Showing posts with label Balkans. Show all posts
Showing posts with label Balkans. Show all posts

May 19, 2016

35,000 year old mtDNA haplogroup U6 from Romania

I wouldn't be very surprised if many of the markers supposedly signifying recent gene flow Africa and Eurasia were actually quite old in Eurasia. The trouble is that reports of such gene flow were often based on simply observing that marker "X" occurs at a higher frequency in Africa than in Eurasia, so a common sense explanation is that it reflects limited recent gene flow between the continents. But, it is now known that common sense is not always the best guide, as e.g., ancient Europeans had mtDNA haplogroup M (in the past considered evidence of Asian admixture), Y-chromosome haplogroup C (ditto), and now U6.

The same should also apply to the Middle East where there has been admixture with Africans since the Islamic period at least. The existence of such admixture does not mean that every single lineage that occurs at low frequency in the Middle East and high frequency in Africa is diagnostic of this later period of admixture. Some of them could well be relics of old Middle Eastern populations. Who knows what people inhabited the presently inhospitable landscape of the Saharan-Arabian desert zone? The living populations can certainly make no claim to being the first ones there, but the genetic heritage of those earlier occupants may still persist in them in traces.

Similarly for the New World; in that case, there is a better case that European-looking lineages are indeed due to the colonization of the Americas over the last five centuries. However, that does not mean that all of them are, and we should be mindful of the possibility of pre-Columbian contact between the Old and New worlds.

Scientific Reports 6, Article number: 25501 (2016)

The mitogenome of a 35,000-year-old Homo sapiens from Europe supports a Palaeolithic back-migration to Africa

M. Hervella et al.

After the dispersal of modern humans (Homo sapiens) Out of Africa, hominins with a similar morphology to that of present-day humans initiated the gradual demographic expansion into Eurasia. The mitogenome (33-fold coverage) of the Peştera Muierii 1 individual (PM1) from Romania (35 ky cal BP) we present in this article corresponds fully to Homo sapiens, whilst exhibiting a mosaic of morphological features related to both modern humans and Neandertals. We have identified the PM1 mitogenome as a basal haplogroup U6*, not previously found in any ancient or present-day humans. The derived U6 haplotypes are predominantly found in present-day North-Western African populations. Concomitantly, those found in Europe have been attributed to recent gene-flow from North Africa. The presence of the basal haplogroup U6* in South East Europe (Romania) at 35 ky BP confirms a Eurasian origin of the U6 mitochondrial lineage. Consequently, we propose that the PM1 lineage is an offshoot to South East Europe that can be traced to the Early Upper Paleolithic back migration from Western Asia to North Africa, during which the U6 lineage diversified, until the emergence of the present-day U6 African lineages.

Link

May 09, 2014

Ancient DNA from the Balkans (Iron Age Thrace)

A new paper in PLoS Genetics presents new data from two Iron Age Thracian individuals and puts the Sardinian-ness of Oetzi in new context. The authors write:
The results of the analyses including additional ancient genomes provide mounting evidence that the Iceman's genetic affinity with Sardinians reflects an ancestry component that was widespread in Europe during the Neolithic. Despite their different geographic origins, both the Swedish farmer gok4 and the Thracian P192-1 closely resemble the Iceman in their relationship with Sardinians, making it unlikely that all three individuals were recent migrants from Sardinia. Furthermore, P192-1 is an Iron Age individual from well after the arrival of the first farmers in Southeastern Europe (more than 2,000 years after the Iceman and gok4), perhaps indicating genetic continuity with the early farmers in this region. The only non-HG individual not following this pattern is K8 from Bulgaria. Interestingly, this individual was excavated from an aristocratic inhumation burial containing rich grave goods, indicating a high social standing, as opposed to the other individual, who was found in a pit [15]. However, the DNA damage pattern of this individual does not appear to be typical of ancient samples (Table S4 in [15]), indicating a potentially higher level of modern DNA contamination. On the other hand, the Swedish and the Iberian hunter-gatherers show congruent patterns of relatedness to the modern populations of Northern Europe, which is consistent with the previous results using those samples.
Also of interest, given previous suggestions that the Iceman had more Neandertal ancestry than modern Europeans:
However, all D-tests involving another non-African population do not significantly deviate from zero, suggesting that the Iceman genome contains levels of archaic ancestry that are comparable to that of other non-African populations.
A model of European history is seen on the left. Some details are probably incorrect (e.g., Sardinian Neolithic probably followed the Cardial/Mediterranean route rather the one shown in C). There are no good ancient DNA from Cardial Neolithic farmers, so the fact that Sardinians are similar to the Iron Age Bulgarian, the Stuttgart LBK German, and the Swedish TRB farmers may mean that the Mediterranean/Cardial farmers were related to the ones that went into Europe following the inland route from the Balkans.

In any case, the fact that there are now data from Bulgaria is great, because it means that southern Europe is not hopeless for ancient DNA preservation and hopefully more is on its way.

UPDATE: Did anyone see a link to the new data? It appears that there are only ~1,000 SNPs in common with the HGDP. [A link to the data will become available at the Bustamante lab website]

PLoS Genet 10(5): e1004353. doi:10.1371/journal.pgen.1004353

Population Genomic Analysis of Ancient and Modern Genomes Yields New Insights into the Genetic Ancestry of the Tyrolean Iceman and the Genetic Structure of Europe

Martin Sikora et al.

Genome sequencing of the 5,300-year-old mummy of the Tyrolean Iceman, found in 1991 on a glacier near the border of Italy and Austria, has yielded new insights into his origin and relationship to modern European populations. A key finding of that study was an apparent recent common ancestry with individuals from Sardinia, based largely on the Y chromosome haplogroup and common autosomal SNP variation. Here, we compiled and analyzed genomic datasets from both modern and ancient Europeans, including genome sequence data from over 400 Sardinians and two ancient Thracians from Bulgaria, to investigate this result in greater detail and determine its implications for the genetic structure of Neolithic Europe. Using whole-genome sequencing data, we confirm that the Iceman is, indeed, most closely related to Sardinians. Furthermore, we show that this relationship extends to other individuals from cultural contexts associated with the spread of agriculture during the Neolithic transition, in contrast to individuals from a hunter-gatherer context. We hypothesize that this genetic affinity of ancient samples from different parts of Europe with Sardinians represents a common genetic component that was geographically widespread across Europe during the Neolithic, likely related to migrations and population expansions associated with the spread of agriculture.

