R1ans still at large (or, the story of India)

Ten ago, in the pre-ancient DNA "Dark Ages" a big debate raged on about the origin of R1a  men in India. The stage had been set even earlier, by the pioneering Eurasian heartland paper which was the first (to my memory) to link M17 with steppe migrations and Indo-Iranians. Yet, there was pushback as the distribution of M17 was better described, and people started using Y-STRs to try to date and place phylogeographically its migrations.

The two poles of the debate were the "Out-of-India", which relied primarily on Y-STR based time estimates that seemed very old (even Paleolithic, if one used the wrong mutation rate) in India, and the "Into-India" which thought that the R1a distribution pointed to its being brought into India by the Indo-Aryans in the conventional ~3,500BC time frame of the "Aryan Invasion Theory" (AIT).

AIT has been much maligned because it has been received as a Western colonialist imposition on Indian history: a way to claim that Indian civilization was not native but European in origin. Europeans were certainly guilty of misusing AIT: for British colonials it represented a precedent for their colonization of India; for German National Socialists it was evidence for the greatness of the Aryan race and its past expansions eastward. It also played into internal Indian politics, espoused by some as a means of furthering their superiority as either descendants of "Aryan conquerors" or as oppressed victims of the same.

Of course, a misuse of a theory does not mean it is wrong, and if a new preprint based on ancient and modern DNA is correct, it means that AIT was basically correct: Indo-Aryans did come to India in the Late Bronze Age, via the steppe, and ultimately from central Europe.

The opposing Out-of-India theory is all but dead, although failed theories often have a long half-life, especially if they are espoused for psycho-political reasons. I would argue that Out-of-India was dead for thousands of years before it was conceived, since even in Homer's time it was known that "India" was not "one thing" but was peopled by Indians in the north and "Eastern Ethiopians" in the south (which differed from their western "actual" Ethiopians of Africa by their possession of straight rather than curly hair). These were the "Ancestral North Indians" and "Ancestral South Indians" that modern science has revealed. Out-of-India is little more than a nationalistic myth functioning as an antidote to this basic dichotomy, a way to imbue India's diverse citizens with a myth of common origins.

Yet, proponents of AIT (who have a non-trivial overlap with R1an enthusiasts) are also scratching their heads because of the 27 ancient South Asian males from South Asia studied in the preprint there is exactly one R1a, who also happened to live after the time of the Buddha and not during the Bronze Age.

Both OIT enthusiasts (who expected copious and abundant R1a in India and its environs since the Paleolithic) and AIT/R1an enthusiasts (who expected to see it come in c. 3,500BC) are bound to be disappointed.

Perhaps the R1a Indo-Aryans did come to South Asia in a conventional AIT time frame and they haven't been sampled. Or, maybe they were, indeed, there, but were not R1ans. Or, maybe both sides missed the bigger story which is that the Indo-Aryans (so closely associated with India today) were simply not there as early as people have thought. 

bioRxiv: doi: https://doi.org/10.1101/292581

The Genomic Formation of South and Central Asia

Vagheesh M Narasimhan, Nick J Patterson et al.

The genetic formation of Central and South Asian populations has been unclear because of an absence of ancient DNA. To address this gap, we generated genome-wide data from 362 ancient individuals, including the first from eastern Iran, Turan (Uzbekistan, Turkmenistan, and Tajikistan), Bronze Age Kazakhstan, and South Asia. Our data reveal a complex set of genetic sources that ultimately combined to form the ancestry of South Asians today. We document a southward spread of genetic ancestry from the Eurasian Steppe, correlating with the archaeologically known expansion of pastoralist sites from the Steppe to Turan in the Middle Bronze Age (2300-1500 BCE). These Steppe communities mixed genetically with peoples of the Bactria Margiana Archaeological Complex (BMAC) whom they encountered in Turan (primarily descendants of earlier agriculturalists of Iran), but there is no evidence that the main BMAC population contributed genetically to later South Asians. Instead, Steppe communities integrated farther south throughout the 2nd millennium BCE, and we show that they mixed with a more southern population that we document at multiple sites as outlier individuals exhibiting a distinctive mixture of ancestry related to Iranian agriculturalists and South Asian hunter-gathers. We call this group Indus Periphery because they were found at sites in cultural contact with the Indus Valley Civilization (IVC) and along its northern fringe, and also because they were genetically similar to post-IVC groups in the Swat Valley of Pakistan. By co-analyzing ancient DNA and genomic data from diverse present-day South Asians, we show that Indus Periphery-related people are the single most important source of ancestry in South Asia — consistent with the idea that the Indus Periphery individuals are providing us with the first direct look at the ancestry of peoples of the IVC — and we develop a model for the formation of present-day South Asians in terms of the temporally and geographically proximate sources of Indus Periphery-related, Steppe, and local South Asian hunter-gatherer-related ancestry. Our results show how ancestry from the Steppe genetically linked Europe and South Asia in the Bronze Age, and identifies the populations that almost certainly were responsible for spreading Indo-European languages across much of Eurasia.

Link

History of extant populations of India

The five components they speak of are ANI, ASI, AAA (Ancestral Austro-Asiatic), ATB (Ancestral Tibeto-Burman), and a distinct fifth ancestry in the Andaman archipelago.

The differentiation of the four main components seems clear enough on the figure (left). The big question is how and in what order the different components got into India. I would wager that ASI was first and I modify my New Year's wish to ask for some ancient DNA from India too.

An interesting bit from the paper:

...that the practice of endogamy was established almost simultaneously, possibly by decree of the rulers, in upper-caste populations of all geographical regions, about 70 generations before present, probably during the reign (319–550 CE) of the ardent Hindu Gupta rulers

How plausible is that to anyone familiar with Indian history?

PNAS doi: 10.1073/pnas.1513197113

Genomic reconstruction of the history of extant populations of India reveals five distinct ancestral components and a complex structure

Analabha Basu, Neeta Sarkar-Roya, and Partha P. Majumder

India, occupying the center stage of Paleolithic and Neolithic migrations, has been underrepresented in genome-wide studies of variation. Systematic analysis of genome-wide data, using multiple robust statistical methods, on (i) 367 unrelated individuals drawn from 18 mainland and 2 island (Andaman and Nicobar Islands) populations selected to represent geographic, linguistic, and ethnic diversities, and (ii) individuals from populations represented in the Human Genome Diversity Panel (HGDP), reveal four major ancestries in mainland India. This contrasts with an earlier inference of two ancestries based on limited population sampling. A distinct ancestry of the populations of Andaman archipelago was identified and found to be coancestral to Oceanic populations. Analysis of ancestral haplotype blocks revealed that extant mainland populations (i) admixed widely irrespective of ancestry, although admixtures between populations was not always symmetric, and (ii) this practice was rapidly replaced by endogamy about 70 generations ago, among upper castes and Indo-European speakers predominantly. This estimated time coincides with the historical period of formulation and adoption of sociocultural norms restricting intermarriage in large social strata. A similar replacement observed among tribal populations was temporally less uniform.

Link

Bronze Age people from Ireland had steppe ancestry and R1b

From the paper:

We were able to deduce that Neolithic Ballynahatty had a dark hair shade (99.5% probability), most likely black (86.1% probability), and brown eyes (97.3% probability) (46). Bronze Age Rathlin1 probably had a light hair shade (61.4%) and brown eyes (64.3%). However, each Rathlin genome possessed indication of at least one copy of a haplotype associated with blue eye color in the HERC2/OCA2 region.

and:

Third, we followed the methods described in Haak et al. (9), which use a collection of outgroup populations, to estimate the mixture proportions of three different sources, Linearbandkeramik (Early Neolithic; 35 ± 6%), Loschbour (WHG; 26 ± 12%), and Yamnaya (39 ± 8%), in the total Irish Bronze Age group. These three approaches give an overlapping estimate of ∼32% Yamnaya ancestry.

PNAS doi: 10.1073/pnas.1518445113

Neolithic and Bronze Age migration to Ireland and establishment of the insular Atlantic genome

Lara M. Cassidy, Rui Martiniano et al.

