Lactase Persistence Over the Last 3,000 Years

As for your bolded comment, there's nothing in that paper or the graphics which indicates anything of the kind. That's pure speculation.

You left out the second text I quoted:


Population history and genetic adaptation of the Fulani nomads: inferences from genome-wide data and the lactase persistence trait (Vicente et al., 2019)

“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 links the Fulani to south-western Europe based on mtDNA haplogroups H1cb1 and U5b1b1b.”

https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6888939/


also:


Internal diversification of non-Sub-Saharan haplogroups in Sahelian populations and the spread of pastoralism beyond the Sahara (Kulichova et al. 2017)

“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).”

https://www.docdroid.net/0mL7YV0/fulani-mtdnay-dna-pdf


Distribution map of the lactase persistence allelle 13910*T, suggesting migration into Africa from western Europe:



Liebert et al., 2017
 
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African cattle mostly belong to the mtDNA haplogroup T1 ...

T1 has been found in Iberian cattle dating from 5200 BC, before the appearance of the earliest cattle in Northwest Africa:


Detecting the T1 cattle haplogroup in the Iberian Peninsula from Neolithic to medieval times: new clues to continuous cattle migration through time (Colominas et al., 2015)

“Our analyses show that haplogroup T3, which dominates among modern domestic European cattle, was the most frequently occurring haplogroup in Iberia since the Neolithic. However, our data also show a presence of T1 haplotypes in Iberia from the Neolithic period through the Bronze Age into the Roman period and the Middle Ages. The presence of T1 haplotypes in Neolithic Iberia, and their chronological and spatial recurrence, clearly demonstrates that T1 was not solely a Muslim or later introduction, as some have argued), though additional and/ or new T1 haplotypes may have been introduced during these periods.

The Neolithic T1 cattle in our data most likely represent populations that expanded out of the Near East in the company of early farmers moving along the Mediterranean coastal route. T1 haplotypes could have reached the coast of eastern Iberia from the Near East as early as ~5400 cal. BC, when human migrations and contacts through the Mediterranean basin are archaeologically attested. Domestic cattle are well documented along the western Mediterranean and coast of Spain at this time, with La Draga being one of the earliest Neolithic Iberian sites where domestic cattle are found. The site of La Draga has close connections with the Mediterranean route of diffusion of the Neolithic, both culturally and due to its location on the eastern coast of the Peninsula. As the earliest cattle in the coastal areas of northwest Africa are dated significantly later than the earliest Neolithic farming sites found there, it is unlikely that cattle could have arrived at La Draga via an African route. The presence of the T1 haplotype at La Draga (northwest Spain, 5200 BC) implies, therefore, that this haplotype travelled with the earliest wave of domestic cattle from the Near East to the north-western Mediterranean. (…)

Our results show the presence of the two most common haplogroups in domestic cattle in Europe (T3) and Africa (T1) in three early Neolithic sites in Iberia. … Our results provide important data about the phylogeny of cattle haplogroup T1 in the Iberian Peninsula, and open new issues about cattle movements through time. Further studies of the geographical distribution of the T1 haplogroup in Europe prior to ~5400 cal. BC, when the Neolithic arrived into the Iberian Peninsula, would shed more light on this topic.”


https://www.docdroid.net/Bpi31Ps/colominas2015-iberia-cattle-pdf
 
African cattle mostly belong to the mtDNA haplogroup T1 ...

T1 has been found in Iberian cattle dating from 5200 BC, before the appearance of the earliest cattle in Northwest Africa:


Detecting the T1 cattle haplogroup in the Iberian Peninsula from Neolithic to medieval times: new clues to continuous cattle migration through time (Colominas et al., 2015)

“Our analyses show that haplogroup T3, which dominates among modern domestic European cattle, was the most frequently occurring haplogroup in Iberia since the Neolithic. However, our data also show a presence of T1 haplotypes in Iberia from the Neolithic period through the Bronze Age into the Roman period and the Middle Ages. The presence of T1 haplotypes in Neolithic Iberia, and their chronological and spatial recurrence, clearly demonstrates that T1 was not solely a Muslim or later introduction, as some have argued), though additional and/ or new T1 haplotypes may have been introduced during these periods.

