Homo sapiens in Arabia by 85,000 years ago

Homo sapiens in Arabia by 85,000 years ago, by Groucutt et al. Nat Ecol. Evol. (2018).

Abstract (emphasis mine):

Understanding the timing and character of the expansion of Homo sapiens out of Africa is critical for inferring the colonization and admixture processes that underpin global population history. It has been argued that dispersal out of Africa had an early phase, particularly ~130–90 thousand years ago (ka), that reached only the East Mediterranean Levant, and a later phase, ~60–50 ka, that extended across the diverse environments of Eurasia to Sahul. However, recent findings from East Asia and Sahul challenge this model. Here we show that H. sapiens was in the Arabian Peninsula before 85 ka. We describe the Al Wusta-1 (AW-1) intermediate phalanx from the site of Al Wusta in the Nefud desert, Saudi Arabia. AW-1 is the oldest directly dated fossil of our species outside Africa and the Levant. The palaeoenvironmental context of Al Wusta demonstrates that H. sapiens using Middle Palaeolithic stone tools dispersed into Arabia during a phase of increased precipitation driven by orbital forcing, in association with a primarily African fauna. A Bayesian model incorporating independent chronometric age estimates indicates a chronology for Al Wusta of ~95–86 ka, which we correlate with a humid episode in the later part of Marine Isotope Stage 5 known from various regional records. Al Wusta shows that early dispersals were more spatially and temporally extensive than previously thought. Early H. sapiens dispersals out of Africa were not limited to winter rainfall-fed Levantine Mediterranean woodlands immediately adjacent to Africa, but extended deep into the semi-arid grasslands of Arabia, facilitated by periods of enhanced monsoonal rainfall.

Related:

Distribution of Southern Iberian haplogroup H indicates exchanges in the western Mediterranean

Recent open access paper The distribution of mitochondrial DNA haplogroup H in southern Iberia indicates ancient human genetic exchanges along the western edge of the Mediterranean, by Hernández, Dugoujon, Novelletto, Rodríguez, Cuesta and Calderón, BMC Genetics (2017).

Abstract (emphasis mine):

Background
The structure of haplogroup H reveals significant differences between the western and eastern edges of the Mediterranean, as well as between the northern and southern regions. Human populations along the westernmost Mediterranean coasts, which were settled by individuals from two continents separated by a relatively narrow body of water, show the highest frequencies of mitochondrial haplogroup H. These characteristics permit the analysis of ancient migrations between both shores, which may have occurred via primitive sea crafts and early seafaring. We collected a sample of 750 autochthonous people from the southern Iberian Peninsula (Andalusians from Huelva and Granada provinces). We performed a high-resolution analysis of haplogroup H by control region sequencing and coding SNP screening of the 337 individuals harboring this maternal marker. Our results were compared with those of a wide panel of populations, including individuals from Iberia, the Maghreb, and other regions around the Mediterranean, collected from the literature.

Results
Both Andalusian subpopulations showed a typical western European profile for the internal composition of clade H, but eastern Andalusians from Granada also revealed interesting traces from the eastern Mediterranean. The basal nodes of the most frequent H sub-haplogroups, H1 and H3, harbored many individuals of Iberian and Maghrebian origins. Derived haplotypes were found in both regions; haplotypes were shared far more frequently between Andalusia and Morocco than between Andalusia and the rest of the Maghreb. These and previous results indicate intense, ancient and sustained contact among populations on both sides of the Mediterranean.

Conclusions
Our genetic data on mtDNA diversity, combined with corresponding archaeological similarities, provide support for arguments favoring prehistoric bonds with a genetic legacy traceable in extant populations. Furthermore, the results presented here indicate that the Strait of Gibraltar and the adjacent Alboran Sea, which have often been assumed to be an insurmountable geographic barrier in prehistory, served as a frequently traveled route between continents.

mtdna-h1-h3-europe-frequency
a, b, c. Interpolated frequency surfaces of clade H and its main sub-clades (H1 and H3). Frequencies (%) are showed in a colour scale. See information about the populations used in Additional files 4 and 5. Map templates were taken from Natural Earth free map repository (http://www.naturalearthdata.com/)

I usually find mtDNA data, especially studies like this one based on modern populations, very difficult to interpret for anthropological purposes. It is well-known that there are important differences in the pattern of Y-DNA and mtDNA expansion and distribution.

A paragraph in this respect caught my attention:

The patterns of variation in the Y-chromosome between western and eastern Andalusians, based on 416 males, have also been investigated for a set of Y-Short Tandem Repeats (Y-STRs) and Y-SNPs [53, 54, 55], Calderón et al., unpublished data] in combination to mtDNA analyses ([18, 19] and present study). In general, for both uniparental makers, Andalusians exhibit a typical western European genetic background, with peak frequencies of mtDNA Hg H and Y-chromosome Hg R1b1b2-M269 (45% and 60%, respectively). Interestingly, our results have further revealed that the influence of African female input is far more significant when compared to male influence in contemporary Andalusians. The lack of correspondence between the maternal and paternal genetic profiles of human populations reflects intrinsic differences in migratory behavior related to sex-biased processes and admixture, as well as differences in male and female effective population sizes related to the variance in reproductive success affected, for example, by polygyny [56, 57].

