Interesting excerpts (emphasis mine, edited for clarity):
On the high frequency of R1b-V88
Our genome-wide data allowed us to assign Y haplogroups for 25 ancient Sardinian individuals. More than half of them consist of R1b-V88 (n=10) or I2-M223 (n=7).
Francalacci et al. (2013) identified three major Sardinia-specific founder clades based on present-day variation within the haplogroups I2-M26, G2-L91 and R1b-V88, and here we found each of those broader haplogroups in at least one ancient Sardinian individual. Two major present-day Sardinian haplogroups, R1b-M269 and E-M215, are absent.
Compared to other Neolithic and present-day European populations, the number of identified R1b-V88 carriers is relatively high.
(…)ancient Sardinian mtDNA haplotypes belong almost exclusively to macro-haplogroups HV (n = 16), JT (n = 17) and U (n = 9), a composition broadly similar to other European Neolithic populations.
On the origin of a Vasconic-like Paleosardo with the Western EEF
(…) the Neolithic (and also later) ancient Sardinian individuals sit between early Neolithic Iberian and later Copper Age Iberian populations, roughly on an axis that differentiates WHG and EEF populations and embedded in a cluster that additionally includes Neolithic British individuals. This result is also evident in terms of absolute genetic differentiation, with low pairwise FST ~ 0.005 +- 0.002 between Neolithic Sardinian individuals and Neolithic western mainland European populations. Pairwise outgroup-f3 analysis shows a very similar pattern, with the highest values of f3 (i.e. most shared drift) being with Neolithic and Copper Age Iberia, gradually dropping off for temporally and geographically distant populations.
In explicit admixture models (using qpAdm, see Methods) the southern French Neolithic individuals (France-N) are the most consistent with being a single source for Neolithic Sardinia (p ~ 0:074 to reject the model of one population being the direct source of the other); followed by other populations associated with the western Mediterranean Neolithic Cardial Ware expansion.
Pervasive Western Hunter-Gatherer ancestry in Iberian/French/Sardinian population
Similar to western European Neolithic and central European Late Neolithic populations, ancient Sardinian individuals are shifted towards WHG individuals in the top two PCs relative to early Neolithic Anatolians Admixture analysis using qpAdm infers that ancient Sardinian individuals harbour HG ancestry (~ 17%) that is higher than early Neolithic mainland populations (including Iberia, ~ 8%), but lower than Copper Age Iberians (~ 25%) and about the same as Southern French Middle-Neolithic individuals (~ 21%).
Continuity from Sardinia Neolithic through the Nuragic
We found several lines of evidence supporting genetic continuity from the Sardinian Neolithic into the Bronze Age and Nuragic times. Importantly, we observed low genetic differentiation between ancient Sardinian individuals from various time periods.
A qpAdm analysis, which is based on simultaneously testing f-statistics with a number of outgroups and adjusts for correlations, cannot reject a model of Neolithic Sardinian individuals being a direct predecessor of Nuragic Sardinian individuals (…) Our qpAdm analysis further shows that the WHG ancestry proportion, in a model of admixture with Neolithic Anatolia, remains stable at ~17% throughout three ancient time-periods.
Steppe influx in Modern Sardinians
While contemporary Sardinian individuals show the highest affinity towards EEF-associated populations among all of the modern populations, they also display membership with other clusters (Fig. 5). In contrast to ancient Sardinian individuals, present-day Sardinian individuals carry a modest “Steppe-like” ancestry component (but generally less than continental present-day European populations), and an appreciable broadly “eastern Mediterranean” ancestry component (also inferred at a high fraction in other present-day Mediterranean populations, such as Sicily and Greece).
From the Bronze Age (~2200–900 BCE), we increase the available dataset from 7 to 60 individuals and show how ancestry from the Pontic-Caspian steppe (Steppe ancestry) appeared throughout Iberia in this period, albeit with less impact in the south. The earliest evidence is in 14 individuals dated to ~2500–2000 BCE who coexisted with local people without Steppe ancestry. These groups lived in close proximity and admixed to form the Bronze Age population after 2000 BCE with ~40% ancestry from incoming groups. Y-chromosome turnover was even more pronounced, as the lineages common in Copper Age Iberia (I2, G2, and H) were almost completely replaced by one lineage, R1b-M269.