The Neolithic and Bronze Age transitions were profound cultural shifts catalyzed in parts of Europe by migrations, first of early farmers from the Near East and then Bronze Age herders from the Pontic Steppe. However, a decades-long, unresolved controversy is whether population change or cultural adoption occurred at the Atlantic edge, within the British Isles. We address this issue by using the first whole genome data from prehistoric Irish individuals. A Neolithic woman (3343–3020 cal BC) from a megalithic burial (10.3× coverage) possessed a genome of predominantly Near Eastern origin. She had some hunter–gatherer ancestry but belonged to a population of large effective size, suggesting a substantial influx of early farmers to the island. Three Bronze Age individuals from Rathlin Island (2026–1534 cal BC), including one high coverage (10.5×) genome, showed substantial Steppe genetic heritage indicating that the European population upheavals of the third millennium manifested all of the way from southern Siberia to the western ocean. This turnover invites the possibility of accompanying introduction of Indo-European, perhaps early Celtic, language. Irish Bronze Age haplotypic similarity is strongest within modern Irish, Scottish, and Welsh populations, and several important genetic variants that today show maximal or very high frequencies in Ireland appear at this horizon. These include those coding for lactase persistence, blue eye color, Y chromosome R1b haplotypes, and the hemochromatosis C282Y allele; to our knowledge, the first detection of a known Mendelian disease variant in prehistory. These findings together suggest the establishment of central attributes of the Irish genome 4,000 y ago.

Link

Into, out of, and across the Eurasian steppe

A new paper in Nature adds to the earlier study in the same journal by presenting data from 101 ancient Eurasians. The year is not yet halfway over, but it seems that the ancient DNA field is moving towards a new norm of studying dozens of individuals at a time and comprehensively tackling the "big problems" that have vexed archaeologists, linguists, and historians for decades if not centuries.

The first conclusion of the new study is the detection of the migration from the steppe to Europe that was the title piece of the earlier study. The authors do not present quantitative estimates of the amount of demographic replacement effected by the Yamnaya-to-Corded Ware migration, so it will be interesting to see if there are any minor significant differences in these. But, the two papers have different Yamnaya and Corded Ware samples, and yet arrive at qualitatively similar conclusions, so at least this part of the story should be considered firmly "settled".

The second conclusion is the migration from the European steppe to the Afanasievo culture of the Altai. This has been long-hypothesized based on the physical type of the Afanasievo people and their possession of a similar pastoralist/wheeled vehicle toolkit that would have allowed them to cover the huge difference between Europe and the Altai. This confirms movement #2 of the Anthony/Ringe model, although I doubt that this migration had anything to do with Tocharians as detailed below. But, it did happen.

The third conclusion is that the later steppe cultures of the Sintashta and Andronovo (putative Indo-Iranians according to some), were not a continuation of the Yamnaya-Afanasievo people, but had extra Neolithic farmer ancestry. So, it seems that Neolithic farmers entered the steppe, and the development of steppe cultures did not happen in isolation. Whether this involved migration of Corded Ware people (as the authors prefer), who were already a mixture of Yamnaya and Neolithic farmers, or some other mixture of Neolithic farmers with steppe populations (e.g., Tripolye plus Yamnaya) remains to be seen.

The fourth conclusion of the paper is that these steppe cultures were also later replaced by people of at least partial East Asian or "Native American"-like ancestry.  So, it seems that movements into the steppe happened both on the western end (as the incursion of Neolithic farmer ancestry into the Sintashta proves), but also on the eastern end, with the Europeoid populations of western origin receiving admixture from the eastern periphery of the Eurasian steppe.

As for the Yamnaya, the authors do not find a very strong signal of admixture (as did the earlier study), which they attribute quite plausibly to the lack of eastern hunter-gatherers in their dataset. On the other hand, they claim that the "Caucasus" genetic component in the steppe populations was of steppe ancestry rather than Near Eastern/Caucasian origin as was claimed in the earlier paper. This is based on the statistic D(Yoruba, Armenia BA; Yamnaya, Corded Ware) that is not significantly different from zero. However, Corded Ware is a mixture of Yamnaya and European Neolithic, so the sign of this statistic is determined by the sign of the statistic D(Yoruba, Armenia BA; Yamnaya, European Neolithic). If Yamnaya was simply a steppe population, descendants of local people without ancestry from the Middle East/Caucasus, then this statistic would be positive because of the shared Middle Eastern ancestry of Armenia BA and European Neolithic. Whereas, if Yamnaya is a mixture of a steppe population and a Middle Eastern/Caucasian one, then the statistic would be positive/negative for the respective parts, which would be consistent with an average not different from zero. I am sure that when the new data is re-analyzed together with the eastern hunter-gatherers it will be clear that the Yamnaya are not a pure steppe population.

Nonetheless, I am quite glad to read a sentence such as this:

Populations in northern and central Europe were composed of a mixture of the earlier hunter-gatherer and Neolithic farmer10 groups, but received ‘Caucasian’ genetic input at the onset of the Bronze Age (Fig. 2).

It seems that my prediction the the West_Asian component would appear in post-5ka Europeans and was related to Indo-Europeans has been adequately confirmed by the last two papers.

Speaking of the Caucasus/Middle East, it seems clear as a first approximation that the Bronze Age Armenians are quite similar to modern Armenians. Whether the genetic continuity of Armenians extends beyond the Bronze Age, or Armenians were formed by mixture in the Bronze Age remains to be seen. The question of Armenian linguistic origins is of course separate as it is commonly understood that the Armenian language is unrelated to Anatolian languages and may have arrived in Armenia from the Balkans at around the Bronze Age-Iron Age transition.

The authors also study some phenotypic traits such as lactase peristence (Yamnaya had some, but overall prevalence was much lower than modern Europeans, hence lots of selection to the present), and skin eye pigmentation. Like Wilde et al., and Mathieson et al., the steppe populations seem to have had brown eyes. Given that so did Neolithic Europeans, and (presumably) ancient Middle Easterners/Caucasians, I think it's a good bet that Proto-Indo-Europeans (whatever solution to the PIE urheimat one accepts) were a brown-eyed people, or in the very least far from the blue-eyed "Aryans" of racial mythology. Even the Bronze Age and Iron Age Asians seem to have been a predominantly brown-eyed people, although the derived HERC2 allele seems to be at a higher frequency in them than in the steppe Europeans.

The story of the Y-chromosomes seems very interesting, although these are not resolved to fine detail. The most interesting aspect of this part of the work is the appearance of haplogroup J in Iron Age samples from Russia, Armenia, and the Altai. This may tie in to the question of the Tocharian origins, which I have claimed were associated with R1b, rather than R1a (as the Indo-Iranians were). The modern Uygurs (who are partially of Tocharian origin) have both J2 and R1b, so were the recipients of West Eurasian elements other than the R1a that so seem to have dominated the eastern steppe, including the Afanasievo. I continue to think there's no evidence that the Afanasievo is Proto-Tocharian, as it's in the wrong place and 3,000 years before the attestation of Tocharian. 

Overall this is an amazing study which adds a lot to what we know about Bronze Age Eurasia. Hopefully there is more to come in the second half of 2015, but for the time being there is plenty to chew on.

Nature 522, 167–172 (11 June 2015) doi:10.1038/nature14507

Population genomics of Bronze Age Eurasia

Morten E. Allentoft, Martin Sikora, Karl-Göran Sjögren, Simon Rasmussen, Morten Rasmussen, Jesper Stenderup, Peter B. Damgaard, Hannes Schroeder, Torbjörn Ahlström, Lasse Vinner, Anna-Sapfo Malaspinas, Ashot Margaryan, Tom Higham, David Chivall, Niels Lynnerup, Lise Harvig, Justyna Baron, Philippe Della Casa, Paweł Dąbrowski, Paul R. Duffy, Alexander V. Ebel, Andrey Epimakhov, Karin Frei, Mirosław Furmanek, Tomasz Gralak, Andrey Gromov, Stanisław Gronkiewicz, Gisela Grupe, Tamás Hajdu, Radosław Jarysz, Valeri Khartanovich, Alexandr Khokhlov, Viktória Kiss, Jan Kolář, Aivar Kriiska, Irena Lasak, Cristina Longhi, George McGlynn, Algimantas Merkevicius, Inga Merkyte, Mait Metspalu, Ruzan Mkrtchyan, Vyacheslav Moiseyev, László Paja, György Pálfi, Dalia Pokutta, Łukasz Pospieszny, T. Douglas Price, Lehti Saag, Mikhail Sablin, Natalia Shishlina, Václav Smrčka, Vasilii I. Soenov, Vajk Szeverényi, Gusztáv Tóth, Synaru V. Trifanova, Liivi Varul, Magdolna Vicze, Levon Yepiskoposyan, Vladislav Zhitenev, Ludovic Orlando, Thomas Sicheritz-Pontén, Søren Brunak, Rasmus Nielsen, Kristian Kristiansen & Eske Willerslev