I think that the greater reduction in patrilineal lineages compared to maternal lineages we usually see during and after prehistoric or historic migrations have more to do with the renown Uí Néill family case and with war-related casualties (since combatants were usually men) than with other more popular explanations, such as enslavement of women or polygyny.

The most successful paternal lines (anywhere in the world) were probably those who remained in power for a long time (be it a patriarchal society based on families, clans, or more complex organizational units), who were richer and thus more capable of having healthy offspring, who in turn were able to survive longer and have more children who inherited power, etc.

In case of recent migrations or population movements that disrupt the previously established organization, after a certain number of generations, successful patrilocal families (usually from incoming lineages) might slowly dominate over a whole region, with poorer families (usually of ‘indigenous’ lineages) suffering a greater – especially perinatal and child – mortality, without any obvious (pre)historic event associated to these gradual changes.

This gradual replacement of paternal lineages is compatible with the adoption of the native language by newcomers. If the number of migrants is greater that the native population, and especially if their technology is more advanced, then a more radical change including ethnolinguistic identification is more likely.

I don’t deny the (pre)historic existence of radical replacement of male populations with continuity of female lineages due to massacres of men, female slavery, or polygyny, but they are probably not the main explanation for most regional differences seen in paternal lineages, and should thus be used with caution.

Gradual replacement and founder effects are also the most logical explanation for why autochthonous continuity myths (that the modern regional prevalence of few successful lineages tended to create in the 2000s) haven’t been corroborated by ancient DNA; e.g. R1b-DF27 in Basques, N1c-M178 in Finnic populations, R1a-Z283 in Slavs, etc. There is nothing different in those areas from other recent founder effects and internal migratory flows seen everywhere in Europe in the past millennia.

Paper discovered via a link by Alberto Gonzalez on Facebook group Iberia ADN

Related:

The uneasy relationship between Archaeology and Ancient Genomics

Allentoft Corded Ware

News feature Divided by DNA: The uneasy relationship between archaeology and ancient genomics, Two fields in the midst of a technological revolution are struggling to reconcile their views of the past, by Ewen Callaway, Nature (2018) 555:573-576.

Interesting excerpts (emphasis mine):

In duelling 2015 Nature papers6,7the teams arrived at broadly similar conclusions: an influx of herders from the grassland steppes of present-day Russia and Ukraine — linked to Yamnaya cultural artefacts and practices such as pit burial mounds — had replaced much of the gene pool of central and Western Europe around 4,500–5,000 years ago. This was coincident with the disappearance of Neolithic pottery, burial styles and other cultural expressions and the emergence of Corded Ware cultural artefacts, which are distributed throughout northern and central Europe. “These results were a shock to the archaeological community,” Kristiansen says.

(…)

Still, not everyone was satisfied. In an essay8 titled ‘Kossinna’s Smile’, archaeologist Volker Heyd at the University of Bristol, UK, disagreed, not with the conclusion that people moved west from the steppe, but with how their genetic signatures were conflated with complex cultural expressions. Corded Ware and Yamnaya burials are more different than they are similar, and there is evidence of cultural exchange, at least, between the Russian steppe and regions west that predate Yamnaya culture, he says. None of these facts negates the conclusions of the genetics papers, but they underscore the insufficiency of the articles in addressing the questions that archaeologists are interested in, he argued. “While I have no doubt they are basically right, it is the complexity of the past that is not reflected,” Heyd wrote, before issuing a call to arms. “Instead of letting geneticists determine the agenda and set the message, we should teach them about complexity in past human actions.”

Many archaeologists are also trying to understand and engage with the inconvenient findings from genetics. (…)
[Carlin:] “I would characterize a lot of these papers as ‘map and describe’. They’re looking at the movement of genetic signatures, but in terms of how or why that’s happening, those things aren’t being explored,” says Carlin, who is no longer disturbed by the disconnect. “I am increasingly reconciling myself to the view that archaeology and ancient DNA are telling different stories.” The changes in cultural and social practices that he studies might coincide with the population shifts that Reich and his team are uncovering, but they don’t necessarily have to. And such biological insights will never fully explain the human experiences captured in the archaeological record.

Reich agrees that his field is in a “map-making phase”, and that genetics is only sketching out the rough contours of the past. Sweeping conclusions, such as those put forth in the 2015 steppe migration papers, will give way to regionally focused studies with more subtlety.

This is already starting to happen. Although the Bell Beaker study found a profound shift in the genetic make-up of Britain, it rejected the notion that the cultural phenomenon was associated with a single population. In Iberia, individuals buried with Bell Beaker goods were closely related to earlier local populations and shared little ancestry with Beaker-associated individuals from northern Europe (who were related to steppe groups such as the Yamnaya). The pots did the moving, not the people.

This final paragraph apparently sums up a view that Reich has of this field, since he repeats it:

Reich concedes that his field hasn’t always handled the past with the nuance or accuracy that archaeologists and historians would like. But he hopes they will eventually be swayed by the insights his field can bring. “We’re barbarians coming late to the study of the human past,” Reich says. “But it’s dangerous to ignore barbarians.”

