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Category Archives: Sardinia

Ancient Italian ape had human-like precission grip

Reconstruction of O. bamboli (Pavel Major / ICP)
Oreopithecus bamboli was primate species, surely a hominine (great ape excluding orangutans) that lived in Tuscany and Sardinia some 8.2-6.7 million years ago.
It has great interest regarding human evolution because it is the oldest known ape to have developed a pad-to-pad precision grip, a characteristic otherwise only found in the human genus.
This trait, hotly debated in the last decades, has been recently confirmed by researchers of the Catalan Institute of Paleontology Miquel Crusafont (ICP). It must be said however that this development is considered convergent evolution and not ancestral to our own precision grip.
O. bamboli fossil
(CC by Ghedoghedo)
I guess that much of the controversy is caused by the old hypothesis that argued that it was the precision grip itself which elicited human brain development, something that obviously did not happen with Oreopithecus.
Other traits of this species are quite different from our own or our australopithecine relatives. They probably walked upright but with different gait (unlike the more human-like Sahelanthropus, of similar age) and their feet were very much unlike ours, with a very open angle for the big toe (hallux).
It seems that their environment was swampy and not strictly forestal.
Sources[es/cat/en]: Pileta, Diari de Girona, Wikipedia.
Ref.: Sergio Almécija et al., The morphology of Oreopithecus bambolii pollical distal phalanx. AJPA 2014. Pay per viewLINK [doi:10.1002/ajpa.22458]
 

Ancient DNA from Eastern Europe and Sardinia

A very interesting doctoral thesis has been known these days (h/t Jean). The thesis by Clio S. I. Dersarkissian (directed by A. Cooper and W. Haak) includes novel ancient mtDNA from North Eastern Europe (Karelia and surroundings) specially and also some Scythian and Sardinian burials from the Metal Ages.
Clio Simone Irmgard Dersarkissian, Mitochondrial DNA in ancient human populations of Europe. University of Adelaide, 2011 (thesis). Freely accessible ··> LINK [identifier: http://hdl.handle.net/2440/74221%5D
The most interesting findings may be those from Karelia:
  • First pre-Neolithic mtDNA H in Northern and Eastern Europe and one of the few findings strongly confirmed in such haplogroup before Neolithic. It clearly reinforces the already well established notion that mtDNA H existed in Europe before the Neolithic.
  • U2e – which might well be descendant or otherwise related to the U2 of Kostenki.
  • C1 – suggesting pre-Neolithic Siberian influences in Northern and Eastern Europe. The specific sublineage (named as “C1f”) has not yet been sequenced elsewhere.
There are some more interesting data regarding ancient NE Europeans, Scythians and Sardinians but let’s see that by parts.

Epipaleolithic peoples from Karelia and Northern Russia

Possibly the most impacting findings of this paper are those regarding two Epipaleolithic sites in Karelia (Uznyi Oleni Ostrov) and nearby parts of Northern Russia (Popovo, in Russia proper but not far from the Karelian border), as well as one more recent site from Sápmi (Lapland).

As I mentioned above, the U2e and C1 (“C1f”) findings are unusual and suggestive of ancestral connections with Kostenki (Early Upper Paleolithic site from Southern Russia with U2 mtDNA) and Central Asia and Siberia. In fact an overall comparison with modern populations, shows strong affinities with West Siberians and Uyghurs for these Epipaleolithic Karelians.
Instead the Bronze Age Sami site shows more generic or distributed Siberian affinities, although there are populations in West Siberia (Nenets?) that also fit well with that mtDNA genetic pool. Bashkirs show similar affinity to both ancient populations (see ch. 1, fig. 3 – p. 103).
Not shown here are the results for the 18th century Sami site of Chalmny-Varre, which look a very modern Sami mtDNA pool, dominated by V7e and complemented by U5b1b1 and U5a1. 

