Incest in dynastic elite in Megalithic Newgrange

Maciamo

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Nature: Incest uncovered at the elite prehistoric Newgrange monument in Ireland

"The huge, elaborate, 5,000-year-old tomb at Newgrange, Ireland, is thought to have been built for a powerful elite. DNA of a man buried there reveals a case of incest. Was this a strategy to maintain a dynastic bloodline?"

Here is the paper: A dynastic elite in monumental Neolithic society, by Cassidy et al. (2020)

Abstract

The nature and distribution of political power in Europe during the Neolithic era remains poorly understood. During this period, many societies began to invest heavily in building monuments, which suggests an increase in social organization. The scale and sophistication of megalithic architecture along the Atlantic seaboard, culminating in the great passage tomb complexes, is particularly impressive. Although co-operative ideology has often been emphasised as a driver of megalith construction, the human expenditure required to erect the largest monuments has led some researchers to emphasize hierarchy—of which the most extreme case is a small elite marshalling the labour of the masses. Here we present evidence that a social stratum of this type was established during the Neolithic period in Ireland. We sampled 44 whole genomes, among which we identify the adult son of a first-degree incestuous union from remains that were discovered within the most elaborate recess of the Newgrange passage tomb. Socially sanctioned matings of this nature are very rare, and are documented almost exclusively among politico-religious elites—specifically within polygynous and patrilineal royal families that are headed by god-kings. We identify relatives of this individual within two other major complexes of passage tombs 150 km to the west of Newgrange, as well as dietary differences and fine-scale haplotypic structure (which is unprecedented in resolution for a prehistoric population) between passage tomb samples and the larger dataset, which together imply hierarchy. This elite emerged against a backdrop of rapid maritime colonization that displaced a unique Mesolithic isolate population, although we also detected rare Irish hunter-gatherer introgression within the Neolithic population.


Y-DNA haplogroups uncovered included 34 I2a individuals and two rare H2a (samples BG72 and JP14). The most common haplogroup within Neolithic Ireland is I2-M284 - some also positive for L1193. I had already listed these clades as belonging to the Megalithic cultures of the British Isles four years ago, so no surprise.

I2a2-tree.png


Table 12 in the supplements shows the skin, hair and eye pigmentation for samples from this study as well as other papers on ancient DNA. British and Irish Neolithic individuals had a variety of skin tones ranging from pale-intermediate to dark. Almost all had brown or black hair. The majority had brown eyes. Only one individual surely had blue eyes and three others might also have had blue eyes.


Cassidy and colleagues also tested two Irish Mesolithic hunter-fishers, who seem to have be genetically distinct from Mesolithic Britons.

"The genetic data obtained from human remains dating to around 4700 BC (from Killuragh Cave, County Limerick, in southwest Ireland) and to around 4100 BC (from Sramore Cave, County Leitrim, in the northwest and Stoneyisland, County Galway, in the west) are the first DNA results for Ireland’s hunter-fisher-forager groups. These Mesolithic Irish people were genetically distinct from their Mesolithic neighbours across the Irish Sea in Britain, suggesting a prolonged period of genetic isolation after these people sailed6 across to Ireland around 8000 BC. In other words, even though they might have ranged widely over Ireland when choosing partners, members of these communities did not sail back to Britain or across to the continent to interact with people there — contrary to what some archaeologists have proposed."

One of those two Mesolithic men (SRA62) belonged to Y-haplogroup I2-L38 and mt-haplogroup U5a2d. Neither haplogroup was found in the Neolithic Irish population (all the Neolithic mtDNA U5 was U5b, not U5a). It's not impossible that these haplogroups did survive to the Neolithic but just weren't among the small number of Neolithic samples tested.
 
this is not the first study that proves how elitarian and paternalistic the megalithic societies were
and probably also non-megalithic middle neolithic societies in Europe, like the Chasséen and Michelsberg cultures

and even though other studies claim that the first Bell Beakers on the British Isles replaced 90 % of these populations, the Y-DNA lineages found in these megalithic societies didn't go extinct, they are well-represented in the extant British population (I2-M284, I2-L161 and others)
we're still missing something here
 
this is not the first study that proves how elitarian and paternalistic the megalithic societies were
and probably also non-megalithic middle neolithic societies in Europe, like the Chasséen and Michelsberg cultures

Actually this study postulates that Newgrange was ruled by an elite that tried to preserve its bloodline through incest, like Egyptian pharaohs.

