Genetic differentiation between contemporary British and Irish populations

 

"This seems to me a thing to be noticed, that just as the men of this country are, during this mortal life, more prone to anger and revenge than any other race, so in eternal death the saints of this land, that have been elevated by their merits, are more vindictive than the saints of any other region." - Gerald of Wales, Topographia Hibernica 

"The Irish hate our order, our civilization, our enterprising industry, our pure religion. This wild, reckless, indolent, uncertain and superstitious race have no sympathy with the English character. Their ideal of human felicity is an alternation of clannish broils and coarse idolatry. Their history describes an unbroken circle of bigotry and blood." - Benjamin Disraeli

"Besides some small islands round about Britain, there is also a large island, Ierne, which stretches parallel to Britain on the north, its breadth being greater than its length. Concerning this island I have nothing certain to tell, except that its inhabitants are more savage than the Britons, since they are man-eaters as well as heavy eaters, and since, further, they count it an honourable thing, when their fathers die, to devour them, and openly to have intercourse, not only with the other women, but also with their mothers and sisters; but I am saying this only with the understanding that I have no trustworthy witnesses for it; and yet, as for the matter of man-eating, that is said to be a custom of the Scythians also, and, in cases of necessity forced by sieges, the Celti, the Iberians, and several other peoples are said to have practised it." - Strabo, Geographica

                                                                                

How different (or similar?) are the Irish and their beloved British neighbours? The answer will more than likely piss off both armchair republicans and loyalists alike. The latter claiming to be the true original inhabitants of the Cruithin only retaking what was always rightfully theirs whilst the former claim to be bastard descendants of Egpytian pharaohs and shipwrecked Basque pescaderos. The sad truth is that both factions are far more interrelated to one another than both would be comfortable admitting regardless of who was what or where going back to antiquity because the British and Irish relationship is far deeper than antiquity. As old to antiquity as antiquity is to us now in 2021. But in the case of the Irish there has been substantially less homogenisation since then. Almost going back as far to the late bronze age (but unfortunately no samples have yet been published from Ireland).

To illustrate this elevated single grave/rhineland bell beaker drift in Ireland (which I believe peaks in Connaught where y-haplogroup R1b reaches its highest frequency in the world) I will be resorting to the Global 25 PCA courtesy of the eurogenes blog and examine samples going all the way from Bronze age to Medieval times.

Chalcolithic/Early Bronze Age

While the G25 is not perfect for mapping NW euro variation
it does a damn fine job of playing out this bronze age
drift.

Middle Bronze Age

MBA samples from Ireland/Wales not available but gradually
the distances to moderns become lower as local neolithic
holdouts are absorbed into the overall population.

Late Bronze Age

Before arrival of Brythonics with Hallstatt C culture.

Iron Age

After arrival of Brythonics there is noticeable increase in EEF.
Even slightly more than modern Irish surprisingly hence the 
higher distance than Scotland_LBA. 
Surely after the arrival of Anglo-Saxons and Vikangs
this close drift with Irish ends?

Early Medieval

Picts narrowly plot closer to modern Scots but quite shockingly

the early medieval samples mostly taken from an 8-9th century

grave from Hinxton, Cambridgeshire plot 

nearest to Irish again so they are heavily intermingled 

with assimilated Brythonics. This would seem to 

indicate that modern Anglos require extra "Southern" drift

that likely arrived from France during the high medieval

era due to the close relations Plantagenets had with French

nobility.

  

Last year's Viking burial paper Sikora et al 2020 goes into this southern drift which I have suspected for some time in the supplementary materials note 11:

"A major 2020 study, which used DNA from Viking-era burials in various regions across Europe, found that modern English samples showed nearly equal contributions from a native British "North Atlantic" population and a Danish-like population. While much of the latter signature was attributed to the earlier settlement of the Anglo-Saxons, it was calculated that up to 6% of it could have come from Danish Vikings, with a further 4% contribution from a Norwegian-like source representing the Norwegian Vikings. The study also found an average 18% admixture from a source further south in Europe, which was interpreted as reflecting the legacy of French migration under the Normans."

18% seems bang on but this turnover did not come from 
Normans themselves who were largely an elite, but rather
from Gascony which was an English possession for centuries
even to the point medieval French chroniclers considered them
to be "Anglais".