The Bronze Age of Eurasia (around 3000–1000 BC) was a period of major cultural changes. However, there is debate about whether these changes resulted from the circulation of ideas or from human migrations, potentially also facilitating the spread of languages and certain phenotypic traits. We investigated this by using new, improved methods to sequence low-coverage genomes from 101 ancient humans from across Eurasia. We show that the Bronze Age was a highly dynamic period involving large-scale population migrations and replacements, responsible for shaping major parts of present-day demographic structure in both Europe and Asia. Our findings are consistent with the hypothesized spread of Indo-European languages during the Early Bronze Age. We also demonstrate that light skin pigmentation in Europeans was already present at high frequency in the Bronze Age, but not lactose tolerance, indicating a more recent onset of positive selection on lactose tolerance than previously thought.

Link

101 ancient genomes from Bronze Age Eurasia

New data has been posted online. This seems related to this earlier post. Hopefully the study linked to this data will appear soon, but genome bloggers can get to it thanks to the early data release.

Investigation of Bronze Age in Eurasia by sequencing from 101 ancient human remains. 

The Bronze Age (BA) of Eurasia (c. 3,000-1,000 years BC, 3-1 ka BC) was a period of major cultural changes. Earlier hunter-gathering and farming cultures in Europe and Asia were replaced by cultures associated with completely new perceptions and technologies inspired by early urban civilization. It remains debated if these cultural shifts simply represented the circulation of ideas or resulted from large-scale human migrations, potentially also facilitating the spread of Indo-European languages and certain phenotypic traits. To investigate this and the role of BA in the formation of Eurasian genetic structure, we used new methodological improvements to sequence low coverage genomes from 101 ancient humans (19 > 1X average depth) covering 3 ka BC to 600 AD from across Eurasia. We show that around 3 ka BC, Central and Northern Europe and Central Asia receive genetic input through people related to the Yamnaya Culture from the Pontic-Caspian Steppe, resulting in the formation of the Corded Ware Culture in Europe and the Afanasievo Culture in Central Asia. A thousand years later, genetic input from North-Central Europe into Central Asia gives rise to the Sintashta and Andronovo Cultures. During the late BA and Iron Age, the European-derived populations in Asia are gradually replaced by multi-ethnic cultures, of which some relate to contemporary Asian groups, while others share recent ancestry with Native Americans. Our findings are consistent with the hypothesised spread of Indo-European languages during early BA and reveal that major parts of the demographic structure of present-day Eurasian populations were shaped during this period. We also demonstrate that light skin pigmentation in Europeans was already present at high frequency during the BA, contrary to lactose tolerance, indicating a more recent onset of positive selection in the latter than previously believed.

Link

Two observations on the ancestry of Armenians

I was thinking a bit on how to interpret the findings of the new Haber et al. preprint, and especially the idea that "29% of the Armenian ancestry may originate from an ancestral population best represented by Neolithic Europeans." I looked at the globe13 proportions, and strangely enough, I had estimated that the three Armenian samples (Armenian_D, Armenians, and Armenians_15_Y) have 28-29% of the Mediterranean component that is modal in Sardinians. This seems like a curious coincidence which has raised my confidence that Haber et al. is picking something real.

Looking back at my inferences of Armenian ancestry, it seems (according to globe13) to come completely from West_Asian, Mediterranean, and Southwest_Asian. The Mediterranean component seems real enough as it seems to match Sardinians/early European farmers well. I am not so sure about the Southwest Asian component which is modal in Yemen Jews and may represent population-specific drift in relatively recent Arabians. The West_Asian component is bimodal in Caucasus and Gedrosia, so it can't be the result of a very drifted population in either region (unless there is spooky action at a distance). 

Another curious finding is the lack of North_European in a latitudinal "column" of populations from the Yemen, through the Levant to the South Caucasus (Georgians and Armenians). It seems that North_European is the only one of the four major Caucasoid components that Armenians lack to any important degree. There is a rather abrupt change between the South Caucasus (~1%) and the North Caucasus (15-20%). It seems that the Greater Caucasus did act like a barrier to gene flow. The K=4 analysis of the same dataset that produced K=13 (globe13) also shows the same barrier: all three Armenian samples and Georgians have ~0% of "Amerindian" (which is surely correlated to "Ancient North Eurasian" ancestry and via it with North_European), but North Caucasians and Europeans have 4-10%.  It's clear that this influence did not cross the Greater Caucasus, as Armenians and Georgians lack it. 

Scandinavian team looking for Indo-Europeans in Kazakhstan

An article in the Astana Times. If anyone has any additional information via Kazakh or Scandinavian media, or can find the press release referred to in the article, feel free to share.

  Scandinavian Team Searches for Indo-European Homeland through Kazakhstan DNA

A Scandinavian team has come to Kazakhstan in search of the common homeland of all Indo-European peoples, collecting bone fragments for analysis in the Centre for Geogenetics at the University of Copenhagen.

The researchers are looking for a genetic connection to match the linguistic connections that have already been drawn, Norwegian historian Sturla Ellingvag of the Explico Historical Research Foundation told The Astana Times on Feb. 20. “We’re trying to find a connection in science, in our DNA, to prove that there is indeed a connection, between, for example, Norwegians and the people in Kazakhstan. And also we are looking for a homeland, which is somewhere on the Caspian steppe, or in Russia, or some say it’s in Armenia or Ukraine. There are many different theories.”

The researchers collected about 120 Bronze and early Iron Age bone samples in total from Pavlodar, Kostanai and Karaganda during their week-long trip to Kazakhstan, from Feb. 14 – 21. Kazakhstan is fascinating, the researcher says, because it contains human remains that are “so far back on the DNA map.”

The 4,000 year old samples they’ve found have been very well preserved, Ellingvag said. “I can only speak from meeting archaeologists in Astana and here in Karaganda, but I’m very much impressed by the professionalism and also by the exhibitions they have,” he said.

The project to search for the ancestral homeland of the Indo-European peoples falls under the umbrella of a large grant from the Danish government and is being supported by the Kon-Tiki Museum in Oslo, Gotenburg University in Sweden and the University of Copenhagen in Denmark, which has one of the best historical DNA analysis labs in the world and which is where the analysis on the Kazakh remains will actually be done. Universities in Karaganda, Pavlodar and Kostanai are also involved.

The Kurgan hypothesis posits that the speakers of proto-Indo-European, the hypothesized common ancestor of the massive Indo-European language group, originally lived on the Pontiac-Caspian steppe, an area of land stretching from the Black Sea to the Caspian Sea and including parts of Russia, Ukraine and northwest Kazakhstan, beginning around the fifth millenium B.C. The hypothesis describes the spread of the language family from the steppe in every direction. “Kurgan” is a term for a type of burial mound common in the Caucasus, across Kazakhstan and beyond.

“Two thousand years ago, we started having Kurgan graves in Scandinavia,” said Ellingvag. The commonalities between burial mounds in Norway and Scythian/Saka mounds in Kazakhstan are striking, he said. “[The Scythian people] had these ornaments, these animal ornaments, which are very, very important in Scandinavian art … and the ornaments are actually quite similar, which is striking, it’s very special.”

The Kurgan hypothesis has been somewhat substantiated by genetic evidence so far, according to a press release by the Kon-Tiki Museum on the project, and advances in the technology for doing historical DNA research over the past few years means it is now possible to get closer to finding this genetic and linguistic starting point for most of the peoples of Europe.

“During the past 15 years, the Y-DNA R1a haplogroup has been characterised as a genetic signal of the Proto-Indo-Europeans. The theory now looks more plausible than ever, thanks to recent discoveries about its structure and phylogeography. Moreover, the Y-DNA R1a haplogroup has been found in numerous ancient remains supposedly belonging to early Indo-Europeans,” the press release explains.