Link

January 08, 2014

6,500-year old tin bronze from Serbia

Antiquity Volume: 87 Number: 338 Page: 1030–1045

Tainted ores and the rise of tin bronzes in Eurasia, c. 6500 years ago

Miljana Radivojević et al.

The earliest tin bronze artefacts in Eurasia are generally believed to have appeared in the Near East in the early third millennium BC. Here we present tin bronze artefacts that occur far from the Near East, and in a significantly earlier period. Excavations at Pločnik, a Vinča culture site in Serbia, recovered a piece of tin bronze foil from an occupation layer dated to the mid fifth millennium BC. The discovery prompted a reassessment of 14 insufficiently contextualised early tin bronze artefacts from the Balkans. They too were found to derive from the smelting of copper-tin ores. These tin bronzes extend the record of bronze making by c. 1500 years, and challenge the conventional narrative of Eurasian metallurgical development.

Link

August 04, 2013

Archaeology: The milk revolution

A couple of interesting quotes from this story:
That next step happened slowly, and it seems to have required the spread of lactase persistence. The LP allele did not become common in the population until some time after it first emerged: Burger has looked for the mutation in samples of ancient human DNA and has found it only as far back as 6,500 years ago in northern Germany. 
 ...
Some of the LeCHE participants are now probing further back in time, as part of a project named BEAN (Bridging the European and Anatolian Neolithic), which is looking at how the first farmers and herders made their way into Europe. Burger, Thomas and their BEAN collaborators will be in Turkey this summer, tracing the origins of the Neolithic using computer models and ancient-DNA analysis in the hope of better understanding who the early farmers were, and when they arrived in Europe.

May 08, 2013

The Geography of Recent Genetic Ancestry across Europe (Ralph and Coop 2013)

This paper first came out last July on the arXiv and went through four versions there before its final form which has now appeared in PLoS Biology. It's great that its early release allowed other people to read it without having to wait for the completion of the peer review process.

I think that this is a good model: journals have the right and obligation to subject papers to close scrutiny according to their own procedures, but this process ought not interfere with the early availability of research results or the ability of anyone other than the chosen reviewers to comment on new results.

PLoS Biol 11(5): e1001555. doi:10.1371/journal.pbio.1001555

The Geography of Recent Genetic Ancestry across Europe

Peter Ralph, Graham Coop

The recent genealogical history of human populations is a complex mosaic formed by individual migration, large-scale population movements, and other demographic events. Population genomics datasets can provide a window into this recent history, as rare traces of recent shared genetic ancestry are detectable due to long segments of shared genomic material. We make use of genomic data for 2,257 Europeans (in the Population Reference Sample [POPRES] dataset) to conduct one of the first surveys of recent genealogical ancestry over the past 3,000 years at a continental scale. We detected 1.9 million shared long genomic segments, and used the lengths of these to infer the distribution of shared ancestors across time and geography. We find that a pair of modern Europeans living in neighboring populations share around 2–12 genetic common ancestors from the last 1,500 years, and upwards of 100 genetic ancestors from the previous 1,000 years. These numbers drop off exponentially with geographic distance, but since these genetic ancestors are a tiny fraction of common genealogical ancestors, individuals from opposite ends of Europe are still expected to share millions of common genealogical ancestors over the last 1,000 years. There is also substantial regional variation in the number of shared genetic ancestors. For example, there are especially high numbers of common ancestors shared between many eastern populations that date roughly to the migration period (which includes the Slavic and Hunnic expansions into that region). Some of the lowest levels of common ancestry are seen in the Italian and Iberian peninsulas, which may indicate different effects of historical population expansions in these areas and/or more stably structured populations. Population genomic datasets have considerable power to uncover recent demographic history, and will allow a much fuller picture of the close genealogical kinship of individuals across the world.

Link

March 18, 2013

New Neandertal remains from Mani

The age and cave origin of these remains may conceivably make them useful for ancient DNA studies.

Journal of Human Evolution doi:10.1016/j.jhevol.2013.02.002

New Neanderthal remains from Mani peninsula, Southern Greece: The Kalamakia Middle Paleolithic cave site

Katerina Harvati et al.

The Kalamakia cave, a Middle Paleolithic site on the western coast of the Mani peninsula, Greece, was excavated in 1993–2006 by an interdisciplinary team from the Ephoreia of Paleoanthropology and Speleology (Greek Ministry of Culture) and the Muséum national d'Histoire naturelle (Paris). The site is dated to between ca. 100,000 and >39,000 years BP (Before Present) and has yielded Mousterian lithics, a rich fauna, and human remains from several layers. The latter include 10 isolated teeth, a cranial fragment and three postcranial elements. The remains represent at least eight individuals, two of them subadults, and show both carnivore and anthropogenic modifications. They can be identified as Neanderthal on the basis of diagnostic morphology on most specimens. A diet similar to that of Neanderthals from mixed habitat is suggested by our analysis of dental wear (occlusal fingerprint analysis) and microwear (occlusal texture microwear analysis), in agreement with the faunal and palynological analyses of the site. These new fossils significantly expand the Neanderthal sample known from Greece. Together with the human specimens from Lakonis and Apidima, the Kalamakia human remains add to the growing evidence of a strong Neanderthal presence in the Mani region during the Late Pleistocene.

Link

February 11, 2013

Human mobility at the start of Balkan Neolithic

Link to press release.

PNAS doi: 10.1073/pnas.1211474110

Strontium isotopes document greater human mobility at the start of the Balkan Neolithic

Dušan Borić and T. Douglas Price

Questions about how farming and the Neolithic way of life spread across Europe have been hotly debated topics in archaeology for decades. For a very long time, two models have dominated the discussion: migrations of farming groups from southwestern Asia versus diffusion of domesticates and new ideas through the existing networks of local forager populations. New strontium isotope data from the Danube Gorges in the north-central Balkans, an area characterized by a rich burial record spanning the Mesolithic–Neolithic transition, show a significant increase in nonlocal individuals from ∼6200 calibrated B.C., with several waves of migrants into this region. These results are further enhanced by dietary evidence based on carbon and nitrogen isotopes and an increasingly high chronological resolution obtained on a large sample of directly dated individuals. This dataset provides robust evidence for a brief period of coexistence between indigenous groups and early farmers before farming communities absorbed the foragers completely in the first half of the sixth millennium B.C.