"Oui oui vee ar caming vor vour vimmin Haroldus"

Taking an average of all Irish/British bronze age 
samples available on G25 we get what I feel is a 
reasonably accurate picture of where this continuity is highest
My thanks to ancestral whispers for the map and reconstruction of
Rathlin 1, pictured on the left. 
Although there are only two Irish BA samples available
currently so the overwhelming majority of these samples are from
Britain. 

The first Irelanders

3D reconstruction of the crania of Louschbour man,
a WHG (Western Hunter Gatherer) who lived around 6000BC
in what is now Belgium, Irish WHG would've looked more or less like this.

Ten thousand years ago, or twelve depending on who you ask, a small band, probably no more than a couple dozen, dark skinned, blue eyed people made landfall on a small island for the first time ever to our knowledge. 

Only a few millennia before the island had been an arctic wasteland that was completely inhospitable to human habitation. Covered in glaciers which gradually melted during the Bølling–Allerød oscillation resulting in the hilly landscape seen here today. The Allerød oscillation resulted in warmer temperatures in the North Atlantic similar to modern levels, before things eventually got cold again, even colder than LGM levels in fact. This cold period is known to us as the 'Younger Dryas' and would've roughly coincided with the arrival of these hunter gatherers to the island we know today as Ireland.

Their ancestors would have originated somewhere in the vicinity of modern day south eastern Europe, belonging to the Epigravettian complex. Rather than the previously hypothesised Franco-Cantabrian regium hypothesis, with the dinosaur Palaeolithic continuist nonsense of R1b cromagnon supermen colonising everything, it seems that Iberia, which was for the latter half of the Upper Palaeolithic a bastion for some of the first Anatomically modern human populations to arrive in Europe during the Aurignacian, however these UP survivors themselves it seems did not survive the expansion of the Epigravettian foragers from the Balkans, with samples from the Epipaleolithic Azilian harbouring significantly higher levels of WHG related genetic ancestry than those hunters of the preceding Magdalenian. The fact that Basques today exhibit the highest WHG genomic ancestry is as it turns out a mere coincidence with virtually all but one Mesolithic and Neolithic samples belonging to paternal lineages other than R1b, such as G2a, I2a and H2a, with the lone exception from El Trocs cave with one farmer belonging to now extinct (except in Sardinian men) R1b-V88. 

This lineage likely made its way into early farming communities from Anatolia somewhere in the Balkans where it was found in Mesolithic samples from Lepenski Vir, Serbia, and the Epigravettian Villabruna individual, the oldest known WHG yet uncovered. 

Front and lateral view of Ripari
Villabruna's skull, oldest as of
yet known individual to possess
the R1b haplogroup, as well as 
blue eyes.
14kya.

Magdalenian hunters (El Miron) already harboured significant levels of WHG ancestry 
as early as 20,000BC WHG had expanded across the Mediterranean. 

With the extinction of most megafauna towards the end of the Pleistocene, early Holocene foragers such as the WHG became dependant on smaller game, with red deer, Irish elk, boars, wild aurochs and rabbits for meat. An emphasis on freshwater fish was also made with most Mesolithic sites being located next to river lanes. As a result of this, WHG were substantially shorter than hunter gatherers that hunted the megafauna that proved plentiful during the UP. With the average Gravettian male standing at 1.83m WHG men on average stood at a much shorter 1.63m. Though were powerfully built.

From Grasgruber et al 2014

For the first time ever we now have two samples from the Mesolithic era from Irish Western Hunter Gatherers, and rather intriguingly they appear to be somewhat inbred, although not to the same extent of that of the "god king" individual buried at Newgrange almost to the point of forming their own unique clade from continental and even British WHG. From the supplement of Cassidy et al 2020: 

"We note a significant decrease in inbreeding through time in ancient populations. Hunter-gatherers show both the largest variance in inbreeding coefficients and the highest median. However, none are estimated to have parents who are more closely related than four degrees. This demonstrates that the maintenance of outbreeding mating networks was common practice in the Upper Palaeolithic and Mesolithic periods of Europe, as already seen for the Sunghir site in Russia."

These people knew about inbreeding, and the problems that would have resulted from it.

The samples were taken from from the remains of two male individuals from Killuragh in Limerick and Sramore in country Leitrim. They both lived around the tail end of the Mesolithic roughly 4000BC. Just as the first wave of farmers were beginning to colonise the island from the Atlantic façade.

After four to six thousand years of isolation they had become so genetically drifted from continental WHG to the point of almost being a separate lineage entirely. At its zenith the largest Mesolithic population would have been no more than a few hundred individuals. Which would explain why Irish Neolithic samples were largely identical in terms of WHG admixture to continental counterparts.

