Focus on Hittite Anatolia

I computed a series of D-statistics on most of the currently available ancient samples from Central Anatolia - dating from almost the Epipaleolithic (Boncuklu_N) to the Hittite era (Anatolia_MLBA) - to try and get a better idea of who the Indo-European-speaking Hittites may have been. The full output as well as details about the key ancient samples used in this analysis are available here. See anything interesting? The most noteworthy statistics, I suppose, are those listed below, because they're significant (Z≥3) and organized chronologically.

However, the thing to keep in mind in regards to D-statistics, and the very similar f4-statistics, when looking for signals of mixture is that they may or may not produce significant Z scores because of several reasons, such as the choice of the outgroup, the choice of the reference samples and the phylogenetic relationship between them, or even the type, quality and density of the data being used.

Perhaps ironically, the D-statistics above suggest that the Neolithic Central Anatolians (Boncuklu_N and Tepecik_Ciftlik_N) were more European-like than those from the Bronze Age, and I suspect that this is one of the main reasons why the idea of Eastern European admixture (from the Pontic-Caspian steppe and/or Balkans) in Hittites is currently being rejected by the geneticists working on the problem. But this dilemma is easy to explain away by the fact that the Neolithic samples carry much higher ratios of Anatolian Epipaleolithic hunter-gatherer admixture and also other types of ancestry shared with and/or closely related to European hunter-gatherers and early farmers.

In other words, I'd say that most of the statistics are being confounded by deep phylogenetic relationships, and thus aren't very useful for solving the Hittite problem. Interestingly, though, that relationship to Europe is reversed somewhat in the D-statistics involving Anatolia_EBA and Anatolia_MLBA, with the latter showing significantly higher affinity to Eastern European Hunter-Gatherers (EHG) and Minoans.

Thus, in my opinion, to get a more complete picture it's also useful to look for patterns in the statistics, even those that, strictly speaking, don't reach significance. One way to do that is with linear models. So here are a few linear models based on some of my D-statistics. The relevant datasheet is available here.

Arguably, the most striking thing about these models is the position at the top of the graphs of the ancient populations from Central Asia and what is now Iran, and the gradually lower position of populations with progressively less of this type of ancestry. The most plausible explanation for this phenomenon is post-Boncuklu_N gene flow into Central Anatolia from the east, possibly as a continuation of something that was happening already since the Epipaleolithic, but becoming more intense during the Neolithic revolution, probably as a result of rapid population growth in and around the Fertile Crescent.

Indeed, I strongly suspect that one of the main reasons why we've been hearing so much lately about Iran as a likely candidate for the Indo-European homeland is this strong eastern signal in Bronze Age Anatolian DNA. If so, then this is likely to be a misunderstanding, because there are better explanations for it than the Indo-Europeans, such as the Hattians and Hurrians.

Another rather obvious outcome in my graphs is the relatively stronger affinity between the Bronze Age Anatolians and the ancient populations from Eastern Europe, including, and especially, those from the Pontic-Caspian steppe, compared to Tepecik_Ciftlik_N. In fact, looking at the Anatolia_EBA vs Tepecik_Ciftlik_N graph, I'd say that steppe admixture was already seeping into Central Anatolia during the Early Bronze Age.

If so, this is an important point that should be taken into account when modeling the ancestry of the Hittite era Anatolians. That's because if Anatolia_EBA already harbored some steppe ancestry, then we'd be shooting ourselves in the proverbial foot if we were to use it as the supposedly unadmixed reference population to try and determine whether Anatolia_MLBA was partly of steppe origin. Hence, to model the ancestry of Anatolia_MLBA, at least in the context of possible migrations from the steppe to Anatolia during the Bronze Age, it might be more useful to use Tepecik_Ciftlik_N as the likely unadmixed reference population.

Let's try that with qpAdm, first on the whole Anatolia_MLBA set, and then on one individual labeled MA2203, who, as far as I can tell, shows an elevated level of steppe ancestry in several different types of analyses. I chose Yamnaya_Kalmykia as the potential mixture source from the steppe because it's likely to be the closest available population in my dataset to the Eneolithic groups of the southern region of the Pontic-Caspian steppe.