I would say that the true barbarians didn’t have a habit or possibility to learn from the higher civilizations they attacked or invaded. Geneticists, on the other hand, only have to do what they expect archaeologists to do: study.

EDIT (30 MAR 2018): A new interesting editorial of Nature, On the use and abuse of ancient DNA.

See also:

Paternal lineages mainly from migrants, maternal lineages mainly from local populations in Argentina

New paper (behind paywall) Genetic variation in populations from central Argentina based on mitochondrial and Y chromosome DNA evidence, by García, Pauro, Bailliet, Bravi & Demarchi, J. Hum. Genet (2018) 63: 493–507.

Abstract (emphasis mine):

We present new data and analysis on the genetic variation of contemporary inhabitants of central Argentina, including a total of 812 unrelated individuals from 20 populations. Our goal was to bring new elements for understanding micro-evolutionary and historical processes that generated the genetic diversity of the region, using molecular markers of uniparental inheritance (mitochondrial DNA and Y chromosome). Almost 76% of the individuals show mitochondrial lineages of American origin. The Native American haplogroups predominate in all surveyed localities, except in one. The larger presence of Eurasian maternal lineages were observed in the plains (Pampas) of the southeast, whereas the African lineages are more frequent in northern Córdoba. On the other hand, the analysis of 258 male samples reveals that 92% of them present Eurasian paternal lineages, 7% carry Native American haplogroups, and only 1% of the males show African lineages. The maternal lineages have high genetic diversity homogeneously distributed throughout central Argentina, probably as result of a recent common origin and sustained gene flow. Migratory events that occurred in colonial and recent times should have contributed to hiding any traces of differentiation that might have existed in the past. The analysis of paternal lineages showed also homogeneous distribution of the variation together with a drastic reduction of the native male population.

argentina-mtdna-haplogroups
Maps showing continental mtDNA haplogroups frequencies in 20 population samples from central Argentina. References for populations abbreviated names are from the tables.

Interesting excerpts:

The immigration waves had less impact in the north–central and northwestern regions, the most populated areas of the country in pre-Hispanic times. The spatial structure of genetic diversity has its origins in historical factors. It is possible to distinguish different stages in migratory processes from abroad, with a heterogeneous regional impact. The genetic composition of central Argentina gives account of these processes. On one hand, the political boundaries between provinces influenced the configuration of the genetic structure of the populations that were formed. In this sense, Córdoba—an important economic and commercial center since colonial times—has a greater component of foreign lineages than the populations of San Luis and Santiago del Estero. On the other hand, the genetic structure of central Argentina also accounts for other processes related to different migration phases and occupations of space over the last 500 years.

argentina-maternal-native-american
Maternal continental contribution (in percentages), and Native American haplogroup frequencies, by population

Similarly, negative values observed in the neutrality tests (Tajima’s D and Fu’s FS), indicate relatively recent population growth, probably associated with technological and organizational changes leading to new lifestyles and important demographic and territorial expansion [75]. In conclusion, the molecular markers of maternal inheritance shows large genetic diversity homogeneously distributed throughout central Argentina, probably as result of a recent common origin and sustained gene flow between sub-populations. In addition, migratory events that occurred in colonial and recent times should have contributed to hiding any traces of differentiation that might have existed in the past. The analysis of paternal lineages showed also homogeneous distribution of the variation across the region but also a drastic reduction of the native male population, with a large prevalence of haplogroups of European origin.

argentina-y-dna
Y chromosome haplogroups frequencies in three provinces from central Argentina and other 19 samples from Argentina, Chile, and Paraguay

Related:

Y-DNA haplogroup R1b-Z2103 in Proto-Indo-Iranians?

chalcolithic_early-asia

We already know that the Sintashta -> Andronovo migrants will probably be dominated by Y-DNA R1a-Z93 lineages. However, I doubt it will be the only Y-DNA haplogroup found.

I said in my predictions for this year that there could not be much new genetic data to ascertain how Pre-Indo-Iranian survived the invasion, gradual replacement and founder effects that happened in terms of male haplogroups after the arrival of late Corded Ware migrants, and that we should probably have to rely on anthropological explanations for language continuity despite genetic replacement, as in the Basque case.

Nevertheless, since we have very few samples, I think we could still see a clear genetic contribution from Yamna to Corded Ware immigrants in the North Caspian region (from Abashevo, in turn a mix of Fatyanovo/Balanovo and Catacomb/Poltavka cultures) in terms of:

  • Ancestral components and PCA in new Sintashta-Petrovka, Andronovo, and/or later samples – similar the ‘steppe’ drift seen in Potapovka relative to Sintashta samples, both formed by incoming Corded Ware migrants – ; and
  • R1b-L23 subclades, either appearing scattered during the Sintashta melting pot (of Abashevo/R1a-Z645 and East Yamna-Poltavka/R1b-Z2103 peoples), or resurging after this period, as we have seen in Pre-Balto-Slavic territory.

