July 14, 2012

Population strata in the West Siberian plain (Baraba forest steppe)

Also from the Population Dynamics in Prehistory and Early History (2012) volume, this is an awesome ancient DNA study which dissects a succession of archaeological cultures stretching from the beginning of the metal ages to the beginning of the Iron Age in a small region of West Siberia. As the authors write:
Our work is devoted to the analysis of human migration processes that occurred during the Bronze Age (4th–early 1st millennium BC) in the forest steppe zone between the Ob and Irtysh rivers (about 800 km from west to east). This area, known as Baraba forest steppe, stretches over 200 km from the taiga zone in the north to the steppes in the south.
The careful examination of the sequence of cultures, combining ancient mtDNA and physical anthropology paints a very compelling picture of the changes that occurred in the span of a few millennia in the Baraba forest steppe. The authors give the map on the left, with the caption: "Fig. 5 | Location of ancient human groups with a high frequency of mtDNA haplogroups U5, U4 and U2e lineages. The area of Northern Eurasian anthropological formation is marked by yellow region on the map (References: 1 Bramanti et al., 2009; 2Malmstrom et al., 2009; 3 Krause et al., 2010; 4 this study)".


The northern Eurasian anthropological formation actually combines eastern and western Eurasian features and may correspond to the Proto-Uralic type. Researchers have clashed about the origins of this population element, with some considering it a third Eurasian race that evolved independently of Caucasoids and Mongoloids, others assigning it to a much diverged branch of one of the two major Eurasian races, and still others considering it the product of admixture between east and west.


All indications are that the type, unlike the Caucasoid-Mongoloid mixtures that took place in Central Asia in the last 2 millennia, is of more ancient vintage, and represents an anthropological element that was indeed of Caucasoid-Mongoloid origins, but in the rather remote past. The authors write with respect to the most ancient periods:

In contrast to the occupation of the southern region of West Siberia, modern humans arrived in the Ob-Irtysh interfluve relatively late, at the end of the Pleistocene, about 13–14 thousand years ago (Okladnikov, Molodin, 1983; Petrin, 1986). The absence of burials dating back to this period in the region does not allow us to conduct a biological investigation of this earliest population. The most ancient anthropological material available is from the Neolithic period (4th–5th millennium BC). 
And, what of the earliest available material?
The anthropological analysis of the material allowed us to detect a specific craniological type in the Baraba population, which was assigned to one of the anthropological formations discovered by V.V. Bunak in 1956 through the analysis of Neolithic materials from the northern forest zone of the East European Plain. Bunak called it the “northern Eurasian anthropological formation” (Bunak, 1956).

This anthropological type developed in a zone that is intermediate to the geographic areas occupied by the classic Caucasoids and the Mongoloids. The exists substantial anthropological evidence showing a wide geographic distribution of this anthropological formation: from the Trans-Urals forest and the Barabian province of Western Siberia in the east to Karelia and the Baltic in the west (Chikisheva, 2010).
The mtDNA evidence seems to support the anthropological assessment:
We have analyzed 18 mtDNA samples from the Ust-Tartas population to date (Fig. 3). The results obtained thus far allow us to draw several preliminary conclusions about the genetic background in the region in the beginning of the Bronze Age. By the Early Metal Period the mtDNA pool structure was already mixed and consisted of both Western and Eastern Eurasian haplogroups in nearly equal proportions. The eastern Eurasian mtDNA cluster was represented by Haplogroups A, C, Z, D, which are most typical of modern and perhaps ancient populations located in the east of the region studied. Haplogroups C and D were predominantly represented by widely distributed root haplotypes. A lineage of Haplogroup A that was detected in two Ust-Tartas samples represents a subcluster that is apparently characteristic of West Siberia and the Volga-Ural Region. The observed presence of Haplogroup Z lineages with a high frequency in the Ust-Tartas group was unexpected, since these lineages are nearly absent in the gene pool of modern indigenous West Siberian populations.

It is worth noting that the Western Eurasian mtDNA haplogroups in the Ust-Tartas series were represented only by Haplogroup U lineages, and specifically by the three subgroups – U2e, U4, U5a1. These results are in agreement with previous data indicating that Haplogroup U lineages (particularly Subgroups U5 and U4) predominated in Eastern, Central and Northern European hunter-gatherer groups from 14000 to 4000 years ago (Bramanti et al., 2009; Malmstrom et al., 2009), and possibly in earlier periods (Krause et al., 2010). The geographic area within which this genetic feature is observed appears to be broad (Fig. 5). Apparently, Baraba was near the eastern periphery of this area.
We now have evidence of the zone of U dominance extending from Iberia in the west and all the way to Lake Baikal in the east. But, this zone is not homogeneous: its western, European, end appears to have lacked the East Eurasian lineages, while starting from Ukraine and to the East the U types were supplemented by the Mongoloid lineages.

