May 09, 2014

Ancient DNA from the Balkans (Iron Age Thrace)

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

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

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

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

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

Martin Sikora et al.

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

Link

31 comments:

Grey said...

The idea of total replacement from the near east (map C) is nonsense.

andrew said...

"There are no good ancient DNA from Cardial Neolithic farmers, so the fact that Sardinians are similar to the Iron Age Bulgarian, the Stuttgart LBK German, and the Swedish TRB farmers may mean that the Mediterranean/Cardial farmers were related to the ones that went into Europe following the inland route from the Balkans."

PCA analysis of Etuscan ancient mtDNA pools show almost perfect identity with LBK individuals, but more significant genetic distance from Sardinians. The easiest way to explain this is to assume that Etruscans are derived from first wave LBK farmers, while Sardinians are derived from first wave CP farmers that while similar to each other vis-a-vis modern populations were still derived from quite genetically distinct populations.

Mark D said...

Why is Sardinia so special for comparisons? Because Cavalli-Sforza chose to test there as part of the Human Genome Diversity Project ten years ago? Only 28 individuals, including 16 men were tested, and as far as I know, there was no verification of genealogical pedigree that's now often done. This from a population of over 1.6 million that has experienced repeated waves of colonization over the centuries from throughout the Mediterranean. Sardinia is no more unique than anywhere else in the region.

Daniel Szelkey said...

If the study didn't study for the y-dna forget about it. Lazy academics cant even say the haplogroup.

truth said...

@ Mark D

268 Sardinians in this study :

http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0043759

mooreisbetter said...

@Andrew: Excellent points. But careful. By pointing out the clear ancientness of Etruscans, you run the risk of running afoul from the "Everything Ancient storytellers said was right" and the "Italians Must All Be Exotic" crowds.

@Mark D: Actually, Sardinians have been shown, both by Cavalli Sforza and others, to be, in layman's terms, so different from other Europeans that they are almost a separate "race." (to use a very poor, unscientific term so you get the gist).

Anima Mortel said...

-"Actually, Sardinians have been shown, both by Cavalli Sforza and others, to be, in layman's terms, so different from other Europeans that they are almost a separate "race." (to use a very poor, unscientific term so you get the gist)"-

No;
It shows that Sardinians are and older (Neolithic) version of Europeans. Akin to Iceman, Gok4 and Stuttgart (Lazaridis et al 2013).

The isolation of Sardinians towards modern day Europeans and any other groups shows that this archaic feature is no longer shared by any modern groups and Sardinians remain closest to Neolithic Europeans.

andrew said...

@moreisbetter:

"By pointing out the clear ancientness of Etruscans, you run the risk of running afoul from the "Everything Ancient storytellers said was right" and the "Italians Must All Be Exotic" crowds."

Etruscans are unquestionably intrusive to Tuscany (in Northern Italy) where the Etuscan label was attached to them, and arrived in Tuscany around the same time as, or a century or two before, Italic peoples migrated to Tuscany. The Etuscans arrived at that time from the general vicinity of the Swiss Alps where they left behind the kindred Raetic people (who do not share of language or ethnicity of the people who now live in the same geographic area where they once did who speak the modern Raetic language which is a Romance language derived from Latin).

Grognard said...

Did they find the y-dna at all? Seems odd it's not included.

Rokus said...

'while Sardinians are derived from first wave CP farmers'

Only on the assumption that Cardial Pottery in their Mediterranean migrations to the west mixed with "European Hunter Gatherers" almost exactly the same way as LBK in Stuttgart. However, evidence of Loschbour- (or Brana 1-) like HG in the Balkan is still missing, and Bulgarian K8 rather correlates with a Russian intrusion - not so strange given the northern cultural links of Thracians.

Hence also the "outrageous" arrow of Figure 4C pointing from central Europe to Sardinia, that doesn't seem to support the traditional assumption on how the Neolithic reached the western Mediterranean bassin by demic diffusion (however, read the nuances made by Cunliffe).

Since both assumptions seem to be wrong and unsupported by the Sardinian evidence, I gather the EEF component should be considered indigenous in southern Europe.

Fanty said...

Hm.
That rings a bell. A million years ago I read that "Futhark" (the Germanic alphabet), commonly better known under the term "runes" (From Germanic "Runa"-->whispering)by their magical use to foretell the future or enchant weaponry by engraving magical spells into them, does greatly resemble Etruscian letters, so that its traditionally thought they are actual adaptions by a contact between Germanic tribes with Etruscans.