The Neolithic T1 cattle in our data most likely represent populations that expanded out of the Near East in the company of early farmers moving along the Mediterranean coastal route. T1 haplotypes could have reached the coast of eastern Iberia from the Near East as early as ~5400 cal. BC, when human migrations and contacts through the Mediterranean basin are archaeologically attested. Domestic cattle are well documented along the western Mediterranean and coast of Spain at this time, with La Draga being one of the earliest Neolithic Iberian sites where domestic cattle are found. The site of La Draga has close connections with the Mediterranean route of diffusion of the Neolithic, both culturally and due to its location on the eastern coast of the Peninsula. As the earliest cattle in the coastal areas of northwest Africa are dated significantly later than the earliest Neolithic farming sites found there, it is unlikely that cattle could have arrived at La Draga via an African route. The presence of the T1 haplotype at La Draga (northwest Spain, 5200 BC) implies, therefore, that this haplotype travelled with the earliest wave of domestic cattle from the Near East to the north-western Mediterranean. (…)

Our results show the presence of the two most common haplogroups in domestic cattle in Europe (T3) and Africa (T1) in three early Neolithic sites in Iberia. … Our results provide important data about the phylogeny of cattle haplogroup T1 in the Iberian Peninsula, and open new issues about cattle movements through time. Further studies of the geographical distribution of the T1 haplogroup in Europe prior to ~5400 cal. BC, when the Neolithic arrived into the Iberian Peninsula, would shed more light on this topic.”


https://www.docdroid.net/Bpi31Ps/colominas2015-iberia-cattle-pdf

it seems like T1 expanded along with Cardial Ware
 
This is the statement to which I objected as being too speculative:

"The evidence indicates that R1b-V88 might have adopted pastoralism in the Balkans and migrated into Africa via western Europe. "

You told me to look at the second paper for the proof of that statement. I don't see any mention of R1b-V88 or that it adopted pastoralism in the Balkans.

Pastoralism, by the way, is a late development in the trajectory from hunter-farmer to agriculture with animals, to primarily herding, to actual "Pastoralism". It's adopted when climatic conditions make farming difficult to impossible.

What we have known for years is that there was migration going both ways across the Straits of Gibraltar. Given the presence of the "European" LP allele in Berbers that trait moved with people going from Iberia to northwestern Africa. There is then what looks "perhaps" like a spread from the Berbers to the Fulani.

I stated all of this upthread. You're preaching to the choir.

That's all well and good. Some of those people may have carried R1b-V88. Perhaps even more important going by the paper you cited is yDna "T", although I'm sure carriers of other y lineages had the LP allele. The group that broke off and went south might have coincidentally carried R1b-V88.

So, no, this isn't a tale of one Mesolithic hunter-gatherer group inventing pastoralism and trail blazing like a comet across all of Europe and into Africa.
 
This is the statement to which I objected as being too speculative:

"The evidence indicates that R1b-V88 might have adopted pastoralism in the Balkans and migrated into Africa via western Europe. "

You told me to look at the second paper for the proof of that statement. I don't see any mention of R1b-V88 or that it adopted pastoralism in the Balkans.

Pastoralism, by the way, is a late development in the trajectory from hunter-farmer to agriculture with animals, to primarily herding, to actual "Pastoralism". It's adopted when climatic conditions make farming difficult to impossible.

What we have known for years is that there was migration going both ways across the Straits of Gibraltar. Given the presence of the "European" LP allele in Berbers that trait moved with people going from Iberia to northwestern Africa. There is then what looks "perhaps" like a spread from the Berbers to the Fulani.

I stated all of this upthread. You're preaching to the choir.

That's all well and good. Some of those people may have carried R1b-V88. Perhaps even more important going by the paper you cited is yDna "T", although I'm sure carriers of other y lineages had the LP allele. The group that broke off and went south might have coincidentally carried R1b-V88.