But, there was structure within the U zone itself: according to Lillie et al. (same volume) in Ukraine during the 6th millennium BC, the West Eurasian types were represented by U1 and U3, a different mix than in the Baraba forest steppe, and haplogroup T was also present, while of the Mongoloid haplogroups only C was present.

As we head into the Bronze Age, the population of the region displayed signs of continuity:

The genetic analysis of the Odinovo and Krotovo groups (10 and 6 samples, respectively) (Fig. 3) did not reveal any differences between them and the previous Ust-Tartas group, such as the presence of new mtDNA haplogroups. The mtDNA pool structure was still mixed. The East Eurasian haplogroups were represented by the D, C, Z (in both the Odinovo and Krotovo groups) and A (in the Krotovo group) haplogroups. The East Eurasian lineages identified were phylogenetically close (lineages of haplogroups A, C, Z) or even identical (D haplogroup, 16223–16362 lineages) to the samples from the Ust-Tartas group. The West Eurasian part of the samples were represented by the U5a1 (Odinovo group) and U2e (Krotovo group) haplogroup lineages.  
Although only a small series of samples have been investigated thus far, the data obtained reveal continuity between the Odinovo and Krotovo populations and the earlier Ust-Tartas group. These findings are consistent with the autochthonous development of the Baraba populations during the Early and the beginning of the Middle Bronze Age, as well as with the anthropological evidence.  
It is during the Middle and Late Bronze ages that we begin to say the first intrusive lineage into the native population mix:
The anthropological analysis of the West Siberian Andronovo population shows at least four craniological types. Three types are related to the Palaeocaucasian race and are represented by proto-European anthropological type variants. The fourth, Mongoloid, component is autochthonous. The most intensive interactions between the Andronovo migrants and the indigenous populations apparently occurred in the Baraba forest steppe and the right bank of the upper Ob River (Chikisheva and Pozdnyakov, 2003).  
To investigate the putative impact of Andronovo migrants on the mtDNA pool structure of the indigenous populations in Baraba, mtDNA samples from the Late Krotovo (n=20) and Andronovo (n=20) groups in this region were analyzed (Fig. 3) and compared to recently published data (n=10) (Keyser et al., 2009) and our own unpublished data (n=6) on mtDNA lineages from West Siberian Andronovo populations located outside the Baraba forest steppe.  
The genetic influence of migrants can be detected by the appearance of a new mtDNA haplogroup that was absent in the populations preceding the migration wave. This new mtDNA haplogroup, a West Eurasian T haplogroup, was detected in the Late Krotovo population. The T haplogroup appears simultaneously (with a 15 % frequency) in the Krotovo and Andronovo groups, but was completely absent in all preceding Baraba populations. We therefore consider the appearance of the Haplogroup T-lineage as the most likely genetic marker of the Andronovo migration wave to the region.  
This assumption is confirmed by mtDNA studies of Andronovo groups from other West Siberian areas. Haplogroup T lineages were found, with a frequency of 25 %, in the samples (n=16) taken from two Andronovo groups from the Krasnoyarsk and upper Ob River areas.  
We also detected another remarkable feature in the mtDNA pool of the Andronovo group from Baraba. Most mtDNA samples belonged to haplogroups, such as the East Eurasian A and C haplogroups, that are typical of preceding Baraba indigenous populations. Still, these haplogroups were not found in the other West Siberian Andronovo groups. Apparently, the Andronovo group from Baraba assimilated the aboriginal Krotovo population, from which it obtained these East-Eurasian mtDNA haplogroups. Obviously, there was reciprocal genetic contact between the migrant and indigenous groups in the region. 

...