Anatolia_MLBA
Seh_Gabi_ChL 0.200±0.043
Tepecik_Ciftlik_N 0.659±0.033
Yamnaya_Kalmykia 0.141±0.022
chisq: 11.425
tail prob: 0.408405
Full output

MA2203
Seh_Gabi_ChL 0.179±0.065
Tepecik_Ciftlik_N 0.622±0.049
Yamnaya_Kalmykia 0.199±0.036
chisq: 12.914
tail prob: 0.299004
Full output

Please note, however, that these mixture models are based on f4-statisctics. So, obviously, they're going to be affected by the same factors as described above that affect f4-statistics. Hence, despite the seemingly statistically sound output, the steppe admixture that you see there might not actually be admixture from the steppe.

In fact, there's a good reason why I'm not shouting from the rooftops that I've just uncovered the presence of steppe ancestry in Bronze Age Anatolia, and thus confirmed the steppe or kurgan hypothesis positing that the Hittite and indeed Indo-European homeland was located in the Pontic-Caspian steppe. That's because I used a mixed bag of UDG-treated capture data and non-UDG-treated shotgun data. This is known to be a serious problem, which can skew the results of even the most robust analyses, and produce spurious statistics and Z scores.

Nevertheless, I'm reasonably confident that my findings will eventually be confirmed with more and higher quality data from ancient Anatolia. Let's wait and see.

See also...

Late PIE ground zero now obvious; location of PIE homeland still uncertain, but...

Global25 workshop 3: genes vs geography in Northern Europe

To produce the intra-North European Principal Components Analysis (PCA) plot below, download this datasheet, plug it into the PAST program, which is freely available here, then select all of the columns by clicking on the empty tab above the labels, and choose Multivariate > Ordination > Principal Components or Discriminant Analysis. This is what you should end up with...

I'd say that the result more or less resembles a geographic map of Northern Europe. Of course, if you're in the possession of your own personal Global25 coordinates, you can add yourself to this plot to check whether your position matches your geographic origin.

Please keep in mind, however, that the vast majority (>90%) of your ancestry must be from north of the Alps, Balkans and Pyrenees to obtain a sensible outcome. Also please ensure that all of the columns in the datasheet are filled out correctly, including the group column, otherwise your position on the plot will be skewed.

See also...

Global25 workshop 1: that classic West Eurasian plot

Global25 workshop 2: intra-European variation

Global25 workshop 4: a neighbour joining tree

Getting the most out of the Global25

Genetic ancestry online store (to be updated regularly)

The South Asian cline that no longer exists

Before the Indo-Europeans and Austroasiatics got to South Asia, probably well within the last 4,000 years, it's likely that all of the genetic variation in the region basically sat along a genetic cline devoid of any Bronze Age steppe and Southeast Asian ancestry, like the one in the Principal Component Analysis (PCA) below running from the Paniya to the "Indus Periphery" ancient sample Shahr_I_Sokhta BA2.

Note that almost all of the South Asian populations, including the Iron Age (IA) Swat Valley groups, are clearly peeling away from the said cline towards the Tajiks, in other words towards Central Asia. This is a reflection of the widespread presence of Sintashta-related steppe admixture among South Asians, especially those speaking Indo-European languages. Moreover, the Bangalis and Burushos are being pushed towards the top left of the plot as a result of East Asian-related ancestry. In the case of the former, this is largely due to gene flow from Austroasiatic groups.

It'll be interesting to see how ancient Harappans behave in this analysis. I'm betting that they'll be very similar to the Indus Periphery trio, although judging by the latest press report on the topic (see here), the Harappan samples from Rakhigarhi might be shifted much closer to the Paniya as a result of a higher ratio of indigenous South Asian ancestry.

The PCA is based on my Global25 test. If you're South Asian and in the possession of Global25 coordinates, you can add yourself to this plot using the datasheet available here. Plug the datasheet into the PAST program (freely available here), select all of the columns, and go Multivariate > Ordination > Principal Components (PCA).