The arrival of East Bell Beakers speaking Indo-European languages involved, nevertheless, the survival of the two non-IE communities isolated from each other – likely stemming from south-western France and south-eastern Iberia – thanks to a long-lasting process of migration and admixture. There are some common misconceptions about ancient languages in Iberia which may have caused some wrong interpretations of the data in the paper and elsewhere:
Iberian languages were spoken at least in the Mediterranean and the south (ca. “1/3 of Iberia“) during the Bronze Age.
Nope, we only know the approximate location of Iberian culture and inscriptions from the Late Iron Age, and they occupy the south-eastern and eastern coastal areas, but before that it is unclear where they were spoken. In fact, it seems evident now that the arrival of Urnfield groups from the north marks the arrival of Celtic-speaking peoples, as we can infer from the increase in Central European admixture, while the expansion of anthropomorphic stelae from the north-west must have marked the expansion of Lusitanian.
Vasconic was spoken in both sides of the Pyrenees, as it was in the Middle Ages.
Wrong. One of the worst mistakes I am seeing in many comments since the paper was published, although admittedly the paper goes around this problem talking about “Modern Basques”. Vasconic toponyms appear south of the Pyrenees only after the Roman conquests, and tribes of the south-western Pyrenees and Cantabrian regions were likely Celtic-speaking peoples. Aquitanians (north of the western Pyrenees) are the only known ancient Vasconic-speaking population in proto-historic times, ergo the arrival of Bell Beakers in Iberia was most likely accompanied by Indo-European languages which were later replaced by Celtic expanding from Central Europe, and Iberian expanding from south-east Iberia, and only later with Latin and Vasconic.
Ligurian is non-Indo-European, and Lusitanian is Celtic-like, so Iberia must have been mostly non-Indo-European-speaking.
The fragmentary material available on Ligurian is enough to show that phonetically it is a NWIE dialect of non-Celtic, non-Italic nature, much like Lusitanian; that is, unless you follow laryngeals up to Celtic or Italic, in which case you can argue anything about this or any other IE language, as people who reconstruct laryngeals for Baltic in the common era do.
a Palaeo-European language (as Villar puts it) expanded into most regions of Iberia in ancient times (he considered at some point the Mesolithic, but that is obviously wrong, as we know now); then
Celts expanded at least to the Ebro River Basin; then
Iberians expanded to the north and replaced these in NE Iberia; and only then
after the Roman invasion, around the start of the Common Era, appear Vasconic toponyms south of the Pyrenees.
Lusitanian obviously does not qualify as Celtic, lacking the most essential traits that define Celticness…Unless you define “(Para-)Celtic” as Pre-Proto-Celtic-like, or anything of the sort to support some Atlantic continuity, in which case you can also argue that Pre-Italic or Pre-Germanic are Celtic, because you would be essentially describing North-West Indo-European…
If Basques have R1b, it’s because of a culture of “matrilocality” as opposed to the “patrilocality” of Indo-Europeans
So wrong it hurts my eyes every time I read this. Not only does matrilocality in a regional group have few known effects in genetics, but there are many well-documented cases of population replacement (with either ancestry or Y-DNA haplogroups, or both) without language replacement, without a need to resort to “matrilineality” or “matrilocality” or any other cultural difference in any of these cases.
In fact, it seems quite likely now that isolated ancient peoples north of the Pyrenees will show a gradual replacement of surviving I2a lineages by neighbouring R1b, while early Iberian R1b-DF27 lineages are associated with Lusitanians, and later incoming R1b-DF27 lineages (apart from other haplogroups) are most likely associated with incoming Celts, which must have remained in north-central and central-east European groups.
NOTE. Notice how R1a is fully absent from all known early Indo-European peoples to date, whether Iberian IE, British IE, Italic, or Greek. The absence of R1a in Iberia after the arrival of Celts is even more telling of the origin of expanding Celts in Central Europe.
I haven’t had enough time to add Iberian samples to my spreadsheet, and hence neither to the ASoSaH texts nor maps/PCAs (and I don’t plan to, because it’s more efficient for me to add both, Asian and Iberian samples, at the same time), but luckily Maciamo has summed it up on Eupedia. Or, graphically depicted in the paper for the southeast:
Does this continued influx of Y-DNA haplogroups in Iberia with different cultures represent permanent changes in language? Are, therefore, modern Iberian languages derived from Lusitanian, Sorothaptic/Celtic, Greek, Phoenician, East or West Germanic, Hebrew, Berber, or Arabic languages? Obviously not. Same with Italy (see the recent preprint on modern Italians by Raveane et al. 2018), with France, with Germany, or with Greece.