A small but informative series of mtDNA samples from the Baraba Late Bronze Age culture population (n=5) was analyzed (Fig. 3), revealing the presence of MtDNA lineages (East Eurasian A and C lineages) that mark the genetic continuity with aboriginal Baraba groups. At the same time, the series includes the Haplogroup-T lineage, which we believe marks the Andronovo migration wave to West Siberia. Our data is therefore consistent with the putative origin of the West Siberian Late Bronze Culture population as the result of interaction between the Baraba indigenous genetic substrate and the newly arrived group.
It is now clear that the Andronovo groups moving into the area possessed mtDNA haplogroup T and assimilated the locals with their U+East Eurasian mix. It is of course interesting that haplogroup T is the only non-U lineage found in the aforementioned study of Mariupol-type cemeteries from Neolithic Ukraine.

The earliest occurrence of haplogroup T is in the Pre-Pottery Neolithic B of the Near East (Tell Hallula), and this haplogroup appears all over the place in Neolithic Europe. While a recent article has suggested a pre-Neolithic dispersal of T subclades into Europe, on the basis of modern populations, this hypothesis is difficult to reconcile with the ancient DNA data.


Pending new discoveries, it appears likely that mtDNA haplogroup T represents a Neolithic entrant into the boreal zone of U dominance. This has, of course, substantial implications in the context of J.P. Mallory's concept of fault lines, as it demonstrates that the steppe populations did not evolve in isolation, but the dominant lineage in the Andronovo groups was a late entrant into the indigenous U-zone of the eastern European plain.


But, the story doesn't end here:

The analysis of mtDNA samples from the Chicha-1 population revealed some interesting patterns. Crucial changes in the composition of mtDNA haplogroups in the gene pool were observed as compared to the earlier Baraba groups studied (Fig. 3). Dominance of Western Eurasian haplogroups and the near absence of East Eurasian were observed. Additionally, several new West Eurasian haplogroups appeared in the region, including Haplogroups U1a, U3, U5b, K, H, J and W.  
The phylogeographic analysis suggests that the distribution and diversification centres of several of these mtDNA haplogroups and specific lineages are located on the west and south west of the Baraba forest steppe region, on the territory corresponding to modern-day Kazakhstan and Western Central Asia (Fig. 10). Apparently, the migration wave from the south strongly influenced the gene pool of the Baraba population in the transitional period from the Bronze to the Early Iron Age. The impact of the northern human groups was probably less evident in the south of the Baraba forest steppe, at least at the mtDNA level. 
The drastic appearance of a purely Caucasoid population at the Iron Age from a southern, east-Caspian origin perhaps corresponds to the arrival of the first steppe Iranians. The vector of proposed migration is reasonable, if we consider both the likely Indo-Iranian homeland east of the Caspian, as well as the literary evidence for Scythian mobility during this period.

All in all, this is commendable research which allows us to intuit a sequence of events:
  • An early mixture zone between Caucasoids and Mongoloids
  • The Bronze Age arrival of mtDNA-T bearing Andronovo groups, the first pastoralists entering the zone of U+East Eurasian boreal hunter-gatherers; these Caucasoid peoples admixed with the natives of the mixture zone.
  • The early Iron Age arrival of a full-blown set of Caucasoid mtDNA lineages from the south paving the way for the Iranian Scytho-Sarmatian period

Human migrations in the southern region of the West Siberian Plain during the Bronze Age: Archaeological, palaeogenetic and anthropological data


Molodin, Vyacheslav I. et al.


In this paper we present archaeological and anthropological data on human migrations in the Western Siberian foreststeppe region during the Bronze Age (4th–beginning of 1st millennium BC). These data, accumulated over forty years of intensive research in the region, are compared to new results showing the diversity of mitochondrial DNA (mtDNA) lineages in this region during that period (92 mtDNA samples from seven ancient human groups). Preliminary analyses have demonstrated the usefulness of ancient DNA in tracing and unravelling patterns of past human migrations.  


Link


Prehistoric populations of Ukraine: Migration at the later Mesolithic to Neolithic transition


Lillie, Malcolm C. et al.


This paper focuses on the identification of population movements during the Mesolithic and Neolithic periods in the Dnieper Basin region of Ukraine. We assess the evidence for migration from the perspective of individual life histories using a combination of palaeoanthropology/pathology, radiocarbon dating, stable isotopic studies of diet, and mtDNA. 


Link

11 comments:

Aaron said...

All of the new information about ancient DNA is very interesting, especially to help explain these great migrations and invading groups.

I must admit that I understand much more about the science of the DNA than I do about the history of people, so I am playing catch up here on human history.