A separate but related project is looking into the DNA of ancient horses. The Kurgan culture is credited with being the first to domesticate the horse.

The research team includes Ellingvag, Danish DNA-scientist Peter Damgaard and Bettina Heyerdahl, daughter of Norwegian archaeologist and explorer Thor Heyerdahl. They are also working with Kazakh researcher Emma Usmanova.

I could also find this Youtube video from this expedition.

Late (not necessarily steppe) split of Proto-Indo-European

This is the paper that I saw referenced in a previous study. As I suspected, the paper does not in fact provide support specifically for the steppe hypothesis, but only for a late split of Proto-Indo-European (that is consistent with the steppe hypothesis but not unique to it).

I don't know how well linguists have figured out how to estimate time depth; the fact that a small tweak in the methodology (compared to Bouckaert et al.) results in a 3,000 year drop in the estimated age of the PIE split does not add to my confidence about the robustness of this field. Regardless of which hypothesis one accepts, the PIE split occurred thousands of years before the first written monuments in any Indo-European language. For the time being, the ball is on the other side of the court, which may accept this finding or come up with another tweak in the methodology that gives yet another date.

In any case, accepting provisionally that the Chang et al. date is accurate then it falsifies the Anatolian farmer/IE language hypothesis. So, steppe aficionados can declare a partial victory, because there is one less opponent to worry about. Falsification of the Anatolian first farmer hypothesis is not a complete victory, as the PIE urheimat question is not a boxing match between Kurganists and Anatolianists, but rather a mêlée with many players holding on to their swords. So, if you're willing to believe that the methodology is mature enough and they finally "got it right", you need only find a PIE split around 6,000 years ago, but you need not find it on the steppe. 

ANCESTRY-CONSTRAINED PHYLOGENETIC ANALYSIS SUPPORTS THE INDO-EUROPEAN STEPPE HYPOTHESIS 

Will Chang et al.

Discussion of Indo-European origins and dispersal focuses on two hypotheses. Qualitative evidence from reconstructed vocabulary and correlations with archaeological data suggest that IndoEuropean languages originated in the Pontic-Caspian steppe and spread together with cultural innovations associated with pastoralism, beginning c. 6500–5500 bp. An alternative hypothesis, according to which Indo-European languages spread with the diffusion of farming from Anatolia, beginning c. 9500–8000 bp, is supported by statistical phylogenetic and phylogeographic analyses of lexical traits. The time and place of the Indo-European ancestor language therefore remain disputed. Here we present a phylogenetic analysis in which ancestry constraints permit more accurate inference of rates of change, based on observed changes between ancient or medieval languages and their modern descendants, and we show that the result strongly supports the steppe hypothesis. Positing ancestry constraints also reveals that homoplasy is common in lexical traits, contrary to the assumptions of previous work. We show that lexical traits undergo recurrent evolution due to recurring patterns of semantic and morphological change.

Link (pdf)

A story of 69 ancient Europeans

A new study on the bioRxiv includes data on 69 ancient Europeans (remember when we got excited in anticipation for the single genome of the Iceman? that was only three years ago) and adds plenty of new info to chew on for those of us interested in prehistory. 

Two Near Eastern migrations into Europe

In 2011, I observed that West Eurasian populations were too close (measured by Fst) to allow for long periods of differentiation between them. By implication, there must have been a "common source" of ancestry uniting them, which I placed in a "womb of nations" of the Neolithic Near East. I proposed that migrations out of this core area homogenized West Eurasians, writing:

In Arabia, the migrants would have met aboriginal Arabians, similar to their next door-neighbors in East Africa, undergoing a subtle African shift (Southwest_Asians). In North Africa, they would have encountered denser populations during the favorable conditions of MIS 1, and by absorbing them they would became the Berbers (Northwest_Africans). Their migrations to the southeast brought them into the realm of Indian-leaning people, in the rich agricultural fields of the Mehrgarh and the now deserted oases of Bactria and Margiana. Across the Mediterranean and along the Atlantic facade of Europe, they would have encountered the Mesolithic populations of Europe, and through their blending became the early Neolithic inhabitants of the Mediterranean and Atlantic coasts of Europe (Mediterraneans). And, to the north, from either the Balkans, the Caucasus, or the trans-Caspian region, they would have met the last remaining Proto-Europeoid hunters of the continental zone, becoming the Northern Europeoids who once stretched all the way to the interior of Asia.

The new paper confirms the last two of these migrations. The remainder involve parts of the world from which no ancient DNA has been studied.

The first migration (early Neolithic) is already uncontroversial, but the paper includes data from Spanish early farmers that are also Sardinian- and LBK-like. The "Sardinian" Iceman was no fluke. It is now proven that not only the LBK but also the Spanish Neolithic came from the same expansion of Mediterranean populations which survives in Sardinia. The authors write:

Principal components analysis (PCA) of all ancient individuals along with 777 present-day West Eurasians4 (Fig. 2a, SI5) replicates the positioning of present-day Europeans between the Near East and European hunter-gatherers4,20, and the clustering of early farmers from across Europe with present day Sardinians3,4,27, suggesting that farming expansions across the Mediterranean to Spain and via the Danubian route to Hungary and Germany descended from a common stock.

The second migration went into eastern Europe:

The Yamnaya differ from the EHG by sharing fewer alleles with MA1 (|Z|=6.7) suggesting a dilution of ANE ancestry between 5,000-3,000 BCE on the European steppe. This was likely due to admixture of EHG with a population related to present-day Near Easterners, as the most negative f3-statistic in the Yamnaya (giving unambiguous evidence of admixture) is observed when we model them as a mixture of EHG and present-day Near Eastern populations like Armenians (Z = -6.3; SI7).

The EHG (Eastern European Hunter-Gatherers) are likely Proto-Europeoid foragers and the Yamnaya (a Bronze Age Kurgan culture) were a mixture of the EHG and something akin to Armenians.The "attraction" of later groups to the Near East is clear in the PCA: hunter-gatherers on the left side, the Near East (as grey dots) on the right side, and Neolithic/Bronze Age/modern Europeans in the middle. The second migration may very well be related to the Uruk expansion and the presence of gracile Mediterranoids and robust Proto-Europeoids in the Yamna:

The Yamna population generally belongs to the European race. It was tall (175.5cm), dolichocephalic, with broad faces of medium height. Among them there were, however, more robust elements with high and wide faces of the proto-Europoid type, and also more gracile individuals with narrow and high faces, probably reflecting contacts with the East Mediterranean type (Kurts 1984: 90).

The authors present a table of Fst values which confirms the homogenizing influence of migrations from the Near East. The WHG group has an Fst=0.086 with Armenians, but the LBK farmers have only 0.023. The EHG group has an Fst=0.067 with Armenians, but the Yamnaya steppe people have only 0.030. Someone might argue that it is the Armenians that are receiving genes from Europe, but the same pattern holds even for the Bedouins, for which admixture with Europeans seems far-fetched: 0.106 to 0.043 and 0.093 to 0.060. It is now clear that the "glue" that did not allow West Eurasian populations to drift very far apart were migrations from the Near East.

The (partial) demise of the farmers

It seems that the legacy of the early farmers suffered two hits, which is why only in Sardinia and (to a lesser degree) in southern Europe that they have persisted as the major component of ancestry. The first blow came during the Neolithic:

Middle Neolithic Europeans from Germany, Spain, Hungary, and Sweden from the period ~4,000-3,000 BCE are intermediate between the earlier farmers and the WHG, suggesting an increase of WHG ancestry throughout much of Europe.

And the coup de grâce after the 5kya mark:

We estimate that these two elements each contributed about half the ancestry each of the Yamnaya (SI6, SI9), explaining why the population turnover inferred using Yamnaya as a source is about twice as high compared to the undiluted EHG. The estimate of Yamnaya related ancestry in the Corded Ware is consistent when using either present populations or ancient Europeans as outgroups (SI9, SI10), and is 73.1 ± 2.2% when both sets are combined (SI10). [...] The magnitude of the population turnover that occurred becomes even more evident if one considers the fact that the steppe migrants may well have mixed with eastern European agriculturalists on their way to central Europe. Thus, we cannot exclude a scenario in which the Corded Ware arriving in today’s Germany had no ancestry at all from local populations.