If that happens in European regions with a known ancient history, why would the recent expansions and bottlenecks of R1b in modern Basques (or N1c around the Baltic, or R1a in Slavs) in the Middle Ages represent an ancestral language surviving into modern times?
The new huge sampling of Sintashta – combined with that of Catacomb, Poltavka, Potapovka, Andronovo, and Srubna – shows quite clearly how this long-term admixture process between Uralic peoples and Indo-Iranians happened between forest-steppe CWC (mainly Abashevo) and steppe groups. The situation is not different from that of Iberia ca. 2500-2000 BC; from Narasimhan, Patterson, et al. (2018):
We combined the newly reported data from Kamennyi Ambar 5 with previously reported data from the Sintashta 5 individuals (10). We observed a main cluster of Sintashta individuals that was similar to Srubnaya, Potapovka, and Andronovo in being well modeled as a mixture of Yamnaya-related and Anatolian Neolithic (European agriculturalist-related) ancestry.
As with Iberia (or any prehistoric region), the details of how exactly this language change happened are not evident, but we only need a plausible explanation coupled with archaeology and linguistics. Poltavka, Potapovka, and Sintashta samples – like the few available Iberian ones ca. 2500-2000 BC – offer a good picture of the cohabitation of R1b-L23 (mainly Z2103) and R1a-Z645 (mainly Z93+): a glimpse at the likely presence of R1a-Z93 within settlements – which must have evolved as the dominant elites – in a society where the majority of the population was initially formed by nomad herders (probably most R1b-Z2103), who were usually buried outside of the main settlements.
Will the upcoming Narasimhan, Patterson et al. (2019) deal with this problem of how R1a-M417 replaced R1b-M269, and how the so-called “Steppe_MLBA” (i.e. Corded Ware) ancestry admixed with “Steppe_EMBA” (i.e. Yamnaya) ancestry in the steppes, and which one of their languages survived in the region (that is, the same the Reich Lab has done with Iberia)? Not likely. The ‘genetic wars’ in Iberia deal with haplogroup R1b-P312, and how it was neither ‘native’ nor associated with Basques and non-Indo-European peoples in general. The ‘genetic wars’ in South Asia are concerned with the steppe origin of R1a, to prove that it is not a ‘native’ haplogroup to India, and thus neither are Indo-Aryan languages. To each region a politically correct account of genetic finds, with enough care not to fully dismiss national myths, it seems.
NOTE. Funnily enough, these ‘genetic wars’ are the making of geneticists since the 1990s and 2000s, so we are still in the midst of mostly internal wars caused by what they write. Just as genetic papers of the 2020s will most likely be a reaction to what they are writing right now about “steppe ancestry” and R1a. You won’t find much change to the linguistic reconstruction in this whole period, except for the most multicolored glottochronological proposals…
The first author of the paper has engaged, as far as I could see in Twitter, in dialogue with Hindu nationalists who try to dismiss the arrival of steppe ancestry and R1a into South Asia as inconclusive (to support the potential origin of Sanskrit millennia ago in the Indus Valley Civilization). How can geneticists deal with the real problem here (the original ethnolinguistic group expanding with Corded Ware), when they have to fend off anti-steppists from Europe and Asia? How can they do it, when they themselves are part of the same societies that demand a politically correct presentation of data?
This is how the data on the most likely Indo-Iranian-speaking region should be presented in an ideal world, where – as in the Iberia paper – geneticists would look closely to the Volga-Ural region to discover what happened with Proto-Indo-Iranians from their earliest to their latest stage, instead of constantly looking for sites close to the Indus Valley to demonstrate who knows what about modern Indian culture:
Iberian cultures, already with a majority of R1b lineages, show a clear northward expansion over previously Urnfield-like groups of north-east Iberia and Mediterranean France (which we now know probably represent the migration of Celts from central Europe). Similarly, Eastern Balts already under a majority of R1a lineages expanded likely into the Baltic region at the same time as the outlier from Turlojiškė (ca. 1075 BC), which represents the first obvious contacts of central-east Europe with the Baltic.
Iberia shows a more recent influx of central and eastern Mediterranean peoples, one of which eventually succeeded in imposing their language in Western Europe: Romans were possibly associated mainly with R1b-U152, apart from many other lineages. Proto-Slavs probably expanded later than Celts, too, connected to the disintegration of the Lusatian culture, and they were at some point associated with R1a-M458 and R1a-Z280(xZ92) lineages, apart from others already found in Early Slavs.
