By sequencing 727 ancient individuals from the Southern Arc (Anatolia and its neighbors in Southeastern Europe and West Asia) over 10,000 years, we contextualize its Chalcolithic period and Bronze Age (about 5000 to 1000 BCE), when extensive gene flow entangled it with the Eurasian steppe. Two streams of migration transmitted Caucasus and Anatolian/Levantine ancestry northward, and the Yamnaya pastoralists, formed on the steppe, then spread southward into the Balkans and across the Caucasus into Armenia, where they left numerous patrilineal descendants. Anatolia was transformed by intra–West Asian gene flow, with negligible impact of the later Yamnaya migrations. This contrasts with all other regions where Indo-European languages were spoken, suggesting that the homeland of the Indo-Anatolian language family was in West Asia, with only secondary dispersals of non-Anatolian Indo-Europeans from the steppe.
Literary and archaeological sources have preserved a rich history of Southern Europe and West Asia since the Bronze Age that can be complemented by genetics. Mycenaean period elites in Greece did not differ from the general population and included both people with some steppe ancestry and others, like the Griffin Warrior, without it. Similarly, people in the central area of the Urartian Kingdom around Lake Van lacked the steppe ancestry characteristic of the kingdom’s northern provinces. Anatolia exhibited extraordinary continuity down to the Roman and Byzantine periods, with its people serving as the demographic core of much of the Roman Empire, including the city of Rome itself. During medieval times, migrations associated with Slavic and Turkic speakers profoundly affected the region.
We present the first ancient DNA data from the Pre-Pottery Neolithic of Mesopotamia (Southeastern Turkey and Northern Iraq), Cyprus, and the Northwestern Zagros, along with the first data from Neolithic Armenia. We show that these and neighboring populations were formed through admixture of pre-Neolithic sources related to Anatolian, Caucasus, and Levantine hunter-gatherers, forming a Neolithic continuum of ancestry mirroring the geography of West Asia. By analyzing Pre-Pottery and Pottery Neolithic populations of Anatolia, we show that the former were derived from admixture between Mesopotamian-related and local Epipaleolithic-related sources, but the latter experienced additional Levantine-related gene flow, thus documenting at least two pulses of migration from the Fertile Crescent heartland to the early farmers of Anatolia.
I haven’t read the supplements, so no major comment from me, except for one: the Greece-focused paper confirms using phenotypic prediction that West Eurasians have been getting lighter-complected since the late Neolithic/Bronze Age. I have no idea why, but some Nazis are offended by this reality and cherry-pick data, but trust me, I open up all the supplements to look at the HIRIS-plex predictions.
The origin, development, and legacy of the enigmatic Etruscan civilization from the central region of the Italian peninsula known as Etruria have been debated for centuries. Here we report a genomic time transect of 82 individuals spanning almost two millennia (800 BCE to 1000 CE) across Etruria and southern Italy. During the Iron Age, we detect a component of Indo-European–associated steppe ancestry and the lack of recent Anatolian-related admixture among the putative non–Indo-European–speaking Etruscans. Despite comprising diverse individuals of central European, northern African, and Near Eastern ancestry, the local gene pool is largely maintained across the first millennium BCE. This drastically changes during the Roman Imperial period where we report an abrupt population-wide shift to ~50% admixture with eastern Mediterranean ancestry. Last, we identify northern European components appearing in central Italy during the Early Middle Ages, which thus formed the genetic landscape of present-day Italian populations.
The inference is basically what I concluded and stated this spring in my piece on ancient Rome:
By 1000 BC, on the eve of the Iron Age, a mixed ancestry derived from both Anatolian farmers and Indo-Europeans from the steppe was present all across continental Italy. Only on isolated islands such as Sardinia was the situation different, where Anatolian farmer societies continued unperturbed by Indo-European migrants, giving rise to the Nuragic civilization, with its massive fortifications. All the length of the Italian mainland, the ancestors of the Etruscans, Sabines, and Romans shared roughly the same genetic heritage, despite their ethno-linguistic differences.
The big novel find is that there is nontrivial Germanic ancestral shift after the fall of Rome. Some commenters have suggested this is actually just from rural Italians, who flooded back into the towns. This could probably be tested.