Confirming the existence of mtDNA H in pre-Neolithic Europe

I really want to underline this, because certain influential people have been dead set into denying the existence of mtDNA haplogroup H in Europe altogether before the Neolithic. Why? Because they have a theory (a hypothesis more properly speaking) and they can’t accept to be wrong about it.
That hypothesis (very popular in some circles) states that European aboriginal hunter-gatherers were very radically annihilated by Neolithic invaders from West Asia (never mind that archaeology alone is much more complicated than that, they don’t seem to like thinking too much, much less looking at the matter from all the angles).
And a central battle they have fought is denying the possibility that mtDNA H (he most common haplogroup today in Western Europe) existed in the continent before Neolithic. The whole haplogroup, in their imaginary reality, could only have arrived with the industrious (and seemingly quite genocidal) farmers from West Asia (who almost never even mixed with anyone aborigine, how odd).
Reality began questioning their findings since 2005 but back in the day only HVS-I or at best HVS-II (control regions of the mtDNA chain) were used, leading to inconclusive results, specially in regards to short-stemmed haplogroup H. So they could still deny and deny…
But, recently, two different new studies have found unmistakable mtDNA H in Magdalenian people from Cantabria and Epipaleolithic people from the Basque Country. The reaction of some such knowledgeable aficionados has been simply unbelievable: they have flatly rejected the results without any reason; these findings are simply too inconvenient truths for their conjectures to be accepted. They are so obsessed with their fantasies that they can’t even accept mounting evidence against them: they have stopped being scientific and begun being fanatics.
Very sad, really.
This finding in Karelia adds to the mounting unquestionable evidence on the matter: mtDNA haplogroup H not only existed in pre-Neolithic Eruope but it was quite extended, roughly through the areas in which is today abundant (and not just SW Europe as I came to suspect for some time). However in most regions was still far less common than it is today (or even totally missing, as seems to be the case in Central Europe).
Said that, it is not too clear yet where does all the improved knowledge of ancient genetics lead us to but what is clear is that mtDNA H is older and specifically older-in-Europe than some (too many) people have been insisting on.
Also it seems more and more obvious that the popular Neolithic farmers did not define the modern genetic landscape of Europe at all. They certainly introduced lineages that surely did not exist before but their overall influence seems limited and it does look like, after an initial burst, they declined also quite abruptly.
This is something that has been in the news these days (but no paper yet) and that I observed also in 2009 in relation to some similar studies (see: here and here). The age that we begin seeing modern-like mtDNA pools actually varies a lot, for example:
  • SW Europe: Basque Country: Neolithic (at least) ··> Hervella 2009 (discussed here).
  • Central Europe: Elbe Basin: Bronze Age or Chalcolithic ··> Schilz 2006[de], Schweitzer 2008.
  • Far North Europe: Sápmi: some time after the Bronze Age and before the 18th century (this study).
  • Central Asia: Iron Age (see below).
I conjecture here that (before the Medieval agricultural revolution) Northern latitudes could in general support lower population densities, being also more susceptible to the effect of climatic fluctuations. But more data is needed before we can have some consolidated certainty.
In any case, I took some time to make a couple of updated maps of the European and North African (1) Late Upper Paleolithic (Magdalenian and Oranian cultures) known ancient mtDNA and (2) Epipaleolithic. With this last one I found some conceptual difficulties so I had to take decisions, which were:
  • A most recent date boundary of 4000 BCE (which already overlaps with Neolithic in most regions since 1500 or more years before). Actually the most recent sites are c. 4200 BCE from Lithuania and c. 4600 BCE from Navarre.
  • No inclusion of any Neolithic data even if contemporary. The only possible exception was Franchti Cave (Greece), which has a sequence beginning in the Epipaleolithic (or Mesolithic) but is largely Neolithic. The exact adscription of the sequenced individual is not known.

The results are:

Late Upper Paleolithic mtDNA from Europe and North Africa
R* and specially R*-CRS can well be H and have often been reported as such but we do not know for sure

Epipaleolithic mtDNA from Europe (until 4000 BCE)
R* and specially R*-CRS can well be H and have often been reported as such but we do not know for sure

Some of these data (and others from more recent periods) can be seen in the dedicated Ancient mtDNA maps page at this blog. It needs some updating however: not much time has passed since I created those maps but new findings do pile up quickly these days. 

Ancient Scythian mtDNA

Another point of interest of the thesis is the ancient Scythian tombs from the Don basin (Iron Age, proto-historical). The results show some greater Eastern genetic influence than modern peoples (Russians) do.