This parallel between Iberia and eastern Europe is no coincidence: as Europe entered the Bronze Age, chiefdom-based systems became common, and thus the connection of ancestry or haplogroups with ethnolinguistic groups became weaker.
What happened earlier (and who may represent the Pre-Balto-Slavic community) will be clearer when we have enough eastern European samples, but basically we will be able to depict this admixture of NWIE-speaking BBC-derived peoples with Uralic-speaking CWC-derived groups (since Uralic is known to have strongly influenced Balto-Slavic), similar to the admixture found in Indo-Iranians, more or less like this:
The Early Scythian period marked a still stronger chiefdom-based system which promoted the creation of alliances and federation-like groups, with an earlier representation of the system expanding from north-eastern Europe around the Baltic Sea, precisely during the spread of Akozino warrior-traders (in turn related to the Scythian influence in the forest-steppes), who are the most likely ancestors of most N1c-V29 lineages among modern Germanic, Balto-Slavic, and Volga-Finnic peoples.
Modern haplogroup+language = ancient ones?
It is not difficult to realize, then, that the complex modern genetic picture in Eastern Europe and around the Urals, and also in South Asia (like that of the Aegean or Anatolia) is similar to the Iron Age / medieval Iberian one, and that following modern R1a as an Indo-European marker just because some modern Indo-European-speaking groups showed it was always a flawed methodology; as flawed as following R1b for ancient Vasconic groups, or N1c for ancient Uralic groups.
Why people would argue that haplogroups mean continuity (e.g. R1b with Basques, N1c with Finns, R1a with Slavs, etc.) may be understood, if one lives still in the 2000s. Just like why one would argue that Corded Ware is Indo-European, because of Gimbutas’ huge influence since the 1960s with her myth of “Kurgan peoples”. Not many denied these haplogroup associations, because there was no reason to do it, and those who did usually aligned with a defense of descriptive archaeology.
However, it is a growing paradox that some people interested in genetics today would now, after the Iberian paper, need to:
accept that ancient Iberians and probably Aquitanians (each from different regions, and probably from different “Basque-Iberian dialects” in the Chalcolithic, if both were actually related) show eventually expansions with R1b-L23, the haplogroup most obviously associated with expanding Indo-Europeans;
acknowledge that modern Iberians have many different lineages derived from prehistoric or historic peoples (Celts, Phoenicians, Greeks, Romans, Jews, Goths, Berbers, Arabs), which have undergone different bottlenecks, the last ones during the Reconquista, but none of their languages have survived;
realize that a similar picture is to be found everywhere in central and western Europe since the first proto-historic records, with language replacement in spite of genetic continuity, such as the British Isles (and R1b-L21 continuity) after the arrival of Celts, Romans, Anglo-Saxons, Vikings, or Normans;
but, at the same time, continue blindly asserting that haplogroup R1a + “steppe ancestry” represent some kind of supernatural combination which must show continuity with their modern Indo-Iranian or Balto-Slavic language from time immemorial.
Behave, pretty please
The ‘conservative’ message espoused by some geneticists and amateur genealogists here is basically as follows:
Let’s not rush to new theories that contradict the 2000s, lest some people get offended by granddaddy not being these pure whatever wherever as they believed, and let’s wait some 5, 10, or 20 years, as long as necessary – to see if some corner of the Yamna culture shows R1a, or some region in north-eastern Europe shows N1c, or some Atlantic Chalcolithic sample shows R1b – to challenge our preferred theories, if we actually need to challenge anything at all, because it hurts too much.
Just don’t let many of these genetic genealogists or academics of our time be unhappy, pretty please with sugar on top, and let them slowly adapt to reality with more and more pet theories to fit everything together (past theories + present data), so maybe when all of them are gone, within 50 or 70 years, society can smoothly begin to move on and propose something closer to reality, but always as politically correct as possible for the next generations.
For starters, let’s discuss now (yet again) that Bell Beakers may not have been Indo-European at all, despite showing (unlike Corded Ware) clearly Yamna male lineages and ancestry, because then Corded Ware and R1a could not have been Indo-European and that’s terrible, so maybe Bell Beakers are too brachycephalic to speak Indo-European or something, or they were stopped by the Fearsome Tisza River, or they are not pure Dutch Single Grave in The South hence not Indo-European, or whatever, and that’s why Iron Age Iberians or Etruscans show non-Indo-European languages. That’s not disrespectful to the history of certain peoples, of course not, but talking about the evident R1a-Uralic connection is, because this is The South, not The North, and respect works differently there.