Modern humans appeared in Europe by at least 45,000 years ago but the extent of their interactions with Neanderthals, who disappeared by about 40,000 years ago, and their relationship to the broader expansion of modern humans outside Africa are poorly understood. Here we present genome-wide data from three individuals dated to between 45,930 and 42,580 years ago from Bacho Kiro Cave, Bulgaria. They are the earliest Late Pleistocene modern humans known to have been recovered in Europe so far, and were found in association with an Initial Upper Palaeolithic artefact assemblage. Unlike two previously studied individuals of similar ages from Romania and Siberia who did not contribute detectably to later populations, these individuals are more closely related to present-day and ancient populations in East Asia and the Americas than to later west Eurasian populations. This indicates that they belonged to a modern human migration into Europe that was not previously known from the genetic record, and provides evidence that there was at least some continuity between the earliest modern humans in Europe and later people in Eurasia. Moreover, we find that all three individuals had Neanderthal ancestors a few generations back in their family history, confirming that the first European modern humans mixed with Neanderthals and suggesting that such mixing could have been common.
There are two primary points of interest:
More evidence of ubiquitous ‘admixture’ between Neanderthals and ‘modern humans’ who may have been in contact with them.
These earliest European modern humans in Southeast Europe seem to be more closely related to (possibly ancestral to?) people in East Asia than modern Europeans.
On the first point, twenty years ago there were some paleoanthropologists who were arguing there was no admixture between Neanderthals and modern humans. That is, no fertile offspring. To not pussyfoot around it, Neanderthals were barely acknowledged as human in the early years of this century by many scholars. This is evident in Richard Klein’s The Dawn of Human Culture, written in 2002. With hindsight much of this is ridiculous. It really didn’t make any sense that the Neanderthal and modern human lineage were not inter-fertile when you dug into the literature on mammalian hybridization. There just wasn’t that much time for them to be totally incompatible.
In 2010 when it was found that Neanderthal ancestry seems to be found in non-Africans, we updated many of our priors. I think it is clear on some level Neanderthal humanization is driven by the fact of Neanderthal admixture. Nevertheless, there was a plausible case this admixture was rare. Perhaps a single Neanderthal tribe was mixed into expanding modern humans? Perhaps a single Neanderthal? These were ideas that were mooted.
With what we know now I’m not sure this is tenable. There are two primary issues. First, there is variation in Neanderthal ancestry among non-African populations which does not seem to be due to African admixture. East Asians have more Neanderthal admixture (~25% more last I checked) than Europeans who have more than West Asians. South Asian Neanderthal admixture is proportional to their distance from East Asians. The less West Eurasian the South Asian, the more Neanderthal.
There are a few ways to understand this. A simple explanation is that there was a secondary admixture event that impacted people migrating to eastern Eurasia. Another hypothesis is that natural selection reduced the Neanderthal fraction over time, but East Eurasians were less impacted by this due to small effective population size. Then, there is the idea that a non-African population, “Basal Eurasians”, that did not have much Neanderthal admixture later mixed into West Asians and Europeans, reducing the original Neanderthal fraction.
Because of the small fractions and the paucity of ancient DNA more than 10,000 years old, there are still arguments around this. These results increase the probability that there were multiple Neanderthal admixture events. At least in my way of thinking.
The authors found that the Neanderthal fraction seems to have decreased from a higher level relatively early on. Rather than a gradual decrease, the authors found suggestions of strong effective selection in the first few generations. Though there is still room for the Basal Eurasian model, I’m not sure it’s quite as necessary now. Along with Oase, and the existence of a Neanderthal-Denisovan hybrid from Denisova cave, the likelihood of frequent admixture between ancient hominin lineages seems pretty high.
I believe the primary issue at this point is that the admixture event seems to be dated to a narrow time period, before the extinction of Neanderthals, but not too ancient (before 60,000 years ago), and the Neanderthal ancestry is quite similar. Perhaps there isn’t the power to detect multiple secondary admixtures at “around the same time.” From the perspective of today, 45,000 v. 50,000 years ago is “around the same time.” But the reality is 5,000 years is a long time.
In simulations, we find that estimates of the mean time of admixture are largely robust to details in gene flow models. In contrast, the duration of the gene flow is much more difficult to recover, except under ideal circumstances where gene flow is recent or the exact recombination rate is known. We conclude that gene flow from Neandertals into modern humans could have happened over hundreds of generations. Ancient genomes from the time around the admixture event are thus likely required to resolve the question when, where, and for how long humans and Neandertals interacted.
The second major finding of this paper is that the very first modern humans to settle in Europe are more genetically close to East Asians than to modern Europeans. This is in contrast to the Oase sample from Romania, just to the north, and a few thousand years later, that was no closer to East or West Eurasians. Or, a new 45,000-year-old sample from Czech Republic, which also shows no connection to any modern people.