"A man buried in the chamber of Newgrange around 5,000 years ago was the offspring of a first-degree incestuous union: his parents were either siblings or parent and child. This finding led the team to speculate that the elite associated with this magnificent monument practised incest as a way of maintaining a dynastic bloodline. Such a strategy, which breaks a near-universal social taboo against incest, was also practised much later by ruling elites in ancient Egypt, in the Inca empire and in ancient Hawaii."
[...]
"However, Cassidy et al. report that those buried between around 3500 and 2500 BC in passage tombs (and also at a different but related type of monument at Millin Bay, County Down, in the north) display familial relationships that extend over considerable geographical distances and span several generations. People buried in passage tombs also seem to be differentiated from those in other kinds of burial monument by having had a particularly meat-rich diet."

"The authors found genetic links between some individuals analysed from the major passage-tomb complexes at Carrowmore and Carrowkeel in County Sligo, and individuals buried 150 kilometres away at Brú na Bóinne (and also at the Millin Bay monument). The authors interpret these links as providing evidence for a non-random selection of partners over large territories, implying a high level of societal complexity. These genetic data support the argument that the trajectory of Irish passage-tomb development over time — generally speaking, going from small and simple to larger and more ostentatious — reflects an increasingly hierarchical society9. The evidence of incest found at Newgrange, suggesting dynastic behaviour, is consistent with this overall picture."


and even though other studies claim that the first Bell Beakers on the British Isles replaced 90 % of these populations, the Y-DNA lineages found in these megalithic societies didn't go extinct, they are well-represented in the extant British population (I2-M284, I2-L161 and others)
we're still missing something here

They aren't extinct, but nevertheless rare today (<5% of the population).
 
Related to Maciamo's post above. In the paper by VanDeLooschbrecht et al 2020 which is pre-print at biorxiv, which has some new samples from Grotta Del Uzzo (Trapani, Sicily). The study covers period from Early Mesolithic (12,250-11,850 BC), Late Mesolithic, to Early Neolithic (5,460-5,220 BC), of the 4 males 1 was Y-DNA I (Sample UZZ_40) and 1 was I2a2 (UZZ_81). These 2 were from the Late Mesolithic period (6,750-5,850 BC). I didn't link the paper here as it is more appropriate for it in the Sicily_Pre Greek thread. If I remember, bicicleur, you mentioned the Grotta De Uzzo site and it looks like we are going to get a new study with those ancient Sicilian Samples that pre-date the ones in Fernandes et al 2020 from earlier this year. On a personal note, the fact that I2a2 is showing up that early in Sicily is interesting since that is my Y-DNA haplogroup. So related to the study on the Irish Neolithic, did I2a2 spread from the Balkans to Ireland and also into Italy/Sicily and spread during Neolithic or is it a surviving HG lineage in both areas, which is what I think most of the research indicates.

If this question is more appropriate for the Y-DNA forum, I will move it there. Thanks
 
all the Neolithic mtDNA U5 was U5b, not U5a

Sample PN112 (Neolithic) was U5b2b.

Egyptian Middle Kingdom nomarch/ruler Djehutynakht (2000 BC) was U5b2b5.

U5b2b5 has also been found in Sardinia (c.2300 BC) and Spain (c.3500 BC).

Neolithic/megalithic Sardinia was dominated by I2a and R1b-V88.