Just don’t talk about how Slavs and Balts enter history more than 1,500 years later than Indo-European peoples in Western and Southern Europe, including Iberia, and assume a heroic continuity of Balts and Slavs as pure R1a ‘steppe-like’ peoples dominating over thousands of kms. in the Baltic, Fennoscandia, eastern Europe, and northern Asia for 5,000 years, with multiple Balto-Slavs-over-Balto-Slavs migrations, because these absolute units of Indo-European peoples were a trip and a half. They are the Asterix and Obelix of white Indo-European prehistory.
Perhaps in the meantime we can also invent some new glottochronological dialectal scheme that fits the expansion of Sredni Stog/Corded Ware with (Germano-?)Indo-Slavonic separated earlier than any other Late PIE dialect; and Finno-Volgaic later than any other Uralic dialect, in the Middle Ages, with N1c.
To sum up: Iberia, Italy, France, the British Isles, central Europe, the Balkans, the Aegean, or Anatolia, all these territories can have a complex history of periodic admixture and language replacement everywhere, but some peoples appearing later than all others in the historical record (viz. Basques or Slavs) apparently cannot, because that would be shameful for their national or ethnic myths, and these should be respected.
Ignorance of the own past as a blank canvas to be filled in with stupid ethnolinguistic continuity, turned into something valuable that should not be challenged. Ethnonationalist-like reasoning proper of the 19th century. How can our times be called ‘modern’ when this kind of magical thinking is still prevalent, even among supposedly well-educated people?
Our analysis of divergence times suggests the population lineage ancestral to modern-day Sardinia was effectively isolated from the mainland European populations ~140–250 generations ago, corresponding to ~4,300–7,000 years ago assuming a generation time of 30 years and a mutation rate of 1.25 × 10−8 per basepair per generation. (…) in terms of relative values, the divergence time between Northern and Southern Europeans is much more recent than either is to Sardinia, signaling the relative isolation of Sardinia from mainland Europe.
We documented fine-scale variation in the ancient population ancestry proportions across the island. The most remote and interior areas of Sardinia—the Gennargentu massif covering the central and eastern regions, including the present-day province of Ogliastra— are thought to have been the least exposed to contact with outside populations. We found that pre-Neolithic hunter-gatherer and Neolithic farmer ancestries are enriched in this region of isolation. Under the premise that Ogliastra has been more buffered from recent immigration to the island, one interpretation of the result is that the early populations of Sardinia were an admixture of the two ancestries, rather than the pre-Neolithic ancestry arriving via later migrations from the mainland. Such admixture could have occurred principally on the island or on the mainland before the hypothesized Neolithic era influx to the island. Under the alternative premise that Ogliastra is simply a highly isolated region that has differentiated within Sardinia due to genetic drift, the result would be interpreted as genetic drift leading to a structured pattern of pre-Neolithic ancestry across the island, in an overall background of high Neolithic ancestry.
We found Sardinians show a signal of shared ancestry with the Basque in terms of the outgroup f3 shared-drift statistics. This is consistent with long-held arguments of a connection between the two populations, including claims of Basque-like, non-Indo-European words among Sardinian placenames. More recently, the Basque have been shown to be enriched for Neolithic farmer ancestry and Indo-European languages have been associated with steppe population expansions in the post-Neolithic Bronze Age. These results support a model in which Sardinians and the Basque may both retain a legacy of pre-Indo-European Neolithic ancestry. To be cautious, while it seems unlikely, we cannot exclude that the genetic similarity between the Basque and Sardinians is due to an unsampled pre-Neolithic population that has affinities with the Neolithic representatives analyzed here.
While we can confirm that Sardinians principally have Neolithic ancestry on the autosomes, the high frequency of two Y-chromosome haplogroups (I2a1a1 at ~39% and R1b1a2 at ~18%) that are not typically affiliated with Neolithic ancestry is one challenge to this model. Whether these haplogroups rose in frequency due to extensive genetic drift and/or reflect sex-biased demographic processes has been an open question. Our analysis of X chromosome versus autosome diversity suggests a smaller effective size for males, which can arise due to multiple processes, including polygyny, patrilineal inheritance rules, or transmission of reproductive success. We also find that the genetic ancestry enriched in Sardinia is more prevalent on the X chromosome than the autosome, suggesting that male lineages may more rapidly trace back to the mainland. Considering that the R1b1a2 haplogroup may be associated with post-Neolithic steppe ancestry expansions in Europe, and the recent timeframe when the R1b1a2 lineages expanded in Sardinia, the patterns raise the possibility of recent male-biased steppe ancestry migration to Sardinia, as has been reported among mainland Europeans at large (though see Lazaridis and Reich and Goldberg et al.). Such a recent influx is difficult to square with the overall divergence of Sardinian populations observed here.