Confirmation of the Bronze Age Indo-European invasion of Europe

In 2012 I had used the paltry data on a handful ancient DNA samples to observe that in ADMIXTURE modern Europeans had a West Asian genetic component (peaking in "Caucasus" and "Gedrosia") that pre-5kya Europeans didn't. I proposed that the Bronze Age migration of the Indo-Europeans spread this component:

But there is another component present in modern Europe, the West_Asian which is conspicuous in its absence in all the ancient samples so far. This component reaches its highest occurrence in the highlands of West Asia, from Anatolia and the Caucasus all the way to the Indian subcontinent. [...] Nonetheless, some of the legacy of the earliest Indo-European speakers does appear to persist down to the present day in the genomes of their linguistic descendants, and I predict that when we sample later (post 5-4kya) individuals we will finally find the West_Asian piece that is missing from the European puzzle.

This prediction is now confirmed:

This pattern is also seen in ADMIXTURE analysis (Fig. 2b, SI6), which implies that the Yamnaya have ancestry from populations related to the Caucasus and South Asia that is largely absent in 38 Early or Middle Neolithic farmers but present in all 25 Late Neolithic or Bronze Age individuals. This ancestry appears in Central Europe for the first time in our series with the Corded Ware around 2,500 BCE (SI6, Fig. 2b, Extended Data Fig. 1).

I was a little puzzled with the "Ancient North Eurasians" recently proposed as a "third ancestral population" for Europeans: it seemed to be a tertium quid that spread after 5kya, but very different geographically than the "West Asian" component. But:

These results can be explained if the new genetic material that arrived in Germany was a composite of two elements: EHG and a type of Near Eastern ancestry different from that which was introduced by early farmers (also suggested by PCA and ADMIXTURE; Fig. 2, SI5, SI6).

So, it seems that there is no contradiction after all and both EHG (which is related to "Ancient North Eurasians") and another type of Near Eastern ancestry (=West_Asian) arrived after 5kya.

1939 strikes back

It is amazing how well this was anticipated by Carleton Coon in 1939. Back then much of West Eurasia was an archaeological/anthropological terra incognita, there was no radiocarbon dating, no DNA, no computers, not even serious multivariate statistics. And yet:

We shall see, in our survey of prehistoric European racial movements, 8 that the Danubian agriculturalists of the Early Neolithic brought a food-producing economy into central Europe from the East. They perpetuated in the new European setting a physical type which was later supplanted in their original home. Several centuries later the Corded people, in the same way, came from southern Russia but there we first find them intermingled with other peoples, and the cul-tural factors which we think of as distinctively Corded are included in a larger cultural equipment. [...] On the basis of the physical evidence as well, it is likely that the Corded people came from somewhere north or east of the Black Sea. The fully Neolithic crania from southern Russia which we have just studied include such a type, also seen in the midst of Sergi's Kurgan aggregation. Until better evidence is produced from elsewhere, we are entitled to consider southern Russia the most likely way station from which the Corded people moved westward.

And in 2015:

Our results support a view of European pre-history punctuated by two major migrations: first, the arrival of first farmers during the Early Neolithic from the Near East, and second of Yamnaya pastoralists during the Late Neolithic from the steppe (Extended Data Fig. 5).

In 1939:

Linguistically, Indo-European is probably a relatively recent phenomenon, which arose after animals had been tamed and plants cultivated. The latest researches find it to be a derivative of an initially mixed language, whose principal elements were Uralic, called element A, and some undesignated element B which was probably one of the eastern Mediterranean or Caucasic languages. 5 The plants and animals on which the Somewhere in the plains of southern Russia or central Asia, the blending of languages took place which resulted in Indo-European speech. This product in turn spread and split, and was further differentiated by mixture with the languages of peoples upon whom it, in one form or other, was imposed. Some of the present Indo-European languages, in addition to these later accretions from non-Indo-European tongues, contain more of the A element than others, which contain more of the B. The unity of the original " Indo- Europeans," could not have been of long duration, if it was ever complete. 

In 2015:

These results can be explained if the new genetic material that arrived in Germany was a composite of two elements: EHG and a type of Near Eastern ancestry different from that which was introduced by early farmers (also suggested by PCA and ADMIXTURE; Fig. 2, SI5, SI6). We estimate that these two elements each contributed about half the ancestry each of the Yamnaya (SI6, SI9), explaining why the population turnover inferred using Yamnaya as a source is about twice as high compared to the undiluted EHG.

The EHG is still flimsy as it's only two individuals from Karelia and Samara who are very similar to each other. It's hard not to imagine that the hunter-gatherer from Russian Karelia (outside any proposed PIE homeland) would be speaking a similar language as his Samara counterpart. Did they both speak "element A" and was PIE formed when the "southern" steppe hunter-gatherers came into contact with "element B" people from the Caucasus? Short of a time machine, we can never say for sure. This might very well be an answer to the conundrum of Uralic/Proto-Kartvelian borrowings. There is simply no geographical locale in which these two language families neighbor each other: Northwest, Northeast Caucasian speakers and the pesky Greater Caucasus intervene. But, maybe there was no such locale, and these borrowings aren't due to some "PIE people" living adjacent to Uralic and Proto-Karvelian speakers but the "PIE people" being a mix of an element A (EHG) that was (or interacted with) Uralic and another element B (Armenian-like) that was (or interacted with) Proto-Kartvelian.

Urheimat (or not?)

The authors of the current paper are agnostic about the PIE homeland:

We caution that the location of the Proto-Indo-European9,27,29,30 homeland that also gave rise to the Indo-European languages of Asia, as well as the Indo-European languages of southeastern Europe, cannot be determined from the data reported here (SI11). Studying the mixture in the Yamnaya themselves, and understanding the genetic relationships among a broader set of ancient and present-day Indo-European speakers, may lead to new  insight about the shared homeland.

Whatever the ultimate answer will be, it seems that Coon was right that "The unity of the original " Indo- Europeans," could not have been of long duration, if it was ever complete." If PIE=EHG (as Anthony and Ringe suggest), then "from the crib", PIE got half its ancestry from a non-IE, Near Eastern source. Conversely, if PIE=Near East (as I suggested) then "from the crib", PIE got half of its ancestry from a non-IE, Eastern European source. The "Yamnaya" seems to max out in Norwegians at around half, which means that they are about a quarter Proto-Indo-European genetically, regardless of which theory is right.

These two possibilities (as well as the third one of PIE being neither-nor, but rather a linguistic mixture of the languages of the EHG and Near East) are testable. The Anthony/Ringe version of the steppe hypothesis predicts pre-Yamnaya expansions from the steppe. Whether these happened and what was their makeup can be tested: if they did occur and they did lack "Near Eastern" ancestry, then the steppe hypothesis will be proven. PIE in the Near East, on the other hand, predicts that some PIE languages (certainly the Anatolian ones) will be a "within the Near East" expansion. If such migrations did occur and they lacked "EHG" ancestry, then some variant of the Gamkrelidze/Ivanov model will be proven. Or, the truth might be that everywhere where Indo-Europeans arrive they carry a blend of "West Asian" and "EHG", supporting the third possibility. Time will tell.

In the interim, I am curious about how much Yamnaya ancestry existed in different parts of Europe (all of the post-5kya samples in this study come from Germany, with a couple from Hungary). In northern Europe, all populations seem to have less Yamnaya ancestry than the Corded Ware: there it must have declined. But, modern Hungarians have more than Bronze Age Hungarians: there it must have increased.