Update 10/08/2018: I managed to almost reproduce my PCA with a graph based on D-stats of the form D(Mbuti,X)(Onge,Ganj_Dareh_N)/D(Mbuti,X)(Ganj_Dareh_N,Sintashta_MLBA). Admittedly, Gonur2_BA didn't want to cooperate by pushing slightly up and away from the ghost South Asian cline. But this may have been due to a lack of data or perhaps minor admixture (keep in mind that this sample is actually from Turkmenistan and not South Asia). However, combining all three of the Indus_Periphery individuals worked well enough. The relevant datasheet is available here.

See also...

Global25 workshop 1: that classic West Eurasian plot

Global25 workshop 2: intra-European variation

Global25 workshop 3: genes vs geography in Northern Europe

Global25 workshop 4: a neighbour joining tree

Getting the most out of the Global25

Genetic ancestry online store (to be updated regularly)

Blast from the past: The Poltavka outlier

The Rakhigarhi ancient DNA paper is coming soon. Very soon.

Yep, you've probably read this sort of thing many times in the last few years, including here. But this time, by all accounts, it's really happening. For the latest Indian press teaser on the topic check out: We Are All Harappans.

At least I don't have to write up a blog post for the occasion, because I already wrote one over two years ago, and it's still current, more or less. Click on the screen cap below to teleport yourselves back to January 2016. And make sure to peruse the comments under the article. Hilarious stuff.

Of course, since then I've written many more posts dealing with South Asian population history, including some that are based on recently published ancient DNA from the region. Here are my picks:

On the doorstep of India

The mystery of the Sintashta people

Yamnaya isn't from Iran just like R1a isn't from India

Indian confirmation bias

The protohistoric Swat Valley "Indo-Aryans" might not be exactly what we think they are

The Out-of-India Theory (OIT) challenge: can we hear a viable argument for once?

Horses may have been ridden in battle as early as the Bronze Age (Chechushkov et al. 2018)

Over at the Journal of Archaeological Science at this LINK. Below is the abstract. Emphasis is mine:

The morphological similarities/dissimilarities between antler and bone-made cheekpieces have been employed in several studies to construct a relative chronology for Bronze Age Eurasia. Believed to constitute a part of the horse bit, the cheekpieces appear in ritual contexts everywhere from the Mycenaean Shaft Graves to the Bronze Age kurgan cemeteries in Siberia. However, these general understandings of the function and morphological changes of cheekpieces have never been rigorously tested. This paper presents statistical analyses (e.g., similarities, multidimensional scaling, and cluster analysis) that document differences in cheekpiece morphology, comparing shield-like, plate-formed, and rod-shaped types in the context of temporal change and spatial variation. We investigated changes in function over time through the use of experimental replicas used in bridling horses. This experimental work supports the hypothesis that these objects served to bridle harnessed (shield-like) or ridden (plate-formed and rod-shaped) horses. Moreover, comparison of use wear on the ancient artifacts with the replicas provides insight into how long the artifacts were used before they were deposited in the funeral contexts or discarded. These observations support that the Sintashta chariots dating back to ca. 2100 BC were ridden and suggest the end of the Late Bronze Age (ca. 1500–1200 BC) as the earliest possible date for horseback riding in warfare. This study highlights changes in horse exploitation and simultaneous shifts in human societies.

Chechushkov et al., Early horse bridle with cheekpieces as a marker of social change: An experimental and statistical study, Journal of Archaeological Science, Volume 97, September 2018, Pages 125-136, https://doi.org/10.1016/j.jas.2018.07.012

See also...