This contribution could better explain the obvious language continuity in the region, beautifully complementing the complex anthropological model we have now of archaeological continuity of Sintashta and Potapovka with the previous Poltavka, seen in a similar material and symbolic culture that survived the arrival of newcomers.

A lot of people seem to be looking like crazy since O&M 2018 for some sort of connection between Corded Ware and Yamna migrants in Eastern and Central Europe (wheter in SNP calls of samples published, or among almost forgotten academic papers), either to support the ideas of the 2015 papers – for those who relied on their conclusions and built (even if only mentally) far-fetched migration models around it – , or just because of some sort of absurd continuity theory involving modern R1a-Z645 subclades:

NOTE. The situation we have seen with the hundreds of samples from O&M 2018, and with the recent additional Eastern European samples, depict an unexpected absolutely clear-cut distinction in Y-DNA haplogroups between Corded Ware and Yamna/Bell Beaker: I really can’t see how the situation could be more obvious for everyone, so I doubt any further samples will make certain people change their minds. Their hope is, I guess, that just one sample may give some more oxygen to infinite pet theories, as we are still surprisingly seeing even with reactionary R1b autochthonous continuists in Western Europe…

However, looking into the most likely future for the field, what we should be expecting right now is continuity of Yamna ancestry and lineages in early Proto-Indo-Iranian territory. Since we only have a few samples from Sintashta-Petrovka, Potapovka, and Andronovo, I think there might be a sizeable number of R1b-Z2103 subclades in the territory inhabited by those who – no doubt – spread the language into Central Asia.

Haplogroup_R1b_(Y-DNA)
Modern Y-DNA haplogroup R1b distribution, by Maulucioni at Wikipedia

While full population replacement by R1a-Z93 lineages in the North Caspian region ca. 2000 BC is not impossible, I don’t think it is very likely, since we already know that there are R1b-Z2103 lineages widely distributed in Indo-Iranian-speaking territory, and Z93 is now known to be an older subclade than YFull’s mean formation date suggested (due to the Ukraine_Eneolithic I6561 sample‘s SNP call), so what we can infer now that actually happened in Sintashta -> Andronovo is not exactly the spread of haplogroup Z93 during its formation, but rather a regional reduction in its variability coupled with the expansion of some of its subclades.

The main question, after the South Asia paper is finally published, will then be:

  1. Given that Yamna peoples were an elite group of patrilineally-related families mainly of R1b-L23 subclades:
  2. Accepting that PCA, ADMIXTURE, and other statistical methods are not relevant (alone) for ethnolinguistic identification: e.g. Yamna ‘outliers’ and East Bell Beaker migrants of R1b-L23 lineages without steppe ancestry; N1c1a1a-L392 lineages and Siberian ancestry unrelated to Uralic speakers; R1a-Z645 and steppe ancestry in North-East Europe related to Uralic-speaking cultures
  3. If we find now, as I expect, genetic continuity of east Yamna in Sintashta -> Andronovo (relative to other late Corded Ware peoples), probably including haplogroup R1b-Z2103 mixed with R1a-Z93 before its further reduction of subclades (e.g. to L657) and expansion during its subsequent spread southward…

bronze_age_early_Asia-andronovo
Diachronic map of migrations in Asia ca. 2250-1750 BC

Why exactly do we need Corded Ware to explain migrations of Late Indo-European speakers?

In other words: if we had the data we have today in 2015, would we have a need for Corded Ware to explain Indo-European migrations from the steppe? Are some people so blinded by their will to (appear to) be right in their past interpretations that they can’t just let go?

NOTE. On a side note, wouldn’t it be nice for this paper to publish some other R1b-L23 (x2103) sample – maybe even R1b-L51 – in Yamna, Andronovo, or Afanasevo territory, to end both autochthonous continuity theories (of North-Eastern and Western Europe) at the same time?

I really hope someone in David Reich’s team understands this matter, or else they will still identify Corded Ware as the (now probably ‘a’ instead) vector of expansion of Indo-European languages, and some of us will still have fun for another 2 or 3 years with such conclusions, until someone in the lab realizes that ancestry ≠ population ≠ ethnic identification ≠ language.

NOTE. It seems rather dull to read how people are discussing in the Twitterverse conventional constructs like ‘human race‘ as found in Reich’s op-ed in The New York Times, as if such grandiose semantic discussions had any practical meaning, when basic anthropological questions actually relevant for Genomics, like the essential ancestral component ≠ people tenet seem not to be of interest for anyone in the field….

Since our Indo-European demic difusion model (and its consequences for our reconstruction of North-West Indo-European) and this blog are becoming more and more popular each day – judging by the constant growth in visits in the past 6 months or so – , I guess the simplemindedness and predictability of certain geneticists is benefitting traditional anthropology directly, driving more and more amateur geneticists to look for sound academic models to answer the growing inconsistencies of genetic research.