So, no, this isn't a tale of one Mesolithic hunter-gatherer group inventing pastoralism and trail blazing like a comet across all of Europe and into Africa.

there is no direct evidence for that but plenty of circumstantial evidence
R1b-V88 adopted pastoralism in the Balkans
they spread along with cardial ware and were also spotted in LBK, tough the LBK line probably went extinct
with cardial ware they arrived in Sardegna and Iberia, probably also in Sicily
they spread in Africa ca 7,6 ka, when the 'Green Sahara' returned and rivers flowed between lake Chad and the Mediterranean
they could very well have spread catlle mtDNA T1 in Iberia and Africa
there was also another branch of pastoralists coming from the Levant into Africa, they were E1b1b1 and the forefathers of the Cushites
T is another branch of Pastoralists, they made it to the Levant, Anatolia, northwestern Morroco, Iberia and amongst LBK, but there is no trace of arriving further into Africa during the neolithic
 
I said that R1b-V88 might have adopted pastoralism in the Balkans, and subsequently migrated into Africa via western Europe, not that they necessarily invented pastoralism. I also said that the spread of the 13910*T allele into Africa is associated with R1b-V88, and possibly T1a (Y-DNA).

According to Marcus et al. 2020: "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.”


The Els Trocs R1b-V88 individual from northeast Spain (5180 BC) belonged to a pastoral population:

"The Els Trocs phytolith record suggests a seasonality of frequentation in accordance with an interest in pastoral and not in cultivation activities. The cave probably also played a role in the rituality of these pastoral groups ... The cave was used as a seasonal (late spring and summer) shelter related to pastoral activities. The presence of bone remains as well as the concentrations of spherulites confirms at least a partial stabling of animals in the cave."

Lancelotti et al. 2012


Some further evidence regarding pastoralism and dairy production:


“This study provides the first isotopic insights into the diet and subsistence economy of Early and Middle Neolithic populations from open-air sites in interior north-central Iberia…. These findings contribute to an understanding of the implementation of an agro-pastoral economy in interior Iberia, suggesting a stronger reliance on animal foods among the first Neolithic groups of inner Iberia than in subsequent periods. (…)

This shift in subsistence, described here for the first time, may tentatively be related to the stronger emphasis on pastoralism increasingly being attributed to the earliest Neolithic communities of interior Iberia, which is consistent with the local palynological and zooarchaeological records. (...)

Unfortunately, it remains unknown whether this potentially greater commitment to a herding economy attributed to the first north-central Iberian Neolithic groups would involve a significant degree of mobility (transhumance, transterminance) or not (e.g. 'from dawn till dusk'). In any case, it is possible that, rather than a fully arable economy, Early and Middle Neolithic subsistence practices in the region emphasized intensive dairying and domestic herbivore meat exploitation complemented by the hunting of ungulates, gathering of wild plants and a small-scale intensive and diversified agriculture based on the cultivation of small cereal plots. This model could be the precursor to more extensive cereal-based farming from the Late Neolithic onwards, as both the isotope evidence and the environmental and archaeological records suggest. (…)

the results show a significant shift in diet between the Early/Middle Neolithic and the Late Neolithic in north-central Iberia, which possibly relates to a subsistence change from mainly pastoral to mixed economies.”

Fernandez-Crespo et al. 2019


"in Europe, milk exploitation varied from east to west along the northern Mediterranean seaboard, as seen in the quasi-absence of dairy residues in ceramic vessels from northern Greece, in contrast to the strong evidence for dairying in the northwestern Mediterranean. The former cannot be solely explained by the potential use of perishable containers for milk processing or mixing with porcine fats, because AtD profiles have shown that husbandry was focused on meat production in these communities. Moving westwards, AtD profiles and lipid residue findings strongly demonstrated that early Neolithic communities were both actively managing animals for milk and processing milk in ceramic vessels. Combined evidence from faunal and lipid residue analyses, therefore, unequivocally show that the production and use of dairy products was widespread across the breadth of the northern Mediterranean, except in mainland Greece, from the onset of agriculture. Milk and dairy products might have been an important staple in early farming communities, and one of the key drivers in the spread and maybe in the adoption of animal domestication."

Spiteri et al. 2016
 
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I said that R1b-V88 might have adopted pastoralism in the Balkans, and subsequently migrated into Africa via western Europe, not that they necessarily invented pastoralism. I also said that the spread of the 13910*T allele into Africa is associated with R1b-V88, and possibly T1a (Y-DNA).

According to Marcus et al. 2020: "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.”