Of horses and men

An early Iranian, obviously

Graeco-Aryan parallels

A closer look at the maternal origins of the Corded Ware people (Juras et al. 2018)

Over at Scientific Reports at this LINK. This is a nice paper, but I'm really looking forward to the Y-DNA and genome-wide data from these new samples. What's the bet that the Yamnaya men from Ukraine will belong to Y-haplogroup R1a-M417? Bring it on soon, please. From the paper, emphasis is mine:

From around 4,000 to 2,000 BC the forest-steppe north-western Pontic region was occupied by people who shared a nomadic lifestyle, pastoral economy and barrow burial rituals. It has been shown that these groups, especially those associated with the Yamnaya culture, played an important role in shaping the gene pool of Bronze Age Europeans, which extends into present-day patterns of genetic variation in Europe. Although the genetic impact of these migrations from the forest-steppe Pontic region into central Europe have previously been addressed in several studies, the contribution of mitochondrial lineages to the people associated with the Corded Ware culture in the eastern part of the North European Plain remains contentious. In this study, we present mitochondrial genomes from 23 Late Eneolithic and Bronze Age individuals, including representatives of the north-western Pontic region and the Corded Ware culture from the eastern part of the North European Plain. We identified, for the first time in ancient populations, the rare mitochondrial haplogroup X4 in two Bronze Age Catacomb culture-associated individuals. Genetic similarity analyses show close maternal genetic affinities between populations associated with both eastern and Baltic Corded Ware culture, and the Yamnaya horizon, in contrast to larger genetic differentiation between populations associated with western Corded Ware culture and the Yamnaya horizon. This indicates that females with steppe ancestry contributed to the formation of populations associated with the eastern Corded Ware culture while more local people, likely of Neolithic farmer ancestry, contributed to the formation of populations associated with western Corded Ware culture.

We investigated the within- and between-group variability using an AMOVA analysis. Concentrating on the eastern and western Corded Ware groups, we found the best variability distribution when the individuals associated with the western Corded Ware culture (CWW in Supplementary Table S5) were grouped together with the Middle Neolithic/Bronze Age Central Europe groups, while individuals associated with the eastern and Baltic Corded Ware culture (CWPlM, CWBal), and Yamnaya horizon groups (YAW and YAE) clustered together with the eastern Europe populations (from the Middle Neolithic-Bronze Age) (4.68% of variability among groups, 3.04% among populations within groups).

By analyzing ancient mitochondrial genomes, we show that people from the eastern and western Corded Ware culture were genetically differentiated. Individuals associated with the eastern Corded Ware culture (from present day Poland and the Czech Republic) shared close maternal genetic affinity with individuals associated with the Yamnaya horizon while the genetic differentiation between individuals associated with the western Corded Ware culture (from present-day Germany) and the Yamnaya horizon was more extensive. This decreasing cline of steppe related ancestry from east to west likely reflect the direction of the steppe migration. It also indicates that more people with steppe-related ancestry, likely both females and males, contributed to the formation of the population associated with the eastern Corded Ware culture. Similarly, closer genetic affinity to populations associated with Yamnaya horizon can be observed in Baltic Corded Ware groups, which confirms earlier indications of a direct migrations from the steppe not only to the west but also to the north, into the eastern Baltic region [18,19,55]. The mitochondrial data further suggests that with increased distance from the source populations of the steppe, the contribution of local people increase, which is seen as an increase of maternal lineages of Neolithic farmer ancestry in individuals associated with the western Corded Ware culture.

...

Interestingly, hg U4c1 found in the Yamnaya individual (poz224) has so-far been found only in two Bell Beaker- associated individuals [61] and one Late Bronze Age individual from Armenia [14], which might suggest a steppe origin for hg U4c1. A steppe origin can possibly also be assigned to hg U4a2f, found in one individual (poz282) but not reported in any other ancient populations to date, and to U5a1- the ancestral lineage of U5a1b, reported for individual poz232, which was identified not only in Corded Ware culture-associated population from central and eastern Europe [55,61] but also in representatives of Catacomb culture from the north Pontic region [24], Yamnaya from Bulgaria and Russia [17,46], Srubnaya [23] and Andronovo [62] -associated groups. Hg U2e, reported for Late Eneolithic individual (poz090), was also identified in western Corded Ware culture-associated individual23 and in succeeding Sintashta14, Potapovka and Andronovo [23] groups, suggesting possible genetic continuity of U2e1 in the western part of the north Pontic region.

