.

This is probably because, even if early Neolithic peoples of the Levant were not quite like them (see here again) they had become almost like them before the Bronze Age because of regional admixture, which I understand was mostly (but not only) north-to-south flow.

Notice that the Boncuklu (Bon) people had very low genetic diversity and they seem to be a dead end rather than directly ancestral. Instead, the Tepecik-Çiftilik (Tep) population seems a good proxy for the ancestors of Neolithic peoples of Western Anatolia and Europe. 

When we think about South Anatolia Neolithic, we usually think first and foremost about the famous Çatalhöyük site. Well, this ancient settlement is in the area of Boncuklu (to the West, both are near Konya) rather than that of Tepecik-Çiftilik (to the East, near Niğde), so it is quite possible that it is another demographic dead end, related but not directly ancestral to mainline European Neolithic. 

Personally I still think they could well have migrated at least partly by boat, along the southern Turkish coast but, until new data comes, I may need to alter my hypothesis of the ultimate origin being in the Northern Levant (Syria, Lebanon, Cyprus even) rather than Anatolia. These people of Tepecik-Çiftilik were, if not direct ancestors at least very closely related to the actual source population, which may well have lived closer to the coast in any case.

Mitochondrial DNA

The newly sequenced South Anatolian farmers had some of the lineages that were later present in Hungary's and Germany's "Danubian Neolithic", notably the now rare N1a1a1, found in 4/9 samples in this study. Also present were K1a (3/9, incl. one K1a12a), U3 (1/9) and N1a1b (1/9).

So it is time to dismiss the hypothesis that claimed N1a1a1 as a European aboriginal lineage: it came with the immigrant farmers and now there can be no doubt about it.

Ancient genomes from Neolithic West Asia

This week we got to know a lot more about the genetics of ancient West Asians, from the Mesolithic, Neolithic and later times. All in a single major study:

Iosif Lazaridis et al., The genetic structure of the world's first farmers. BioRxiv 2016. Freely accessible (pre-pub) → LINK [doi: http://dx.doi.org/10.1101/059311]

Abstract

We report genome-wide ancient DNA from 44 ancient Near Easterners ranging in time between ~12,000-1,400 BCE, from Natufian hunter-gatherers to Bronze Age farmers. We show that the earliest populations of the Near East derived around half their ancestry from a 'Basal Eurasian' lineage that had little if any Neanderthal admixture and that separated from other non-African lineages prior to their separation from each other. The first farmers of the southern Levant (Israel and Jordan) and Zagros Mountains (Iran) were strongly genetically differentiated, and each descended from local hunter-gatherers. By the time of the Bronze Age, these two populations and Anatolian-related farmers had mixed with each other and with the hunter-gatherers of Europe to drastically reduce genetic differentiation. The impact of the Near Eastern farmers extended beyond the Near East: farmers related to those of Anatolia spread westward into Europe; farmers related to those of the Levant spread southward into East Africa; farmers related to those from Iran spread northward into the Eurasian steppe; and people related to both the early farmers of Iran and to the pastoralists of the Eurasian steppe spread eastward into South Asia.

Highlights:

• There were (at least) two clearly distinct populations in West Asia in the Mesolithic and Early Neolithic times.
• Both populations contributed to the West Anatolian farmers that are precursors of the settlers of Neolithic Europe.
• The so-called "Basal Eurasian" component is not yet clarified if it is something local or admixture with Africans or both. However it is clear that it is associated with reduced Neanderthal admixture.
• West Eurasian genetic composition can be now understood quite well as the mixture from four sources: two West Asian ones, favored by the Neolithic revolution, and two Paleo-European ones.

This graphic shows pretty well how the ancient populations of West Eurasia are expressed as a mixture of those four founder populations:

That is if you can get through the nomenclature, which is inherited in many cases from a long array of recent studies. I'm not even sure myself in many cases what samples exactly and where from are thrown in each category. But the most important part is that Iran_N and Levant_N are the two Neolithic-specific founder populations of the Fertile Crescent (yeah, N stands for "Neolithic", not "North") and that the other two founder populations from pre-Neolithic Europe are WHG (Epi-Magdalenian peoples from Western and Central Europe) and EHG (Eastern European hunter-gatherers, of Epigravettian culture and maybe even proto-Uralic in one case).

Then we see in the case of Europe how:

1. Anatolia_N (precursors of mainline European Neolithic) are a mix of both West Asian farmer groups, plus a sizable fraction of Western Paleo-european ancestry already.

2. This fraction of Western Paleoeuropeanness increases as the farmers expanded into Europe (EN) and then as there was probably some backflow of Western origins in relation to Megalithism and Bell Beaker (MNChL). But in general remains the same basic genetic composition and in no known case incorporates any Eastern Paleoeuropean component at all, not yet.

3. It is only with the Indoeuropean ("Kurgan") invasions reflected in the category LNBA, when the EHG component begins feeling very important in Europe. If I'm correct, all those samples are from Germany other areas of Central and North Europe, with the Iberian and Italian ones of similar chronology placed in the MNChL tag instead. The LNBA/MNChL contrast is not a strictly chronological analysis but an analysis by categories of ancestry that do overlap in time.

4. In Armenia instead, we see a decrease of the minor EHG component but then an increase in the MLBA ("middle and late Bronze Age") when Armenians arrive from the Balcans and Phrygia, conquering the pre-existing Hurro-Urartean peoples (whose language was probably related to Chechen and other NE Caucasian languages), which should correspond to the formation of Urartu and more specifically to the Hayasa-Azzi and Shupria stages, both considered Urartean (Hurrian). The WHG and Levant-N components we see since the Chalcolithic is similar to what we see in West Anatolia and probably reflect interactions corresponding to Central-Eastern Anatolia, Kurdistan and Syria, for which we have no direct ancient data yet.

Ancient samples (colored and labeled) projected on a PCA of modern West Eurasian populations (in gray):

For a reference on which are the modern populations in gray, a good reference is this older but fully labeled PCA by Olalde.