The Els Trocs R1b-V88 individual from northwest Spain (5180 BC) belonged to a pastoral population:

"The Els Trocs phytolith record suggests a seasonality of frequentation in accordance with an interest in pastoral and not in cultivation activities. The cave probably also played a role in the rituality of these pastoral groups ... The cave was used as a seasonal (late spring and summer) shelter related to pastoral activities. The presence of bone remains as well as the concentrations of spherulites confirms at least a partial stabling of animals in the cave."

Lancelotti et al. 2012


Some further evidence regarding pastoralism and dairy production:


“This study provides the first isotopic insights into the diet and subsistence economy of Early and Middle Neolithic populations from open-air sites in interior north-central Iberia…. These findings contribute to an understanding of the implementation of an agro-pastoral economy in interior Iberia, suggesting a stronger reliance on animal foods among the first Neolithic groups of inner Iberia than in subsequent periods. (…)

This shift in subsistence, described here for the first time, may tentatively be related to the stronger emphasis on pastoralism increasingly being attributed to the earliest Neolithic communities of interior Iberia, which is consistent with the local palynological and zooarchaeological records. (...)

Unfortunately, it remains unknown whether this potentially greater commitment to a herding economy attributed to the first north-central Iberian Neolithic groups would involve a significant degree of mobility (transhumance, transterminance) or not (e.g. 'from dawn till dusk'). In any case, it is possible that, rather than a fully arable economy, Early and Middle Neolithic subsistence practices in the region emphasized intensive dairying and domestic herbivore meat exploitation complemented by the hunting of ungulates, gathering of wild plants and a small-scale intensive and diversified agriculture based on the cultivation of small cereal plots. This model could be the precursor to more extensive cereal-based farming from the Late Neolithic onwards, as both the isotope evidence and the environmental and archaeological records suggest. (…)

the results show a significant shift in diet between the Early/Middle Neolithic and the Late Neolithic in north-central Iberia, which possibly relates to a subsistence change from mainly pastoral to mixed economies.”

Fernandez-Crespo et al. 2019


"in Europe, milk exploitation varied from east to west along the northern Mediterranean seaboard, as seen in the quasi-absence of dairy residues in ceramic vessels from northern Greece, in contrast to the strong evidence for dairying in the northwestern Mediterranean. The former cannot be solely explained by the potential use of perishable containers for milk processing or mixing with porcine fats, because AtD profiles have shown that husbandry was focused on meat production in these communities. Moving westwards, AtD profiles and lipid residue findings strongly demonstrated that early Neolithic communities were both actively managing animals for milk and processing milk in ceramic vessels. Combined evidence from faunal and lipid residue analyses, therefore, unequivocally show that the production and use of dairy products was widespread across the breadth of the northern Mediterranean, except in mainland Greece, from the onset of agriculture. Milk and dairy products might have been an important staple in early farming communities, and one of the key drivers in the spread and maybe in the adoption of animal domestication."

Spiteri et al. 2016

this is correct, and it is very consistent with the R1b-V88 pedigree and it's TRMCA's

https://www.yfull.com/tree/R-V2219/

Y8451 split during the Cardial Ware expansian into a Sardinian and African branch
7,6 ka is after the 8,2 ka climate event, when the Green Sahara reapeared

340


https://www.yfull.com/tree/R-Y8451/
 
there is no direct evidence for that but plenty of circumstantial evidence
R1b-V88 adopted pastoralism in the Balkans
they spread along with cardial ware and were also spotted in LBK, tough the LBK line probably went extinct


mtDNA from Els Trocs (c.5200 BC) might have come from central European LBK:

"it cannot be ruled out that the phase I individuals from Els Trocs came from the north via the Rhone Valley. The grounds for this assumption are the genetic profile set of the group. One adult male from Els Trocs (CET 5) exhibits mtDNA haplogroup N1a1a1, which was common in early Neolithic Central Europe but previously unknown in Spain. He represents the oldest early Neolithic individual on the Iberian Peninsula matching a Central European Neolithic mtDNA haplogroup. Only recently has new evidence been published of other N1a individuals along the hypothetical migration route from Central to Southwest Europe through France."