Germany and a slice of Hungary is a very narrow window through which to see the whole of Europe and these results must be tested by looking at samples from beyond the "heartland". I do hope that some kind of Moore's law operates in the world of ancient DNA, and in three more years we'll be reading studies about thousands of ancient individuals.

bioRxiv doi: http://dx.doi.org/10.1101/013433
Massive migration from the steppe is a source for Indo-European languages in Europe

Wolfgang Haak , Iosif Lazaridis , Nick Patterson , Nadin Rohland , Swapan Mallick , Bastien Llamas , GuidoBrandt , Susanne Nordenfelt , Eadaoin Harney , Kristin Stewardson , Qiaomei Fu , Alissa Mittnik , Eszter Banffy ,Christos Economou , Michael Francken , Susanne Friederich , Rafael Garrido Pena , Fredrik Hallgren , ValeryKhartanovich , Aleksandr Khokhlov , Michael Kunst , Pavel Kuznetsov , Harald Meller , Oleg Mochalov ,Vayacheslav Moiseyev , Nicole Nicklisch , Sandra L. Pichler , Roberto Risch , Manuel A. Rojo Guerra , ChristinaRoth , Anna Szecsenyi-Nagy , Joachim Wahl , Matthias Meyer , Johannes Krause , Dorcas Brown , DavidAnthony , Alan Cooper , Kurt Werner Alt , David Reich

We generated genome-wide data from 69 Europeans who lived between 8,000-3,000 years ago by enriching ancient DNA libraries for a target set of almost four hundred thousand polymorphisms. Enrichment of these positions decreases the sequencing required for genome-wide ancient DNA analysis by a median of around 250-fold, allowing us to study an order of magnitude more individuals than previous studies and to obtain new insights about the past. We show that the populations of western and far eastern Europe followed opposite trajectories between 8,000-5,000 years ago. At the beginning of the Neolithic period in Europe, ~8,000-7,000 years ago, closely related groups of early farmers appeared in Germany, Hungary, and Spain, different from indigenous hunter-gatherers, whereas Russia was inhabited by a distinctive population of hunter-gatherers with high affinity to a ~24,000 year old Siberian6. By ~6,000-5,000 years ago, a resurgence of hunter-gatherer ancestry had occurred throughout much of Europe, but in Russia, the Yamnaya steppe herders of this time were descended not only from the preceding eastern European hunter-gatherers, but from a population of Near Eastern ancestry. Western and Eastern Europe came into contact ~4,500 years ago, as the Late Neolithic Corded Ware people from Germany traced ~3/4 of their ancestry to the Yamnaya, documenting a massive migration into the heartland of Europe from its eastern periphery. This steppe ancestry persisted in all sampled central Europeans until at least ~3,000 years ago, and is ubiquitous in present-day Europeans. These results provide support for the theory of a steppe origin of at least some of the Indo-European languages of Europe.

Link

Strong (?) linguistic and archaeological evidence for steppe Indo-Europeans

In a new paper in Annual Review of Linguistics, David Anthony and Don Ringe make the archaeological and linguistic case for the steppe IE homeland hypothesis. It is very useful to see the evidence presented concisely in this way. Of course, I don't think that the evidence for the steppe IE hypothesis is "so strong" as it is said to be in the paper's abstract.

The authors first discuss the phylogeny of IE languages:

It seems clear that the ancestor of the Anatolian subgroup (which includes Hittite) separated from the other dialects of PIE first, so from a cladistic point of view Anatolian is half the IE family (e.g., Jasanoff 2003). Within the non-Anatolian half, it appears that the ancestor of the Tocharian subgroup (whose attested languages were spoken in Xinjiang, today in western China, until approximately the tenth century ce) separated from the other dialects before the latter had diverged much (e.g., Winter 1998, Ringe 2000). It follows that an item inherited by two or more of the daughter subgroups can be reconstructed for “early” PIE only if it is attested in at least one Anatolian language and at least one non-Anatolian language, and such an item can be reconstructed for the ancestor of the non-Anatolian subgroups only if it is attested in one or both of the Tocharian languages and in some other IE language. 

This doesn't seem to be evidence for the steppe hypothesis, but rather for the Anatolian one. The authors hypothesis a pre-4000BC Out-of-Steppe migration into the Balkans (migration 1 left), but that takes you only into the Balkans and not into all the places in Anatolia where IE languages were spoken historically (right). The hypothesis that pre-4000BC Proto-Anatolians migrated from the steppe must bridge quite a lot of ground to reach the historical Anatolians of the 2nd and 1st millennium BC. It must also explain that the physical anthropological change in Anatolia in the Chalcolithic and the Bronze Age is associated with migration of brachycephals which seems incompatible with movements from either the Copper Age Balkans or the Pontic-Caspian steppe that were occupied by gracile and robust dolicho-mesocephals respectively. Migration 1 is of course possible but it is hardly a better explanation for the first-order split of Anatolian vs. post-Anatolian Indo-European than expansions from the Neolithic "womb of nations" that included Anatolia. Arrows 1 and 2 do harmonize with the proposed linguistic phylogeny. But, why would steppe Indo-Europeans first migrate southwest, then east, and then west? What was the cause of this particular sequence of migrations (which is invoked to harmonize the the linguistic evidence)? I am perfectly willing to consider linguistic replacement in Anatolia (it happened at least twice in recorded history, first with Greek and then with Turkish), but the case is not particularly strong that it happened from a pre-4000BC Out-of-Steppe movement via the eastern Balkans. As for the Tocharians, their recorded language of the 1st millennium AD is 4ky removed from movement 2 to the Altai, so even if future discoveries convincingly prove this movement, the yawning gap of 4 thousand years will remain. My analysis of modern Uygurs shows that the Caucasoid element in the Turkic inhabitants of Eastern Turkestan (which presumably includes the Indo-European element) is complex, and perfectly compatible with a non-steppe, but rather West Asian ultimate origin of Tocharians.

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The authors next discuss the wheel vocabulary (left). The idea is that wheeled vehicles weren't known when farmers colonized Europe, so if PIE has terminology for wheeled vehicles, then it has to be later. I don't find these arguments convincing, because words change meanings, and PIE doesn't have a word for wheeled vehicle, but for a variety of its components, each of which might have meant something else originally.

But, even if one concedes wheeled vehicles, it's still the case that neither Anatolian nor Tocharian has a repertoire of many terms for wheeled vehicles. So, all the wheel vocabulary proves (if one accepts it at all) is that post-Anatolian or even post-Tocharian languages had wheeled vehicles. This then provides a chronological constraint to the spread of post-Anatolian (or post-Tocharian) IE languages after the invention of wheeled vehicles, but tells us absolutely nothing about where they spread from (and hence the steppe hypothesis) but only when they spread. Thus, the wheel vocabulary is consistent with a whole number of theories that propose a spread of IE languages after the invention of wheeled vehicles and provides no special support for the steppe hypothesis over others.

The authors next discuss the lexical borrowings into Proto-Uralic and Proto-Finno-Ugrian. But, this only shows that early Uralic and Finno-Ugrian speakers came into contact with early Indo-Europeans, not that Indo-Europeans originated near Uralic and Finno-Ugrian speakers. Moreover, the authors also mention that PIE shows evidence of contacts with Proto-Kartvelian speakers. By the same line of argument, the PIE homeland should be close to Georgia where Proto-Kartvelian languages were presumably spoken. Indeed, Uralic languages are very widely spoken from Europe to eastern Siberia and the Uralic geographical constraint is much weaker than the Kartvelian one, as there are many parts of Eurasia (including the Pontic-Caspian steppe) that Uralic languages may have been spoken of, but a very small part of Eurasia (the southern Caucasus and northeastern Anatolia) for which any evidence of Kartvelians exists. The authors suggest that a steppe PIE explains both Proto-Uralic and Proto-Kartvelian borrowings as the steppe is between presumable speakers of these two language families. True, but Northeast and Northwest Caucasian speakers lie between Proto-Kartvelians and the steppe. A PIE origin on the steppe must bypass the NW/NE Caucasian speakers to bring Indo-Europeans in contact with Kartvelians, and a PIE origin in highland West Asia must bypass the same to bring them into contact with Uralic speakers. In sum, I don't see at all how the evidence from lexical borrowings favors the steppe hypothesis strongly.

The authors next discuss the archaeological implications of placing the homeland on the steppe and discuss how it may have been carried out. I continue to think that the spread of metallurgy is the most obvious candidate for an enabling factor. Having the capacity to build and trade metal objects, or kill people with metal weapons gives one obvious advantages that are unquestionable in every circumstance in the way that flocks of cattle and sheep, horses, and wheeled vehicles are not. Not only Indo-Europeans but also Semitic speakers may have been enabled by metallurgy to spread their languages. Neither Indo-Europeans nor Semites may have been master metallurgists, but metallurgical progress enabled language spread just as other technological innovations (e.g., stirrups, firearms, ocean-worthy boats) did in more recent history.

The authors then criticize past work on phylogenetic modeling of languages and end their article with this sentence:

Work currently in progress by the team of Chang, Hall, Cathcart, and Garrett promises to fill that gap.