The most interesting difference however is the complete different order of the letters in the alphabet plus the concept of writing one line from left to right and the next from right to left.

Any contact of Germanic people with Etruscans would need the Etruscans to have lived further north, right? Also, they should have been the Celts between them?

Simon_W said...

The ADMIXTURE analysis is particularly interesting. Of course I'm very keen to know how West Asian and how North European these Iron Age Bulgarians were. But because of the limited data set of modern populations used, a component that's clearly similar to the Dodecad K7b or Globe13 or MDLP World-22 West Asian component shows up only at K=8, it's the pink component which makes up one half of the Adygei and the entirety of some Druze individuals.

So the early Iron Age Bulgarian is about as West Asian as the modern French are, and just a little more than Ötzi. On the whole he resembles Gok4; he has just a little more of the Palestinian, Druze and Mozabite components, and a little less of the Sardinian component, what's natural, since he was closer to the entry point of the non-European early farmers.

The late Iron Age Bulgarian has only very slight West Asian admixture. And interestingly, up to K=4, he's very similar to the hunter-gatherers, he seems to have only a little bit of early farmer admixture. (At K=5 and more he also has a lot of the Sardinian components etc, but the same holds true for the hunter-gatherers, so that's because of the hunter-gatherer ancestry in Sardinians etc, and not the other way round.)

These results, if valid, are very interesting, also regarding the question of IE origins, and they give me food for thought. Isn't it striking that hunter-gatherer-like individuals were roaming around in the late Iron Age and that this individual belonged to the upper class?

I would also like to point out that these individuals were not necessarily both Thracians. Not all of the ancient tribes known from the Thracian region were definitely Thracians, partly they may even have been non-IE and related with the pre-Greek Lemnians.

Judging from Hellenthal et al. 2014, most of the strong West Asian admixture in modern Tuscans dates from Medieval admixture. The ancient Tuscans, and by inference the Etruscans, seem to have been more similar to the modern French. But if some early Iron Age Bulgarians were Sardinian-like and only as West Asian as the modern French, then the ancient Etruscans may well have had an eastern origin, in the Aegean. The Protovillanovan wave from the north may have been Oscan-Umbrian rather than Etruscan. Again, we must be careful not to conflate the Protovillanovan culture with the later Villanovan culture. And we must not forget, the Etruscan language seems to be closer to Lemnian than to Raetic.

Dr Rob said...

@ Mark D

"Why is Sardinia so special for comparisons? Because Cavalli-Sforza chose to test there as part of the Human Genome Diversity Project ten years ago? Only 28 individuals, including 16 men were tested, and as far as I know, there was no verification of genealogical pedigree that's now often done. This from a population of over 1.6 million that has experienced repeated waves of colonization over the centuries from throughout the Mediterranean. Sardinia is no more unique than anywhere else in the region."

Totally.
Same with Basques, Saami, etc

Dr Rob said...

@Andrew: "Etruscans are unquestionably intrusive to Tuscany (in Northern Italy) where the Etruscan label was attached to them, and arrived in Tuscany around the same time as, or a century or two before, Italic peoples migrated to Tuscany. The Etuscans arrived at that time from the general vicinity of the Swiss Alps where they left behind the kindred Raetic people (who do not share of language or ethnicity of the people who now live in the same geographic area where they once did who speak the modern Raetic language which is a Romance language derived from Latin)."

Andrew, that is all, 100% speculation. No credible scholar, at least recently, would even claim that any theory about the origins of Etruscan would be anything other than an educated guess. The entire flimsy theory rests on Pliny's supposed ethnographic knowledge about the Etruscans, Raeti, etc Of course, his knowledge was actually zilch, nor did he really care about where the Vindeleci or Raeti really came from. He just invented an origin, by linking them to a long extinct people, to explain their existence to people back in Rome.

Whatever the case, the language has nothing directly to do with the observed genetic pattern. There is no reason to assume that the Etruscans are particularly "ancient". The Etruscans, as a people, only really formed in the late 8th - 7th century through a coalescence of local communities. Essentially, they were a new creation.

eurologist said...

I see I need to emphasize again that SE Europe (from much of Italy through the Balkans and all-the-way to the Pontic), between the Gravettian and the early Neolithic (i.e., ~20,000 years) was largely separated from Europe at large, and more closely connected to Anatolia and the Levant geographically and by known seafaring and trade.