Briefly: Natufians fall on top of modern Palestinians, their slightly admixed Neolithic descendants fall between Palestinians and Jews, Middle Neolithic European Farmers fall on top of Sardinians, the so-called Europe-Steppe continuum (early Western Indoeuropeans) fall between Central Europe, France and the Balcans, most Western Europeans do not overlap with ancient samples but appear to have even greater Paleoeuropean admixture instead, etc.

Y-DNA Haplogroups

Iranian Mesolithic and Neolithic samples carried the following patrilineages:

• Mesolithic: J(xJ2a1b3,J2b2a1a1)
• Ganj Dareh Neolithic: P1(xQ,R1b1a2,R1a1a1b1a1b,R1a1a1b1a3a,R1a1a1b2a2a) and an undefined CT
• Late Neolithic: G2a1(xG2a1a)

Meanwhile Palestinian Mesolithic and Neolithic samples carried: 

• Natufian (Mesolithic): E1b1b1b2(xE1b1b1b2a,E1b1b1b2b), E1b1(xE1b1a1,E1b1b1b1), E1b1b1b2(xE1b1b1b2a,E1b1b1b2b), plus two undefined CT.
• Pre-Pottery Neolithic B/C: H2, E(xE2,E1a,E1b1a1a1c2c3b1,E1b1b1b1a1,E1b1b1b2b), E1b1b1, T(xT1a1,T1a2a), E1b1b1(xE1b1b1b1a1,E1b1b1a1b1,E1b1b1a1b2,E1b1b1b2a1c), plus three ill-defined CT.

CT is the main pan-Eurasian macro-haplogroup and is not informative, except in Palestine because it implies exclusion of E.

Otherwise we see an important presence of E (mostly E1b1b) a lineage we know was carried by early farmers into Europe and that has ultimately African origins. It probably indicates migration of NE Africans into Palestine in the Mesolithic, something also supported by Archaeology. However these NE Africans were surely already mixed with Eurasian ancestry, which probably arrived to the Nile Basin in the early LSA, some 50-40 Ka ago. So it's a complex story of multiple admixture events in the continental crossroads that is Egypt and also Palestine and other nearby areas.

We also see G2a1 in Late Neolithic Iran, and this one is the main lineage brought to Europe by the early farmers if we are to judge on known ancient sequences (today it is not more important that E1b but it is maybe more evenly distributed). However we only see it in the Late Neolithic, so it may have originated further west.

We see too little J, only J(xJ1a,J2a1,J2b) in Chalcolithic Iran and in Bronze Age Jordan: J(xJ1,J2a,J2b2a) again and J1(xJ1a). I guess that a lot remains to be researched on this issue because J is by far nowadays the most common haplogroup of West Asia, and also impacted Europe and South Asia (J2) and North and NE Africa (J1).

On the issue of "Basal Eurasian": African or West Asian?

The question remains unanswered, as I said before but there are two clues: on one side the presence of E1b in Mesolithic and Neolithic Palestine clearly supports a direct NE African influence, also backed by archaeological evidence. But there is some nuance in the issue of FST distances that I want to highlight.

The distances are available in a very extensive supplementary table, so I took just a few to get a better understanding, not only of this issue but in general of the genetic distances of the four founder populations:

Quite ironically it is not the Natufians who are the closest to the African reference population (Yoruba) but the CHG, Iran-N and Levant-N groups. In fact the Natufians are the most distant ones after the WHG population. However this is tricky because the affinity to Yoruba may also be caused by the "ghost" Basal Eurasian population, claimed first of all by Lazaridis 2014, which would be a remnant of the Out of Africa Migration (not strictly African but close enough and impossible to discern from true African admixture in most analyses).

So we may imagine that the "Highlander" (CHG and Iran-N) populations were somehow influenced by that Basal Eurasian ghostly population, which might have survived in the Persian Gulf oasis, for example. Or whatever else.

The presence of the same or similar element in Levant-N reflects possibly admixture with Iran-N or a similar population, something that is implicit in the table above but I'll address below more explicitly.

If there is (and there must be, because of Y-DNA E1b) some African admixture in the Natufian population, it was very diluted already in the autosomal (general DNA) aspect before farming began.

Update (Jul 2): all the four paragraphs above are possibly misleading to some extent because, as several commenters have rightfully pointed out, generic drift alone just causes the effect of increased distance to general reference populations like Yoruba and Han, this genetic drift is caused by relative isolation, so it seems that Magdalenian Europeans (WHG) and Natufian Palestinians (Natufian) were both more isolated populations in general terms than the Iran-Caucasus-Eastern Europe ones, whose sheer numbers apparently kept them more similar to the generic root of Humankind, less endogamous. 

However, per archaeology, such "sheer numbers" are not to be expected in that area, rather the opposite (Western Europe and Palestine are much more richer areas in terms archaeological, suggesting denser populations). So the question remains open as far as I can tell but it should be discerner with more precise tools than mere FST.

A visual of smallest genetic distances between (each "-" represents 0.01 in the table above):

a) Ancient West Asians:

CHG-----IrN-------LeN----Nat

Neolithic peoples of West Asia, even if different, are closer among them than their pre-Neolithic precursors.

b) Pre-Neolithic West Eurasians:

WHG--------EHG----------CHG--------------Nat

The distances between Natufians and everyone else are comparable to those with Han Chinese, however only in the case of the populations that appear to have extra affinity to East Asia (Iran, Caucasus and Eastern Europe), otherwise it is smaller.

All four populations were distant enough from each other to be considered clearly distinctive. Even EHG and WHG were quite dissimilar.

c) The four West Eurasian founders considered above:

WHG--------EHG----------IrN-------LeN

There is much greater similitude between Iran and Levant Neolithic peoples than between their Mesolithic precursors. This implies some sort of intense admixture as agriculture and herding developed. Not enough to erase the differences but enough to blur them significantly.

Genetic influence from East Asia or a related population is also apparent in all Northeastern populations but even more so in Iran Neolithic. Why?

There is much more in the study and supp. materials but I can only review so much.