African R1b-V88 appears to have come from Spain or Sardinia.

“A west Eurasian origin for R1b-V88 is supported by a recent phylogenetic analysis that puts modern Sardinian carrier haplotypes basal to the African R1b-V88 haplotypes. The putative coalescence times between the Sardinian and African branches inferred there fall into the Neolithic Subpluvial (“green Sahara”, about 7,000 to 3,000 years BCE). Previous observations of autosomal traces of Holocene admixture with Eurasians for several Chadic populations provide further support for a hypothesis that at least some amounts of Early European Farmer (EEF) ancestry crossed the Sahara southwards. […] Overall, our analysis provides evidence that R1b-V88 traces back to eastern European Mesolithic hunter gatherers and later spread with the Neolithic expansion into Iberia and Sardinia [and from there into Africa].”

https://static-content.springer.com.../MediaObjects/41467_2020_14523_MOESM1_ESM.pdf
 
[/FONT]One of those two Mesolithic men (SRA62) belonged to Y-haplogroup I2-L38 and mt-haplogroup U5a2d. Neither haplogroup was found in the Neolithic Irish population (all the Neolithic mtDNA U5 was U5b, not U5a). It's not impossible that these haplogroups did survive to the Neolithic but just weren't among the small number of Neolithic samples tested.

Cheddar man was also pré-I2-L38, a clade now extinct :
https://www.yfull.com/tree/I-Y10705/
 
Sample PN112 (Neolithic) was U5b2b.

Egyptian Middle Kingdom nomarch/ruler Djehutynakht (2000 BC) was U5b2b5.

U5b2b5 has also been found in Sardinia (c.2300 BC) and Spain (c.3500 BC).

Neolithic/megalithic Sardinia was dominated by I2a and R1b-V88.

You may be interested in checking my mtDNA haplogroup pages. I listed the ancient samples found so far under each clade as a reference for everyone. Here is the U5 page.

African R1b-V88 appears to have come from Spain or Sardinia.

“A west Eurasian origin for R1b-V88 is supported by a recent phylogenetic analysis that puts modern Sardinian carrier haplotypes basal to the African R1b-V88 haplotypes. The putative coalescence times between the Sardinian and African branches inferred there fall into the Neolithic Subpluvial (“green Sahara”, about 7,000 to 3,000 years BCE). Previous observations of autosomal traces of Holocene admixture with Eurasians for several Chadic populations provide further support for a hypothesis that at least some amounts of Early European Farmer (EEF) ancestry crossed the Sahara southwards. […] Overall, our analysis provides evidence that R1b-V88 traces back to eastern European Mesolithic hunter gatherers and later spread with the Neolithic expansion into Iberia and Sardinia [and from there into Africa].”

https://static-content.springer.com.../MediaObjects/41467_2020_14523_MOESM1_ESM.pdf

My opinion is that R1b-V88 people originated in the Levant and propagated the Neolithic herder lifestyle throughout Africa from there. They would have crossed from North Africa to Iberia and Sardinia.
 
I am pressed for time and can't really go through this, can someone tell me how these individuals compare genetically with the mainland Megalith culture? I seem to remember that the Atlantic megalith culture were mainly Farmer genetically, but with a male-skewered influx of WHG, does the WHG in these individuals resemble Irish-type WHG or non-Irish?
 
My opinion is that R1b-V88 people originated in the Levant and propagated the Neolithic herder lifestyle throughout Africa from there. They would have crossed from North Africa to Iberia and Sardinia.

Has R1b-V88 been found in any ancient samples from the Middle East?
 
My opinion is that R1b-V88 people originated in the Levant and propagated the Neolithic herder lifestyle throughout Africa from there. They would have crossed from North Africa to Iberia and Sardinia.

Do you still think the megalithic culture came to western Europe from the Levant?
 
Has R1b-V88 been found in any ancient samples from the Middle East?