Hgs W3a1 and W3a1a, found in two Yamnaya individuals from this study (poz208 and poz222), were also identified in Yamnaya-associated individuals from the Russia Samara region [17] and later in Únětice and Bell Beaker groups from Germany [61,63], supporting the idea of an eastern European steppe origin of these haplotypes and their contribution to the Yamnaya migration toward the central Europe. The W3a1 lineage was not identified in Neolithic times and, thus, we assume that it appeared in the steppe region for the first time during the Bronze Age. Notably, hgs W1 and W5, which predate the Bronze Age in Europe, were found only in individuals associated with the early Neolithic farmers from Starčevo in Hungary (hg W5)64, early Neolithic farmers from Anatolia (hg W1-T119C) [23], and from the Schöningen group (hg W1c)61 and Globular Amphora culture from Poland (hg W5) [45].

Juras et al., Mitochondrial genomes reveal an east to west cline of steppe ancestry in Corded Ware populations, Scientific Reports, 02 August 2018, DOI: https://doi.org/10.1038/s41598-018-29914-5

See also...

The staging point, obviously

Late PIE ground zero now obvious; location of PIE homeland still uncertain, but...

A Mycenaean and an Iron Age Iranian walk into a bar...

What do they have in common? The same type of Near Eastern ancestry? From Iran? Nope, that's a joke. Obviously, they share the same type of steppe ancestry. This probably has some very important linguistic implications.

The relevant Principal Component Analysis (PCA) datasheet is available here. Below are two pairs of formal mixture models that support my inferences from the PCA.

Mycenaean
Srubnaya_MLBA 0.266±0.029
Tepecik_Ciftlik_N 0.734±0.029
taildiff: 0.588000631
Full output

Mycenaean
Minoan_Lasithi 0.790±0.023
Srubnaya_MLBA 0.210±0.023
taildiff: 0.187709803
Full output

...

Turkmenistan_IA
Namazga_CA 0.528±0.040
Srubnaya_MLBA 0.472±0.040
taildiff: 0.561330411
Full output

Turkmenistan_IA
Dzharkutan1_BA 0.530±0.037
Srubnaya_MLBA 0.470±0.037
taildiff: 0.485083377
Full output

But seriously, what's the direct link between populations like Tepecik_Ciftlik_N/Minoan_Lasithi and Namazga_CA/Dzharkutan1_BA, except some exceedingly distant farmer ancestry from the Fertile Crescent?

See also...

An early Iranian, obviously

Graeco-Aryan parallels

Yamnaya isn't from Iran just like R1a isn't from India

An early Iranian, obviously

Today, the part of Asia between the Caspian Sea and the Altai Mountains, known as Turan, is largely a Turkic-speaking region. But during the Iron Age it was dominated by Iranian speakers. Throughout this period it was the home of a goodly number of attested and inferred early Iranic peoples, such as the Airya, Dahae, Kangju, Massagetae, Saka and Sogdians.

Indeed, the early Iron Age Yaz II archaeological culture, located in southwestern Turan, is generally classified as an Iranian culture, and even posited to have been the Airyanem Vaejah, aka home of the Iranians, from ancient Avestan literature.

That's not to say that Iranian speakers weren't present in this part of the world much earlier. They probably were, and it's likely that we already have their genomes (see here). But the point I'm making is that Turan can't be reliably claimed to have been an Iranian realm until the Iron Age.

Ergo, any ancient DNA samples from Turan dating to the Iron Age, as opposed to, say, the Bronze Age, are very likely to be those of early Iranian speakers. One such sample is Turkmenistan_IA DA382 from Damgaard et al. 2018.

Below is a screen cap of the "time map" from homeland.ku.dk, with the slider moved to 847 BC, showing the location of the burial site where the remains of DA382 were excavated. The site is marked with the Z93 label because DA382 belongs to the Eastern European-derived Y-chromosome haplogroup R1a-Z93. Interestingly, his burial was located in close proximity to archaeological sites associated with the above mentioned and contemporaneous Yaz II culture.