NOTE. I am not saying the rejection of Corded Ware as spreading Indo-European is definitive. Maybe more samples within some years will depict a clear ancient expansion of Early or Middle Proto-Indo-Europeans from Khvalynsk to the forest-steppe and forest zone, and later with certain Corded Ware migrants into Central Europe, over whose territory a Late Indo-European dialect from Bell Beakers became the superstrate, as some have proposed in the past – e.g. to explain Krahe’s Old European hydronymy. I really doubt you could demonstrate such an old ethnolinguistic identification with a clear, unbroken archaeological trail, though, and we know now that this old hydronymy is probably of Late Indo-European nature (possibly even more recent).

What I am saying is: with the data we have now, it does not make any sense to keep the anthropological models invented by geneticists ex nihilo in 2015, and the hundred different alternative Late Indo-European migration models that arebornwitheachnewpaper.

These Yamna -> Corded Ware migration models didn’t have any sense for me since early 2016, but now after O&M 2017, and especially O&M 2018, I don’t think any geneticist with a little knowledge in Linguistics or Archaeology (if they are decent about their quest for truth in describing ancient European migrations) would buy them, if not for some sort of created ‘tradition’. So let’s ditch Corded Ware as Late Indo-European-speaking, let’s accept that late Corded Ware migrants should most likely be identified as early Uralic speakers, and then future data will tell if we are – again – wrong.

Please, don’t let Genomics become another pseudoscience based solely on Bioinformatics like glottochronology: let anthropologists (preferably mainstream archaeologists, but also the true Indo-Europeanists, linguists) help you interpret your raw data. Don’t deceive yourselves thinking that you have read enough about the Indo-European question, or that you know enough Indo-Europeanists (say what?) to derive your own conclusions.

Use the South Asia paper to begin expressly retracting the Corded Ware mess.

Please pretty please with sugar on top?

Related:

For commenters: this post concerns an anthropological question, and deals with the expansion of Late Proto-Indo-European speakers from Yamna, and the controversy surrounding the role of Corded Ware migrants that a handful of academics propose spread from it, based on a renewed model of Gimbutas’ outdated Kurgan theory and on the so-called ‘Yamnaya’ ancestry.

It happens so that the discussion has turned lately mainly to ancient Y-DNA haplogroups, because they help confirm previous mainstream anthropological models of cultural diffusion and migration. It is obviously not reasonable to judge prehistoric ethnolinguistic migrations from ca. 5,000 years ago based on historical nation-states and ethnic or religious concepts invented since the Middle Ages, coupled with “your” people’s main modern (or your own) paternal lineage.

EDIT (27 MAR 2018): Minor corrections and post made shorter.

Agricultural origins on the Anatolian plateau

anatolia-neolithic-agriculture

New paper (behind paywall) Agricultural origins on the Anatolian plateau, by Baird et al. PNAS (2018), published ahead of print (March 19).

Abstract (emphasis mine):

This paper explores the explanations for, and consequences of, the early appearance of food production outside the Fertile Crescent of Southwest Asia, where it originated in the 10th/9th millennia cal BC. We present evidence that cultivation appeared in Central Anatolia through adoption by indigenous foragers in the mid ninth millennium cal BC, but also demonstrate that uptake was not uniform, and that some communities chose to actively disregard cultivation. Adoption of cultivation was accompanied by experimentation with sheep/goat herding in a system of low-level food production that was integrated into foraging practices rather than used to replace them. Furthermore, rather than being a short-lived transitional state, low-level food production formed part of a subsistence strategy that lasted for several centuries, although its adoption had significant long-term social consequences for the adopting community at Boncuklu. Material continuities suggest that Boncuklu’s community was ancestral to that seen at the much larger settlement of Çatalhöyük East from 7100 cal BC, by which time a modest involvement with food production had been transformed into a major commitment to mixed farming, allowing the sustenance of a very large sedentary community. This evidence from Central Anatolia illustrates that polarized positions explaining the early spread of farming, opposing indigenous adoption to farmer colonization, are unsuited to understanding local sequences of subsistence and related social change. We go beyond identifying the mechanisms for the spread of farming by investigating the shorter- and longer-term implications of rejecting or adopting farming practices.

anatolia-neolithic
Map of central Anatolia showing the principal sites mentioned in the text.

Interesting excerpts:

The persistence of foraging and rejection of farming at Pınarbaşı is also worthy of further consideration. Pınarbaşı’s longevity as a settlement locale in the early Holocene appears to have been based on hunting of wild mammals, wetland exploitation, and significant focus on nut exploitation, all afforded by its ecotonal setting between the hills, plain, and wetland. Perhaps this existing diversity, including nutritious storable plant resources, was a key factor in a lack of interest in adopting cultivation. Another factor may have been a conscious desire to maintain traditional identities and long-standing distinctions with other communities, in part reflected in its particular way of life and its specific connections with particular elements in landscape, for example the almond and terebinth woodlands whose harvests underwrote the continuity of the Pınarbaşı settlement.

The variability in response to the possibilities of early food production in a relatively small geographical area demonstrated here is notable and provides an example useful in evaluating the spread of farming in other regions. It shows the possible role of indigenous foragers, the potential patchwork and diffuse nature of the spread of farming, the lack of homogeneity likely in the communities caught up in the process, the probability of significant continuities in local cultural traditions within the process, and the potentially long-term stable adaptation offered by lowlevel food production. The strength of identities linked to exploitation of particular foods and particular parts of the landscape may have been a major factor contributing to rejection or adoption of food production by indigenous foragers.