Alt et al. 2020
 
mtDNA from Els Trocs (c.5200 BC) might have come from central European LBK:
"it cannot be ruled out that the phase I individuals from Els Trocs came from the north via the Rhone Valley. The grounds for this assumption are the genetic profile set of the group. One adult male from Els Trocs (CET 5) exhibits mtDNA haplogroup N1a1a1, which was common in early Neolithic Central Europe but previously unknown in Spain. He represents the oldest early Neolithic individual on the Iberian Peninsula matching a Central European Neolithic mtDNA haplogroup. Only recently has new evidence been published of other N1a individuals along the hypothetical migration route from Central to Southwest Europe through France."
Alt et al. 2020
that is a possibility
but EN Els Trocs and Chaves had both R1b-V88 and I2a1b
and I2a1b was already present in the Alpes-Maritimes as cardial ware pastoralist 7,5 ka, 200 years earlier
I think that is a more likely origin for the Els Trocs pastoralists

Ancient genome-wide DNA from France highlights the complexity of interactions between Mesolithic hunter-gatherers and Neolithic farmers

Maïté Rivollat*, Choongwon Jeong, Stephan Schiffels, İşil Küçükkalıpçı, Marie-Hélène Pemonge,
Adam Benjamin Rohrlach, Kurt W. Alt, Didier Binder, Susanne Friederich, Emmanuel Ghesquière, Detlef Gronenborn,
Luc Laporte, Philippe Lefranc, Harald Meller, Hélène Réveillas, Eva Rosenstock, Stéphane Rottier, Chris Scarre,
Ludovic Soler, Joachim Wahl, Johannes Krause, Marie-France Deguilloux, Wolfgang Haak*

5480-5370 (6445±40; GrA-26893) Pendimoun Alpes-Maritimes, France PEN France_EN_PEN_A F K1a1b1
5480-5360 (6440±40; GrA-26894) Pendimoun Alpes-Maritimes, France PEN France_EN_PEN_B F HV0+195
5480-5337 (6450±40; GrA-32061) Pendimoun Alpes-Maritimes, France PEN France_EN_PEN_A M I2a1a2 M423 Dolman J1c3

soon after they expanded in France & the Pyrenées :
https://www.yfull.com/tree/I-L161/

check Pendimoun in the supplements of this study :

Pendimoun (Castellar, Alpes-Maritimes, Provence-Alpes-Côte-d’Azur, France)
Contact: Didier Binder, Henri Duday
Pendimoun (PEN) is a rock-shelter discovered in 1955, which is 4 km north of
Menton, very close to the French-Italian border. In 1956, preliminary field work provided a
male burial attributed to the Cardial period, but this individual was not considered in this
study due to contradictory dates from chemical preservatives (105). More extensive
excavations performed between 1985 and 2006 (106, 107) revealed a detailed
archaeological sequence from the Early Mesolithic (Sauveterrian) to modern times, with a
specific important sequence attributed to the Impresso-Cardial complex, dated to between
5750 and 5200/5150 cal BCE. The latter includes three inhumations - two females and one
male (107) - dated to the Early Cardial stage, which postdates the Impressa stage. Direct
radiocarbon dates and stable isotopes on human remains provide a narrow range between
5480 and 5330 cal BCE for this phase, placing them amongst the oldest Neolithic individuals
analysed in this part of the Mediterranean coast. The burials are primarily deposited in
individual pits, with distinctive stone architectures for the females and the male. All three
individuals show signs of pathologies or trauma, raising questions about a selective
recruitment.
Unpublished DNA analyses from the ancient DNA labs in Bordeaux and Madrid by
M.-F. Deguilloux, E. Fernandes, C. Gamba, E. Arroyo-Pardo and M.-H. Pemonge have
established that the mitochondrial DNA (mtDNA) was preserved.
We sampled for this study one petrous bone for each of the three individuals.
Calibrated date intervals are given at 2-sigma level.
• PEN001_real1 / Pendimoun F1; 5480-5370 cal BCE (6445±40BP; GrA-26893)
• PEN001_real2 / Pendimoun F2; 5480-5360 cal BCE (6440±40BP; GrA-26894)
• PEN003 / Pendimoun H2; 5480-5337 cal BCE (6450±40BP; GrA-32061)
 