DA382 didn't get much of a run in the Damgaard et al. paper, and little wonder because the authors also analyzed 73 other ancient samples. So let's take a close look at this individual's genetic structure to see whether there's anything particularly Iranian about it.

Damgaard et al. did mention that DA382 was partly of Middle to Late Bronze Age (MLBA) steppe origin. And indeed, my own mixture models using qpAdm confirm this finding with very consistent results and strong statistical fits. Here are a couple of two-way examples...

Turkmenistan_IA
Namazga_CA 0.528±0.040
Srubnaya_MLBA 0.472±0.040
taildiff: 0.561330411
Full output

Turkmenistan_IA
Dzharkutan1_BA 0.530±0.037
Srubnaya_MLBA 0.470±0.037
taildiff: 0.485083377
Full output

The fact that the MLBA Srubnaya samples from the Pontic-Caspian steppe can be used to model DA382's ancestry (alongside Bronze and Copper Age populations from Turan) with such ease shouldn't be surprising, considering the he belongs to R1a-Z93, which is the dominant Y-haplogroup in the Srubnaya and all other closely related MLBA steppe peoples.

Now, Srubnaya is generally regarded to be the proto-Iranian archaeological culture. How awesome is that considering those qpAdm fits? But, admittedly, this is just an inference, even if a robust one, based on genetic, archaeological and historical linguistics data. So apart from the fact that DA382 comes from Iron Age Turan, an Iranian-speaking realm, is there any other way to link him directly to Iranians?

Well, he's very similar in terms of overall genetic structure to some of the least Turkic-admixed Iranian speakers still living in Turan, and might well be ancestral to them.

For instance, below is a Principal Component Analysis (PCA) featuring a wide range of ancient and present-day West Eurasian samples. Note that, in line with the qpAdm models, DA382 clusters about half-way between the populations of the MLBA steppe and pre-Kurgan expansion Turan, and amongst present-day Yaghnobi and Pamiri Tajiks. In fact, he clusters at the apex of a southeast > northwest cline made up of Tajiks that appears to be pulling towards Europeans.

Needless to say, Tajiks, especially Pamiri Tajiks, also pack a lot of Srubnaya-related ancestry. I've talked about this plenty of times at this blog (for instance, see here). But what happens if I try to model Pamiri and Yaghnobi Tajiks with DA382?

Tajik
Turkmenistan_IA 0.892±0.023
Han 0.108±0.023
taildiff: 0.794566182
Full output

Wow, it's an awesome fit! My mind's made up: DA382 was probably an Iranian speaker and, more specifically, an Eastern Iranian speaker. Who disagrees and why? Feel free to let me know in the comments (unless you're banned, in which case, f*ck off).

See also...

A Mycenaean and an Iron Age Iranian walk into a bar...

Late PIE ground zero now obvious; location of PIE homeland still uncertain, but...

New PCA featuring Botai horse tamers, Hun and Saka warriors, and many more...

"The Homeland: In the footprints of the early Indo-Europeans" time map

Click HERE to view the interactive time map, and give it some time to load if you're on a slow connection. Use the slider tool to explore different time periods from 6385 BC to 1 BC. It's still a work in progress, so feel free to let the author, Mikkel Nørtoft, know if anything should be added, tweaked and/or generally improved.

Below is a screen cap of the map with the slider moved to 3618 BC. Note the sheep in the North Pontic steppe and the wheels just west of it. Wink, wink, nudge, nudge...

See also...

Late PIE ground zero now obvious; location of PIE homeland still uncertain, but...

SMBE 2018 abstracts

Abstracts from the upcoming SMBE 2018 conference (8-12 July) are now available HERE. Below are a few that I found interesting. Emphasis is mine. Feel free to post your own favorites in the comments.

The first Epipaleolithic Genome from Anatolia suggests a limited role of demic diffusion in the Advent of Farming in Anatolia

Feldman et al.