Goat genetics suggest that two populations were domesticated

Quickies

Licia Colli, Hovirang Lancioni et al., Whole mitochondrial genomes unveil the impact of domestication on goat matrilineal variability. BMC Genomics 2015. Open access → LINK [doi:10.1186/s12864-015-2342-2]

Abstract

Background

The current extensive use of the domestic goat (Capra hircus) is the result of its medium size and high adaptability as multiple breeds. The extent to which its genetic variability was influenced by early domestication practices is largely unknown. A common standard by which to analyze maternally-inherited variability of livestock species is through complete sequencing of the entire mitogenome (mitochondrial DNA, mtDNA).

Results

We present the first extensive survey of goat mitogenomic variability based on 84 complete sequences selected from an initial collection of 758 samples that represent 60 different breeds of C. hircus, as well as its wild sister species, bezoar (Capra aegagrus) from Iran. Our phylogenetic analyses dated the most recent common ancestor of C. hircus to ~460,000 years (ka) ago and identified five distinctive domestic haplogroups (A, B1, C1a, D1 and G). More than 90 % of goats examined were in haplogroup A. These domestic lineages are predominantly nested within C. aegagrus branches, diverged concomitantly at the interface between the Epipaleolithic and early Neolithic periods, and underwent a dramatic expansion starting from ~12–10 ka ago.

Conclusions

Domestic goat mitogenomes descended from a small number of founding haplotypes that underwent domestication after surviving the last glacial maximum in the Near Eastern refuges. All modern haplotypes A probably descended from a single (or at most a few closely related) female C. aegagrus. Zooarchaelogical data indicate that domestication first occurred in Southeastern Anatolia. Goats accompanying the first Neolithic migration waves into the Mediterranean were already characterized by two ancestral A and C variants. The ancient separation of the C branch (~130 ka ago) suggests a genetically distinct population that could have been involved in a second event of domestication. The novel diagnostic mutational motifs defined here, which distinguish wild and domestic haplogroups, could be used to understand phylogenetic relationships among modern breeds and ancient remains and to evaluate whether selection differentially affected mitochondrial genome variants during the development of economically important breeds.

Note: "Southeastern Anatolia" should read Northern Kurdistan, as the Turkish official concept of Anatolia wildly goes beyond the actual Anatolia or Asia Minor peninsula into Upper Mesopotamia. Also Anatolia Peninsula was not involved, as far as we know, in the Early Neolithic and only cow domestication, which is of a later date, can be tracked to that region. The oldest known goats are from the M'lafatian culture of the Zagros (Jarmo and such). The same happens with sheep and pigs.

Goat domestication took place in the Zagros

Quickies

Genetic data seems to support the archaeological notion of domestication of the goat (also appliable to several other animals) in the Zagros mountains, roughly what we now call Kurdistan.

L. Colli, K. Lancioni et al., Whole mitochondrial genomes unveil the impact of domestication on goat matrilineal variability. BMC Genomics 2015. Open access → LINK [doi:10.1186/s12864-015-2342-2]

Abstract

Background

The current extensive use of the domestic goat (Capra hircus) is the result of its medium size and high adaptability as multiple breeds. The extent to which its genetic variability was influenced by early domestication practices is largely unknown. A common standard by which to analyze maternally-inherited variability of livestock species is through complete sequencing of the entire mitogenome (mitochondrial DNA, mtDNA).

Results

We present the first extensive survey of goat mitogenomic variability based on 84 complete sequences selected from an initial collection of 758 samples that represent 60 different breeds of C. hircus, as well as its wild sister species, bezoar (Capra aegagrus) from Iran. Our phylogenetic analyses dated the most recent common ancestor of C. hircus to ~460,000 years (ka) ago and identified five distinctive domestic haplogroups (A, B1, C1a, D1 and G). More than 90 % of goats examined were in haplogroup A. These domestic lineages are predominantly nested within C. aegagrus branches, diverged concomitantly at the interface between the Epipaleolithic and early Neolithic periods, and underwent a dramatic expansion starting from ~12–10 ka ago.

Conclusions

Domestic goat mitogenomes descended from a small number of founding haplotypes that underwent domestication after surviving the last glacial maximum in the Near Eastern refuges. All modern haplotypes A probably descended from a single (or at most a few closely related) female C. aegagrus. Zooarchaelogical data indicate that domestication first occurred in Southeastern Anatolia. Goats accompanying the first Neolithic migration waves into the Mediterranean were already characterized by two ancestral A and C variants. The ancient separation of the C branch (~130 ka ago) suggests a genetically distinct population that could have been involved in a second event of domestication. The novel diagnostic mutational motifs defined here, which distinguish wild and domestic haplogroups, could be used to understand phylogenetic relationships among modern breeds and ancient remains and to evaluate whether selection differentially affected mitochondrial genome variants during the development of economically important breeds.

Fig. 4

Spatial frequency distributions of goat mtDNA haplogroups in different geographic areas based on different datasets: modern breeds (C. hircus) a ; wild goats (C. aegagrus) b; and ancient goat remains c. See Additional file 2 (Table S5) for more information

Notice that the term "Southeastern Anatolia" is clearly being used in the Turkish imperialist ideological frame and actually must be read as Northern Kurdistan, not at all in the Anatolian Peninsula but rather Upper Mesopotamia.

Caucasus and Swiss hunter-gatherer genomes

I know I'm late for the party but better late than never, right?

A recent study sequenced three hunter-gatherer genomes from Georgia and one from Switzerland, expanding our understanding of the pre-Neolithic genetic landscape of West Eurasia.

Eppie R. Jones et al., Upper Palaeolithic genomes reveal deep roots of modern Eurasians. Nature Communications 2015. Open access → LINK [doi:10.1038/ncomms9912]

Abstract

We extend the scope of European palaeogenomics by sequencing the genomes of Late Upper Palaeolithic (13,300 years old, 1.4-fold coverage) and Mesolithic (9,700 years old, 15.4-fold) males from western Georgia in the Caucasus and a Late Upper Palaeolithic (13,700 years old, 9.5-fold) male from Switzerland. While we detect Late Palaeolithic–Mesolithic genomic continuity in both regions, we find that Caucasus hunter-gatherers (CHG) belong to a distinct ancient clade that split from western hunter-gatherers ~45 kya, shortly after the expansion of anatomically modern humans into Europe and from the ancestors of Neolithic farmers ~25 kya, around the Last Glacial Maximum. CHG genomes significantly contributed to the Yamnaya steppe herders who migrated into Europe ~3,000 BC, supporting a formative Caucasus influence on this important Early Bronze age culture. CHG left their imprint on modern populations from the Caucasus and also central and south Asia possibly marking the arrival of Indo-Aryan languages.