Any theory about Cardium and LBK differences (or the lack thereof) needs to take not of this.

Mer said...

We're still in the process of putting the files on SRA - will post link on our lab website (bustamantelab.stanford.edu) when it's up.

Grey said...

@Fanty
"Any contact of Germanic people with Etruscans would need the Etruscans to have lived further north, right? Also, they should have been the Celts between them?"

Pure speculatin' for fun but an alternative explanation might be the two populations used to be adjacent and the Germans moved west to the north of the Carpathians while the Etruscans moved west using the southern maritime route.

.

"Everything Ancient storytellers said was right"

Off-topic but speaking of ancient tales of red-haired, green-eyed white devils:

http://www.dailymail.co.uk/femail/article-1103391/Mixed-race-couple-birth-black-white-twins--second-time.html

Seems to me mixed race couples where the white half has red hair (i.e. two copies of the red hair gene) and the non-white half is carrying a recessive copy of the red hair gene might be able to prove if MC1R represents an earlier depigmentation.

(Maybe IRF4 as well.)

(Although being recessive that might mean the truth is actually odder i.e. 1/4 white, 3/4 brown.)

The thing about ancient story-tellers imo is not that they should be taken at face value but mined for clues.

V Robazza said...

@Dr Rob

The 2013 academic paper on etruscans, state they originate from the Alps/southern germany as part of the rhaetic people.
Although it also states that they came to the alps from Anatolia, but that was 2000 years before hand.

so, Rhaetics created etruscans and not Etruscans created rhaetic

They most likely took the name etruscans from the land they arrived in, Etruria

Ryan said...

It's been suggested that Germanic Runes were derived from Etruscan, so contact between the isn't that crazy.

Simon_W said...

@ Mark D

Over 400 Sardinians in this study:
http://dienekes.blogspot.com/2014/05/ancient-dna-from-balkans-iron-age-thrace.html

@ Anima Mortel

Since modern Europeans are inbetween Mesolithic hunter-gatherers and early farmers (and an eastern, MA1-related element) it's wrong to call the special early farmer affinity of the Sardinians archaic. (The Mesolithic came before the Neolithic.) Basically, what distinguishes modern Europeans from the early farmers are the increased North European and West Asian autosomal components, and both components are still very weak in Sardinians. In contrast, the Mediterranean component which peaks in Sardinia, is still present all over Europe, especially in the South and West.

@ Andrew

Afaik it's not quite certain that Raetic tribes were also present in what is now Switzerland, but it's possible. In any case, the main area where Raetic inscriptions have been found, is focused on Trentino-South Tyrol. From there it extends into immediately adjacent parts of Veneto (Schio, Verona, Feltre), and through northern Tyrol up into southernmost Bavaria. Isolated Raetic finds have been found as far east as Slovenia (Vace, Negova).

@ Dr Rob

You surely mean Livy, not Pliny.
Apart from that I mostly agree, though I do have my substantiated opinions on Etruscan origins - but the fact that I often change them is evidence for the uncertainty in the field. I think at least the Villanovan culture, starting around 900 BC, was probably ethnically Etruscan – the geographical overlap with the later Etruscan area is telling.

Mark D said...

@ Simon W "Basically, what distinguishes modern Europeans from the early farmers are the increased North European and West Asian autosomal components, and both components are still very weak in Sardinians. In contrast, the Mediterranean component which peaks in Sardinia, is still present all over Europe, especially in the South and West."

The paper that "truth" had linked also reported that Sardinians had only a small amount of Northern European. Could Sardinia then be representative of a Mediterranean-wide grouping? I still have trouble reconciling its supposed "ancient DNA" with its history of colonization from around the Mediterranean, including Phoenician, Carthaginian, Roman(with their slaves), Vandals, Byzantines, and Aragonese. The island even has somewhat different languages from north to south, and R1b is reported to be around 20%, similar to what's found in Aragon. There's just too much historic admixture to mark it as "Neolithic"

Simon_W said...


@ V Robazza

Which 2013 paper do you mean?

(Sorry for the double post, I didn't see V Robazza's comment before.)

Fanty said...

"It's been suggested that Germanic Runes were derived from Etruscan, so contact between the isn't that crazy."