The results are also relevant for understanding the processes that underpinned the initial development of farming within the Fertile Crescent itself: that is, the region in which the wild progenitors of the Old World founder crops and stock animals are found. Recent research has rejected the notion of a core area for farming’s first appearance in southwest Asia and demonstrated that farming developed in diverse ways over the Fertile Crescent zone from the southern Levant to the Zagros, very analogous to the situation just described for Central Anatolia (2). Cultivation, herding, and domestication developed in that region, and it seems inescapable that exchange of crops and herded animals occurred between communities (2), involving a spread of farming within the Fertile Crescent, leading eventually to the Neolithic farming package that was so similar across the region and which spread into Europe (5). Central Anatolia was clearly linked to the Fertile Crescent, with significant evidence of exchange and some shared cultural traditions from at least the Epipaleolithic (22). The evidence presented here demonstrates very clearly the movement of crops between settlements and regions in early phases of the Neolithic through exchange, and thus allows us to identify episodes of crop exchange that were probably taking place within the Fertile Crescent itself, but are difficult, if not impossible, to distinguish due to the presence of crop progenitors across much of the region.

A very interesting read in combination with 14C-radiometric data and climatic conditions showing potential triggers of dispersal of Neolithic lifeways from Turkey to Southeast Europe, e.g. Dispersal of Neolithic Lifeways: Absolute Chronology and Rapid Climate Change in Central and West Anatolia, by Lee Clare & Bernhard Weninger, in The Neolithic in Turkey, Vol.6 (2014), Edited by Mehmet Özdogan, Nezih Basgelen, Peter Kuniholm.

anatolian-neolithic-aegean
The Late Neolithic (6600-6000 cal. BC) witnesses the rapid westward dispersal of Neolithic communities, apparently reaching the Aegean in the space of a very short time (ca. 6600 cal. BC). This process is linked to the demand of individuals, groups, and communities for less vulnerable conditions in the face of climate fluctuation associated with RCC. Coastal areas not only offered respite from more frequently occurring physical impacts (extreme winters and high drought risk) in inner Anatolia, they may also have provided refuge for weaker (more vulnerable) social groups (…).

Featured image, from the latter: “In the Early Pottery Neolithic (7000-6600 cal. BC) there occurs a clear break with precedeing (PPN) traditions, attested by abandonment and decreasing size of settlements, albeit that evidence for migration of groups westwards towards the Aegean is still ambiguous (black arrows: human migrations; white arrows: Anatolian obsidian)”

See also:

Genetic ancestry of Hadza and Sandawe peoples reveals ancient population structure in Africa

Open access paper Genetic Ancestry of Hadza and Sandawe Peoples Reveals Ancient Population Structure in Africa, by Shriner, Tekola-Ayele, Adeyemo, & Rotimi, GBE (2018).

Abstract (emphasis mine):

The Hadza and Sandawe populations in present-day Tanzania speak languages containing click sounds and therefore thought to be distantly related to southern African Khoisan languages. We analyzed genome-wide genotype data for individuals sampled from the Hadza and Sandawe populations in the context of a global data set of 3,528 individuals from 163 ethno-linguistic groups. We found that Hadza and Sandawe individuals share ancestry distinct from and most closely related to Omotic ancestry; share Khoisan ancestry with populations such as ≠Khomani, Karretjie, and Ju/’hoansi in southern Africa; share Niger-Congo ancestry with populations such as Yoruba from Nigeria and Luhya from Kenya, consistent with migration associated with the Bantu Expansion; and share Cushitic ancestry with Somali, multiple Ethiopian populations, the Maasai population in Kenya, and the Nama population in Namibia. We detected evidence for low levels of Arabian, Nilo-Saharan, and Pygmy ancestries in a minority of individuals. Our results indicate that west Eurasian ancestry in eastern Africa is more precisely the Arabian parent of Cushitic ancestry. Relative to the Out-of-Africa migrations, Hadza ancestry emerged early whereas Sandawe ancestry emerged late.

Excerpts:

Introduction
In the Hadza population, the distribution of Y chromosomes includes mostly B2 haplogroups, with a smaller number of E1b1a haplogroups, which are common in Niger-Congo-speaking populations, and E1b1b haplogroups, which are common in Cushitic populations (Tishkoff, et al. 2007). In the Sandawe population, E1b1a and E1b1b haplogroups are more common, with lower frequencies of B2 and A3b2 haplogroups (Tishkoff, et al. 2007).

Conclusion
We found that Hadza ancestry diverged early, rather than late. We found evidence for contributions of Cushitic and Niger-Congo ancestries in Tanzania, consistent with the movements of herding and cultivating Cushitic speakers ~4,000 years ago and agricultural Niger-Congo speakers ~2,500 years ago (Newman 1995). However, we did not find evidence of a substantial contribution of Nilo-Saharan ancestry that might have resulted from movement of pastoralist Nilo-Saharan speakers (Newman 1995). We also identified west Eurasian ancestry in eastern and southern African populations more precisely as the Arabian parent of Cushitic ancestry. Finally, our ancestry analyses support the hypothesis that Omotic, Hadza, and Sandawe languages group together, rather than Omotic languages belonging to the Afroasiatic family and Hadza and Sandawe languages belonging to the Khoisan family.