Figure 1: Genetic structure of ancient Europe.

 (a). Principal component analysis. Ancient data from Bichon, Kotias and Satsurblia genomes were projected¹¹ onto the first two principal components defined by selected Eurasians from the Human Origins data set¹. The percentage of variance explained by each component accompanies the titles of the axes. For context we included data from published Eurasian ancient genomes sampled from the Late Pleistocene and Holocene where at least 200 000 SNPs were called^{1, 2, 3, 4, 5, 6, 7, 9} (Supplementary Table 1). Among ancients, the early farmer and western hunter-gatherer (including Bichon) clusters are clearly identifiable, and the influence of ancient north Eurasians is discernible in the separation of eastern hunter-gatherers and the Upper Palaeolithic Siberian sample MA1. The two Caucasus hunter-gatherers occupy a distinct region of the plot suggesting a Eurasian lineage distinct from previously described ancestral components. The Yamnaya are located in an intermediate position between CHG and EHG. (b). ADMIXTURE ancestry components¹² for ancient genomes (K=17) showing a CHG component (Kotias, Satsurblia) which also segregates in in the Yamnaya and later European populations.

The Swiss one (Bichon, Jura) is maybe less of a novelty, roughly falling within the already known parameters for Western European hunter-gatherers of Magdalenian tradition (WHG in the jargon) but the three samples from the Caucasus (CHG) are really a much needed new data-point, different from everything else that what we knew and surprisingly close to modern Caucasus populations. 

They are however very distant from all other known ancient West Eurasian samples. Fig. 2 shows an estimated divergence with early Neolithic Europeans (EEF, Stuttgart) dating from before the Last Glacial Maximum, to 24,000 years ago. The divergence of this composite West Asian macro-population (EEF's Paleoeuropean admixture is accounted for separately) with the pre-Neolithic Europeans seems to be of c. 46,000 years, what is consistent with early Upper Paleolithic (the large error margin allows for a secondary Gravettian genesis contact anyhow). On the other hand the divergence between Bichon and Lochsbour seems to fit with the Magdalenian time-frame as one would expect.

CHG are surprisingly close to modern Caucasus population, particularly to Georgians. CHG also appear to be an excellent candidate population for the formation of the early Indoeuropean Yamna people, which fit best as a mix of CHG and EHG (Eastern European hunter-gatherers). 

Figure 4: The relationship of Caucasus hunter-gatherers to modern populations.

a). Genomic affinity of modern populations¹ to Kotias, quantified by the outgroup f₃-statistics of the form f₃(Kotias, modern population; Yoruba). Kotias shares the most genetic drift with populations from the Caucasus with high values also found for northern Europe and central Asia. (b). Sources of admixture into modern populations: semicircles indicate those that provide the most negative outgroup f₃ statistic for that population. Populations for which a significantly negative statistic could not be determined are marked in white. Populations for which the ancient Caucasus genomes are best ancestral approximations include those of the Southern Caucasus and interestingly, South and Central Asia. Western Europe tends to be a mix of early farmers and western/eastern hunter-gatherers while Middle Eastern genomes are described as a mix of early farmers and Africans.

I find notable that the CHG component (do not confuse with the African one of similar color) is still apparent in the Indian subcontinent, something that was already detected in other analyses. The CHG component seems to be the core of the so called "ancient North Indian" (ANI) component, also known as "Gedrosian" or "Caucaso-Baloch". What they call in the above analysis "South Asian" would be approximately the also known as "ancient South Indian" (ASI) component, which is presumably pre-Neolithic. 

"Farmer" means European Early Farmer (EEF) and already implies some Paleolithic European admixture, until we have some Levant and Mesopotamian genuine first Neolithic samples, we should not assume that all the Fertile Crescent Neolithic people were just like that, although some may have been close. In fact, I tend to think that the CHG or a similar "highlander" component was probably important in the Zagros Neolithic and consequently in the Mesopotamian and Iranian one, reaching eventually to South Asia. See here for more details on how the Neolithic expansion in Europe and India were largely parallel but not identical at all in source populations. 

To illustrate this early Neolithic complexity, still apparent to some extent in West Asian genetics and, as I just said, in European and South Asian ones, the following archaeo-cultural map should help:

Source: Eleni Asouti 2006 (red color annotation is mine)

I strongly recommend to read the full source study, because it is very informative about what were some of our ancestors¹ doing when farming and herding were being developed in West Asia, but the map above alone gives a very good glimpse of the ethno-cultural complexity of these ancient West Asian populations. 

My understading is that the mainline (Thessalian or Aegean) European Neolithic founders must have originated within the PPNA/B complex, although uncertain about which specific culture within it (most likely not Harifian because that one is surely at the origin of Semitic languages but almost any other one would do, notably those close to the Mediterranean coast: Sultanian, Aswadian and Mureybetian). Instead the populations affecting Eastern European, Mesopotamian-Iranian (Sumer and Elam) and Indian Neolithic are most probably rather linked to what is here called as M'lafatian or Zagros Neolithic, which in turn were most likely linked one way or the other to Caucasus hunter-gatherers and in general to the "highlander" West Asian element apparent in other studies in contrast to a more EEF-like "lowlander" one. 

________________________________________

¹ Sure: I'm thinking mostly of Euro-Mediterranean and Central-South Asian peoples but even if you are East Asian or Tropical African it's still very probable that some random ancestor comes from this crucial paleo-historical knot (or almost from anywhere else: admixture never ends and we are all related, even if thinly, within the last millennium or so).

The oldest known plague

New ancient DNA research has identified the bacterium Yersinia pestis, which causes the deadly and epidemic illness known as plague, in Chalcolithic and Bronze Age Central Eurasia, from Altai to Poland and the Caucasus.