Link

February 07, 2013

Balanica BH-1: 397-525 thousand years old

The occurrence of derived Neandertal traits in Sima vs. their absence in penecontemporaneous samples from  southeastern Europe is fairly interesting. It might suggest that the Neandertal suite of traits first appeared in western Europe.

We tend to think of Old World H. heidelbergensis as a parental species which produced -at least in the western part of the Old World- two descendant species, sapiens and neanderthalensis, but clearly that is not the whole story. Unrelated to the current paper, but perhaps worthy of note is that these two descendant species make their appearance far apart in time, with Neandertaloid traits already in evidence very early in Europe, and modern human ones late in east Africa. We may wonder about what was taking place in the temporal gap between 600 and 200 thousand years ago, and the spatial gap between Europe and Africaa.

PLoS ONE 8(2): e54608. doi:10.1371/journal.pone.0054608

New Radiometric Ages for the BH-1 Hominin from Balanica (Serbia): Implications for Understanding the Role of the Balkans in Middle Pleistocene Human Evolution

William J. Rink et al.

Newly obtained ages, based on electron spin resonance combined with uranium series isotopic analysis, and infrared/post-infrared luminescence dating, provide a minimum age that lies between 397 and 525 ka for the hominin mandible BH-1 from Mala Balanica cave, Serbia. This confirms it as the easternmost hominin specimen in Europe dated to the Middle Pleistocene. Inferences drawn from the morphology of the mandible BH-1 place it outside currently observed variation of European Homo heidelbergensis. The lack of derived Neandertal traits in BH-1 and its contemporary specimens in Southeast Europe, such as Kocabaş, Vasogliano and Ceprano, coupled with Middle Pleistocene synapomorphies, suggests different evolutionary forces acting in the east of the continent where isolation did not play such an important role during glaciations.

Link

January 17, 2013

Moldavian Y-chromosomes

The most salient feature is probably the absence of E-V13 (which is modal in Balkan populations) in Ukrainians. The position of Moldavians in the MDS plot of haplogroup frequencies is as expected, with a clear differentiation of "southern" populations from Italy, Greece/Albania, and Anatolia, and "northern" ones from the west Balkans (ex-Yugoslavs) and eastern Europe: Moldavians occupy an intermediate position along this second group of populations.


PLoS ONE 8(1): e53731. doi:10.1371/journal.pone.0053731

Paleo-Balkan and Slavic Contributions to the Genetic Pool of Moldavians: Insights from the Y Chromosome

Alexander Varzari et al.

Moldova has a rich historical and cultural heritage, which may be reflected in the current genetic makeup of its population. To date, no comprehensive studies exist about the population genetic structure of modern Moldavians. To bridge this gap with respect to paternal lineages, we analyzed 37 binary and 17 multiallelic (STRs) polymorphisms on the non-recombining portion of the Y chromosome in 125 Moldavian males. In addition, 53 Ukrainians from eastern Moldova and 54 Romanians from the neighboring eastern Romania were typed using the same set of markers. In Moldavians, 19 Y chromosome haplogroups were identified, the most common being I-M423 (20.8%), R-M17* (17.6%), R-M458 (12.8%), E-v13 (8.8%), R-M269* and R-M412* (both 7.2%). In Romanians, 14 haplogroups were found including I-M423 (40.7%), R-M17* (16.7%), R-M405 (7.4%), E-v13 and R-M412* (both 5.6%). In Ukrainians, 13 haplogroups were identified including R-M17 (34.0%), I-M423 (20.8%), R-M269* (9.4%), N-M178, R-M458 and R-M73 (each 5.7%). Our results show that a significant majority of the Moldavian paternal gene pool belongs to eastern/central European and Balkan/eastern Mediterranean Y lineages. Phylogenetic and AMOVA analyses based on Y-STR loci also revealed that Moldavians are close to both eastern/central European and Balkan-Carpathian populations. The data correlate well with historical accounts and geographical location of the region and thus allow to hypothesize that extant Moldavian paternal genetic lineages arose from extensive recent admixture between genetically autochthonous populations of the Balkan-Carpathian zone and neighboring Slavic groups.

Link

December 30, 2012

Trojan pottery across the Bronze/Iron Age boundary

TROY POTTERY HOLDS A KEY TO THE GREAT BRONZE AGE COLLAPSE
Before the sack of Troy, the city looked east towards the powerful Hittite Empire. But this political powerhouse collapsed around the time that Troy was destroyed. Grave says the post-conflict pottery is Balkan in style because the Trojans were keen to align themselves with the people there, who had become the new political elite and powerbase in the region.

The collapse of the Late Bronze Age political and economic structures of the eastern Mediterranean undercut elite production spheres serving this network at Troy.

The people of Early Iron Age Troy shifted their focus to elaborating their own household ceramic traditions to re-establish their role in newly configured social and economic networks that now looked to the Balkans rather than the Hittite Anatolia.
Related:



Journal of Archaeological Science Available online 12 November 2012

Cultural dynamics and ceramic resource use at Late Bronze Age/Early Iron Age Troy, northwestern Turkey

Peter Grave et al.

Changes in resource use over time can provide insight into technological choice and the extent of long term stability in cultural practices. In this paper we re-evaluate the evidence for a marked demographic shift at the inception of the Early Iron Age at Troy by applying a robust macroscale analysis of changing ceramic resource use over the Late Bronze and Iron Age. We use a combination of new and legacy analytical datasets (NAA and XRF), from excavated ceramics, to evaluate the potential compositional range of local resources (based on comparisons with sediments from within a 10 kilometer site radius). Results show a clear distinction between sediment-defined local and non-local ceramic compositional groups. Two discrete local ceramic resources have been previously identified and we confirm a third local resource for a major class of EIA handmade wares and cooking pots. This third source appears to derive from a residual resource on the Troy peninsula (rather than adjacent alluvial valleys). The presence of a group of large and heavy pithoi among the non-local groups raises questions about their regional or maritime origin.

Link

December 10, 2012

Roma origins once more (Moorjani et al. 2012)

I had first noticed that this new paper by Moorjani et al. was referenced by Loh et al., and it has now been posted on arXiv. In the last week, a couple of other papers on the same topic (Mendizabal et al. on autosomal DNA and Rai et al. on a Y-chromosome founder lineage) have also appeared.