there is no direct evidence for that but plenty of circumstantial evidence
R1b-V88 adopted pastoralism in the Balkans
they spread along with cardial ware and were also spotted in LBK, tough the LBK line probably went extinct
with cardial ware they arrived in Sardegna and Iberia, probably also in Sicily
they spread in Africa ca 7,6 ka, when the 'Green Sahara' returned and rivers flowed between lake Chad and the Mediterranean
they could very well have spread catlle mtDNA T1 in Iberia and Africa
there was also another branch of pastoralists coming from the Levant into Africa, they were E1b1b1 and the forefathers of the Cushites
T is another branch of Pastoralists, they made it to the Levant, Anatolia, northwestern Morroco, Iberia and amongst LBK, but there is no trace of arriving further into Africa during the neolithic

Excuse my ignorance... but does this mean that these R1b-V88 people drank milk around the time they adopted pastoralism, ie did they drink milk during their Balkan stay?
From what I read the drinking of milk and consumption of lactose rich foods predates the evolution of the lactose persistence gene(?).

I read contradictive information. One states that lactose persistance developed ~4k Years ago, from the previous studies I shared in my earlier post.
Yet here it states 2k to 20k years ago the lactose persistence gene was developed. I mean 2-20k and 4k are not mutually exclusive. Would just like your opinion on this.

"But around 8,000 years ago in what's now Turkey — just when humans were starting to milk newly domesticated cows, goats and sheep — mutations near the gene that produces the lactase enzyme started becoming more frequent. And around the same time, adult lactose tolerance developed. The mutation responsible for that may be between 2,000 and 20,000 years old; estimates vary."

"
But now that doesn't happen for most people of Northern and Central European descent and in certain African and Middle Eastern populations. This development of lactose tolerance took only about 20,000 years — the evolutionary equivalent of a hot minute — but it would have required extremely strong selective pressure."

https://www.npr.org/sections/thesal...eral thousand years," as scientists call it).

I thought Angela said Balkan people can digest milk thanks to Slavic genes xD??? Yet V88 might have/has developed pastoralism in the Balkans, and 8-10k years ago in Anatolia we have evidence of milk consumption and statements saying "around 8,000 years ago in what's now Turkey — just when humans were starting to milk newly domesticated cows, goats and sheep — mutations near the gene that produces the lactase enzyme started becoming more frequent".


You are misinformed, again.
Please read post number 26.

If Albanians can drink milk with no ill effects, perhaps you have your Slavic ancestry to thank. That's one northern invasion we missed after the fall of the Empire.
I mean either I am misinterpreting what Angela said, or I am wrong that Slavs migrated to the Balkans much later than this lactose persistence ordeal started.
 
R1b-V88 did pick up pastoralism in the Balkans ca 8 ka, 200 years after farmers from Anatolia arrived in that area
whether they had developped lactase persistence, I don't know but that seems to early to me
however, if the farmers from Anatolia had lactase persistence, R1b-V88 would have inherited it too, as they mixed with farmers daughters
 
Lactase persistence and 13910*T in Iran:


The evolutionary genetics of lactase persistence in seven ethnic groups across the Iranian plateau (Charati et al. 2019)

"The LP (lactase persistence) frequency distribution ranged from 0 to 29.9% in the seven Iranian ethnic groups with an average value of 9.8%. The variants, − 13910*T and − 22018*A, were significantly associated with LP phenotype in Iranians. (...) Our results indicate the distribution of LP in seven ethnic groups across the Iranian plateau. Soft selective sweep rather than hard selective sweep played a substantial role in the evolution of LP in Iranian populations. (...)

The recent genetic screening for Iranians showed the occurrence of − 13910*T at 10%
. The genotype-phenotype correlation was high, suggesting that − 13910*T might explain LP in Iranian population (…) The prevalence of − 13910*T was found in most populations (4.16–7.69%) with the exception of the Gilaks. (...)


All − 13910*T alleles were observed on haplotype A that is agreement with previous studies. (...)