Anatolia was home to some of the earliest farming communities, which in the following millennia expanded into Europe and largely replaced local hunter-gatherers. The lack of genetic data from pre-farming Anatolians has so far limited demographic investigations of the Anatolian Neolithisation process. In particular, it has been unclear whether farming was adopted by indigenous hunter-gatherers in Central Anatolia or imported by settlers from earlier farming centers. Here we present the first genome-wide data from an Anatolian Epipaleolithic hunter-gatherer who lived ~15,000 years ago, as well as from Early Neolithic individuals from Anatolia and the Levant. By using a comparative dataset of modern and ancient genomes, we estimate that the earliest Anatolian farmers derive over 90 percent of their ancestry from the local Epipaleolithic population, indicating a high degree of genetic continuity throughout the Neolithic transition. In addition, we detect two distinct waves of gene flow during the Neolithic transition: an earlier one related to Iranian/Caucasus ancestry and a later one linked to the Levant. Finally, we observe a genetic link between Epipaleolithic Near-Easterners and post-glacial European hunter-gatherers that suggests a bidirectional genetic exchange between Europe and the Near East predating 15,000 years ago. Our results suggest that the Neolithisation model in Central Anatolia was demographically similar to the one previously observed in the southern Levant and in the southern Caucasus-Iran highlands, further supporting the limited role of demic diffusion during the early spread of agriculture in the Near East, in contrast to the later Neolithisation of Europe.

...

Demographic processes in Estonia from Bronze Age through Iron Age to Medieval times

Metspalu et al.

N3 and R1a are the two most common Y chromosome haplogroups among modern Estonians. R1a appears with Corded Ware culture but the arrival of hg N has not been determined. To this end we have extracted and studied aDNA from teeth of 18 individuals bracketing the changes in the material culture in the end of the Bronze and early Iron Age. We find N3 in Iron Age but not in Bronze Age. Due to the small sample size we cannot refute the existence of hg N in the latter. In genome wide analyses the Bronze Age and especially Iron Age samples appear very similar to modern Estonians implying population continuity. Christianization (13 cc AD) established a new elite of West European origin, which presumably had an impact on the genetic structure of the local population. To investigate this we extracted DNA from teeth of 35 individuals, who have been uncovered from both rural (considered local Estonian population) and town (likely of West European origin) cemeteries of Estonia. We compared the low coverage genomes with each other and with relevant modern and ancient Estonian and other European populations. We find that there is a clear discontinuity between the elite and common people, where the former group genetically with modern German samples and the latter with modern Estonians. We do find three individuals of mixed genetic ancestry. But importantly we do not see a steady shift of either local population strata, which suggests limited contact between the elite and the common people.

...

Genetic transition in the Swiss Late Neolithic and Early Bronze Age

Furtwaengler et al.

Recent studies have shown that the beginning of the Neolithic period as well as final stages of the Neolithic were marked by major genetic turnovers in European populations.The transition from hunter-gatherers to agriculturalists and farmers/farming in the 6 th millennium BP coincided with a human migration from the Near East. In the 3 rd millennium BP a second migration into Central Europe occurred originating from the Pontic steppe linked to the spread of the Corded Ware Complex ranging as far southwest as modern day Switzerland. These genetic processes are well studied for example for the Middle-Elbe-Saale region in Eastern Germany, however, little is known from the regions that connect Central and Southern Europe. Here, we investigate genomic data from 69 individuals from the Swiss Plateau and Southern Germany that span the transition of the Neolithic to the Bronze Age (5500 to 4000 BP). Our results show a similar genetic process as reported for the Middle-Elbe-Saale region suggesting that the migration from the Pontic steppe reached all the way into the Swiss plateau. The high quality of the ancient genomic data also allowed an analysis of core families within multiple burials, the determination and qualification of different ancestry components and the determination of the migration route taken by the ancestors of the Late Neolithic populations in this region. This study presents the first comprehensive genome wide dataset from Holocene individuals from the Swiss plateau and provides the first glimpse into the genetic history of this genetically and linguistically diverse region.