All three studies appear to converge on NW India as the place of origin of the European Roma, and on a recent admixture between this "Proto-Roma" population and Europeans. It will be interesting to see if there are any substantial differences between Moorjani et al. and Mendizabal et al. in the reconstruction of Roma origins. There is also an appendix on updates to rolloff and other topics of a technical nature that ought to be useful to readers irrespective of their interest in this particular population.

It'll probably take me a while to digest everything in this paper, but I will make one quick observation after (virtually) leafing through the article; the observation that {CEU, ANI} form a clade with Adygei as an outgroup is used to infer admixture proportions. I recently had a blog post on the differential relationship of ANI to Caucasus populations, in which I showed that while D(CEU, Adygei; South Asian, Onge) was positive, and significant in some cases -- indicating CEU being more closely related to ANI (Ancestral North Indians) than Adygei -- the reverse was the case for D(CEU, Georgian/Lezgin; South Asian, Onge).

A second observation was inspired by the following figure:


High IBD sharing with Romanians makes sense, because there is good evidence (e.g., presence of Y-haplogroup E-V13) that the Roma picked up European ancestry in the Balkans. So, I'm fairly sure that we are seeing a real signal that the Roma have Romanian-like recent European ancestors. But, we ought to be vigilant, because it is possible that some Romanians may have Roma ancestry too!  This was the case in a couple of individuals from the Romanian sample of Behar et al. (2010).

This is a more general issue: IBD sharing occasionally involves strictly -or mostly- unidirectional gene flow,  e.g., sharing between European and African Americans largely went EA->AA way, so an AA sharing with a EA more often than not involves EA->AA gene flow.

But, in other cases, the direction of gene flow is more obscure (so, e.g., sharing between German, Magyar, and Slavic speakers, and Jews in the old Austro-Hungarian Empire). This issue often comes up in the genealogical community, with a typical example being a couple of individuals (let's call them Klaus and Mikolaj) discovering a shared IBD segment, and Klaus thinking he's found a Polish ancestor, and Mikolaj a German one.

In any case, as the authors themselves note it will be interesting to use more European reference populations, and this might indicate whether they picked up European ancestry in one particular region, carrying it with them as they expanded into the Balkans and beyond, or whether they picked it up by interacting with different host populations (e.g., Greek Gypsies with Greeks, Romanian Gypsies with Romanians, and so on).


arXiv:1212.1696 [q-bio.PE]

Reconstructing Roma history from genome-wide data

Priya Moorjani et al.

The Roma people, living throughout Europe, are a diverse population linked by the Romani language and culture. Previous linguistic and genetic studies have suggested that the Roma migrated into Europe from South Asia about 1000-1500 years ago. Genetic inferences about Roma history have mostly focused on the Y chromosome and mitochondrial DNA. To explore what additional information can be learned from genome-wide data, we analyzed data from six Roma groups that we genotyped at hundreds of thousands of single nucleotide polymorphisms (SNPs). We estimate that the Roma harbor about 80% West Eurasian ancestry-deriving from a combination of European and South Asian sources- and that the date of admixture of South Asian and European ancestry was about 850 years ago. We provide evidence for Eastern Europe being a major source of European ancestry, and North-west India being a major source of the South Asian ancestry in the Roma. By computing allele sharing as a measure of linkage disequilibrium, we estimate that the migration of Roma out of the Indian subcontinent was accompanied by a severe founder event, which we hypothesize was followed by a major demographic expansion once the population arrived in Europe.

Link

December 06, 2012

Romani origins and admixture (Mendizabal et al.)

I will comment on the paper in this space after I read it. For the time being here's a link to the press release.
"From a genome-wide perspective, Romani people share a common and unique history that consists of two elements: the roots in northwestern India and the admixture with non-Romani Europeans accumulating with different magnitudes during the out-of-India migration across Europe," Kayser said. "Our study clearly illustrates that understanding the Romani's genetic legacy is necessary to complete the genetic characterization of Europeans as a whole, with implications for various fields, from human evolution to the health sciences."
The results seem to complement a recent Y-chromosome study of the major founder lineage of European Roma H-M82.

Current Biology dx.doi.org/10.1016/j.cub.2012.10.039

Reconstructing the Population History of European Romani from Genome-wide Data

Isabel Mendizabal et al.

The Romani, the largest European minority group with approximately 11 million people [1], constitute a mosaic of languages, religions, and lifestyles while sharing a distinct social heritage. Linguistic [2] and genetic [3, 4, 5, 6, 7 and 8] studies have located the Romani origins in the Indian subcontinent. However, a genome-wide perspective on Romani origins and population substructure, as well as a detailed reconstruction of their demographic history, has yet to be provided. Our analyses based on genome-wide data from 13 Romani groups collected across Europe suggest that the Romani diaspora constitutes a single initial founder population that originated in north/northwestern India ∼1.5 thousand years ago (kya). Our results further indicate that after a rapid migration with moderate gene flow from the Near or Middle East, the European spread of the Romani people was via the Balkans starting ∼0.9 kya. The strong population substructure and high levels of homozygosity we found in the European Romani are in line with genetic isolation as well as differential gene flow in time and space with non-Romani Europeans. Overall, our genome-wide study sheds new light on the origins and demographic history of European Romani.

Link

December 02, 2012

Talk by Christina Papageorgopoulou on Mesolithic/Neolithic Greek DNA

The talk is in Greek but the slides are mostly in English. This is related to the BEAN Project which I have covered before in this blog.

She discusses various aspects of DNA preservation (poorer than Central Europe/Siberia due to higher temperature, and potentially because non-freshly excavated samples were tested) which appears to be variable across sites.

A screenshot of haplogroups detected so far (X, K, J, H, T). Interestingly, the X haplogroup only comes from Franchthi cave, although Dr. Papageorgopoulou cautions against very simple interpretations of this fact.



She mentions a complete absence of haplogroup U in all her samples so far (either Mesolithic or Neolithic), in contrast to Central European Mesolithic samples where it predominates.

Fst values between Greek Mesolithic/Neolithic samples and Central European Mesolithic ones are high (~0.2), but with Linearbandkeramik they are very low (0.03-0.05).



I'm looking forward to the first publication of these important results, and hopefully they will be soon supplemented with results from Bulgaria and Anatolia that are also to be studied as part of the BEAN Project.