Not yet, but R1b-V88 is present in the Levant and Egypt today and African herders that carry R1b-V88 today like the Hausa and the Fulani have no European autosomal DNA but do have Middle Eastern one.
 
Do you still think the megalithic culture came to western Europe from the Levant?

Yes, they came with Neolithic farmers. The oldest Megalithic sites in Europe are in Malta and Iberia, then they spread northwards. Somewhere in Iberia and/or France local I2a hunter-gatherers integrated Neolithic farming societies and their paternal lineages spread to Britain and Ireland.
 
I am pressed for time and can't really go through this, can someone tell me how these individuals compare genetically with the mainland Megalith culture? I seem to remember that the Atlantic megalith culture were mainly Farmer genetically, but with a male-skewered influx of WHG, does the WHG in these individuals resemble Irish-type WHG or non-Irish?

You will find a summary of European Neolithic Y-DNA and mtDNA here.
 
Not yet, but R1b-V88 is present in the Levant and Egypt today and African herders that carry R1b-V88 today like the Hausa and the Fulani have no European autosomal DNA but do have Middle Eastern one.


“Based on genome-wide analyses we propose that ancestors of the Fulani population experienced admixture between a West African group and a group carrying both European and North African ancestries. This admixture was likely coupled with newly adopted herding practices, as it resulted in signatures of genetic adaptation in contemporary Fulani genomes, including the control element of the LCT gene enabling carriers to digest lactose throughout their lives. The lactase persistence (LP) trait in the Fulani is conferred by the presence of the allele T-13910, which is also present at high frequencies in Europe. We establish that the T-13910 LP allele in Fulani individuals analysed in this study lies on a European haplotype background thus excluding parallel convergent evolution. (…) Our findings suggest that Eurasian admixture and the European LP allele was introduced into the Fulani through contact with a North African population/s. (…)

The T-13910 allele is reported to be the key variant regulating maintenance of LCT gene expression in European adults. This variant is generally not detected in most East African and Middle Eastern populations, where other LP variants are observed instead. Fulani populations living mainly in the western Sahel/Savannah belt, however, carry the European-LP mutation with frequencies ranging from 18 to 60%. (…)


European admixture in Fulani genomes has been reported in previous studies (…)

We started by investigating the genetic affinities of the Fulani from Ziniaré in Burkina Faso using a set of comparative populations from Africa, Europe and Near East. The principal component analysis, PCA, clusters the Fulani groups with other West Africans while displaying some genetic affinity to Eurasians. This prevalent West African component was also visible in population structure analysis, where the Fulani from Ziniaré in Burkina Faso have ancestry fractions of 74.5% West African, 21.4% European and 4.1% East African origin at K = 3. We observe a similar genetic structure among all other Fulani groups in our dataset, except for the Fulani from Gambia. We notice that some individuals in this group display a higher European ancestry component than others, suggesting some degree of sub-structure in this population. (…)


To examine which particular source population was a likely candidate for this postulated European contact, we extracted all European-like segments across the Fulani genomes. … The European-like segments showed the highest shared drift with Sardinians and French Basque populations … A previous study has reported a Mozabite-like (i.e. Berber-like) component in the Fulani from Burkina Faso and Niger, raising the possibility that the source population for the European admixture fraction (and LP mutation) could be of North African origin. … the ancestry components of the Mozabite group could explain the non-West African genetic variation in the Fulani. ... The identification of the specific ancestry fragments flanking European-like segments, supervised admixture and model based analyses support the view that the European ancestry in Fulani genomes is coupled to their North African component (...)

Fregel and colleagues (2018) linked the diffusion of people across Gibraltar to Neolithic migrations and the Neolithic development in North Africa. This trans-Gibraltar mixed ancestry was previously observed in the Fulani mitochondrial gene-pool that link the Fulani to south-western Europe based on mtDNA haplogroups H1cb1 and U5b1b1b.”