If you read the first paper, you know that years ago the GoyetQ116-1 sample from Belgium that dates to 35,000 years before the present, and which seems to have some ancestral connection to the later Magdalenian people of Pleistocene Europe, had an East Eurasian affinity. Additionally, earlier mtDNA work showed eastern affinities in some Pleistocene Europeans. The authors above make the connection between these eastern-affinity early Europeans and the “Initial Upper Paleolithic” (IUP), which seems to have expanded across a broad swath of Eurasian, from Europe all the way to western Mongolia.
The implication then is that some of the ancestry of East Eurasians comes from a migration that took a route through Europe, and around the Black Sea littoral. This is a bit more complicated than a pure “southern route”, but it’s not crazy, and has long been proposed.
I think at this point we need to take a step back, and acknowledge that the period between 40 and 60 thousand years ago is important, but we look through the glass darkly. Something happened, as all the ancient Eurasian hominin lineages were absorbed by the “Out of Africa” population (which may not have been expanding out of Africa!), and the phylogenetic relationships of some of these people do not make sense in light of the phylogeography of the present.
It is entirely feasible to me that the ancient non-African or proto-non-African populations had already started to develop some internal structure. The putative Basal Eurasian v. “the rest” is one key bifurcation, but there may already have been divisions between the western and eastern branches of Eurasians in northeast Africa or the Near East. We just don’t know yet. Basically, ancient substructure that is getting “blown up” with rapid radiation.
In 2014 when BEu was created as a construct to explain the greater affinities of Mesolithic European hunter-gatherers with East Eurasians than modern Europeans*, I probably would have been a little surprised that a mostly BEu individual had still not been discovered in the ancient DNA. After all, we had seen Ma’lta boy answer the question of who the mysterious “Siberians” were that left an imprint all over West Eurasia. Ma’lta was part of an ancient Paleo-Siberian group, the Ancient North Eurasians (ANE), who no longer exist in “pure” form, but contributed ancestry (or more precisely were related to groups that did so) to hunter-gatherers in Eastern Europe and the ancestors of New World populations.
No such luck with BEu. In fact, BEu as a construct is so vaguely understood that research groups can estimate that populations are 10% BEu, or 40% BEu, contingent upon various specifications in their model. In the middle 2010’s one of the scientists involved in the groups working in this space and using BEu as a construct even told me that this population may never have existed in pure form anyhow (not Iosif Lazaridis to be clear).
Basically, there is still not a lot of clarity. BEu seems to have diverged before Neanderthal admixture into the non-African lineage ~55,000 years ago. It also seems to have been subject to the long bottleneck of all non-Africans. A very plausible model that BEu occupied the southern Middle East, while non-basal Eurasians occupied the northern Middle East, seem to be the highest probability to me. The authors of this preprint argue that the Basal Eurasian ur-heimat might be in and around the Persian Gulf. I laughed when I read that because it reminds me of the early 20th century “Lost Civilization Underwater” motif. But it could be true.
I would hazard to guess dates, but I think the separation and later (early) admixture of BEu and non-BEu people in the Near East is probably strongly conditional on the paleoclimate data, which I am not fluent in.
* BEu ancestry came into Europe with Neolithic farmers, so affinities between Mesolithic hunter-gatherers and East Asians is higher than between modern Europeans and East Asians.
The Middle and Late Bronze Age Near East, a period roughly spanning the second millennium BC (ca. 2000-1200 BC), is frequently referred to as the first ‘international age’, characterized by intense and far-reaching contacts between different entities from the eastern Mediterranean to the Near East and beyond. In a large-scale tandem study of stable isotopes and ancient DNA of individuals excavated at Tell Atchana (Alalakh), situated in the northern Levant, we explore the role of mobility at the capital of a regional kingdom. We generated strontium isotope data for 53 individuals, oxygen isotope data for 77 individuals, and added ancient DNA data from 9 new individuals to a recently published dataset of 28 individuals. A dataset like this, from a single site in the Near East, is thus far unparalleled in terms of both its breadth and depth, providing the opportunity to simultaneously obtain an in-depth view of individual mobility and also broader demographic insights into the resident population. The DNA data reveals a very homogeneous gene pool, with only one outlier. This picture of an overwhelmingly local ancestry is consistent with the evidence of local upbringing in most of the individuals indicated by the isotopic data, where only five were found to be ‘non-local’. High levels of contact, trade, and exchange of ideas and goods in the Middle and Late Bronze Ages, therefore, seem not to have translated into high levels of individual mobility detectable at Tell Atchana.