November 23, 2012

The comings and goings of Near Eastern and European domestic pigs (Ottoni et al. 2012)

This is an excellent paper whose findings re: pig domestication seem to parallel many of my own observations regarding the flow of human populations. It is open access, so you can read it for yourselves, but the following figure illustrates the situation admirably:


The left-right arrangement of the columns corresponds to a west-east longitude across West Asia. It can be easily seen that some of the early domestic samples (yellow, bottom row) are concentrated in the west (Y1 haplotype), while others (blue, Arm1T) in the east.

Neolithic European samples possessed the Y1 haplotype, but lacked the Arm1T one. So, the authors conclude that:
The ancient Anatolian data presented here reveal that both wild and possibly domestic Neolithic pigs (identified using traditional metrics) possessed Y1 haplotypes ... The presence of these lineages corroborates the supposition that the earliest domestic pigs in Europe originated from populations originally domesticated in the Near East, conclusively linking the Neolithization of Europe with Neolithic cultures of western Anatolia (Larson et al. 2007a; Haak et al. 2010).
I have repeatedly highlighted the "puzzle" of the early European Neolithic: the signature Y-haplogroup G2a was unaccompanied by other common Near Eastern lineages, and the modal "West Asian" ancestral component in present-day West Asian populations seems to have been absent in early Neolithic samples, which were dominated by a "Sardinian-like" population. I have argued that this meant that the European Neolithic was drawn from a limited founder source that was more "Mediterranean/Southern" autosomally than "West Asian", at least in terms of the components identified by the Dodecad Project.

In Europe itself, the early Near Eastern domestic pigs were replaced by European ones:

Ancient DNA extracted from early Neolithic domestic pigs in Europe resolved this paradox by demonstrating that early domestic pigs in the Balkans and central Europe shared haplotypes with modern Near Eastern wild boar (Larson et al. 2007a). The absence of Near Eastern haplotypes in pre-Neolithic European wild boar suggested that early domestic pigs in Europe must have been introduced from the Near East by the mid 6th millennium BC before spreading to the Paris basin by the early 4th millennium BC (Larson et al. 2007a). 
By 3,900 BC, however, virtually all domestic pigs in Europe possessed haplotypes  from an indigenous European domestication process (Larson et al. 2007a) only found in European wild boar. This genetic turnover may have resulted from the accumulated introgression of local female wild boar into imported domestic stocks, or from an indigenous European domestication process (Larson et al. 2007a).
We have seen that early Neolithic domestic pigs came from Western Anatolia, but apparently these did not last, but were replaced in Europe by pigs carrying mtDNA of European wild boar. An additional possibility is that the European wild boar were better adapted to local conditions in Europe, so the stock of European farmers gradually became "local" due to artificial/natural selection favoring the local "European" type. It might also be that in accordance with Bergmann's rule, European-descended pigs were simply bigger, and thus more economically productive.

In any case, the interesting thing is that pigs carrying the "European" haplotype went the other way, crossing from Europe to Asia. The beginning of this process seems to have occurred in the Middle Bronze Age:

The temporal and geographic distribution of genetic haplotypes presented in our study demonstrates that the first AMS dated pig with European ancestry (haplotype A) appeared almost 1,000 years earlier than the Armenian samples in a Late Bronze Age context (~1,600-1,440 BC) at Lidar Höyük (fig. 1). An even earlier Middle Bronze Age specimen from the same site also possessed a European signature, but a direct
AMS date for this specimen could not be obtained.
I have written how increased mobility and long-range networks associated with the new metallurgical class facilitated commerce during the Bronze Age. The authors suggest the possibility of Minoan-Mycenaean/Hittite involvement during the Bronze Age, which are certainly plausible conduits for European pigs to have crossed the Aegean at this time. But, as you can see from the figure, the "European" pigs are still outliers during the Middle and Bronze Ages, but become common in the Iron Age sample from Lidar Höyük, and eventually replacing local types throughout Anatolia and Armenia, but, apparently, not Iran:
The frequency of pigs with European ancestry increased rapidly from the 12th century BC, and by the 5th century AD domestic pigs exhibiting a Near Eastern genetic signature had all but disappeared across Anatolia and the southern Caucasus. Though we did not detect European signatures in the ancient Iranian samples (fig. 1), the eastward spread of European lineages may have continued into Iran later than the Iron Age since European lineages have been found in wild caught modern Iranian samples (Larson et al. 2007a).
Of course a 12th century BC increase in European domestic pigs is entirely consistent -chronologically- with the Phrygian/Armenian settlement in Anatolia, and this association is further reinforced by the lack of European signatures in pigs from Iran where Phrygo-Armenians did not settle. The increase in European pigs could later be mediated by the Greek colonization, and the increase in trade during antiquity, just as trade would later introduce East Asian pig DNA into Europe.

The beautiful temporal transect presented in the Figure may also prove useful for students of ancient human DNA. I'd love to see how humans living close to sites #14-16, dominated by Arm1T haplotypes throughout history might differ from those of Neolithic West Anatolia, and whether the "mixed" Iron Age sample from Lidar Höyük shows evidence of the arrival of European-like human populations to accompany the European pigs.

Mol Biol Evol (2012) doi: 10.1093/molbev/mss261

Pig domestication and human-mediated dispersal in western Eurasia revealed through ancient DNA and geometric morphometrics

Claudio Ottoni et al.

Zooarcheological evidence suggests that pigs were domesticated in Southwest Asia ∼8,500 BC. They then spread across the Middle and Near East and westward into Europe alongside early agriculturalists. European pigs were either domesticated independently or appeared so as a result of admixture between introduced pigs and European wild boar. These pigs not only replaced those with Near Eastern signatures in Europe, they subsequently also replaced indigenous domestic pigs in the Near East. The specific details of these processes, however, remain unknown. To address questions related to early pig domestication, dispersal, and turnover in the Near East, we analyzed ancient mitochondrial DNA and dental geometric morphometric variation in 393 ancient pig specimens representing 48 archeological sites (from the Pre-Pottery Neolithic to the Medieval period) from Armenia, Cyprus, Georgia, Iran, Syria and Turkey. Our results firstly reveal the genetic signature of early domestic pigs in Eastern Turkey. We also demonstrate that these early pigs differed genetically from those in western Anatolia that were introduced to Europe during the Neolithic expansion. In addition, we present a significantly more refined chronology for the introduction of European domestic pigs into Asia Minor that took place during the Bronze Age, nearly 1,000 years earlier than previously detected. By the 5th century AD, European signatures completely replaced the endemic lineages possibly coinciding with the demographic and societal changes during the Anatolian Bronze and Iron Ages.