Vicente et al. 2019


“We show that age estimates of the maternal lineage H1cb1, occurring almost exclusively in the Fulani, point to the time when the first cattle herders settled the Sahel/Savannah belt. Similar age estimates were obtained for paternal lineage R1b-V88, which occurs today in the Fulani but also in other, mostly pastoral populations. Maternal clade U5b1b1b, reported earlier in the Berbers, shows a shallower age, suggesting another possibly independent input into the Sahelian pastoralist gene pool. (…)

The fact that haplotypes having –22,018*G and –13,910*T LCT alleles are shared by the Fulani from Northern Cameroon, the Mozabite Berbers from Algeria and European populations (Lokki et al., 2011; Ranciaro et al., 2014) can be taken as another signal of an ancient pastoralist migration from outside Africa (…)

In the present study, we demonstrate that the age estimates of West Eurasian uniparental haplogroups U5b and H1cb – which, in the Sahel/Savannah belt, occur more often in pastoralists (especially the Fulani) – are chronologically concordant with the introduction of livestock to the region (as documented independently by archaeological data).

Inferring the geographical location of the ancestral West Eurasian population that was the source of these Holocene inputs to the present-day Sahel/Savannah uniparental gene pool can be approached from a phylogeographical perspective. … It is well attested that H1 emerged in Iberia/South Europe (Pereira et al., 2005b) … The parental lineage U5b1b1, aged 9 ka, is highly dispersed from the Saami in Scandinavia to Berbers in North Africa, indicating the Franco-Cantabrian refuge area as the most parsimonious source of late-glacial expansions of hunter-gatherers repopulating the entire region heading both north and south (Achilli et al., 2005). The most probable hypothesis is, therefore, that the U5b1b1b found in the Fulani today is derived from North African U5b1b1 ancestors. Both H1cb and U5b1b1b seem to be excellent markers of European – and not Near Eastern – ancestry inputs into the contemporary Fulani population. (…)

Apart from the results of the analysis of uniparental haplogroups in this study, there is other support for European inputs into the Sahelian population in the literature. A couple of genome-wide datasets have also revealed an outlier position and/or a European/West Eurasian ancestry of the Fulani (Henn et al., 2012; Tishkoff et al., 2009). In addition, other analyses of the Fulani collected in the Sahel have detected signatures of positive selection for Eurasian alleles in the TAS2R genes responsible for detection of natural alkaloids such as quinine and strychnine (Triska et al., 2015).”

Kulichova et al. 2017


“Although the T-13910 variant [associated with Lactase Persistence] was absent across most of Africa, we detected this polymorphism in the Mozabite from Algeria (northern Africa) at a frequency of 27.3% and in a few central African populations, namely the Arabic Baggara and Fulani from Cameroon (8% and 22.9% frequency, respectively) and the Bulala from Chad (2.2% frequency). The highest frequency of the T-13910 variant outside of Africa was found in northern Europeans (66.7%), and the lowest frequency occurred in the Middle East (2.2%–4.2%). (…)

The T-13910 and A-22018 variants [associated with Lactase Persistence] occurred on the same haplotype background, consistent with previous studies of European populations. Furthermore, some African populations with the T-13910 and A-22018 variants, such as the Bulala from Chad and the Fulani from Cameroon, shared the same haplotype background with Europeans. However, the Fulani also shared a distinct T-13910/A-22018 microsatellite haplotype with the Arabic Baggara from Cameroon. Further, we noted that the Mozabite from Algeria had the T-13910 and A-22018 variants on the same haplotype background as did the Palestinians from the Middle East. (…)

Although the T-13910 variant is present in central and northern African pastoralist populations, such as the Fulani and Arabic Baggara from Cameroon and the Mozabite from Algeria, it was not observed in other sub-Saharan African populations. This distribution of the T-13910 allele in Africa and the shared haplotype background of chromosomes containing this variant in Africans and non-Africans suggest that the T-13910 variant was introduced through gene flow from outside of Africa into northern and/or central Africa.”