Yeah. Question is WHEN.
There is talk about a "PAN Germanic Futhark",wich means, ALL Germanic tribes apear to have use that alphabet. If there wasnt massive contact between the various Germanic tribes, the alphabet should have been introduced quiet early? -.-

I mean, "Germanic" is an umbrella term for up to 7 cultures, wich some archeolists claim that they are that different that TODAY they would not use a single umbrella term on all those people, if it hadnt existed in the first place. (claims is: Much more cultural variation than in what is called "Celts")

If the alphabet is one of the shared stuff (along with variations of the "Norse" religion and most likely the language family, thats a pretty interesting thing.

Uh wait... isnt it sometimes claimed the Etrustcans originate in Anatolia? The Franks used to claim Anchestry in the people of ancient Troy (wich was in Anatolia too) but I never read about: How did they even know someting like Troy existed? Who told them the story that they fell in love with, to claim them for their origins? They are Barbarians from the forrests and swamps of Germania and Troy was long gone?

Dr Rob said...

WRT Etruscan origins. Anything is possible of course, however, I'd be weary of any theory which claims to have 'proven' their origin.

What we do know is that during the Iron Age Italy was a very dynamic society, with significant exchange and even small-scale immigration of groups into it, and within it. And the closest analogies point to the Balkans- so much so that archaeologists speak of an 'Adriatic cultural koinon".

This is not to say that Italic speakers came from the Balkans, as Italic developed (as linguistic currently favour) *within * Italy.

Whatever the case, the linguistic diversity of Iron Age Italy has been underestimated. Even in Etrusca - Etruscan was the language of the elites, often used for religious ceremonies. I am sure that many other languages were spoken.

Ezr said...

Those speculating about contacts between Etruscans and Germanic peoples should remember two things:

1) The consensus is that the Etruscan alphabet (from which the Latin alphabet emerged) is derived from Greek alphabet variants used in southern Italy. It was adopted by the Etruscans themselves at a relatively late time and certainly after they were firmly established in central Italy. Besides, there were other northern Italian alphabets derived from similar sources that could have influenced Germanic runes.

2) There is solid archeological record of ancient trade routes such as the Amber road going all the way from Scandinavia and the Baltic to the Mediterranean, leading to all sorts of cultural, technological and presumably genetic exchanges. This alone can easily explain those slight similarities without having to posit speculative theories of ethnogenesis.

Ben Overboord said...

"Uh wait... isnt it sometimes claimed the Etrustcans originate in Anatolia? The Franks used to claim Anchestry in the people of ancient Troy (wich was in Anatolia too) but I never read about: How did they even know someting like Troy existed? Who told them the story that they fell in love with, to claim them for their origins? They are Barbarians from the forrests and swamps of Germania and Troy was long gone? "

First, the Franks were quite intergrated in Roman culture. Romans had Frankish generals as early as the 3rd century. Secondly, the Troy claim does not come from the Franks directly but comes from a genealogy that Gregory of Tours provides.

I read somewhere the theory that he may have heard a story about a kingdom on the eastern part of the Rhine with Xanten as central city. That was called Colonia Ulpia Traiana in Roman times.

Simon_W said...

@ Mark D

Judging from Hellenthal et al. 2014, Sardinians are a mix of a French-like population (but at the same time akin to Basques, Welsh and Mozabites) that admixed around 634 AD with 24% of Egyptian-like people, reminiscent of Eastern Mediterranean and North African populations. So maybe their similarity with Neolithic Europeans is rather young?

Simon_W said...

Ups, I've overlooked this, @ Mark D: R1b only 20% in Aragon? I think nowhere on the Iberian peninsula is R1b this rare. According to Eupedia, Aragon has 60.5% R1b.

agiering said...

@ Mark D and Simon_W

According to the ADMIXTURE graph from the Lazaridis et al paper on Mal'ta, Sardinians appear quite similar to the Stuttgart neolithic farmer sample. Compared to most other European groups both populations seemed to be overwhelmingly of a particular Mediterranean component, a fairly small hunter-gatherer component, and none of the South Asian component. It does seem like the most likely explanation is that Sardinians are descended from the neolithic farmers (like Stuttgart) and received relatively little admixture since, perhaps because of its isolation as an island.

Arch Hades said...

So what was the genetic analysis of the Thracians? What modern populations did they show strongest ties to?

Simon_W said...

Interesting, Genetiker has run the DNA files of the two Thracians and of two additional ones through various calculators from Dodecad and the MDLP:

http://genetiker.wordpress.com/2014/06/11/analyses-of-iron-and-bronze-age-bulgarian-genomes/

It seems like I was correct with my assessment of individual P192-1 here, but quite wrong with K8. The latter had a very strong West Asian component.