S. Rasmussen, M.E. Alentoft et al., Early Divergent Strains of Yersinia pestis in Eurasia 5,000 Years Ago. Cell 2015. Open access → LINK [doi:http://dx.doi.org/10.1016/j.cell.2015.10.009]

Summary

The bacteria Yersinia pestis is the etiological agent of plague and has caused human pandemics with millions of deaths in historic times. How and when it originated remains contentious. Here, we report the oldest direct evidence of Yersinia pestis identified by ancient DNA in human teeth from Asia and Europe dating from 2,800 to 5,000 years ago. By sequencing the genomes, we find that these ancient plague strains are basal to all known Yersinia pestis. We find the origins of the Yersinia pestis lineage to be at least two times older than previous estimates. We also identify a temporal sequence of genetic changes that lead to increased virulence and the emergence of the bubonic plague. Our results show that plague infection was endemic in the human populations of Eurasia at least 3,000 years before any historical recordings of pandemics.

The bacterium was yet unable to cause the bubonic form of the plague and could not spread by means of fleas either. Instead it probably caused pneumonic and spticemic plague and was propagated by coughing and sneezing, much like the flu. 

Figure 1

Archaeological Sites of Bronze Age Yersinia pestis
(A) Map of Eurasia indicating the position, radiocarbon dated ages and associated cultures of the samples in which Y. pestis were identified. Dates are given as 95% confidence interval calendar BC years. IA: Iron Age.
(B) Burial four from Bulanovo site. Picture by Mikhail V. Khalyapin. See also Table S1.

In spite of the hype, the prevalence of the plague was low: only 7 out of 101 samples tested positive for the bacterium, ranging from c. 2800 BCE (Altai) to c. 1000 BCE (Southern Azerbaijan, Iran). A 7% prevalence is still significant but it's also obvious that 93% of the people in the period studied did not die from the plague, so let's not exaggerate, alright?

The exaggeration is already seeded in the study with passages as this one:

These early plagues may have been responsible for the suggested population declines in the late 4^th millennium BC and the early 3^rd millennium BC (Hinz et al., 2012, Shennan et al., 2013).

Neither of the referenced studies (see here and here) deals with Eastern Europe, West Asia or Central Asia, and the analyzed dates only slightly overlap with the period in which Y. pestis is found, so I fail to see the logic. It is true that there could be a coincidence in the case of Little Poland, where both Y. pestis and a demographic decline are apparent c. 2000 BCE but in all other cases it really needs a good deal of imagination to make any association.

In any case, it is clear that even the most virulent plague ever known, the Black Death, only managed to make a dent in the European demography and its consequences were not those of demographic re-expansion of the less affected populations (Polish, Basques) but a double socio-economic transition in two phases:

1. The lack of manpower in the decades after the Black Death allowed the lower classes to renegotiate their situation in various ways. It was the period in which the feudal system was most dramatically eroded, with peasants gaining control on their farmed lands and lords losing large shares of their exploitative profits, while being forced to compete against each other for whom offered best working conditions to the now scarce farmers, who, legally or not, migrated from the worst places to the ones offering better conditions. 
2. A reaction by profit-jealous landlords that largely replaced farming by husbandry, which requires less manpower. A well known case were the English enclosures, which would, slowly but steadily, set the foundations of Capitalism.

This is explained by historian, actor and director Terry Jones in the following video:

So the consequences and context of these epidemics must be considered adequately and not distorted nor simplified unduly in the line of the infamous Guns, Germs and Steel book, which greatly exaggerates the consequences of natural epidemics and is one of the favorite books of Eurocentric reactionaries with a distorted and overly simplistic view of things. One of the "virtues" of the epidemic hype is that it somehow absolves the winners from their historical crimes, blaming them on nature almost alone: instead of genocide, they use these deformations of reality to blame the mass destruction of whole nations and populations, be them Neanderthals or Native Americans, on "natural causes". 

Hypocrisy!

Instead I wonder about the real demographic impact, not yet well assessed, of the epidemics, and also about the real socio-economic consequences of such demographic declines. Of course, they could have allowed for localized migrations in the aftermath of the epidemics (or whatever other causes of demographic declines) but they should also have favored at least short-term renegotiations of the social order in favor of the suddenly scarce working classes (farmers). This "upward mobility" and partial "leveling" of what was already in many cases a feudal-like caste society should have dramatic effects in the constitution of the nations of the Bronze Age, regardless of later re-adjustments and expansions, which one can imagine as an imperfectly cyclical process.

Neolithic genomes from Northwestern Turkey

Or yet another ancient European DNA study, with some quirks and, critically, the first ancient farmer sample from the Eastern Mediterranean, specifically Northwestern Anatolia, near Yenişehir (Bursa province).

Iain Mathieson et al. Eight thousand years of natural selection in Europe. BioRxiv (pre-pub), 2015. Freely accessible → LINK [doi: http://dx.doi.org/10.1101/016477]

Abstract

The arrival of farming in Europe around 8,500 years ago necessitated adaptation to new environments, pathogens, diets, and social organizations. While indirect evidence of adaptation can be detected in patterns of genetic variation in present-day people, ancient DNA makes it possible to witness selection directly by analyzing samples from populations before, during and after adaptation events. Here we report the first genome-wide scan for selection using ancient DNA, capitalizing on the largest genome-wide dataset yet assembled: 230 West Eurasians dating to between 6500 and 1000 BCE, including 163 with newly reported data. The new samples include the first genome-wide data from the Anatolian Neolithic culture, who we show were members of the population that was the source of Europe's first farmers, and whose genetic material we extracted by focusing on the DNA-rich petrous bone. We identify genome-wide significant signatures of selection at loci associated with diet, pigmentation and immunity, and two independent episodes of selection on height.

As you can see from the title and the abstract, much of the study is focused on more or less debatable selection signatures. Interesting, of course, but not what my greatest interest, less so as I perceive that there is missing data that may be crucial for the understanding of some of such selection, notably the mainstream European LCT 13910-T allele.