An article by these authors is listed in the journal Language website, albeit without the actual text of the article yet:

Ancestry-constrained phylogenetic analysis supports the Indo-European steppe hypothesis 

Will Chang, Chundra Cathcart, David Hall and Andrew Garrett, University of California, Berkeley 

Discussion of Indo-European origins and dispersal focuses on two hypotheses. Qualitative evidence from reconstructed vocabulary and correlations with archaeological data suggest that Indo-European languages originated in the Pontic-Caspian steppe and spread together with cultural innovations associated with pastoralism, beginning c. 6500–5500 BP. An alternative hypothesis, according to which Indo-European languages spread with the diffusion of farming from Anatolia, beginning c. 9500–8000 BP, is supported by statistical phylogenetic and phylogeographic analyses of lexical traits. The time and place of the Indo-European ancestor language therefore remain disputed. Here we present a phylogenetic analysis in which ancestry constraints permit more accurate inference of rates of change, based on observed changes between ancient or medieval languages and their modern descendants, and we show that the result strongly supports the steppe hypothesis. Positing ancestry constraints also reveals that homoplasy is common in lexical traits, contrary to the assumptions of previous work. We show that lexical traits undergo recurrent evolution due to recurring patterns of semantic and morphological change.

It's not clear what the article itself shows. If this is simply a criticism of the "old" dates of the first PIE split proposed by Gray/Atkinson and Bouckaert et al., then this does not really support the steppe hypothesis uniquely, but rather argues against the Neolithic Anatolian one. However, there are many hypotheses other than these two (including my Bronze Age expansion of Indo-European languages hypothesis from West Asia which is somewhat related to that of Stanislav Grigoriev) that can accommodate a later split of PIE.

To conclude, I think that archaeology and linguistics have failed to make a convincing case for steppe Proto-Indo-Europeans. It is certainly a respectable and popular theory but the evidence for it is hardly "so strong" as to create serious problems for other hypotheses. Hopefully, archaeogenetics will succeed where archaeology and linguistics (despite their valiant efforts) have failed.

Annual Review of Linguistics Vol. 1: 199-219 (Volume publication date January 2015) DOI: 10.1146/annurev-linguist-030514-124812

David W. Anthony1 and Don Ringe2

The Indo-European Homeland from Linguistic and Archaeological Perspectives

Archaeological evidence and linguistic evidence converge in support of an origin of Indo-European languages on the Pontic-Caspian steppes around 4,000 years BCE. The evidence is so strong that arguments in support of other hypotheses should be reexamined.

Link

Indo-Europeans preceded Finno-Ugrians in Finland and Estonia

According to an abstract of a Ph.D thesis (below). This would appear to work well with the dating of the signature Y-chromosome haplogroup of Finno-Ugrians. 

Bidrag till Fennoskandiens språkliga förhistoria i tid och rum (Heikkilä, Mikko)

My academic dissertation "Bidrag till Fennoskandiens språkliga förhistoria i tid och rum" ("Spatiotemporal Contributions to the Linguistic Prehistory of Fennoscandia") is an interdisciplinary study of the linguistic prehistory of Northern Europe chiefly in the Iron Age (ca. 700 BC―AD 1200), but also to some extent in the Bronze Age (ca. 1700―700 BC) and the Early Finnish Middle Ages (ca. AD 1200―1323). The disciplines represented in this study are Germanistics, Nordistics, Finnougristics, history and archaeology. The language-forms studied are Proto-Germanic, Proto-Scandinavian, Proto-Finnic and Proto-Sami. This dissertation uses historical-comparative linguistics and especially loanword study to examine the relative and absolute chronology of the sound changes that have taken place in the proto-forms of the Germanic, Finnic and Samic languages. Phonetic history is the basis of historical linguistics studying the diachronic development of languages. To my knowledge, this study is the first in the history of the disciplines mentioned above to examine the systematic dating of the phonetic development of these proto-languages in relation to each other. In addition to the dating and relating of the phonetic development of the proto-languages, I study Fennoscandian toponyms. The oldest datable and etymologizable place-names throw new light on the ethnic history and history of settlement of Fennoscandia. For instance, I deal with the etymology of the following place-names: Ahvenanmaa/Åland, Eura(joki), Inari(järvi), Kemi(joki), Kvenland, Kymi(joki), Sarsa, Satakunta, Vanaja, Vantaa and Ähtäri. 

My dissertation shows that Proto-Germanic, Proto-Scandinavian, Proto-Finnic and Proto-Sami all date to different periods of the Iron Age. I argue that the present study along with my earlier published research also proves that a (West-)Uralic language – the pre-form of the Finnic and Samic languages – was spoken in the region of the present-day Finland in the Bronze Age, but not earlier than that. In the centuries before the Common Era, Proto-Sami was spoken in the whole region of what is now called Finland, excluding Lapland. At the beginning of the Common Era, Proto-Sami was spoken in the whole region of Finland, including Southern Finland, from where the Sami idiom first began to recede. An archaic (Northwest-)Indo-European language and a subsequently extinct Paleo-European language were likely spoken in what is now called Finland and Estonia, when the linguistic ancestors of the Finns and the Sami arrived in the eastern and northern Baltic Sea region from the Volga-Kama region probably at the beginning of the Bronze Age. For example, the names Suomi ʻFinlandʼ and Viro ʻEstoniaʼ are likely to have been borrowed from the Indo-European idiom in question. (Proto-)Germanic waves of influence have come from Scandinavia to Finland since the Bronze Age. A considerable part of the Finnic and Samic vocabulary is indeed Germanic loanwords of different ages which form strata in these languages. Besides mere etymological research, these numerous Germanic loanwords make it possible to relate to each other the temporal development of the language-forms that have been in contact with each other. That is what I have done in my extensive dissertation, which attempts to be both a detailed and a holistic treatise.

Ancient central European mtDNA across time (Brandt, Haak et al. and Bollongino et al.)

Two important new papers appeared in Science today. In the first one (Brandt, Haak et al.), researchers compiled mtDNA results from 364 prehistoric central Europeans from the early Neolithic to the early Bronze Age, spanning about four millennia of history. Importantly they uncover not a smooth transition between early Neolithic farmers and modern Europeans, but a punctuated series of haplogroup frequency changes that cannot really be explained by genetic drift in a single European population evolving over time. Hopefully this kind of research can be repeated in other parts of the world, as it provides a way to see evolution and migration as it happens.

Earlier work has disproved the hypothesis that modern Europeans are simply "acculturated" hunter-gatherers, and this newer research disproves the idea that they are simply the descendants of early farmers, little modified since the beginning of the Neolithic.

I am sure that myself and others will spend some time trying to digest the wealth of information present in the paper and its supplementary materials. Yet, one conclusion can already be made, that migrationism is alive and well. Anyone adhering to a "pots not people" paradigm will find difficult to explain the sharp discontinuities found in the genetic record. European foragers contrast with the earliest farmers, who, in turn, contrast with and the Late Neolithic copper cultures that supplanted them a few thousand years later and spawned the Bronze Age world. If pots aren't people, it's strange that archaeological cultures defined largely by pots (right) also appear to mark genetic contrasts.

These discontinuities are most evident in Figure 3 from the paper:

You may follow the grey line to see how central Europe, once populated exclusively by hunter-gatherers, experienced a virtual disappearance of their matrilineages for almost two thousand years after the advent of farming.  Then, between the Middle to Late Neolithic, around five thousand year ago, the hunter-gatherers make their re-appearance before their lineages converge to their modern (minority) frequency. The authors present a model of migration to explain these events, illustrated in a movie in the supplementary material, and also in the figure on the left.

Of particular interest is a set of haplogroups marked by the yellow line (I, U2, T1, R) and are most strongly represented in the Unetice and Corded Ware samples before reverting to a small minority in the present-day. These may be potentially very informative to understand the c. 5,000-year old ago upheaval. I reproduce below three of the genetic distance maps from the supplement for the three latest cultures (CWC: Corded Ware; BBC: Bell Beaker; and UC: Unetice):

I note the European-ness of Bell Beaker (probably due to elevated frequencies of haplogroup H) and the eastern European-ness/west Asian-ness of Corded Ware/Unetice.