...

Genome-Wide Ancient DNA Portrays the Forming of the Finnish Population Along a 1400-Year Transect

Majander et al.

The Finnish population has long been a subject of interest for the fields of medical and population genetics, due to its isolation-affected genetic structure and the associated unique set of inherited diseases. Recent advances in ancient DNA techniques now enable the in-depth investigation of Finland's demographic past: the impact of migrations, trade and altering livelihood practices. Here we analyse genome-wide data from over 30 individuals, representing ten archaeological burial sites from southern Finland, that span from the 5th to 19th century. We find the historical individuals to differ genetically from Finns today. Comparing them with surrounding ancient and modern populations, we detect a transition from genotypes generally connected with prehistoric hunter-gatherers, and specifically resembling those of the contemporary Saami people, into a more East-Central European composition, associated with the established agricultural lifestyle. Starting from the Iron Age and continuing through the Early Medieval period, this transition dates remarkably late compared to the respective changes in most regions of Europe. Our results suggest a population shift, presumably related to Baltic and Slavic influences, also manifested in the archaeological record of the local artefacts from the late Iron Age. Our observations also agree with the archaeological models of relatively recent and gradual adoption of farming in Finland.

...

Population migration and dairy pastoralism on the Bronze Age Mongolian steppe

Warinner et al.

The steppe belt that extends across Eurasia was the primary corridor of Eneolithic and Bronze Age migrations that reshaped the genetics of Europe and Asia and dispersed the Indo-European language family. Beginning in the Eneolithic, a new and highly mobile pastoralist society formed on the Western Steppe. These steppe herders expanded both westwards, contributing to the Corded Ware culture of Eastern and Central Europe, and eastwards, contributing to the mobile pastoralist Afanasevo, Sintashta, Andronovo, and Okunevo cultures in Central Asia. The eastern extent of this Western Steppe herder expansion is not well defined. Here we investigate genome-wide ancestry data obtained from 20 Late Bronze Age (16th-9th century BCE) khirigsuur burials from Khovsgol, Mongolia and further investigate evidence for dairy pastoralism by LC-MS/MS analysis of dental calculus. Overall, we observe limited Western Steppe gene flow into Late Bronze Age Mongolia, but adoption of Western ruminant dairying by ca. 1500 BCE.

...

The Transition to Farming in Eneolithic (Copper Age) Ukraine was Largely Driven by Population Replacement

Schmidt et al.

The transition to a farming-based economy during the Neolithic happened relatively late in southeastern Europe. Material changes occurred through pottery manufacture, but it wasn't until the sixth millennium BCE that farming was adopted by the Cucuteni-Trypillian archaeological complex (4800-3000 BCE). In many parts of Europe, early farmers who were descended from Anatolian migrants slowly admixed with local hunter-gatherers over the course of the Neolithic. In Eastern Europe and the Balkans, this process may have been more complex since early farmers would likely have admixed with local groups prior to spreading into continental Europe. Studies from the Baltic and Estonia suggest little genetic input from early farmers or continuous admixture with hunter-gatherers. Here, we investigate the impact of Trypillian migrations into Ukraine through the analyses of 19 ancient genomes (0.6 to 2.1X coverage) from the site of Verteba Cave. Ceramic typology and radiocarbon dating of the cave indicate continuous occupation from the Mesolithic to the Medieval Period, with peak occupation coinciding with the middle to late Tripolye. We show that the Trypillians replaced local Ukrainian Neolithic cultures. Also, hunter-gatherers contributed very little ancestry to the Trypillians, who are genetically indistinct from early Neolithic farmers. The one exception is a female that has mostly steppe-related ancestry. Direct radiocarbon dating of this individual places her in the the Middle Bronze Age (3545 years before present). Her lack of farmer ancestry suggests abrupt population replacement resulting perhaps from inter-group hostilities or plague that spread through Europe during the Late Neolithic.

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Ahead of the pack

Genetic borders are usually linguistic borders too

Yamnaya isn't from Iran just like R1a isn't from India