The results, which place ancient Scythians closer to modern Central Asians than to Eastern Europeans are consistent with other recent studies that show an inflow of Eastern Asian mtDNA lineages into Central Asia even before the Turkic invasions of the Roman period and early Middle Ages.

Bronze Age Sardinian mtDNA

Finally the thesis deals with Sardinians from the Bronze Age (Nuraghic period). The sites are both from the most central parts of Sardinia, so they may be more representative of an early refuge population than to the overall Bronze Age of the island but still they are curious and interesting:

Dersarkissian argues that this suggests continuity but with many doubts, partly because the source of the genetic data (isolated teeth) did not allow for any certain identification of individuals. Still the resulting mtDNA pool (no matter how you look at it) is not really modern but rather reminds of Central European and Mediterranean Neolithic sites. 
The may well be some of the last Neolithic immigrants, who, instead of replacing the hunter-gatherer aborigines all around (as some imagined too dearly) were the ones taking refuge in this turbulent period in the highlands of Sardinia.
Who knows?!
 

Italian and Sardinian autosomal genetics

A new paper investigates the genetic structure of Italy:
Cornelia di Gaetano et al., An Overview of the Genetic Structure within the Italian Population from Genome-Wide Data. PLoS ONE, 2012. Open access ··> LINK [doi:10.1371/journal.pone.0043759]
The results confirm that Sardians are a very distinct population and show that Italians essentially seem to cluster with mainland Europeans (NW Europeans in principle but Iberian or Balcanic comparisons are missing), West Asians and Sardinians in this order.
There is some N-S gradient in the Peninsula and Sicily but it’s mostly determined by an increasing West Asian affinity in the South. Central Italians stand between North and South but clearly closer to the North but many individuals from NW Italy (Liguria and Piedmont, as well as some Sardinians) actually cluster with Southern Italians as well as with “mixed” Sardinians (those Sardinians who stand between the main insular cluster and the peninsular one).

Let’s see:

Figure 1. SNP-Based PC of 1,262 individuals from 10 sub-populations.
The Italian population plotted onto the first two principal components defined by the European HGDP-CEPH populations and CEU HapMap data. Scatter plot of the first two principal components, obtained using R software (prcomp). Analysis based on 125,799 autosomal SNPs. Individuals included belong to Northern Italy (N-IT): black dots, Central Italy (C-IT): red dots, Southern Italy (S-IT): green dots, Sardinian (SAR): blue dots… 
[the original legend does not explain well the other populations (too many blatant errors in the text) but it’s obvious that the group to the top-right corner are other Europeans (French, CEU), while the group to the center-left are West Asians (Druze, Palestinian, Bedouins) and Mozabites. Larger images can be downloaded from the paper].

In this first characterization we see a primary duality between Europe and West Asia (the Paleo-Neolithic dichotomy probably) and a secondary one between Sardinia and mainland Europe.
Figure 2. SNP-Based PC of 1,014 individuals from the Italian dataset.
A. A Scatter Plot of the Italian population of the first two principal components obtained via R software (prcomp). Individuals included belong to Northern Italy : black dots, Central Italy : red dots, Southern Italy : green dots, Sardinian: blue dots.
B. Italian population without the Sardinian-projected scatter plot of the first two principal components obtained via the R software (prcomp)
[larger images can be downloaded from the paper]
Here we see (A) a main dichotomy between Sardinia and Peninsular Italy (with Sicily) and a secondary N-S gradient. However in (B) it becomes more obvious that to some extent there are two distinct clusters: Southern and Central-North Italy with certain clear separation.
However, and this is quite interesting some North Italians strongly cluster with Southern Italians. Razib mentions this fact as signature of internal Italian migrations but individual migrations would not look that way because the genetic distinction would have diluted in the meantime, appearing at most as intermediate. What we see instead is preserved genetic identity, not too diluted or not diluted at all, with Southern Italy in many Northern Italians.
Who are these Northern Italians, I wondered then. The answer is in the supplements:

Hidden population structure within the Italian dataset. Scatter plot of the first two eigenvectors based on 125,799 autosomal SNPs and 1,012 individuals. Colors represent the four different macro-areas; green- Southern Italy (Apulia, Calabria/Sicily, Campania, Basilicata), red- Central Italy (Tuscany, Lazio, Emilia Romagna and Abruzzo/Marche), black- Northern Italy (Piedmont,Liguria, Aosta Valley and Lombardy), blue- Sardinia (these samples were labeled for the linguistic area). Subjects are symbol- labeled by municipality. Information on municipality was not used for calculations.
[click to expand]

In this image we can appreciate how all Northern Italians clustering with Southern Italians are from two specific regions: Liguria and Piedmont (Piemonte), the Northwestern regions of Italy, bordering France. What do these two regions have in common? All I can think is that, in ancient times they were mostly inhabited by the Ligures, a pre-Indoeuropean people plausibly descendant from the first Neolithic colonization (Cardium Pottery, via the Chassey-Cortaillod-La Lagozza cultural complex). 

Roman region of Liguria (Regio IX)

We are also provided with a bayesian cluster analysis, for which K=4 seems the most valid result (K=3 and K=5 also give low cross-validation values but do not seem more informative):

Figure 3. Clustering of the European, Northern African and Middle Eastern individuals by the Structure software.
Model-based
ancestry analysis based on a subset of HGDP-CEPH and HapMap CEU data
using the merged data of 126K autosomal SNPs. Ancestry for each
individual was inferred using ADMIXTURE [50] at K = 4. Abbreviations as in Figure 1.

This confirms four clusters: Main European (green), Sardinian (red), West Asian (blue) and North African (purple). 
I tend to consider the West Asian component as the main Neolithic input in Europe, although, of course other DNA sections may well have traveled around in that period or later on. 
I also find notable that Sardinian affinity exists among Italians, French and North Africans (surely via Iberia) but almost not among North American Euro-descendants (CEU) of NW European origin and West Asians, who instead do sport some notable Mainline European affinity. 
It’s also interesting that CEU are among the most North African related of all European populations.


Some prehistoric and proto-historic speculation

IF, and only IF, the affinity of Ötzi with Sardinians can be considered representative of how most Italy was in the Chalcolithic (and not a random fluke specific of that man or his mountain community), then, we should consider two further waves into Italy: (1) of West Asian affinity (maybe from the Agean since the Bronze Age or even before) and (2) of mainland European affinity (Indoeuropeans: Italics, Celts).
IF this is correct then the Ligures would not be so much descendant genetically from La Lagozza-Chassey, as I said above but from the “Aegean” wave. This would also be consistent with some individual Tuscans clustering with Southern Italians as well (historical Etruscans are one of the culminations of these Aegean waves together with the Greek colonies). 
But sincerely, I am not aware of any such Aegean flow arriving to the proto-historical Liguria, are you?
So I must consider that there is another possibility: that the Sardinian element represents only one of several Neolithic (or maybe even Paleolithic but nothing clear here) elements in Italy, maybe associated to Y-DNA I2a (strong in Croatia, Bosnia, etc.), while the other, the one most akin to West Asia, would be related to Y-DNA E1b-V13 (strong in Greece and Albania) and maybe other patrilineages from the Eastern Mediterranean like J2b, etc. Both E1b-V13 and I2a are know from ancient DNA from the Neolithic of the Western Mediterranean, so they did indeed take part in these migrations.

Then the “Greek” or “Aegean” (or “Albanian” if you wish) component was reinforced by Bronze Age flows while the “Dalmatian” one was diluted instead by the successive Indoeuropean (Kurgan) waves.

I’ll leave it this way until more evidence comes forward.

 

On and around with Ötzi’s genome

As you’re probably more than aware by now there’s a new paper on the market (yeah, 32 bucks – but worry not that I already got my hands on it) on the most loved mummy of Europe: Ötzi the Iceman.

A. Keller et al., New insights into the Tyrolean Iceman’s origin and phenotype as inferred by whole-genome sequencing. Nature 2012. Pay per view.