In particular, the occurrence of − 13910*T in two distinguished LP haplotypes was observed in Iranians. Both haplotypes in Iranians were dated within 3000 years that was much later than the dairying practices spread from the West Asia to Europe, raising the possibility that the − 13910*T alleles might be introduced into Iran recently."

https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6371433/
 
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Lactase persistence and 13910*T in India:


Herders of Indian and European Cattle Share Their Predominant Allele for Lactase Persistence (Romero et al. 2012)

“we sought to determine whether lactase persistence has evolved independently in the Indian subcontinent. Here, we present the results of the first comprehensive survey of the LCT enhancer region in south Asia. Having genotyped 2,284 DNA samples from across the Indian subcontinent, we find that the previously described west Eurasian -13910 C>T mutation accounts for nearly all the genetic variation we observed in the 400- to 700-bp LCT regulatory region that we sequenced. Geography is a significant predictor of -13910*T allele frequency, and consistent with other genomic loci, its distribution in India follows a general northwest to southeast declining pattern, although frequencies among certain neighboring populations vary substantially. We confirm that the mutation is identical by descent to the European allele and is associated with the same >1 Mb extended haplotype in both populations. (...)

the derived -13910*T allele has the highest frequency among the observed mutations as well as the widest distribution throughout the Indian subcontinent. Its frequency ranges from 0.8% among the Tibeto-Burman speakers to 18.4% among Indo-European speakers, with west India showing the highest frequency of the derived allele; it has an average countrywide frequency of 10.3%. (...)

We grouped populations by their linguistic or geographic affiliation and carried out Kruskal–Wallis tests to assess the effect of these factors on -13910*T allele frequencies. Both geography and language exhibit significantly nonrandom association with allelic frequencies, whereby groups from west India, or those that speak Indo-European languages are more likely to carry the -13910*T allele than any other group. It is difficult to quantify the degree of autocorrelation between our two main explanatory variables, but they are well known to covary; a further Kruskal–Wallis test applied only on data from Indo-European speakers showed that geography remains significant even when controlling for language. (...) at a countrywide level pastoralism is significantly associated with elevated frequencies of -13910*T (...)

a key question is whether the Indian -13910*T allele is identical by descent to the European one or the product of convergent evolution. … All -13910*T alleles in our southern Asian sample are found on the previously defined European A haplotype background, strongly arguing for a single common origin. (...)

Indo-European–speaking groups were repeatedly identified as being significantly associated with lactase persistence in our data (...)

Population-level -13910*T allele frequencies within our samples range from 0 to 0.489 (48.9%) (...)


Short- and long-range haplotype data point strongly toward a single shared origin of the -13910 C>T mutation in Europe and India. Given the distribution of the -13910*T allele, the observed degree of haplotype conservation indicates a recent origin followed by a rapid rise in frequency, most likely driven by strong positive selection.

Comparison of the European and Indian history of cattle keeping suggests that the context for the selective sweep was in place in Europe roughly 2,000 years earlier than in south Asia (...) It is therefore plausible that from Europe, the allele subsequently spread into Central Asia, the Near East, Pakistan, and India (...)

Interestingly, a similar small-scale introgression event may also explain the presence of the 13910*T allele in central African populations. The overall low frequency of the -13910*T allele in the subcontinent, and its absence from roughly a third of the population samples, suggests that the selective advantage it confers has been largely restricted to pastoralist groups. Three of the five sampled pastoralist populations have -13910*T allele frequencies vastly greater than the sample mean; of those, the Ror of Haryana are responsible for the frequency peaks visible in figure 1, a pattern suggestive of adaptation to a lifestyle tightly associated with the consumption of milk products by these populations.

Populations from north or west India that speak Indo-European languages have, in general, substantially higher frequencies of the -13910*T allele than Dravidian speakers from the south. ... and within the Indo-European speaking groups, there is a clear west to east frequency decline ... This geographically contiguous patterning of variation in the locus is consistent with other genetic data from both autosomal and uniparentally inherited marker loci in India. (...)

haplotype analyses indicate that the -13910*T allele in India is identical by descent to that found in Europe and western Asia, whereas examination of the pattern of haplotype block structure in the context of the archaeological history of herding across this intercontinental region suggests that the -13910*T allele was introduced to India from the west."