Like the vast majority of WHG the uniparentals for these two were fairly typical. Both belonged to subclades of mtDNA haplogroup U5b and Y-DNA haplogroup I2a. With Killuragh6 belonging to I2a1a2 and Sramore62 to I2a1b2, the same haplotype as that of the most famous WHG, Cheddar man. Who made the headlines in 2018 for a somewhat subpar reconstruction.

 A more recent and quite frankly 

probably more accurate reconstruction 

compared to that created by 

the Natural History museum of London.

Artist’s impression of a mesolithic Iberian from
La Braña, Arintero from around 5000BC
just as the first wave of cardial farmers arrived
from across the Mediterranean. 
Basques today have amongst the highest proportion of 
WHG admixture in Europe.

Killuragh cave, a site which appears to 

have been frequented by mesolithic hunters

for thousands of year with many 

bone fragments being recovered from here.

Rather then being buried within the cave systems, bones were placed at the mouth of the cave, perhaps for ritualistic reasons which were then washed away by natural processes. 

                                                                                                                                       

"Closest" modern populations

As per the conclusion of Cassidy et al 2020, the two Mesolithic individuals shared highest drift with British and other Epigravettian offshoot populations from the continent. 

3D PCA

2D PCA

When modelled with Epigravettian WHG (Villabruna)
Irish samples do not score any Iberian HG as previously thought

As for introgression by Irish WHG into oncoming neolithic communities that established themselves on the island roughly around 4000BC, this signal seems to be nonexistent as the vast majority of Neolithic samples from the island do not harbour significantly higher levels compared to contemporaneous samples from the continent, specifically the cardial Mediterranean region where we now know they spread from. 

Genocide?

Given that the foragers would've numbered no more than a few hundred, and the farmers in the tens of thousands, it's logical to think that Irish WHG more or less met their end by the beginning of the Neolithic transition, as did most WHG did in general with some straggler groups surviving in northern Scandinavia in isolated pockets as recently as 2000BC.

For the majority of or at least the first half of its populated history, as much time has transpired since the beginning of the Neolithic to now as has the beginning of the Mesolithic to its end, if we take the twelve thousand year date into consideration.

Conflicts over land probably did occur, with local WHG probably hunting imported domesticated livestock and sparking the occasional scrap, but I don't see any reason to suspect any semblance of a violent genocide, with peaceful integration probably occurring over several centuries as cultural boundaries between the two began to fade as it would do again in the later Bronze age. 

Cosy

3D model of a Bronze Age Irishman’s skull

Rathlin 1 buried in supine position c.1906BC
The foetal like position is thought to symbolise
rebirth.
Sloan et al 2006

While this blog certainly isn’t one dedicated to 20th century anthropology, (there are plenty of Facebook groups for that) I quite recently came across this rather excellent specimen on sketchfab and felt I just had to post it here. The crania appears to be in excellent condition, although a piece of the maxilla is missing. Along with pieces of the parietal bone, perhaps being caved in by a cudgel like implement? Was this man murdered? Interestingly Rathlin island was a rich source of porcellanite, used for making stone axe heads which were exported to Britain. Despite being technically the Bronze Age stone tools and weaponry did not immediately fall out of fashion. As late as 1200BC men were still wielding cudgels as evidenced by the Tollense river valley battle site. Only local elites would’ve had the knowledge to smelt bronze. 

The skull appears to be that of Rathlin 1, the oldest and best preserved of the three food vessel culture men uncovered in 2006 and sequenced later by trinity college geneticists. He recently made a cameo appearance in part 1 of an Irish language documentary series which I was pleasantly surprised by, having been quite interested in this individual myself. Unlike most present day Irish his craniometric measurements appears to be brachycephalic as was that of most bell beaker people. He also has the stereotypical flat occiput which is not common in Irish skulls of today which tend to be rounded.

Although I am not particularly knowledgable in the field of anthropology, I would personally classify him as a Dinarised Borreby, using Carleton Coon’s own classification system. 

All credit of course goes to the centre of archaeological fieldwork for this great upload.

Are Irish the most Indo-European Europeans?

 

Migration route of R1b-L51 men with single grave culture. By 2800BCE these communities
had already established themselves in the Rhineland, assimilating the local funnelbeaker population. 
From here their mixed descendants overwhelm the western remnants of “old Europe”. 