Once again, haplogroup R1b1a2 (M269), and only R1b1a2, related to male-biased, steppe-related Indo-European migrations…just sayin’.
NOTE. In fact, the increase in Neolithic ancestry found in south-west Ireland with expanding Bell Beakers (likely Proto-Beakers), coupled with the finding of I2a subclades in Megalithic cultures of western Europe, would support this replacement after the Cardial and Epi-Cardial expansions, which were initially associated with G2a lineages.
I am not convinced about a survival of Palaeo-Sardo after the Bell Beaker expansion, though, since there is no clear-cut cultural divide (and posterior continuity) of pre-Beaker archaeological cultures after the arrival of Bell Beakers in the island that could be identified with the survival of Neolithic languages.
We may have to wait for ancient DNA to show a potential expansion of Neolithic ancestry from the west, maybe associated with the emergence of the Nuragic civilization (potentially linked with contemporaneous Megalithic cultures in Corsica and in the Balearic Islands, and thus with an Iberian rather than a Basque stock), although this is quite speculative at this moment in linguistic, archaeological, and genetic terms.
Nevertheless, it seems that the association of a Basque-Iberian language with the Neolithic expansion from Anatolia (see Villar’s latest book on the subject) is somehow strengthened by this paper. However, it is unclear when, how, and where expanding G2a subclades were replaced by native I2 lineages.
Blevins (to appear) presents a new reconstruction of Proto-Basque, the mother of Basque and Aquitanian, based on standard methods in historical linguistics: the comparative method and the method of internal reconstruction. Where all previous reconstructions of Proto-Basque assume a contrast between two sibilants, *s, a voiceless apical sibilant, and *z a voiceless laminal sibilant (Martinet 1955; Michelena 1977; Lakarra 1995; Trask 1997), this proposal is unique in positing only a single sibilant *s. Under this account, all instances of Common Basque /z/ are derived from *s. More specifically, in syllable coda, *Rs > *Rz (R a sonorant) while in the syllable onset, *sT > *zT (T an oral stop). The true split of *s into /s/ vs. /z/ occurs when clusters like *Rz or *zT are further simplified to /z/.
In this talk, internal evidence for a single sibilant, *s, in Proto-Basque is presented, and sound changes underlying the sibilant split are examined within the context of Evolutionary Phonology (Blevins 2004, 2006, 2015, 2017). Similar sound changes are identified in other languages with similar cluster types (e.g. Kümmel 2007:232), and the phonetic basis of the sibilant split is informed by recent studies of sibilant retraction (e.g. Stevens and Harrington 2016; Stuart-Smith et al. 2018).
I was never convinced, not just about a genetic connection, but about the very possibility of discovering it if there was any, mainly because such a link would be quite old, and Basque is known to have been greatly influenced by surrounding IE prestige languages for millennia until it was first attested in the 16th century. Internal reconstruction can only avail a gross reconstruction of few aspects up to a certain point in time, probably not very far beyond the Pre-Roman period, and that only thanks to the available Aquitanian inscriptions.
This haplogroup is also found in Sardinia, which may be connected to the expansion of V88 subclades (which I have speculatively proposed could be linked to Afro-Asiatic) into Africa through Italy and the Green Sahara; although it could also be linked to a speculative Vasconic-Iberian – Palaeo-Sardo group.
Without knowing the exact Pre-Proto-Indo-European stage at which Blevins would place the Basque separation, it is difficult to know how it could fit within any macro-language proposal – and thus potential ancestral population expansion.
Seeing how many Basque nationalists (obviously obsessed with racial purity) are still rooting for an autochthonous Palaeolithic origin of R1b lineages (especially P312) linked with the Basque language and dat huge Vasconic Western Europe; and now, after Olalde & Mathieson 2018, how some are also suggesting a Neolithic link of R1b with the Neolithic expansion and Sardinians, for lack of further modern genetic differences with other Western Europeans… I wonder how a lot of people inclined to believe this nonsense today, and mentally linking Vasconic with haplogroup R1b, will be paradoxically necessarily tied precisely to this kind of macro-family proposals in the future.