Link

November 10, 2012

Investigating East Asian admixture in Balkans/Anatolia/Caucasus

I used ALDER with a dataset of populations from the Balkans, Anatolia, and Caucasus, using the She, Japanese, Miaozu, and Dai as East Asian references. A few caveats for this analysis:

  1. Some populations may possess "South Asian" admixture which may be mistaken for East Asian
  2. Populations differ in the number of SNPs used in the analysis; for example, the Armenian_D sample includes mostly Family Finder data which has a smaller overlap with the SNP set used
  3. Populations differ in the number of individuals used, from a low of 5 (e.g., Turkish_Cypriot) to a high of 45 (Armenian_D)
I have also added the HGDP Europeans to the analysis. The results can be seen below; I have made bold those rows where all estimates are at least one standard error above zero, and bold/big those where the estimates are they are two standard errors above zero. I consider the latter to be the most reliable.



I have already discussed the Turkish signal of admixture at length elsewhere. I will note that the Iranian_D sample produces similar or younger admixture dates, which would make sense, given the fact that the Iranians came under control of the Mongols, while the successes of the latter in Anatolia were short-lived.

A very interesting signal is that of the North Ossetians which show admixture ~9-10 centuries ago. This seems to have occurred a little after the foundation of the kingdom of Alania, and I think it makes excellent sense to view it as the signal of Eurasian nomads (who must have carried some East Asian admixture at that time) intermingling with pre-Iranic local Caucasus populations, Two other populations from the Caucasus, the Georgians and Lezgins (and also the Abkhaz and Chechens) show earlier admixture signals that could very well date to the period of east-west Eurasian migrations inaugurated by the Huns, although a possible Sassanian origin of such influence cannot be overlooked.

The Kurds are another interesting case where the Dodecad sample and the Yunusbayev et al  sample produce very different dates. The different number of SNPs may be at play here, or it may be that some Dodecad participants have recent Turkish ancestry that cause the admixture date average to appear lower, although the globe13 analysis suggests that the "East Asian" found here may be in fact "South Asian".

It seems to me that with large, dense, and well-curated sample sets from several of these populations, their admixture dynamics will become more distinct.

August 24, 2012

Proto-Indo-European homeland in Neolithic Anatolia (Bouckaert et al.)

A new paper in Science uses Bayesian phylogeographic methods to model the spatial expansion of Indo-European languages from their Anatolian homeland. An informative video shows how the authors estimate the process took place across space and time:



There is also a podcast with Q.D. Atkinson on the new study, as well as a website by the authors on their research; the FAQ/Controversies section seems particularly useful.

I don't hold high hopes that, despite the mounting evidence, this will dissuade people from arguing for a steppe PIE origin. And, it shouldn't. Only a vigorous debate will resolve the issue conclusively. And, since IE languages appear on the archaeological record long after their split under any scenario, this may be one of those problems that will never be solved to everyone's satisfaction.

I don't agree with all the details of the authors' model, but certainly they place the PIE homeland near to where I believe it was. Resistance to an Anatolian origin will become more convincing if adherents of different homeland solutions manage to put their ideas in quantitative form. Expert opinion is valuable, but very knowledgeable linguists and/or archaeologists have placed the PIE homeland all the way from Central Europe to Bactria-Sogdiana and from the Pontic-Caspian steppe to Mesopotamia. So, one has to wonder why expert opinion has such a high variance, but every quantitative effort to solve the problem has come up with a single solution.

As I wrote recently:
My own working hypothesis would derive the earliest Proto-Indo-Europeans with groups living in Neolithic eastern Anatolia and northern Mesopotamia. There are details to be fleshed out, such as when this group of people reached the Balkans (pending ancient DNA from the region), and how they interfaced with the populations living in the north of the Black and Caspian seas (e.g., via a trans-Caucasus movement or a counterclockwise spread around the Caspian).
The current paper suggest a slightly different origin, in Southern Anatolia, perhaps influenced by the distribution of the historical Anatolian languages in the area when they were first put down in writing. But, I suspect that the transposition of Anatolian languages into the areas where they were first attested may have happened late in prehistory. In any case, whether the PIE homeland was in Southern or Eastern Anatolia, the results of this paper explicitly reject the Kurgan Pontic steppe hypothesis.

From the paper:
The distribution for the root location lies in the region of Anatolia in present-day Turkey. To quantify the strength of support for an Anatolian origin, we calculated the Bayes factors (21) comparing the posterior to prior odds ratio of a root location within the hypothesized Anatolian homeland (11) (Fig. 1, yellow polygon) with two versions of the steppe hypothesis—the initial proposed Kurgan steppe homeland (6) and a later refined hypothesis (7) (Table 1). Bayes factors show strong support for the Anatolian hypothesis under a RRW model.  
... 
As the earliest representatives of the main Indo-European lineages, our 20 ancient languages might provide more reliable location information. Conversely, the position of the ancient languages in the tree, particularly the three Anatolian varieties, might have unduly biased our results in favor of an Anatolian origin. We investigated both possibilities by repeating the above analyses separately on only the ancient languages and only the contemporary languages (which excludes Anatolian). Consistent with the analysis of the full data set, both analyses still supported an Anatolian origin (Table 1). 

The West Asian origin of the Proto-Indo-Europeans makes excellent sense in the light of the genetic evidence. But, as I hint at the above paragraph, the tempo of their expansion into Europe remains to be clarified. I strongly suspect, on the basis of the Iceman and Swedish Neolithic TRB farmer (Gok4) whose DNA has been published that the earliest Neolithic was not Indo-European, because these individuals lack the "West Asian" autosomal component.