Knowing that this post will illicit some controversy and probably various accusations of racism,  I am going to just get it out of the way that I am in no part making this post out of some weird sense of ethno-chauvinism, but rather out of curiousity in exploring the genetic makeup of the Irish genotype as this blog’s name would suggest. This is the elephant in the room that needs to be addressed as I feel it has gone ignored for far too long now.

Are the Irish the genetically purest Indo-European ethnic group in terms of WSH/Steppe_EMBA ancestry?

I believe the answer is yes, and I’ll be exploring why with various Global 25 models you will see below. Does that mean Irish are the closest living breathing ethnic group to the earliest Corded ware population that are more than likely responsible for spreading Indo-European languages across Eurasia?

No. 

As confusing as that might sound to some of you as it did to me at one point in time. All homogenous populations are drifted from one another, but some are more drifted than others. The higher EEF affinity of NW Europeans relative to the more farmer devoid, hunter gatherer rich ancestry of Balto-Slavic  speakers. 

qpAdmixture stats revealing ancient ancestry
proportions of Northern and Eastern Europeans 
The green “eneolithic steppe” component represents
early proto-Indo-European input. Irish outscore all but Latvians and certain Russians. 
But what about just WSH/Steppe_EMBA itself? Who will come out on top?

Globular amphora from Poland represents EEF. Though I must caution

some of this “yellow” component is not from GAC itself, but darker farmer 

women of neolithic western Europe who were somewhat more devoid of the

high WHG signal found in GAC samples. I will perhaps address in a later

post whether our farmer ancestry is mostly Danubian (LBK) or Mediterranean

(Impresso-Cardial) derived. GAC being mostly derived from the former. 

Overall they were around 70% ANF, 30% WHG. Hence the drift with Basques. 

My contention is that while the steppe component of Balto-Slavs is indeed high. The higher drift these populations show with early corded samples may very well be inflated by this higher forager signal. Of which we know was a homogenised blend of mostly western but also eastern European foragers. The latter already making up at least half of the genome of proto Indo-Europeans the other half coming from from forager women of the Caucasus mountains that were present in the north Caucasus piedmont just south of where this admixture event took place. Possibly the initial mixing between the two as early as 5000BC. Two millennia before the formation of the corded ware horizon. I believe this EHG rich baltic forager admixture inflates the drift modern Eastern Europeans have with these earliest Indo-European speakers. Due to hunter gatherer populations living in relatively small numbers, higher genetic drift is observed. A similar phenomenon is observed in the Basque population. The mountainous region providing a refugium for these WHG survivors. Basques possess levels of WHG on par with Balts. 

To illustrate this drift. Look carefully. These samples came from roughly the same time period.

What happened here?

Oh so that’s what happened.
By subtracting CWC_Baltic from the Bronze Age Lithuanian I was able to create a ghost proxy for
baltic hunter gatherers that mixed with the corded ware agropastoralists. 

Ghosts are not as reliable as the real thing however so do not take these results too literally.

I have yet to see any convincing evidence outlining why this drift occurs with NE Europeans, but not with NW Europeans. The Global 25 is extremely sensitive to this drift and cannot simply be chalked up as noise or whatever some other copes I’ve seen online claim. The NE cluster does certainly harbour more EHG but not WSH. 

I can think of other examples other than just Balto-Slavs, such as Dagestani Avars and Tajiks. Neither of whom posses any amount of steppe ancestry higher than that of NW/NE European cluster. But ultimately they are closer than we are due to their zagrosian neolithic component being much closer to that of the CHG like signal found in Yamnaya related groups than that of our anatolian neolithic component which lacks the Ancient North Eurasian ancestry that CHG/Iran_N possesses.  

For the first model I included more or less every population from the northern half of Europe, for the first run I use Yamnaya for steppe, globular amphora for farmer, and the ghost Balt hunter subtracted from Baltic_LTU_BA. I also decided to include Scandinavian hunter gatherers from Norway, whose EHG rich base elevate the drift Scandinavians have with the main CWC cluster, though SHG played nowhere near as a significant a role in the ethnogenesis of Scandinavians as the BHG did with Balts and to a lesser extent Slavs. It is also plausible this SHG like component is actually just BHG that arrived later on into Scandinavia due to Viking raids in the Baltic. Sweden in particular being affected. 

I expect the upcoming SGC samples to be not too different from this.
Essentially the locus of a good chunk of European genetic intra variation.