How can you do an analysis of selection on this allele while ignoring the first known carriers of it: Chalcolithic (proto-)Basques and Swedes (Gökhem particularly)?

Luckily there are other highlights...

Northwestern Anatolian ancient genetics

The new ancient Neolithic samples come from two sites: Menteşe Höyük (n=5) and Barcın Höyük (n=21), both located in the Yenişehir plain, southeast of Istanbul across the Marmara Sea. The archaeological context of the samples, as well as that of many other European ones, resequenced for this study with new technology, is discussed in the Supplementary Information section.

These ancient Northwest Anatolian farmers have shown to be very similar to early European farmers. The authors estimate that these were only some 10% admixed with Paleoeuropeans, relative to the Anatolian samples, although later individuals from the West did of course had further Paleoeuropean admixture.

I must emphasize the adjective "Northwestern" because Anatolia Peninsula is a large territory where the Neolithic had differential implementation in time and cultures. Critically we cannot be certain that there is any identity between these Western Anatolian first farmers and those from South-Central Anatolia, for example those of the world-famous Çatal Höyük village. This is because the Neolithic of South-Central Anatolia is much older and there are archaeological indications that suggest that the settlement of Western Anatolia and Greece took place via coastal migration. The origin of this coastal migration probably involved Cyprus, which in turn was more directly related to the Neolithic of the Levant (PPNB) than to that of South-Central Anatolia. Some genetic data also seem to suggest that the precursors of early European farmers were from the Levant, rather than from further North. But of course the full resolution of this mystery will have to await for ancient DNA from the relevant regions, something that may be aided by the recent technological breakthroughs but that will also require peace, so geneticists and archaeologists can do their field work (there are of course many other much more excruciating reasons to hope for peace and normalization in West Asia, naturally, don't get me wrong).

In any case we finally have a reference genome for what can be termed the Aegean Neolithic and it seems it was even closer to European derivatives. We cannot anyhow discard that there was some backflow from Greece or other parts of the Balcans to Western Anatolia because there was indeed some interaction across the Aegean. However a much more clear cut cultural divide has been argued to exist between the cultures of the Marmara Sea and those of inland Thrace, so, if there was any such backflow, it probably happened before the expansion of Thessalian Neolithic northwards.

Principal Component Analysis

This is the Principal Component Analysis provided by this study (fig. 1B). The modern samples are in gray with no labeling whatsoever but I guess most readers will approximately identify them easily, as the basic layout has been repeated in so many recent aDNA studies:

Figure 1: Population relationships of samples. (...) B: Principal component analysis of 777 modern West Eurasian samples (grey), with 221 ancient samples projected onto the first two principal component axes and labeled by culture. Abbreviations: [E/M/L]N Early/Middle/Late Neolithic, LBK Linearbandkeramik, [E/W]HG Eastern/Western hunter-gatherer, [E]BA [Early] Bronze Age, IA Iron Age.

As in Olalde 2015 or Haak 2015, or even Lazaridis 2014, WHGs appear located rather "towards the South", unlike in some other PCAs, particularly Europe-only ones. I do find this to be interesting and potentially informative, at least while we await for Atlantic European ancient nuclear DNA.

Therefore, you'll forgive me for the redundancy of reusing the above image a couple of times in order to make a couple of points.

The thesis that most of these studies are pushing for is a simplistic triangular scenario for the formation of modern Europeans with a formula that can be described as {x.EEF+y.WHG+z.Kurgan}. I don't deny that this is quite approximative but I am also quite certain that it is missing important clues. In fact, the triangular thesis seems to fail to explain most Northern European genetic makeup, while the origins of Basques also remain somewhat unexplained by it. Let's see:

Annotations on the PCA: triangular thesis fails, extra HG (EHG?) is needed.

It would seem quite apparent that the triangular thesis (described on the PCA by a slashed line) fails to explain most of Western and Northern European genetic makeup, which is clearly much more deviated towards Paleoeuropean hunter-gatherers than it allows.

Just including on the equation Eastern European hunter-gatherers (dotted line) would be enough to solve most of the problem, although it does of course arises other questions about how and when this extra Paleoeuropean blood was incorporated.

This solution would still leave Basques outside of it. It requires instead of a Western hunter-gatherer extra admixture on top of a simple Neolithic cluster basis:

Annotations on the PCA: Basques can be explained (?) as simple {Neo-European + WHG admixture}

Of course that the actual sources of Paleoeuropean admixture can be more complex, as suggested by some studies, like Günther & Valdiosera 2015, who claimed Scandinavian HG admixture not just in Gökhem farmers but also in Ötzi ("Iceman" in the above graph). These did not use EHG samples but in any case, if correct, it is a pre-Kurgan admixture from the Northeast of the subcontinent.

A key excerpt from the Supplementary Information 2 section that someone (Simon, I think) used to argue for steppe ancestry in Basques in a discussion at Eurogenes blog:

The Iberian Chalcolithic population lacks steppe ancestry, but Late Neolithic central and northern Europeans have substantial such ancestry (Extended Data Fig. 3E) suggesting that the spread of ANE/steppe ancestry did not occur simultaneously across Europe. All presentday Europeans have less steppe ancestry than the Corded Ware5, suggesting that this ancestry was diluted as the earliest descendants of the steppe migrants admixed with local populations. However, the statistic f4(Basque, Iberia_Chalcolithic; Yamnaya_Samara,Chimp)=0.00168 is significantly positive (Z=8.1), as is the statistic f4(Spanish, Iberia_Chalcolithic; Yamnaya_Samara, Chimp)=0.00092 (Z=4.6). This indicates that steppe ancestry occurs in present-day southwestern European populations, and that even the Basques cannot be considered as mixtures of early farmers and hunter-gatherers without it4.

What does this say in fact? It says nothing about Western Hunter-Gatherers, only that Basques appear as more Yamna-like than the Iberian Chalcolithic sample. I see no reason why this cannot be caused by simple extra WHG admixture, although it can also imply other Paleoeuropean such as SHG or EHG inflow. What I do see from other studies (and again for all I can discern in this one) is that Basques do lack any clear Yamna signature and notably their Caucasus or Northern West Asian component (always present where Kurgan admixture is unmistakable and therefore a clear indicator of it) is effectively zero (some individuals may have tiny non-zero such component, all very normal).