From a DNA perspective two notable things about this preprint. First, it confirms that there was a massive pulse of Iranian/Caucasus ancestry into the western Fertile Crescent between 5000 and 2000 BC. We don’t have any idea what was going on here, but my own suspicion is that the Uruk period, 4000 to 3100 BC, has something to do with this genetic turnover and assimilation. We don’t know what happened during the Uruk period because there’s no real writing of narrative history (there is proto-cuneiform), but this is when city life really expanded in Mesopotamia. Additionally, there were replica copies of Mesopotamian style towns to the west, and even into Anatolia. The Uruk period was arguably the peak of Mesopotamian political power and influence before the rise of Assyria thousands of years later.
The end of the Uruk period was characterized by a massive collapse. Some archaeologists hypothesize that the catastrophe literature of the Sumerians may reflect memories of the end of the Uruk civilization. In some ways, the Akkadians and Sumerians may have lived in the shadows of their forebears.
The second issue is “the lady in the well”. This is a woman who seems to have been pushed into a well and fallen to her death. She is the major genetic outlier from this site: “Individual ALA019 – the Well Lady – takes up an extreme outlier position in the PCA closest to sampled individuals from Bronze Age Iran/Turkmenistan/Uzbekistan/Afghanistan.”
Her positioning is unambiguous. Unless we’re missing something this woman is of “steppe” heritage. Probably Indo-Iranian. But, the best guess from the isotope work is that she was raised in the region. The data they have indicate she lived between 1550-1600 BC. She was almost certainly from an Indo-Iranian group that had settled in the region. In fact, she was almost certainly associated with the Indo-European element in Mitanni. But in this region of the world the Dasa in their walled cities overcame the free people and swallowed them up culturally and demographically.
The major lacunae in this paper, and all such papers, is Iraq. There is ancient DNA from Turkey, the Levant, but what about Iraq? I’m sure people are working on it, but this is a region that would give us a better sense of “donor” populations.
Upward Sun River 1, an individual from a unique burial of the Denali tradition in Alaska (11500 calBP), is considered a type representative of Ancient Beringians who split from other First Americans 22000-18000 calBP in Beringia. Using a new admixture graph model-comparison approach resistant to overfitting, we show that Ancient Beringians do not form the deepest American lineage, but instead harbor ancestry from a lineage more closely related to northern North Americans than to southern North Americans. Ancient Beringians also harbor substantial admixture from a lineage that did not contribute to other Native Americans: Amur River Basin populations represented by a newly reported site in northeastern China. Relying on these results, we propose a new model for the genomic formation of First American ancestors in Asia.
Read the preprint. I’ll note three things
– The authors suggested that differentiation between American native lineages occurred in eastern Siberia, not Beringia. In other words, there’s a lot of ancient structure in the New World that dates to the Last Glacial Maximum
– Speaking of ancient structure, using the new methods in this paper they detect more pervasive “Australo-Melanesian” ancestry in lower quality ancient remains. It seems clear ta this point that this isn’t about Australo-Melanesian, as much as genetic variation in East Eurasia during the Pleistocene that we have a poor grasp of at this point
– In Europe ancient DNA quickly converged on the trihybrid model of Mesolithic hunter-gatherers, and early farmers, and steppe pastoralists. The tsunami of ancient DNA has fleshed out a lot of details, but the basic framework has been in place for five years. The situation in other parts of the world seems to be more unsettled and dynamic. More sophisticated models and more ancient DNA keeps returning surprise on the margin
The power of ancient DNA in terms of human evolution at this point is to a large extent the ability to understand the arc of human cultural history as reflected in our genealogies. Archaeologists have long attempted to infer aspects of social and cultural practice from material remains. Now, geneticists are getting into this game, with mixed results.
Today the Bradley lab in Ireland published a paper about the genetics of Neolithic Ireland (i.e., before the arrival of Bell Beakers ~2,500 BC). The first author is Lara Cassidy, who I interviewed last year. There is a reason this paper is in Nature.