I then decided to use other samples as a stand in for WSH and was able to obtain a slightly improved fit using Afanasievo herders who appear to be closer to proto-CWC than Yamanya. Irish again come out on top.

Rus_Ust_Ida_EBA represents east asian admixture from 
various Turkic and Uralic groups that affected mostly just Russia.

I went for one last run using the Poltavka volga-ural successor culture to yamnaya and again Irish come out once more on top of the rest of Europe in terms of Indo-European. I invite anyone capable of a more fine scale analyses to debunk my own using formal stats or otherwise. 

A Genomic Compendium of an Island

Y-haplogroup R1b and mental health

"Hurrrr but it's most common there so it must be from there
hurrddddurrrrrrrrrrrrrrrr"

In light of recent publications over the past couple of months of studies featuring genotyped male individuals mostly of WSH origin bearing R1b-M269 markers from across Bronze Age Eurasia, it is time we ask ourselves why is it the same usual suspects continue to hold the now increasingly untenable position that the widespread haplogroup, in this case R1b-L51 did not spread with late eneolithic migrations out of the pontic steppe, despite spreading both west AND as it turns out east with kurgan bell beaker and afanasievo horizons respectively. 

In a post back in October of last year, I discussed the find of a male individual found buried within the context of a corded ware associated burial (there are those who dispute this but they are of course wrong) belonging to a paternal haplotype (R1b-L151) unusual amongst men of that culture (typically various subclades of R1a-M417).

He was of course not the first CWC man found belonging to this lineage (RISE 1) though he was the first to be of reasonably high coverage and with an identical autosomal profile to other corded samples. But of course, absolutely nothing to do with corded ware. 

At all.

Whatsoever.

Not only do they deny its western distribution had anything to do with kurgan related expansions, but some of the even more mentally unstable clowns amongst them deny any involvement of R1b-L51 with yamnaya like groups at all and that its phylogeny came about from local founder effects with traditions native to neolithic Western Europe and that the simultaneous spread of steppe ancestry across the region at the exact same time as this rapidly expanding lineage was merely coincidental, and that it only underwent this massive founder effect with the arrival of said steppe ancestry. But not because of it. (Yeah seriously)

Unfortunately for these individuals who frequent certain genomics blogs spreading this somewhat unlikely theory. The past couple of months has not been very kind to them with several new ancient DNA samples from Afanasievo and Corded ware sites in Mongolia and Poland respectively.

As you can see nothing to do with the steppe.
Yamnaya and Bell beakers were definitely not related.
It’s all a big coincidence they share a clade together.
It’s what the kurganists want you to believe.

With the recent release of the preprint entitled The Genomic Formation of Human Populations in East Asia and the surprising (albeit pleasantly surprising) discovery of a male individual of the Afansievo “Chermuchek” culture now confirmed to have belonged to the R1b-L52 Y-SNP found so far to the east in Mongolia of all places, the eneolithic steppe pre-Yamnaya origin of the up until now elusive haplotype ancestral to the lineages that swept across Western Europe with bell beaker migrations is now looking like a pretty done deal. The Palaeolithic continuity nonsense is dead and it needs to stay this way. A few dead end extinct subclades of R1b turning up in the occasional epigravettian or cardial sample isn’t going to save you. It’s not 2008 anymore.

No doubt a proto-Tocharian speaker living this close to the Altai will cause
conniptions amongst turanist LARPers.

And of course the bell beaker phenomenon is largely yet to be fully answered by mainstream academia with regards to its ultimate origins but with the recent publication of several more corded ware individuals in the recently published “Corded Ware cultural complexity uncovered using genomic and isotopic analysis from south-eastern Poland” from Łubcze, Mirocin and Święte confirmed to belong to derived R1b-L51 associated calls, it’s now looking to be pretty much a done deal at this point. Any denial of BBC spreading with single grave migrations across the north Central European would take a very special sort of mental gymnastics to pull off the logic on how it arrived in our part of Europe simultaneously with steppe atDNA. And if that wasn’t bad enough.

Another R1b-L151 confirmed sample then turned up in a late neolithic male individual labelled “Aesch 25” published in “Ancient genomes reveal social and genetic structure of late neolithic Switzerland”  buried in a dolmen in Switzerland. Unfortunately for the steppe naysayers he wasn’t a neolithic farmer, take a look at who and where he plots with below. 

The nail in the coffin.