But, when did the Indo-Europeans first set foot on Europe? Were they already present at the time of Dimini and Vinča in the Balkans? I tend to think that a reasonable proposition, because the 8.2 kiloyear event may have transposed a second set of Neolithic farmers into Europe, of Halafian origin. Or, did they appear later, during the Copper and Bronze Ages with the spread of metallurgy? Until we get ancient DNA from the Balkans and Anatolia, we won't know for sure. But, Y-haplogroups J2, and R1 so conspicuously absent from Neolithic Europe down to 5ka (and in the case of J2, completely missing from the record altogether) must have entered Europe at some point. Did they take the fast train into Europe post-5ka, or did they lurk in both Anatolia and Europe pre-5ka? Thanks to the BEAN project we might find out.

The idea that ~5ka something happened in Europe is also supported by the paper:
Despite support for an Anatolian Indo- European origin, we think it unlikely that agriculture serves as the sole driver of language expansion on the continent. The five major Indo-European subfamilies—Celtic, Germanic, Italic, Balto-Slavic, and Indo-Iranian—all emerged as distinct lineages between 4000 and 6000 years ago (Fig. 2 and fig. S1), contemporaneous with a number of later cultural expansions evident in the archaeological record, including the Kurgan expansion (5–7). 
So, while the deepest prehistory of Indo-European is firmly rooted in Anatolia during the early Neolithic, this is not inconsistent with something important happening in Europe during c. 5ka. But this was a secondary phenomenon, not the earliest seat of the Indo-Europeans. Also, I would not particularly relate this to the Kurgan expansion, but more probably to the arrival of metallurgical "guilds" with higher social complexity.

Both horses and wheeled vehicles quickly spread far and wide because of their simplicity and utility; if they were first adopted by a particular people, they quickly spread beyond it. Metallurgy, on the other hand, requires specialized knowledge about a variety of technical subjects, as well as a complex network of people with distinct roles: miners, metalworkers, traders, warriors, administrators. As such, the people who invented it would have had a distinct advantage until their trade secrets were leaked, or too many Bronze weapons were in the hands of their enemies. During the Bronze Age, more and more people got access to weaponry, and by the end of it, wars were raging all across Western Eurasia.

We tend to think of the Neolithic farmers, but it is quite likely that people kept coming into Europe since its initial colonization. After all, the people who came to the Americas in 1492 were the vanguard of many others who followed them. The same must have happened in Europe as well: a continuous process of settlement by various groups at different times, at least until the Bronze and Iron Ages when everyone, all over West Eurasia, seem to have become very quarrelsome and more than willing to use their swords, spears, axes, and arrows to dissuade newcomers who ventured into their territory.

Coverage of the new paper elsewhere: NY Times, Nature, Gene Expression, John Hawks.

Science 24 August 2012: Vol. 337 no. 6097 pp. 957-960 DOI: 10.1126/science.1219669

Mapping the Origins and Expansion of the Indo-European Language Family 

Remco Bouckaert et al.

ABSTRACT

There are two competing hypotheses for the origin of the Indo-European language family. The conventional view places the homeland in the Pontic steppes about 6000 years ago. An alternative hypothesis claims that the languages spread from Anatolia with the expansion of farming 8000 to 9500 years ago. We used Bayesian phylogeographic approaches, together with basic vocabulary data from 103 ancient and contemporary Indo-European languages, to explicitly model the expansion of the family and test these hypotheses. We found decisive support for an Anatolian origin over a steppe origin. Both the inferred timing and root location of the Indo-European language trees fit with an agricultural expansion from Anatolia beginning 8000 to 9500 years ago. These results highlight the critical role that phylogeographic inference can play in resolving debates about human prehistory.

Link

July 26, 2012

A look at Y chromosomes of Romania via Count Dracula

In short: researchers tried to see whether they could identify a specific Y chromosome lineage associated with the House of Basarab in Romania, the most famous member of which is Vlad the Impaler, an inspiration for the mythical Count Dracula. To do this, they tested Basarab-surnamed individuals, as well as the general Romanian population.

The whole exercise was, in a sense, a failure, since it neither disclosed a Basarab-specific lineage, nor resolved the historical question about the origin of the House of Basarab (Vlach or Cuman). But, it gave us some wonderful new data on Romania that is, of course, quite welcome.

This seems like a good candidate for a future ancient DNA study, assuming of course, that Vlad and his family are still in their final resting place, and there are brave enough researchers to disturb them (j/k).

On a more serious note, the authors correctly state that even if the Basarab house was originally Turkic, they could still have carried West Eurasian chromosomes, since incoming Turkic groups in Europe were not purely Mongoloid like their more remote ancestors. On the other hand, I note that most of the Basarab-surnamed individuals belonged to E-V13, I-P37.2, J-M241 all of which are almost certainly native Romanian. If one of them carries the original chromosome, then the odds are in favor of a Romanian origin, although nothing short of ancient DNA work can resolve the issue, assuming that's possible.

Table S1 contains the new Romanian data, and Table S2 data from surrounding populations (Hungary, Bulgaria, Ukraine).

PLoS ONE 7(7): e41803. doi:10.1371/journal.pone.0041803

Y-Chromosome Analysis in Individuals Bearing the Basarab Name of the First Dynasty of Wallachian Kings

Begoña Martinez-Cruz et al.

Vlad III The Impaler, also known as Dracula, descended from the dynasty of Basarab, the first rulers of independent Wallachia, in present Romania. Whether this dynasty is of Cuman (an admixed Turkic people that reached Wallachia from the East in the 11th century) or of local Romanian (Vlach) origin is debated among historians. Earlier studies have demonstrated the value of investigating the Y chromosome of men bearing a historical name, in order to identify their genetic origin. We sampled 29 Romanian men carrying the surname Basarab, in addition to four Romanian populations (from counties Dolj, N = 38; Mehedinti, N = 11; Cluj, N = 50; and Brasov, N = 50), and compared the data with the surrounding populations. We typed 131 SNPs and 19 STRs in the non-recombinant part of the Y-chromosome in all the individuals. We computed a PCA to situate the Basarab individuals in the context of Romania and its neighboring populations. Different Y-chromosome haplogroups were found within the individuals bearing the Basarab name. All haplogroups are common in Romania and other Central and Eastern European populations. In a PCA, the Basarab group clusters within other Romanian populations. We found several clusters of Basarab individuals having a common ancestor within the period of the last 600 years. The diversity of haplogroups found shows that not all individuals carrying the surname Basarab can be direct biological descendants of the Basarab dynasty. The absence of Eastern Asian lineages in the Basarab men can be interpreted as a lack of evidence for a Cuman origin of the Basarab dynasty, although it cannot be positively ruled out. It can be therefore concluded that the Basarab dynasty was successful in spreading its name beyond the spread of its genes.