I don’t like linguistic assumptions from admixture analysis; especially from scarce modern samples, as in this case.

Nevertheless, these papers may help clarify the different nature of Omotic and Cushitic among Afroasiatic languages, and thus leave the origin of Afroasiatic either:

a) To the east, with the traditionalist Afroasiatic – Semitic/Hamitic homeland association.

afroasiatic-homeland
Expansion of Afroasiatic

b) To the west, near modern Chadic languages (associated with the expansion of R1b-V88 subclades through a Green Sahara), as I suggested.

Related:

Genomic analysis of Germanic tribes from Bavaria show North-Central European ancestry

antiquity-europe

New open access paper Population genomic analysis of elongated skulls reveals extensive female-biased immigration in Early Medieval Bavaria, by Veeramah, Rott, Groß, et al. PNAS (2018), published ahead of print.

First, a bit of context on the Bavarii:

Europe experienced a profound cultural transformation between Late Antiquity and the Middle Ages that laid the foundations of the modern political, social, and religious landscape. During this period, colloquially known as the “Migration Period,” the Roman Empire gradually dissolved, with 5th and 6th century historiographers and contemporary witnesses describing the formation and migration of numerous Germanic peoples, such as the Goths, Alamanni, Gepids, and Longobards. However, the genetic and social composition of groups involved and the exact nature of these “migrations” are unclear and have been a subject of substantial historical and archaeological debate

In the mid 6th century AD, the historiographer Jordanes and the poet and hagiographer Venantius Fortunatus provide the first mention of a group known as the Baiuvarii that resided in modern day Bavaria. It is likely that this group had already started to form in the 5th century AD, and that it emanated from a combination of the romanized local population of the border province of the former Roman Empire and immigrants from north of the Danube (2). While the Baiuvarii are less well known than some other contemporary groups, an interesting archaeological feature in Bavaria from this period is the presence of skeletons with artificially deformed or elongated skulls.

bavarii-pca
Procrustes-transformed PCA of ancient samples using pseudohaploid calls based on off-target reads using an imputed POPRES modern reference dataset. Blue, green, and red male or female symbols are ancient Bavarian individuals with normal, intermediate, and elongated skulls, respectively. Orange circles are Anglo-Saxon era individuals. Large circles are medians for regions, dots are individuals. CE, central Europe; EE, eastern Europe; NE, northern Europe; NEE, northeastern Europe; NEW, northwestern Europe; SE, southern Europe; SEE, southeast Europe; WE, western Europe. Percentage of variation explained by PCs 1 and 2 for modern populations only is 0.25% and 0.15%.

Abstract (emphasis mine):

Modern European genetic structure demonstrates strong correlations with geography, while genetic analysis of prehistoric humans has indicated at least two major waves of immigration from outside the continent during periods of cultural change. However, population-level genome data that could shed light on the demographic processes occurring during the intervening periods have been absent. Therefore, we generated genomic data from 41 individuals dating mostly to the late 5th/early 6th century AD from present-day Bavaria in southern Germany, including 11 whole genomes (mean depth 5.56×). In addition we developed a capture array to sequence neutral regions spanning a total of 5 Mb and 486 functional polymorphic sites to high depth (mean 72×) in all individuals. Our data indicate that while men generally had ancestry that closely resembles modern northern and central Europeans, women exhibit a very high genetic heterogeneity; this includes signals of genetic ancestry ranging from western Europe to East Asia. Particularly striking are women with artificial skull deformations; the analysis of their collective genetic ancestry suggests an origin in southeastern Europe. In addition, functional variants indicate that they also differed in visible characteristics. This example of female-biased migration indicates that complex demographic processes during the Early Medieval period may have contributed in an unexpected way to shape the modern European genetic landscape. Examination of the panel of functional loci also revealed that many alleles associated with recent positive selection were already at modern-like frequencies in European populations ∼1,500 years ago.

bavarii-admixture
Supervised model-based clustering ADMIXTURE analysis for ancient samples based on phased haplotypes for individual 1,000 bp loci from the 5-Mb neutralome. Analysis is based on the best of 100 runs for K = 8, but NC_EUR is the ancestry summed across 1000 Genomes CEU, 1000 Genomes GBR, and GoNL populations (i.e., it represents a northern/central European ancestry). Blue, green, and red male or female symbols are ancient Bavarian individuals with normal, intermediate, and elongated skulls, respectively.

There is no Y-DNA data to keep confirming the North-Central origin of certain modern European subclades in Central and South-Central Europe.

The potential Ostrogothic sample from Crimea was probably Hunnic, as the paper itself suggests, and both Ostrogoths and Gepids are known to have been allies of the Huns for a long time. It is also a well-known fact that East Germanic tribes migrated south- and eastward through eastern Europe, and then from the steppe westward.