Ranciaro et al. 2014


"Almost certainly, any population with a sizable level of the 13910*T allele [T-13910] has relatively recent (post-Mesolithic) ancestry from Europe. In this context, note the presence of 13910*T in South Asia and North Central Africa. Populations in these regions also show high frequencies of two Y-chromosome haplogroups that are present in samples from Mesolithic Eastern Europe: R1a and R1b-V88, respectively. It's hard to imagine that this is a coincidence."

http://eurogenes.blogspot.com/2017/10/global-distributions-of-lactase.html




 
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Ok. Point taken. There are a few papers I hadn't seen. However the Lactase Persistence allele (T-13910) is not a proof of Europeanness in itself as it could just as well have originated in the Middle East before spreading to Europe. As for mt-haplogroups H1 and U5b they have been present in North Africa at least since the Neolithic and so could have been picked up there by incoming Near Eastern herders before they moved south to the Sahel (probably after the Green Sahara of the Neolithic dried up). Or the Neolithic herders might have come to Iberia first, picking up H1 and U5b there, then moving together to North Africa. Whichever it is, it is 100% certain that animal domestication and the whole Neolithic lifestyle originated in the Near East and not in Iberia, and all R1b-V88 tribes in Africa are farmers or herders. AFAIK these African tribes also carry the near Eastern Y-haplogroup T1a, but no European I2.
 
animal domestication and the whole Neolithic lifestyle originated in the Near East and not in Iberia

I'm not saying it originated in Iberia.

“By inspecting our reference panel of western Eurasian ancient individuals, we identified R1b-V88 markers in 10 mainland European ancient samples, all dating to before the Steppe expansion (>3k years BCE). Two very basal R1b-V88 (with several markers still in the ancestral state) appear in Serbian HGs as old as 9,000 BCE, which supports a Mesolithic origin of the R1b-V88 clade in or near this broad region. The haplotype appears to have become associated with the Mediterranean Neolithic expansion – as it is absent in early and middle Neolithic central Europe, but found in an individual buried at the Els Trocs site in the Pyrenees (modern Aragon, Spain), dated 5,178-5,066 BCE and in eleven ancient Sardinians of our sample. Interestingly, markers of the R1b-V88 subclade R1b-V2197, which is at present day found in Sardinians and most African R1b-V88 carriers, are derived only in the Els Trocs individual and two ancient Sardinian individuals (MA89, 3,370-3,110 BCE, MA110 1,220-1,050 BCE). MA110 additionally carries derived markers of the R1b-V2197 subclade R1b-V35, which is at present-day almost exclusively found in Sardinians.

This configuration suggests that the V88 branch first appeared in eastern Europe, mixed into Early European farmer individuals (after putatively sex-biased admixture), and then spread with EEF to the western Mediterranean. Individuals carrying an apparently basal V88 haplotype in Mesolithic Balkans and across Neolithic Europe provide evidence against a previously suggested central-west African origin of V88. A west Eurasian R1b-V88 origin is further supported by a recent phylogenetic analysis that puts modern Sardinian carrier haplotypes basal to the African R1b-V88 haplotypes. The putative coalescence times between the Sardinian and African branches inferred there fall into the Neolithic Subpluvial (“green Sahara”, about 7,000 to 3,000 years BCE). Previous observations of autosomal traces of Holocene admixture with Eurasians for several Chadic populations provide further support for a speculative hypothesis that at least some amounts of EEF ancestry crossed the Sahara southwards. Genetic analysis of Neolithic human remains in the Sahara from the Neolithic Subpluvial would provide key insights into the timing and specific route of R1b-V88 into Africa - and whether this haplogroup was associated with a maritime wave of Cardial Neolithic along Western Mediterranean coasts and subsequent movement across the Sahara.