Let me quote the abstract first:
…The scale and sophistication of megalithic architecture along the Atlantic seaboard, culminating in the great passage tomb complexes, is particularly impressive…Although co-operative ideology has often been emphasized as a driver of megalith construction…the human expenditure required to erect the largest monuments has led some researchers to emphasize hierarchy…Here we present evidence that a social stratum of this type was established during the Neolithic period in Ireland. We sampled 44 whole genomes, among which we identify the adult son of a first-degree incestuous union from remains that were discovered within the most elaborate recess of the Newgrange passage tomb. Socially sanctioned matings of this nature are very rare, and are documented almost exclusively among politico-religious elites—specifically within polygynous and patrilineal royal families that are headed by god-king…We identify relatives of this individual within two other major complexes of passage tombs 150 km to the west of Newgrange, as well as dietary differences and fine-scale haplotypic structure (which is unprecedented in resolution for a prehistoric population) between passage tomb samples and the larger dataset, which together imply hierarchy. This elite emerged against a backdrop of rapid maritime colonization that displaced a unique Mesolithic isolate population, although we also detected rare Irish hunter-gatherer introgression within the Neolithic population.
There’s a lot of moving parts in this research, and the most interesting element isn’t the genetics, but the social structure that you can infer from the genetics. It seems entirely likely that the “Megalithic civilization” of Atlantic Europe was hierarchal. But this pretty much confirms it. As noted in the paper violations of first-order incest taboos as a cultural norm (as opposed to deviancy) are strongly associated with very stratified preliterate or semiliterate societies (with the possible exception of Zoroastrianism). Additionally, this research highlights that a set of individuals, likely paternally related, seem to be enriched in elite burials.
In a few circles, there are ideas that Neolithic Europeans were peaceful and matrilineal. The existence of stratification like this and the likelihood of ‘god-kings’ makes that very unlikely in Ireland. Though there was no doubt some variation in Neolithic Europe, the existence of “long-houses” in Germany from contemporary cultures is ominous. The matrilineal element is distinct. There are matrilineal societies that are quite warlike (e.g., the Nairs of Kerala or the Iroquois). But the possibility of common Y chromosomes suggests that this was a patrilineal society, which is, on the whole, more common anthropologically.
To me this is the most awesome part of the paper:
The Brú na Bóinne passage tombs appear in Medieval mythology that relates their construction to magical manipulations of the solar cycle by a tribe of gods, which has led to unresolved speculation about the durability of oral traditions across millennia…Although such longevity seems unlikely, our results strongly resonate with mythology that was first recorded in the eleventh century AD, in which a builder-king restarts the daily solar cycle by copulating with his sister…Fertae Chuile, a Middle Irish placename for the Dowth passage tomb (which neighbours Newgrange), is based on this lore, and can be translated as ‘Hill of Sin’ or ‘Hill of Incest’…
This is incredible. Unless you are set in your ways I think it is hard to deny that the medieval Irish were passing on a recollection in their myth from an encounter between Bell Beakers and the late descendants of the Newgrange people. In The Isles Norman Davies argues that the Irish, unlike the English and the British more generally (Brythonic), kept their own mythology, and so have a sense of their past in a way that is uncommon among Northern European peoples. The Irish legends imply that there were multiple waves of people, and it is assumed that the people who live in the great mounds are the Tuatha Dé Danann, who became ancient Irish demigods.
I suspect that the early Bell Beakers viewed the monuments of the Tuatha Dé Danann, the Newgrange people, like how some Americans view “Indian graveyards.” Even among modern people, there is superstition, so what can we expect from the ancients? The Greeks forgot their heritage and assumed that the cyclopean citadels of their ancestors were built by giants. No doubt agro-pastoralist Bell Beakers looked at the massive ruins, and perceived the work of the gods.
And we shouldn’t underestimate the ability of oral tradition to recall events for premodern people. Mount Mazama blew 7,700 years ago, but native people in the area have legends of that explosion. Australian Aboriginals have myths that clearly outline and detail landmarks that are now underwater due to rising sea levels. The point is that it’s plausible that facts that are 3,000 years old could persist down to Christian Ireland, and be recorded by priests.
There are lots more I could say about this paper in the details. They found an infant with Down syndrome! They show recent introgression of hunter-gatherer ancestry into some Neolithic individuals. Also, they show hunter-gatherer substructure (Irish hunter-gatherers do not have ancestry from Magdelanian populations, only Villabruna). But the biggest aspect here is that this paper now sets a standard for how you can synthesize ancient DNA with archaeology and mythology.
In the East and South Asian groups, the data suggest a very recent arrival of Denisovan DNA (mainly <15,000 YBP). In non-Africans, Neanderthal sharing remains high for branches with lower-end age younger than ~30,000 YBP. These dates are only lower bounds on the introgression time, and an accurate arrival date of Neanderthal DNA would require estimating a joint genealogy that requires further work….