A single grave pioneer in the flesh. Dating to roughly 2854-2501 BC. Just before the BBC expansions which swept across Western, and to a lesser extent Eastern Europe as well replacing the CWC which was driven further to the east across the middle Dnieper.

From Ireland to Poland a relatively uniform homogenous population spanned across Europe. 

And over the next several centuries, Eurasia.  

Pcw 160, polish CWC man deemed to be R1b-M269, Mirocin, Malopolska c.2459-2352 BC

A look at some new neolithic Irish genomes

Main passage tomb Listoghil, Carrowmore.

Samples were taken from 2km away in Primrose grange dolmens.
The individual buried here at Carrowmore 4 was deemed to be
the father of one of the Primrose males. 

With the recent publication of genotypes on the Reich lab’s Harvard website featured in last year’s paper on several neolithic samples from megalithic tombs across Northern Europe including several from Primrose and Carrowmore in County Sligo (you can read it here), I decided to take a look at some of them using the Global 25 tool courtesy of the eurogenes blog.

By far the best for modelling ancient populations.

Due to the lack of autosomal DNA samples from ancient sites in Ireland in the past few years, and with the Genomic Compendium of an Island embargoed for another two months. I decided to pass the time and boredom by playing around with a few of the Neolithic genomes featured in last year’s paper whose genotypes were only released quite recently. They’re not as high quality as the one shotgun sequenced ballynahatty sample from the 2015 Cassidy et al paper.

But beggars cannot be choosers I suppose.

In line with recent genetic findings the neolithization of Ireland was a demic event carried out predominantly by cardial agriculturalists spreading from the Aegean via a Mediterranean route eventually reaching the North Atlantic as opposed to the Danubian pioneer populations moving from a more direct Central European route. I wrote an article about which route Ireland was populated from last year concluding with the evidence it was the former and not the latter. The Danubian farmers were generally more devoid of WHG related ancestry as opposed to the cardial populations who assimilated the various Epigravettian and Magdalenian offshoot groups that lived across Southern Europe. The presence of mtDNA haplogroups in Irish neolithic samples that were common in cardial Iberian mitogenomes also provides further evidence (K1a,HV0,H1,T2)

Even more significantly all of the male samples belong to the same Y-DNA haplotype (I2a2a1a1) snowing evidence of patrilineality. The researchers came to the conclusion that these individuals all belonged to the same kindred that lived over the course of twelve generations, allowing thirty years per generation.

D-stats from Cassidy et al 2015 showing Irish neolithic samples sharing
 more drift with cardial related farmers

But how do we know Ireland was populated from the so called Atlantic facade for certain? How can we be sure this drift isn’t just some artefact from similar ratios of WHG admixture to cardial groups rather than actually coming from them? How do we know Ballynahatty’s population didn’t come from a Danubian route and assimilated some hunter gatherer bands along their way?

We know by taking a closer look at the HG component of the Neolithic Irish genomes, which seems to possess ancestry related to Magdalenian complex samples from Iberia rather than just plain old WHG from Central Europe. Magdalenian related ancestry provides a tracer dye for movement of Neolithic groups. They were the descendants of the first modern humans to reach Europe 40kya and sought refuge in Iberia during the LGM before eventually being assimilated by incoming WHG foragers towards the end of the Pleistocene. Both eventually being assimilated by anatolian farmers. 

Looking each of the six samples available to me this type of ancestry is self evident, for my model I will use early neolithic samples from Atapuerca (Iberia_N) and Stuttgart (DEU_LBK) respectively.

For their HG ancestry I will use the 19kya Magdalenian from El Miron.

Starting off with the most high resolution sample from dolmen 002,
the presence of Danubian farmer ancestry could be an artefact or
perhaps real authentic introgression from them did occur with
cardial groups in the Paris basin.

At least three of the Primrose grange samples can be feasibly modelled
sans any Danubian farmer ancestry and are completely Iberian derived
Iberia_N = 90% ANF + 10% WHG

Primrose 016 appears to be an outlier as he harbours roughly one third
of his genome from Danubian EEF, it is difficult to distinguish
such autosomally similar populations so I decided to make another
model just for him.

Despite the improved fit with samples from the Michelsburg culture in

the Paris basin he still requires the same proportion of Danubian
making him the purest Neolithic sample from this site.

Average of all six samples.

PCA revealing variation between Irish neolithic samples and contemporary populations from
across Europe at the time.
Primrose 12 appears to be equidistant between Sardinian and Basque clusters respectively.