Figure 1:




https://academic.oup.com/mbe/article/29/1/249/1749245
 
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I think the point is that this mutation wasn't positively selected until recently, which doesn't really tell us much about its origin. The spread is obviously linked to cultures who practiced animal husbandry. It's certainly possible, if not likely, these European cultures were rich in R1b, even if that male lineage had nothing to do originally with cattle domestication, or goat or some other milking animal..

How did V88 get to Africa? Ancient Spain? Sardinia? Heck even ancient Syria is possible. (a sizeable V88 population exists there too)
 
So what was dairy consumption like back then?

-yogurt?
-strained yogurt?
-milk?
-cheese?
-cream?
-clotted cream?
-butter?
 
Toubou DNA:


Chad Genetic Diversity Reveals an African History Marked by Multiple Holocene Eurasian Migrations (Haber et al. 2016)

“the Toubou have 20%–30% Eurasian ancestry (…) Previous studies have suggested that the Eurasian backflow into East Africa came from a population related to early Neolithic farmers. We wanted to know whether the Eurasian ancestry we found in the Toubou can be traced to the same source populations … the most significant result was for present-day Sardinians. … early Neolithic farmers were the most significant contributors (…) Among modern populations, Sardinians showed the highest genetic affinity to both the Toubou ... Ancient Eurasians also showed correlated affinity to both the Toubou... the early Neolithic LBK (Linearbandkeramik, or Linear Pottery) population (∼5,000 BCE) had the highest affinity. (...)

(in the Toubou) we found signals of selection on MCM6 rs4988235 (13910*T), a variant associated with the lactase-persistence phenotype (ability to digest milk). This SNP was previously found to be under strong positive selection in Europeans, where it was probably advantageous to individuals living in pastoralist societies. The frequency of this variant in the Toubou is 2%… Although this SNP appears to be a candidate for selection, we suggest that it has probably drifted neutrally in the Toubou after the Eurasian gene flow: the Toubou have ~30% Eurasian ancestry from a population similar to the Greeks, who have 13% derived alleles at rs4988235, suggesting an expectation of ~3.9% of the derived allele simply from admixture. We similarly found in the Toubou signals at HERC2 rs1129038 a major contributor to blue eye color in Europeans, as well as a signal at SLC24A5 rs1834640, a major contributor to (light) pigmentation.”

https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5142112/


“We find in Chad a high frequency of Y-haplogroup R1b (R1b-V88) in all ethnic groups examined. … We also find the Eurasian haplogroup T (T1a) in Toubou with Toubou having a high frequency (31%) of their studied males belonging to this haplogroup… instances of this haplogroup in examined ancient populations are in the Linearbandkeramik (LBK) population which we found to be the most significant reference for the Eurasian ancestry in Toubou”

(Haber et al, 2016 supplementary material)


Is there actually any close relationship between the T1a found in the Toubou and in the Linearbandkeramik?


Fulani also have about 18% Y-DNA T1a.

https://en.wikipedia.org/wiki/Fula_people#Genomic_studies
https://en.wikipedia.org/wiki/Haplogroup_T-M184#Africa
https://www.eupedia.com/europe/Haplogroup_T_Y-DNA.shtml


"there are 119 lineages in the macro-haplogroup K-M9 (which includes haplogroups K1-K4 and L to T). Of these lineages, only two have been observed in sub-Saharan Africa at appreciable frequencies: T-M70 (T1a) and R-V88 (R1b-V88)"

Cruciani et al. 2010


This connection between R1b-V88 and T1a is very interesting. As far as I'm aware the only ancient culture in which R1b-V88 and T have been found together so far is the Varna culture, in Bulgaria (4500 BC).





"The numerous archaeological materials [in the Varna necropolis] make it possible to try to explain some aspects of the spiritual culture of the time. … When comparing the stock of grave 43 (a rich grave containing the skeleton of a man) and other collected data, we could suggest that the symbolic graves of this type are designed either as a glorification of the dead or as a cult of the ancestors – founders of the clan and tribe. That the god is a man is judged by the presence of a sceptre and some typically male in function objects… In this grave the prevailing part of the finds implies that the god could be that of stock-breeding. The main evidence for this are the two animal figurines and 30 heads of animals made of flat gold plates.”

Ivanov 1982 p.21
 

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