Overall, our analysis provides evidence that R1b-V88 traces back to eastern European Mesolithic hunter gatherers and later spread with the Neolithic expansion into Iberia and Sardinia.”

Marcus et al. 2020, supplementary material


 
Yes, they came with Neolithic farmers. The oldest Megalithic sites in Europe are in Malta and Iberia, then they spread northwards.

"radiocarbon results suggest that megalithic graves emerged within a time interval of 200 y to 300 y in the second half of the fifth millennium cal BC in northwest France, the Mediterranean, and the Atlantic coast of the Iberian Peninsula. Northwest France is, so far, the only megalithic region in Europe which exhibits a premegalithic monumental sequence and transitional structures to the megaliths, suggesting northern France as the region of origin for the megalithic phenomenon. (...) the maritime skills, knowledge, and technology of these societies must have been much more developed than hitherto presumed."

Paulsson 2019


“There is great debate over the origin of megaliths and burial in stone chambered tombs, which emerge during the fifth millennium BC (Scarre 2007). The oldest theory of the origin of megaliths is that they represented the spread of a religious cult by megalithic missionaries. This was, however, ruled out by the impact of radiocarbon dating, which showed the Atlantic megaliths to be much older than their supposed Mediterranean forebears. The rejection of a Mediterranean inspiration for megaliths led to suggestions of a local origin, pointing to the presence of burials in late Mesolithic Brittany. At Teviec and Hoedic these occur in shell middens: 23 burials at Teviec and 14 at Hoedic. Men, women, and children were interred together in stone-lined pits, in the most elaborate examples covered by small heaps of stones (cairns) and in one case marked by a small upright stone. The burials date to the period 5500-5000 BC (Schulting 1999b). Ideas of multiple burial therefore already existed among gatherer-hunters of Brittany before the emergence of monuments, and with them the possibility of a purely local development of megaliths”

Fowler et al. 2015


“However far back the origins of seafaring may be, it is clear that the period 12,000–7000 BC saw the emergence of seafaring communities where livelihood was predicated on maritime resources all around the Mediterranean and along the Atlantic coasts of Europe. These maritime foragers seem largely to have developed patterns of restricted mobility within stabilized local systems. But there were exceptions; for example, the spread of foragers from the North European Plain first to Sweden and later along the entire Atlantic coast of Norway. These were maritime adventurers. ... (p.117)

One of the most striking features of the Atlantic facade in the fifth and fourth millennia was the way in which the disparate communities honoured their ancestors by depositing their remains in collective tombs often built of large stones (megaliths). The earliest manifestations of this phenomenon are found in Portugal, Galicia, and Brittany in the period 4700–4500 BC, but the rite of collective burial in graves sometimes lined with stone slabs begins in the Mesolithic period and is evident in shell midden sites found in the Muge and Sado valleys in Portugal and on the islands of Hoedic and Teviec off the south coast of Brittany." (p.119)

Cunliffe 2017




Paulsson 2018
 
Once again I missed that Marcus et al. (2020) paper. I knew about the Mesolithic R1b-V88 in the Iron Gates and Ukraine, but assumed that R1b-V88 had a wide distribution around the Black Sea back in the Mesolithic (after all it was a nomadic age) and that Mesolithic R1b-V88 tribes from Anatolia eventually moved south to the Levant and North Africa and west along the Mediterranean as far as Iberia with the Cardium Pottery culture.

Marcus et al. say that R1b-V88 was picked up in Southeast Europe by Anatolian farmers on their way to Iberia.

What is certain is that all Africa and Middle Eastern V88 today is also derived for V2197. According to Yfull V2197 formed c. 7500 BCE and all present individuals descend from a common ancestor who lived c. 5600 BCE. That interval is exactly when the Neolithic spread across Anatolia and the Balkans, so that does not help us determine whether R1b-V2197 was among the original Anatolian farmers or was assimilated early in the southern Balkans en route to the Western Mediterranean.