The most notable conclusion would seem to make Ötzi closest in all to Sardinians or more like   Corsicans, at least by Y-DNA. This one has been described now as G2a-L91, what is per ISOGG 2012 G2a2b (although the authors use the old nomenclature G2a4) and is most commonly found in Southern Corsica and the Corsican-speaking parts of Sardinia (Gallura).
The autosomal DNA has been compared with an all-Europe sample (the Behar 2010 one, I think based on the nomenclature used), to which a Sardinian sample was added. The result (right) does suggest a Sardinian (or Corsican) affinity of Ötzi.
Notice please that in the supplemental material the Ötzi dot achieves three different positions depending on the level of refinement: while all place Ötzi to the bottom left corner, he’s exact position varies quite a bit – it’s not like PC analysis (nor genetics overall) is rocket science, you know.
Also another caveat I have with this kind of analysis is that all it says is that Sardinians and Ötzi are very negative for both PC components, th Northern and the Eastern ones. The only association at the bottom left corner is a negative one: neither Nordic nor Greek, and this is not too informative.
Yes, the Y-DNA points to an association with Corsica (rather than Sardinia), what reinforces the suggestion posited by the autosomal DNA basic (but negative) analysis, still it would be nice if the authors would have bothered to do some ‘Admixture’ type of analysis as complement. 
At the moment all we have is a negative: Ötzi, who belonged to a Cardium Pottery derived cultural group (Bocca Quadrata or La Lagozza, can’t recall right now) and bears a quite clear Neolithic marker such as Y-DNA G2a, shows up as strongly non-Balcanic, unlike most modern Italians (Europe S sample).

It looks odd indeed… but it might be explained if we assume that from that time on, secondary (post Neolithic) Bronze Age flows from the Aegean (and Central Europe) altered gradually the genetic composition of Italy. This is supported by archaeology as far as I know: even before Mycenaean Greeks, the Aegean was influencing Southern and Central Italy more and more. This trend was reinforced in the late Bronze Age (Mycenaean colonization in the South, Etruscan migration in the Center) and the Iron Age (classical Greek colonization of Magna Graecia).

Before the Romans Italy was all or most of the time a recipient of cultural influences (from the Balcans, from SW Europe and from Central Europe) and did not, as far as I can tell, export culture except as secondary trampoline (the Cardium Pottery Case notably). Excepting the Cardium Pottery case, it acted more as a buffer between West and East and dead end than what its central Mediterranean position would suggest. Even in the Heraklean myth, original Greek version, the route to the fabled Hesperides does not go through Italy but North Africa. Only later, as the Romans rose to prominence, was Hercules made to journey back through Italy, something not specified in the original version. 
I’m saying all this because it may explain why the Europe S (Italy) component tends so strongly towards the Balcans (and to lesser extent Northern Europe) but neither Ötzi nor Sardinians do, even if they look Neolithic-blooded to some extent.
 

Escargots crossed the Mediterranean via Sardinia hitchhiking human ships

Snails of the Tudorella sulcata species, which lives on land, are found all around the Western Mediterranean: in France, Spain, Sardinia, Algeria and Malta. A new genetic study deals on how the snails, original from SW Europe, made it to Sardinia and Algeria.

Sampling sites (color-coded by region)

Statistical parsimony networks with based on 604 bp of the mitochondrial COI gene (a) and 219 bp of the nuclear hsp70 gene (b).

As often happens, the haploid (not recombined) mitochondrial genome is the most informative: the snails traveled to Africa mostly via Sardinia. 
The authors estimate (???) ages between 10 and 3,000 years ago, with a m.l. date of 8000 BP and claim this would have some sort of relation with Neolithic spread. However this is far from clear: on one side there is no known Sardinia-North Africa interaction before Megalithism (since c. 6-5000 years ago, dying out gradually only around 3000 years ago), which would also include Malta in the equation. On the other side, Iberia, a possible alternate origin/route has been neglected in the research.
 

Echoes from the Past (Oct 14) – the genetic isolation of humankind

I’m planning an entry on Paleolithic and Neolithic navigation but meanwhile, here it goes some stuff (mariner or not) that I find interesting.
Homo genus became genetically isolate thanks to natural spermicide
H. erectus (female) reconstruction
A critical change in a immune system molecule, from Neu5Gc to Neu5Ac, made our ancestors effectively isolated from our cousins from the Pan genus and probably also from the then common australopithecines. 
This change would simply kill any non-human sperm in the uterus or, would it manage to succeed, the resultant fetus. This incompatibility with other hominins may have been critical in the process of speciation of the first Homo species such as Homo erectus, Homo habilis or maybe A. sediba. 
··> Science Daily, Darius Ghaderi et al. at PNAS (PPV for six months or freely accessible in some world regions).