Admixture analysis with two and three source populations

The authors find that, while many populations can be modeled as product of simple two-way admixture, many need a three-way model, notably from the late Chalcolithic onwards:

Extended Data Figure 2: Early isolation and later admixture between farmers and steppe populations. A [actually B]: Mainland European populations later than 3000 BCE are better modeled with steppe ancestry as a 3rd ancestral population. B [actually A]: Later (post-Poltavka) steppe populations are better modeled with Anatolian Neolithic as a 3rd ancestral population. C: Estimated mixture proportions of mainland European populations without steppe ancestry. D: Estimated mixture proportions of Eurasian steppe populations without Anatolian Neolithic ancestry. E: Estimated mixture proportions of later populations with both steppe and Anatolian Neolithic ancestry. [F is below]

However this varies, because notably the Iberian Chalcolithic sample can still be modeled as a two-way admixture, what is in conformity with the consideration that the increase in the complexity took place not in any single event but rather first in Central and Eastern Europe and only later further West. This is in full conformity with the Kurgan model of Indoeuropean expansion, although it may require some refinement here and there.

For example it is becoming quite obvious that there was not only a westward movement of Eastern European populations but also a subsequent eastward backflow of the resulting admixed Central European ones. This is discussed in the supplementary materials, from page 43 onwards.

Extended Data Figure 2: (...) F: ADMIXTURE plot at k=17 showing population differences over time and space.
(click to expand)

To this I must add my conviction that the triangular model is not enough to actually explain modern European genetics and that greater Paleoeuropean genetic input in Northern and Western Europe is required as well. The great challenge in this regard is to sample Atlantic (and Baltic) Europe properly and extract whatever consequences that ancient genomes from these areas may provide. 

Naturally there is also some other research to be done in West Asia, where a good deal of the European (and also West Asian, naturally) ancestors lived once upon a time. That is the other major challenge. In this sense this study must be commended for its breakthrough in sampling ancient Northwestern Anatolians, which is a step in the right direction.

There are other blank zones to be researched as well in Southern Europe (Italy, Balcans) that may well provide complementary information.

Alleged selection

The authors claim to have found evidence for selection in twelve different alleles. I remain mildly skeptic because it is hard to judge if this was all selection or founder effect was involved as well. 

Some of the alleged targets of selection are:

Lactase persistance: rs4988235, also known as 13910-T, already mentioned above. The authors mention that the allele’s earliest appearance in our data is in a central European Bell Beaker sample (individual I0112) that lived between approximately 2300 and 2200 BCE. Older signals from the Chalcolithic Basque Country (fixated in a subpopulation) and Sweden are totally ignored. Of course it is a draft so far but it is clear that key information, widely available, is being ignored.

A light skin allele known as rs16891982 (in the gene SLC45A2). This allele was low in the studied ancient populations (but again might have been higher in the blank under-researched areas, I can't say). Unlike it, the derived allele of gene SLC24A5, was fixated in Neolithic NW Anatolians, as well as derived European ancient populations, being a clear case of founder effect (although it may have also helped with adaption to the low vitamin D diet caused by transition to agriculture). There are other pigmentation genes that may have been selected in complex interaction, as they seem to be partly correlated with latitude and are hard to explain based on ancient populations alone.

An important datum here is that: unlike closely related western hunter-gatherers, the Motala samples have predominantly derived pigmentation alleles at SLC45A2 and SLC24A5. So... is there another source of these light skin alleles (there are others and much is unknown anyhow) that is not from Neolithic farmers?

Another selection target is in the TLR1-TLR6-TLR10 gene cluster, which seems related to resistance to mycobacteria such as those causing leprosy, tuberculosis, etc. Regarding this complex cluster, I rather quote:

The strongest signal is at rs2269424 near the genes PPT2 and EGFL8 but there are at least six other apparently independent signals in the MHC (Extended Data Fig. 3); and the entire region is significantly more associated than the genome-wide average (residual inflation of 2.07 in the region on chromosome 6 between 29-34 Mb after genome-wide genomic control correction). This could be the result of multiple sweeps, balancing selection, or background selection in this gene-rich region.

The EDAR gene, related to tooth morphology (remember Pippi?) and hair thickness, as well denser sweat and mammary glands, in East Asians is also listed. Curiously enough, half of the Motala individuals (Epipaleolithic Sweden), carried the derived allele of rs3827760. Modern Scandinavians often have this derived allele, although the authors believe that it is because of more recent admixture:

The EDAR derived allele is largely absent in present-day Europe except in Scandinavia, plausibly due to Siberian movements into the region millennia after the date of the Motala samples.

Uh, really? How can you be so sure? I am very skeptic here again and would rather suspect a more complex pattern of partial Paleolithic (or at least Epipaleolithic) continuity, which may indeed have been brought from East Asia with the proto-Uralic migrations (or whatever). 

Another trait for which the authors claim selection is what they call "genetic height", i.e. height not measured from the actual individuals but from alleles that are believed to influence it. They argue for selection for lower height in Neolithic and Chalcolithic Iberia and for greater height in the Steppe instead, both being corrected to greater height in modern populations. Without objective measures to control for the assumed "genetic height", among other reasons, I find the whole story a bit hard to believe but who knows?

The production of this entry took a whole 8 hours working journey, including a half hour break but not preliminary reading and related discussions. If you liked it and have some coin to spare, consider donating. Thank you.

Vasco-Nubian?

This is something I've been chewing on for more than a year now and yet never got myself to blog about (although I have mentioned in private or in comments here and there). Impelled by the minor but quite apparent NE African influence, genetic and cultural, on the Neolithic peoples of the Levant, whose offshoots eventually landed in Greece triggering the European Neolithic, I decided in the Spring of 2014 to explore, via mass-lexical comparison, if Basque language (and by extension the wider Vasconic family, which I believe now to be that of mainline European Neolithic) might have any relation with Nubian languages. I did not expect to find anything but noise but to my surprise the number of apparent cognates is quite significant. 