Comparing YRI, GBR, BEB (Bengali in Bangladesh) and CHB to expectations under panmixia, we observe a strong excess of mutations on deep branches with lower coalescence age <40,000 YBP in all cases, which is almost entirely explained by Neanderthals/Denisovans in the non-African populations, but not in YRI (Fig. 4d, Methods). In panmictic simulations with matched population size histories, we observed no such excess (Supplementary Fig. 6). This gives evidence for ancient but uncharacterized population structure within Africa, as suggested elsewhere…Figure 4b shows one example consistent with an introgression event in YRI, not involving a closely relative of Neanderthals.
The inference of deep structure within Africa is not a great surprise. But note that their estimate for Denisovan admixture is that on the whole, it could be half as old as Neanderthal admixture. I don’t put much stock in the specific estimate of the date, but rather the relative time.
In the 1970s A. J. Ammerman and L. L. Cavalli-Sforza argued for the validity of a model of Neolithic expansion of farmers into Europe predicated on a “demic diffusion” dynamic. This is in contrast to the idea that farming spread through the diffusion of ideas, not people. The formal theory is inspired by the Fisher wave model, but empirically just imagine two populations with very different carrying capacities due to their mode of production, farmers, and hunter-gatherers. In a Malthusian framework, the farmer carrying capacity in a given area of land might be ~10× greater than that of hunter-gatherers. Starting at the same initial population, the farmers will simply breed the hunter-gatherers out of existence.
As the farmers reaching their local carrying capacity, migration outward will occur in a continuous and diffusive process. For all practical purposes, the farmers will perceive the landscape occupied by hunter-gatherers as “empty.” This is due to the fact that hunter-gatherers often engage in extensive, not intensive, exploitation of resources. In contrast, even slash and burn agriculturalists leave a much bigger ecological footprint. They swarm over the land.
The beauty of the demic diffusion process is that that it’s analytically elegant and tractable. Families or villages engaged in primary production to “fill up” a landscape through simple cultural practices which manifest on the individual scale that allow for aggregate endogenous growth. And this model underlies much of the work by Peter Bellwood in First Farmers and Colin Renfrew’s theories about the spread of Indo-European langauges. You can call it the Walder Frey theory of history.
I didn’t really think deeply about this theory because I didn’t have much empirical knowledge until I read Lawrence Keeley’s War Before Civilization. In this book, Keeley observes that the archaeological record suggests that there was violent conflict between the first farmers and hunter-gatherers in northwestern Europe, near the North Sea. He reports that there seems to have been a broad front of conflict, presumably a prehistoric “no man’s land.” Not only that, but Keeley claims that the spread of agriculture stopped for a period. The barrier between hunter-gatherer occupation and farmer territory was not permeable. Not diffusion.
As a stylized fact, the demic diffusion framework treats all farmers as interchangeable and all hunter-gatherers as interchangeable. On the face of it, we know that this is wrong. But the assumption is that to a first approximation this axiom will allow us to capture the main features of the dynamics in question. This may be a false assumption. The fact is we know that some hunting and gathering populations can engage in intensive resource extraction and remain sedentary.
Most “old hands” in the discipline of historical population genetics remember when grand narratives were constructed out of Y chromosomal haplogroup distributions. One of the most distinctive ones is that of haplogroup R1b, which exhibits very high frequencies in the west of Europe, as high as more than 80% among the Basques. Because the Basques are the only non-Indo-European population which exists today in Western Europe, it was presumed that they are more ancient than other groups. And, their high frequency of R1b (along with other peculiarities such as a high frequency of Rh-), was taken to indicate that they reflected the genetics of Europe’s aboriginal hunter-gatherers when farming arrived.
This turned out to be wrong in a lot of details. Genetically the Basques are quite like the European farmers from Anatolia who replaced the original hunter-gatherers. Less so than the Sardinians, as they have more hunter-gatherer ancestry. But instead of being the language of European hunter-gatherers, it seems plausible that the Basque language descends from that of the Cardial culture.
There’s a lot of moving parts in this research, and the most interesting element isn’t the genetics, but the social structure that you can infer from the genetics. It seems entirely likely that the “Megalithic civilization” of Atlantic Europe was hierarchal. But this pretty much confirms it. As noted in the paper violations of first-order incest taboos as a cultural norm (as opposed to deviancy) are strongly associated with very stratified preliterate or semiliterate societies (with the possible exception of 