Human thumb (Neanderthal or H. heidelbergensis) found in Sardinia
The finding of a thumb bone in Sardinia, dated to 250-300,000 years ago, may help break the fantasy of ancient humans not being able to navigate. This finding adds to those of Crete (c. 190 Ka ago and the famous Flores hominin), all of which must have crossed vast spans of sea in order to get to their destinations, implying at least some level of navigation. 
In the discussion at NeanderFollia, David indicated further evidence of archaic navigation I was unaware of: H. erectus must have reached Flores c. 900,000 years ago, in what is probably the most ancient navigation feat we can confirm ··> John Hawks, Environmental Grafitti, Adam Brumm et al. at Nature (PPV).
Also there is at least some uncertainty of H. ergaster or some other human species maybe crossing to Europe via the Strait of Gibraltar at similar dates as in Flores or maybe even earlier, but, because of the various possible routes involved this is less conclusive. Instead, Flores, Sardinia and Crete have not been connected to the mainland at any time in the biological history of the genus Homo.
Art workshop found in South Africa
A number of shells with indications of having held ochre have been found in the important site of Blombos Cave, South Africa. The shells had holes which suggest that they were used as containers. Other tools, such as hammers and knives, to work the clay, have also been found.

Babies know justice instictively
While actual perception and interest on fairness varies, a good deal of human babies (15 months old) clearly show interest in fair sharing and will actively share. Other babies have less interest in fairness however but they will share anyhow, even if in a less generous manner. 
Malaria research casts doubt on mitochondrial DNA ‘molecular clock’

It seems that the molecular clock is not on streak. Recently it was radically challenged for Y-DNA and it seems obvious that it will not survive in general, at least without radical revisions. A crucial assumption for the molecular clock hypothesis is that the clock ticks regularly or almost so. 
Well, it does not seem to be the case of mtDNA either: certainly not for the primate parasite Plasmodium sp

The use of fossils from the host as absolute calibration and the assumption of a strict clock likely underestimate time when performing molecular dating analyses on malarial parasites. Indeed, by exploring different calibration points, we found that the time for the radiation of primate parasites may have taken place in the Eocene, a time consistent with the radiation of African anthropoids. The radiation of the four human parasite lineages was part of such events. 

Celtic astronomical kurgan found in Germany
Dated to the 7th century BCE, the plan of a burial mound (or kurgan) of the Hallstatt period in the early Celtic area of Southern Germany has been reported. Allegedly the disposition of the wooden posts around the mound inform about the astronomy of the Moon, primarily, and the Sun and they may even describe constellations.

Altamira at risk on short-sighted tourism greed

Millán Mozota denounces at his blog, echoing other researchers, the short-sighted attitude of the Cantabrian authorities who have decided to open the Altamira cave to the public again in spite of the dramatic risk for the art in it.

In the last decade, considerable attention has been paid to the deterioration of the caves that house the world’s most prominent Paleolithic rock art. This is exemplified by the caves of Lascaux (Dordogne, France) (1) and Altamira (Cantabria, Spain), both declared World Heritage Sites. The Altamira Cave has been closed to visitors since 2002. Since 2010, reopening the Altamira Cave has been under consideration. We argue that research indicates the need to preserve the cave by keeping it closed in the near future.

The public can enjoy a replica of part of the cave at the nearby museum.
Iberian Neolithic idols
While in Spanish language, I can’t but call your attention to this fifth article of Neolítico de la Península Ibérica on the diverse array of idols known from the Neolithic and Chalcolithic of Iberia. Even if you can’t read any Spanish, you will no doubt gather some information and visual recreation from simply watching the many images and maps included in this blogpost. For example:

Orange ovals: “eyed” idols (oculados), brown ovals: “plate” idols (ídolos placa)

··> Neolítico de la Península Ibérica[es].

Last minute news:  some iris pattern genetics unveiled ··> The Spitoon.