My primary analysis was this one but now I have combined it with a comparison with Proto-Indoeuropean (PIE), which is also very probably related to the roots of Vasconic: LINK (open office spreadsheet). 

The synthesis is as follows:

Of course the "cognates" are only apparent cognates at this stage of the research and the evaluation is necessarily subjective. But judge yourselves. 

If we discard the "weak" apparent cognates, the vocabulary correlation between Basque and Nubian and between Basque and PIE is pretty similar. But, in my understanding, both are well above the noise threshold, an example of which could be the PIE-Nubian apparent cognates, which are many many less. 

I must say anyhow that the oblique apparent cognates, that is when one word sounds much not like its strict synonym but a related one (for example words meaning hot and fire), look all very solid and most intriguing. 

Also, when attributing probabilities to origins of Basque words, Nubian appears to be at the origin of almost double the words (26%) that can be attributed to PIE (15%). Of course, for lack of data or because they actually have other origins, the unknown origins apply to the majority of words (56%), double than the Nubian origin ones.

However Nubian here is constituted of three different languages (Dilling, Nobiin and Midob), while PIE is just a single theoretical construct. This last must be done this way because many modern and historical IE languages, notably in Europe, have other Vasconic substrate influences, which must be studied separately from general PIE-Vasconic shared vocabulary. This kind of late Vasconic influence is very much unlikely in the case of Nubian instead. In any case I don't know of any a proto-Nubian Swadesh list readily available. 

Finally I must mention that because the PDF format is horrible for copy-pasting, I chose to re-transcribe the Nubian words according to my best approximation using a normal keyboard (not always the same characters that the original list uses).

Strongest Basque-Nubian apparent simple cognates

• Basque - Nubian languages (English)
  
• azal - àzì, àzzì-di (bark)
• haragi - árízh (meat)
• odol - ógór, èggér (blood)
• buru - úr (head)
• oin - ó:y (foot)
• esku - ish-i, ès-sì (hand)
• hil* - di-ìl (to die)
• euri - are, ara, áwwí, áré, árí, áró (rain)
• harri - kugor, kakar (stone) [notice also the pre-IE root *kharr- speculated to be at the origin of Karst, etc.] 
• lur - gùr (soil, ground)
• haize - irsh-i, éss-í (wind)

There are some others that are shared with Indo-European and with similar subjective "weight", not listing them here to keep things clear. There are also other apparent cognates that are arguably less clear like bat - be (one) that I'm also skipping here but you can find in the spreadsheet.

*Hil (meaning both to die and to kill in Basque, which can't be confused because they conjugate differently) seems ancestral to English ill and kill (this one via a Germanic precursor).

The intriguing oblique cognates

Notice that these words do not mean the same, yet their meanings seem strikingly related.

• Nubian (English) - Basque (English)
• hor, koy, kà:r (tree) - harri (stone) [notice that zuhaitz (tree) can be interpreted etymologically as zur-haitz = wood-rock, so the relation is not that weird]
• ok-i, og (breast) - ogi (bread)
• a-l (heart) - ahal (can (verb), potential, power)
• azh, àz-ír, àzza (to bite) - (h)ortz (tooth), aitz (rock, peak) [some argue that originally "to cut", present in many cutting tool names: aizkor = axe, aitzur = hoe, aizto = knife, etc.*]
• shu, zhúù (to walk) - joan (to go) [often pronounced jun or shun]
• é:zhi (water) - heze (wet) [also archaic particle *iz-, meaning "water" by all accounts: itxaso = sea, izurde = dolphin, izotz = ice, and common in Vasconic river toponymy]
• zhuge (to burn) - su (fire) 
• zhùg, sù, sú:w (hot) - su (fire)
• úr-i, úrúm (black) - urdin (blue) [archaic also green, grey]

*This one is an obvious and very prevalent Vasconic substrate infiltrator in Western IE languages: axe, adze, azada (hoe in Spanish), etc.

How can this be possible?

It is of course a mere working hypothesis and ultimately you judge but I find it hard to disdain. However there is no apparent connection, notably no significant genetic connection, between Basques and Nubians. So how can we explain this?

I have it reasonably clear myself, so I made a map to explain it:

Basque is after all just the last survivor of a once much larger family (Vasconic), a family that most likely corresponds to the languages spoken by the early European farmers (mainline Neolithic of Aegean roots). As that expansion was largely done in about a mere thousand years, I estimate that when both branches met near the Rhine, the two peoples could still understand each other, even if with some difficulty. Only the Southern/Western branch(-es) survived long enough to leave historical evidence, so it is hard to guess how the Northern branch evolved anyhow.

The Nubian linguistic connection is anyhow not the only thing that requires the Levant or Palestinian Neolithic step, also Y-DNA E1b-M78 (mostly V13 in Europe, attested in some early farmers and still very important among Greeks and Albanians particularly) and probably the so-called "Basal Eurasian" component that Lazaridis detected among early European farmers and that could well be the signature of African genetics from the Nile.

Linguistically, also the very notorious presence of Semitic (an Afroasiatic branch) in West Asia is surely another legacy of the same African influences in the Mesolithic Levant. Before this research, I thought it was the only one but now I strongly suspect that at some point Nubian (Nilo-Saharan) languages were also present in the region. Maybe one (Nubian evolving towards Vasconic) corresponded to Natufian proper and the other (proto-Semitic) to Harifian, the semi-desert pastoralist facies of the same wider culture. Can't say for sure.

The chain was once long but now only some of the most distant links remain unbroken. It is difficult to imagine that they were ever connected at all...

To do...

A lot remains to be done, of course:

• These mass lexical comparisons only apply to a few families in the region and the rest should also be tested for. My energies are limited and so are my qualifications as "linguist", so I encourage others, hopefully more energetic and knowledgeable, to expand.
• Grammatical features cannot be analyzed by this methodology. Again my means are limited. 
• Anthropological research would be an interesting complement. So far the only shared cultural trait I could spot would be the use of bells attached to ankles for dancing but there could be others. 
• ...