The Yfull phylogeny shows that the clades found in Egypt, Chad, Cameroon, Benin are all 7600 years old (5600 BCE), so the spread to Africa was very fast and slightly predate the arrival of Neolithic farmers in Iberia. But between Y-DNA age estimates and radiocarbon age estimates there is plenty of leeway, so I wouldn't say it is conclusive evidence that Neolithic R1b-V2197 definitely first spread to Africa.

For the moment, what we can say is that R1b-V88 was present among Mesolithic people in the Balkans and Ukraine (and possibly Anatolia too) and that R1b-V2197 spread with Early Neolithic farmers around the Mediterranean, including North Africa.
 
"radiocarbon results suggest that megalithic graves emerged within a time interval of 200 y to 300 y in the second half of the fifth millennium cal BC in northwest France, the Mediterranean, and the Atlantic coast of the Iberian Peninsula. Northwest France is, so far, the only megalithic region in Europe which exhibits a premegalithic monumental sequence and transitional structures to the megaliths, suggesting northern France as the region of origin for the megalithic phenomenon. (...) the maritime skills, knowledge, and technology of these societies must have been much more developed than hitherto presumed."

Paulsson 2019


“There is great debate over the origin of megaliths and burial in stone chambered tombs, which emerge during the fifth millennium BC (Scarre 2007). The oldest theory of the origin of megaliths is that they represented the spread of a religious cult by megalithic missionaries. This was, however, ruled out by the impact of radiocarbon dating, which showed the Atlantic megaliths to be much older than their supposed Mediterranean forebears. The rejection of a Mediterranean inspiration for megaliths led to suggestions of a local origin, pointing to the presence of burials in late Mesolithic Brittany. At Teviec and Hoedic these occur in shell middens: 23 burials at Teviec and 14 at Hoedic. Men, women, and children were interred together in stone-lined pits, in the most elaborate examples covered by small heaps of stones (cairns) and in one case marked by a small upright stone. The burials date to the period 5500-5000 BC (Schulting 1999b). Ideas of multiple burial therefore already existed among gatherer-hunters of Brittany before the emergence of monuments, and with them the possibility of a purely local development of megaliths”

Fowler et al. 2015


“However far back the origins of seafaring may be, it is clear that the period 12,000–7000 BC saw the emergence of seafaring communities where livelihood was predicated on maritime resources all around the Mediterranean and along the Atlantic coasts of Europe. These maritime foragers seem largely to have developed patterns of restricted mobility within stabilized local systems. But there were exceptions; for example, the spread of foragers from the North European Plain first to Sweden and later along the entire Atlantic coast of Norway. These were maritime adventurers. ... (p.117)

One of the most striking features of the Atlantic facade in the fifth and fourth millennia was the way in which the disparate communities honoured their ancestors by depositing their remains in collective tombs often built of large stones (megaliths). The earliest manifestations of this phenomenon are found in Portugal, Galicia, and Brittany in the period 4700–4500 BC, but the rite of collective burial in graves sometimes lined with stone slabs begins in the Mesolithic period and is evident in shell midden sites found in the Muge and Sado valleys in Portugal and on the islands of Hoedic and Teviec off the south coast of Brittany." (p.119)

Cunliffe 2017




Paulsson 2018

That study contradicts the information from Wikipedia.

Mesolithic


  • c. 9500 BC: Construction in Asia Minor (Göbekli Tepe); from proto-Hattian or else a yet-to-be-discovered culture (the oldest religious structure in the world).
  • c. 7400 BC: A 12 m long monolith probably weighing around 15,000 kg found submerged 40 m under water in the Strait of Sicily south-west of Sicily. Its origin and purpose are unknown.[12]
Neolithic



Based on this timeline, we see a clear diffusion from East to West, staring in Anatolia very close to the onset of the Neolithic, then to Israel and Sicily, then Portugal and Andalusia, and only much later in Brittany.
 

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