Obviously, the PCA of a late Gepid sample – after a certain number of generations and admixture events with ‘local’ populations during the migrations – , and of a Crimean sample without a clear cultural identification, are of limited value today, until more samples are available.

Hence sadly no valid data yet to add to the debate of East Germanic nature, which mainly concerns its traditionally described origin in Scandinavia – i.e. close to North Germanic dialects – against a different origin (and dialectal branch) within Proto-Germanic territory.

NOTE. Just to be clear for future papers on Germanic tribes, I would expect East Germanic males to show either:
a) mainly R1b-U106, I1, and R1a-Z645 subclades, and to cluster closely to samples of Scandinavia during Antiquity, which would support a Scandinavian origin – a predominance of typically Scandinavian R1a-Z284 subclades would be more indicative of this origin, of course;
b) or mainly R1b-U106, R1b-P312, and I1 subclades and a PCA cluster close to West Germanic tribes, which would challenge its traditional dialectal identification.

I agree with the authors in that a few samples are able to describe certain migratory events, though, such as the emphasized female-biased long-distance migration in Bavaria, as well as the diverse ancestry of women versus men.

Related:

Population turnover in Remote Oceania shortly after initial settlement

oceania-ancient-migration

Open Access article Population Turnover in Remote Oceania Shortly after Initial Settlement, by Lipson, Skoglund, Spriggs, et al. (2018), based on the recent preprint at bioRxiv.

Summary:

Ancient DNA from Vanuatu and Tonga dating to about 2,900–2,600 years ago (before present, BP) has revealed that the “First Remote Oceanians” associated with the Lapita archaeological culture were directly descended from the population that, beginning around 5000 BP, spread Austronesian languages from Taiwan to the Philippines, western Melanesia, and eventually Remote Oceania. Thus, ancestors of the First Remote Oceanians must have passed by the Papuan-ancestry populations they encountered in New Guinea, the Bismarck Archipelago, and the Solomon Islands with minimal admixture [ 1 ]. However, all present-day populations in Near and Remote Oceania harbor >25% Papuan ancestry, implying that additional eastward migration must have occurred. We generated genome-wide data for 14 ancient individuals from Efate and Epi Islands in Vanuatu from 2900–150 BP, as well as 185 present-day individuals from 18 islands. We find that people of almost entirely Papuan ancestry arrived in Vanuatu by around 2300 BP, most likely reflecting migrations a few hundred years earlier at the end of the Lapita period, when there is also evidence of changes in skeletal morphology and cessation of long-distance trade between Near and Remote Oceania [ 2, 3 ]. Papuan ancestry was subsequently diluted through admixture but remains at least 80%–90% in most islands. Through a fine-grained analysis of ancestry profiles, we show that the Papuan ancestry in Vanuatu derives from the Bismarck Archipelago rather than the geographically closer Solomon Islands. However, the Papuan ancestry in Polynesia—the most remote Pacific islands—derives from different sources, documenting a third stream of migration from Near to Remote Oceania

oceania-vanuatu
The population of Vanuatu in the Pacific was largely replaced2,900–2,300 years ago
This second wave of migrants came from New Britain, east of New Guinea
A third wave spread different ancestry to the far-flung islands of Polynesia

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The preferred northwest passage to Scandinavia

Pontus Skoglund writes (and shares publicly) his perspective on early postglacial migrations of hunter-gatherers into Scandinavia, in Northwest Passage to Scandinavia (Nat. Ecol. Evol.): an initial migration from the south and a second coastal migration north of the Scandinavian ice sheet.

He sums up the recently published Open Access paper Population genomics of Mesolithic Scandinavia: Investigating early postglacial migration routes and high-latitude adaptation, by Günther, Malmström , Svensson, Omrak, et al. PLoS Biol (2018) 16(1): e2003703, based on preprint at BioRxiv Genomics of Mesolithic Scandinavia reveal colonization routes and high-latitude adaptation (2017).

Abstract:

Scandinavia was one of the last geographic areas in Europe to become habitable for humans after the Last Glacial Maximum (LGM). However, the routes and genetic composition of these postglacial migrants remain unclear. We sequenced the genomes, up to 57× coverage, of seven hunter-gatherers excavated across Scandinavia and dated from 9,500–6,000 years before present (BP). Surprisingly, among the Scandinavian Mesolithic individuals, the genetic data display an east–west genetic gradient that opposes the pattern seen in other parts of Mesolithic Europe. Our results suggest two different early postglacial migrations into Scandinavia: initially from the south, and later, from the northeast. The latter followed the ice-free Norwegian north Atlantic coast, along which novel and advanced pressure-blade stone-tool techniques may have spread. These two groups met and mixed in Scandinavia, creating a genetically diverse population, which shows patterns of genetic adaptation to high latitude environments. These potential adaptations include high frequencies of low pigmentation variants and a gene region associated with physical performance, which shows strong continuity into modern-day northern Europeans.

The ice sheet distribution – which did not improve nuch for thousands of years – was clearly the greatest barrier for potential migrations in the region.

baltic-sea-10000-bc
Baltischer Süßwassersee Vorläufer der Ostsee vor 12.000 Jahren, by Juschki and Koyos at Wikipedia

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