 

Paleo-Sardinian language: a relative of Basque?

Basque linguist J.M. Elexpuru discusses today at Noticias de Álava[es] the possibility that the lost pre-Romance language of Sardinia could be related to Basque, following the steps of Catalan linguist Eduardo Blasco Ferrer, who just published a book titled Paleosardo, le radici linguistiche de la Sardegna neolítica (Paleo-Sardinian, the linguistic roots of Neolithic Sardinia).
Ruins of a Sardinian nuraghe
Sardinia belonged to the Carthaginian Empire since the 6th century BCE and then passed to the Roman one in the 3rd century, remaining since then in the Romance linguistic area. However little is known of the history of the island before, except the famous nuraghe forts (similar to SE Spanish motillas) and that it was colonized (after some ill-known Epipaleolithic episode) within the Cardium Pottery culture in the Neolithic, probably from Central Italy.
However Dr. Elexpuru synthesizes this way the position of Blasco Ferrer:

… there was a migration from the Basque area in the Mesolithic (8000-5000 b.C.) which settled the island. There were surely other flows later on. Genetic research on mitochondrial DNA have revealed that haplogroup V, originary from the Basque-Cantabrian area, is very high in the central region. The language carried by the settlers, named Paleo-Sardinian by linguists, was the one spoken through all the Neolithic and Bronze Age in the island and still survived for some centuries to Roman domination in the central region, which was known as Barbaria. In some parts of the island the density of pre-Roman toponyms is well above 40%. 

He concludes mentioning some of the river and settlement names that are quite obviously Vasconic:  
  • River names: (h)aran, ardi, baso, berri, bide, ertz, goni (goi), gorri, iri, istil, iz, lats, lur, mando, on, orri, (h)osto, (h)otz, (h)obi, (i)turri, ur, zuri.
  • Village names: Aritzo, Ardaule, Asuni, Goni, Loiri, Luras, Olzai, Orgósolo, Ortueri, Osini, Turri, Ulassai, Uras, Uri, Urzulei…
I must say that all this would make better sense if Iberian and Ligurian could be somehow integrated in the picture. One reason is that haplogroup V is now known to be much more frequent and probably original not from the modern Basque Country nor even Gascony but from farther East: Catalonia probably. However now and again there are other rare or somewhat common lineages that appear shared between Iberia and/or the Basque Country and Sardinia (and sometimes also North Africa). 
One of the most common ones is Y-DNA I2a, a West Mediterranean and Pyrenean clade extremely common in Sardinia, which, if of Neolithic origin, would be the only such lineage quite frequent among Basques. But it could also be pre-Neolithic. 
Frequency of Y-DNA I2a, from Rootsi 2004
For the record, it was discussed  earlier in this blog (also here) the striking similitude of Basque and Sardinian (and some other European) carnival performances, all this in relation to the apparent paleo-European veneration of the bear and the continentally widespread shared root for this animal (hartz in Basque, almost the same in proto-Indoeuropean).
Also for the record I must mention that, in my not so humble opinion, the very word Sardinia seems to have a Basque etymology. Obviously it is derived from the pan-European word sardine but this term only makes etymological sense in Basque: sarda (fish school) + -gin (suffix of doing/making < egin) + -e/-a (nominative declension, like the article “the”).  It needs of a loss of a syllabe (would make sardagina) but I still think it’s plausible that sardine (and hence Sardinia) means school-doer or school-maker in Basque or a related language from old.
Whatever the case it is extremely difficult to deny the Basqueness of the toponyms listed above, even if I am sure that soon someone will come and contest such obviousness, based not on common sense but on twisted and ill-explained elaborations.
But what I still do not have fully clear is in which direction the Vasconic language spread. Of course the default hypothesis of an expansion from the Franco-Cantabrian region makes good sense but it is difficult to completely discard a Neolithic spread of the language family in the context of Cardium Pottery culture (and loosely related Atlantic ones, including Megalithism).
I also think that this Vasconic substrate is not something peculiar of Sardinia and that anyone who looks around with a keen eye and a half-decent knowledge of Basque language can’t but stumble once and again on Basque-like toponimy all around the western half of the continent.
Article found via Ostraka Euskalduna[eu]