June 27, 2012

BEAN – Bridging the European and Anatolian Neolithic

The BEAN project has been on my radar for a while now, and it's great that it seems to be alive and well. Hopefully it won't be too long until it starts producing results.

The origins of human settlement: Mainz University coordinates a new EU project for young researchers
Junior researcher Zuzana Fajkošová passes international selection procedure and begins her doctorate in the Palaeogenetics Group at the JGU Institute of Anthropology

27.06.2012

"BEAN – Bridging the European and Anatolian Neolithic" is the name of a new multinational educational network which has received funding from the European Commission for the next four years. It is classified as a so-called Initial Training Network (ITN) in the EU Marie Curie Actions program, which allows young scientists early access to research activity at top international institutions. A basic requirement for funding is that the researchers involved leave their home country and conduct their research in another European country.

The BEAN Network consists of several European partners in England, Switzerland, France, Germany, Serbia, and Turkey, and has set itself the goal of enhancing the skills of a new generation of researchers in the subjects of anthropology, pre-history, population genetics, computer modeling, and demography. Many different disciplines are participating in the initiative. An important associate partner on the German side is the German Federal Statistical Office in Wiesbaden. The common focus of the project partners centers around questions associated with the origin of first farmer settlements, which were established some 8,000 years ago in West Anatolia and the Balkans. Where did they come from? Were they migrants from the Middle East? Are they our ancestors?

Anthropologists at Johannes Gutenberg University Mainz (JGU) have been meticulous in their preparation of the project over the last years and have entered into various cooperations to underpin it. Seven research institutions and two commercial companies are now working together on the BEAN project. Two leading researchers serve the network in an advisory capacity. These are archaeologist Ian Hodder from Stanford, who established his reputation with his excavations in Catal Höyük, and Hermann Parzinger, President of the Prussian Cultural History Foundation, who spent many years excavating and researching in European Turkey.

As of July 2012, doctoral candidate Zuzana Fajkošová, who completed her undergraduate studies at Masaryk University in Brno and at Charles University in Prague in the Czech Republic, will be the first of two BEAN researchers to start work at JGU's Institute of Anthropology and in the new palaeogenetic laboratory, which is currently in the final stages of construction on the edges of the university's Botanic Garden. She will analyze DNA from the bones of the last hunter-gatherers and the first settled farmers in the region between West Anatolia and the Balkans using the new cutting-edge technology of Next Generation Sequencing (NGS). Together with her colleagues in Dublin, London, and Geneva, she will use the genomic data to compile a model for the settlement of Europe.

"It is both a great honor and a huge opportunity for me that I can work together with such renowned researchers. I'm looking forward to Mainz, the university and the institute's new building," comments Fajkošová, who turned down a number of other offers in order to work at JGU. "A major factor leading to her appointment was the fact that besides mastering biomolecular techniques she also has good programming skills,” explains Professor Dr. Joachim Burger, the Network Coordinator. "A few years ago we more or less founded the discipline of Neolithic Palaeogenetics single-handedly. However, undertaking genomic projects is possible only with the help of international colleagues. That is why we are so pleased that such networks give us and our colleagues the chance to train young research talents."

Besides academic training, the young researchers will be able to do practical work for the two commercial companies within the network and thereby gain work experience in a non-university environment. "This is important as not all of the candidates will opt for a pure research career," explains Karola Kirsanow, who moved from Harvard to Mainz last year and now administrates the network together with Burger. "Our young colleagues have to attend many workshops, courses, and internships, most of them abroad. While this makes for a very tough program, we believe that it significantly enhances the quality of the training and similarly enhances candidates' career prospects."

June 18, 2012

Is Burushaski Indo-European?

I try to shy away from linguistic controversies, and I doubt that the idea of the Indo-European origin of Burushaski will go down without a fight. The JIES is a little behind the times when it comes to online access, so I can't quite comment on the substantive part of Prof. Casule's theory.

We'll have to wait to see how much of this is press release hype and how much is new evidence. This idea has been in circulation for a while, but if there is indeed an issue of JIES dedicated to it, it probably deserves our attention.

Here is a paper critical of the thesis. Here is a seminar abstract by Prof. Casule with some more citations.

Cracking the code on the origins of a new European language
There is strong evidence to support the discovery of a new European language.

Macquarie University historical linguistics researcher, Associate Professor Ilija Casule, discovered that the language, known as Burushaski, which is spoken by about 90,000 people who reside in a remote area of North West Pakistan, is Indo-European in origin, not Indo-Iranian.

Professor Casule’s discovery, which has now been verified by a number of the world’s top linguists, has excited linguistics experts around the world. An entire issue of the eminent international linguistics journal The Journal of Indo-European Studies is devoted to a discussion of his findings later this month.

More than 50 eminent linguists have tried over many years to determine the genetic relationship of Burushaski. But it was Casule’s painstaking research, based on a comprehensive grammatical, phonological, lexical and semantic analysis, which established that the Burushaski language is in fact an Indo-European language most likely descended from one of the ancient Balkan languages. Professor Casule believes that language is most probably ancient Phrygian.

The Phrygians migrated from Macedonia to Anatolia (today part of Turkey) and were famous for their legendary kings who figure prominently in Greek mythology such as King Midas who turned whatever he touched into gold. They later migrated further east, reaching India. Indeed, according to ancient legends of the Burushashki people, they are descendants of Alexander the Great.

Tracing the historical path of a language is no easy task. Professor Casule said he became interested in the origins of Burushaski more than 20 years ago.

“People knew of its existence but its Indo-European affiliation was overlooked and it was not analysed correctly. It is considered a language isolate – not related to any other language in the world in much the same way that the Basque language is classified as a language isolate,” he said.

The remoteness of the area that was independent until the early 1970s when it became part of Pakistan, ensured Burushaski retained certain grammatical and lexical features that led Professor Casule to conclude it is a North-Western Indo-European language, specifically of the Paleobalkanic language group and that it corresponds most closely with Phrygian.

Dr Casule’s work is groundbreaking, not only because it has implications for all the Indo-European language groups, but also provides a new model for figuring out the origins of isolate languages – where they reside in the linguistic family tree and how they developed and blended with other languages to form a new language.

Image: Map of Burushashki speaking areas