Moving on to the next shorter paper by Bollongino et al. which produces evidence for an interesting hypothesis: that hunter-gatherers did not disappear in central Europe after the introduction of farming, but some of their descendants persisted for at least two thousand years afterwards:

In summary, the results of 14C and stable isotope analysis, together with the DNA evidence, suggest that the Blätterhöhle individuals are sampled from three distinct populations: (i) Mesolithic hunter-gatherers, (ii) Neolithic farmers, and (iii) Neolithic fisher-hunter-gatherers (special-izing in freshwater fish). The latter two notably date to the fourth mil-lennium BC, which is around 2000 years after the introduction of farming to Central Europe.

I was reminded of an older paper about first contact between farmers and hunter-gatherers. An important consequence of the second paper is that hunter-gatherer lineages in modern Europeans may have come not only from outlying areas where foragers persisted in greater numbers, but also from within the farming realm itself.

Science 11 October 2013: Vol. 342 no. 6155 pp. 257-261 DOI: 10.1126/science.1241844

Ancient DNA Reveals Key Stages in the Formation of Central European Mitochondrial Genetic Diversity 

Guido Brandt, Wolfgang Haak et al.

The processes that shaped modern European mitochondrial DNA (mtDNA) variation remain unclear. The initial peopling by Palaeolithic hunter-gatherers ~42,000 years ago and the immigration of Neolithic farmers into Europe ~8000 years ago appear to have played important roles but do not explain present-day mtDNA diversity. We generated mtDNA profiles of 364 individuals from prehistoric cultures in Central Europe to perform a chronological study, spanning the Early Neolithic to the Early Bronze Age (5500 to 1550 calibrated years before the common era). We used this transect through time to identify four marked shifts in genetic composition during the Neolithic period, revealing a key role for Late Neolithic cultures in shaping modern Central European genetic diversity.

Link

Science DOI: 10.1126/science.1245049

2000 Years of Parallel Societies in Stone Age Central Europe

Ruth Bollongino et al.

Debate on the ancestry of Europeans centers on the interplay between Mesolithic foragers and Neolithic farmers. Foragers are generally believed to have disappeared shortly after the arrival of agriculture. To investigate the relation between foragers and farmers, we examined Mesolithic and Neolithic samples from the Blätterhöhle site. Mesolithic mitochondrial DNA sequences were typical of European foragers, whereas the Neolithic sample included additional lineages that are associated with early farmers. However, isotope analyses separate the Neolithic sample into two groups: one with an agriculturalist diet and one with a forager and freshwater fish diet, the latter carrying mitochondrial DNA sequences typical of Mesolithic hunter-gatherers. This indicates that the descendants of Mesolithic people maintained a foraging lifestyle in Central Europe for more than 2000 years after the arrival of farming societies.

Link

Multiple sources of European barley

This seems to parallel quite well with my hypothesis of a secondary expansion into Europe of the component I've labeled "West Asian", after the early Neolithic. From the paper:

The wild progenitor of barley, Hordeum spontaneum, is typically a winter-germinating species, responsive to day length (Turner et al. 2005), but recent work (H. Jones et al. 2008) has demonstrated that the non-responsive allele also occurs in wild barley in Israel, Jordan and Iran, in regions where this allele is favoured by the climatic conditions. There is strong evidence to suggest that the non-responsive allele in European cultivars (Group A) is genetically more similar to the allele in Iranian wild barleys than to the allele in wild barleys from Israel and Jordan.

and:

 Detailed genetic comparisons between groups A, B and C (H. Jones 2008; H. Jones et al. 2008) suggest that Group A represents a more recent introduction of day-length non-responsive barley into Europe, rather than the selection of non-responsiveness from within populations that diffused into Europe in the Early Neolithic. In addition, three of the nine principal groups identified on the basis of the neutral SSR markers (groups 1–3) are genetically closely related and very similar in their phenotypic characteristics. They are located in north-west and Central Europe (Figure 3b), and are made up almost entirely of  day-length non-responsive barleys (98 per cent) with a spring growth habit (98 per cent) (H. Jones et al. 2011). The distinctiveness of the landraces making up groups 1–3 suggests that the responsive/non-responsive difference between Southern and Northern Europe is not simply a reflection of current selective pressure but rather is one aspect of a more fundamental genetic difference between barley populations. These day-length non-responsive cultivars have therefore been interpreted as representing a later spread of barley into Northern Europe from the eastern part of the Fertile Crescent or beyond. This raises the issue of when they spread into Europe and whether or not they were introduced via the same route as the initial  spread of agriculture.

and:

A possible channel for the introduction of day-length non-responsive barley into Northern Europe from the east at this time could have been through early exchange networks with metal-working communities of south-eastern Europe (Sherratt 1976; Bogucki 1999: 220–21). The Chalcolithic societies of the Balkans also maintained connections further east with the west Eurasian Steppes, Anatolia, and the Caucasus in the fifth and early fourth millennia, and copper was exchanged from west to east across the steppes north of the Black Sea (Kohl 2007: 31–39). Day-length non-responsive barley could, therefore, have travelled from east to west along the same route, north of the Black Sea via the Caucasus or, perhaps more likely given its agricultural nature, south of the Black Sea through Anatolia, possibly via a coastal route around the Black Sea, and then from the Carpatho-Balkan metallurgical region into Northern Europe (Figure 3b). A similar suggestion has been made for the later appearance of the oil plant Lallemantia, which was apparently introduced into Europe in the third millennium BC, possibly along the same trade routes as tin-bronzes (Valamoti & Jones 2010). 

Antiquity Volume: 87 Number: 337 Page: 701–713

DNA evidence for multiple introductions of barley into Europe following dispersed domestications in Western Asia

G. Jones et al.

It has long been recognised that the Neolithic spread across Europe via two separate routes, one along the Mediterranean coasts, the other following the axis of the major rivers. But did these two streams have a common point of origin in south-west Asia, at least with regard to the principal plant and animals species that were involved? This study of barley DNA shows that the domesticated barley grown in Neolithic Europe falls into three separate types (groups A, B and C), each of which may have had a separate centre of origin in south-west Asia. Barley was relatively rarely cultivated by the early Linearbandkeramik farmers of Central and Northern Europe, but became more common during the fifth and fourth millennia BC. The analysis reported here indicates that a genetic variety of barley more suitable for northern growing conditions was introduced from south-west Asia at this period. It also suggests that the barley grown in south-eastern Europe at the very beginning of the Neolithic may have arrived there by different routes from two separate centres of domestication in south-west Asia. The multiple domestications that this pattern reveals imply that domestication may have been more a co-evolutionary process between plants and people than an intentional human action.

Link

The place of Armenian in the Indo-European language family

Journal of Language Relationship • Вопросы языкового родства • 10 (2013) • Pp. 85—137

The place of Armenian in the Indo-European language family: the relationship with Greek and Indo-Iranian*

Hrach Martirosyan

The main purpose of this paper is to present lexical correspondences that unite Armenian with Greek and/or Indo-Iranian. They include shared innovations on the one hand, and isolated lexemes on the other. These two lexical corpora — lexical innovations on an inherited basis and isolated words — can be placed within the same temporal and spatial framework. After the Indo-European dispersal Proto-Armenian would have continued to come into contact with genetically related Indo-European dialects. Simultaneously, it would certainly also have been in contact with neighbouring non-Indo-European languages. A word can be of a substrate origin if it is characterized by: (1) limited geographical distribution; (2) unusual phonology and word formation; (3) characteristic semantics. The material presented here, albeit not exhaustive, allows to preliminarily conclude that Armenian, Greek, (Phrygian) and Indo-Iranian were dialectally close to each other. Within this hypothetical dialect group, Proto-Armenian was situated between Proto-Greek (to the west) and Proto-Indo-Iranian (to the east). The Indo-Iranians then moved eastwards, while the Proto-Armenians and Proto-Greeks remained in a common geographical region for a long period and developed numerous shared innovations. At a later stage, together or independently, they borrowed a large number of words from the Mediterranean / Pontic substrate language(s), mostly cultural and agricultural words, as well as animal and plant designations. On the other hand, Armenian shows a considerable number of lexical correspondences with European branches of the Indo-European language family, a large portion of which too should be explained in terms of substrate rather than Indo-European heritage.

Link (pdf)