The introduction of the Southern or Geodressian component of DNA within the last 4-5k years really fascinates me. It seems that the spread of this genetic component across Europe occurred quickly, and that it impacted nearly all groups except for some groups like Sardinia, Wales and Finland.

Could someone help me out with my theory on how this could happen? Basically I want you to punch holes in my theory because I don't know enough about human history, and I just want some more information, I have no agenda for this theory and whether I am right or wrong does not matter.

My Theory:

Southern Component - Northern Tribes of Israel

Geodressian - Assyrian/Babylonian DNA

The Assyrians deported the Northern tribes between 800-700 BC and they mixed with them so that the northern tribes would loose their identity. These people would be similar to the Samaritans today.

Around the same time as the expulsion of the ten tribes, there is a large increase in the population of Europe and the steppes, new ethnic groups appear on the archeological record with unknown origins such as the Cimmarians and the Scythians.

The Cimmarians disappear around 500 BC which corresponds to the same time as the appearance of the Celtic people.

These Indo-European speaking people bring new languages throughout Europe and are fairly barbaric, and it sounds like they may have admixed quite a lot. These are the only invading groups that I am aware of that had such a widespread effect across Eurasia that could explain the shift in genes that occurred during the last 4-5k years.

Is this a plausible theory? I would love to find out more.

Ponto said...

Interesting how Europe's borders seem to be extended further east into Asia with every report on European genetics. West Siberia is Asia and appears to be the original home of the oldest mtDNA haplogroups belonging to U.

This reminds me of the study showing the geographical placement of modern populations compared with their genetic placement. All the Europeans and most of the South Asians ended up genetically in the area north of the Caspian Sea heading towards the Arctic. The only exception was the Mozabites that ended up in the north Levant.

Maybe it is about time to drop the European origins of most of the ancestors of modern Europeans and the separation of "European" and "Asian" haplogroups as they all seem to originate in the zone between the two parts of Eurasia. Kostenki Man belonged to mtDNA U2.

wagg said...

"It is of course interesting that haplogroup T is the only non-U lineage found in the aforementioned study of Mariupol-type cemeteries from Neolithic Ukraine"

I think this is a mistake.

http://dienekes.blogspot.fr/2012/06/ancient-mtdna-from-neolithic-ukraine.html

"Although the majority of identified mtDNA haplogroups belonged to the traditional West Eurasian lineages of H and U (...)"

Dienekes said...

"It is of course interesting that haplogroup T is the only non-U lineage found in the aforementioned study of Mariupol-type cemeteries from Neolithic Ukraine"

I think this is a mistake.


You got the wrong paper, the link to Lillie et al. is in the text. They are not the same sample, Nikitin et al. has samples that are about 500 years older and lack T.

This actually seems like a very decent confirmation about T being a late entrant in Ukraine.

terryt said...

"Is this a plausible theory? I would love to find out more"

The timing is wrong for some of it:

"Southern Component - Northern Tribes of Israel"

The Southern Component is several thousand years older than the dispersal of the Northern Tribes. 'The Assyrians deported the Northern tribes between 800-700 BC.' At other posts Dienekes has shown that Oetzi, the alpine mummy, belongs to that component, and it is some 4000 BC.

http://dienekes.blogspot.co.nz/2012/03/another-look-at-oetzi-with-euro7-and.html

"Around the same time as the expulsion of the ten tribes, there is a large increase in the population of Europe and the steppes, new ethnic groups appear on the archeological record with unknown origins such as the Cimmarians and the Scythians".

I think what you're seeing here is the sudden expansion of writing rather than a sudden expansion of new people. These sorts of expansions and contractions have probably been continuous throughout our history.

pconroy said...

Dienekes,

Yes, your observation on mtDNA T is exactly what I previously surmised based on the Pala paper. But not just T, T1a and more especially T1a1a1 - I think this lineage may have been one of those carried by the Indo-European culture bearers.

mtDNA T1a was previously found in the Tarim Mummies, so the Andronovo T is probably T1a also.

From the Pala paper:


70% of the samples in T1a1 fall within T1a1a1 (Table S2).
The geographic distribution of T1 is extraordinary—lineages
are distributed, albeit at varying frequencies, across
its range throughout the tree, from northwestern Africa
throughout Europe, the Caucasus, and the Near East,
into western India, and across central Asia into Siberia.
The South Asian lineages tend to cluster with or match
Near Eastern ones in the HVS-I network, but common
HVS-I types frequently match across an extremely wide
range. Indeed, the root type of T1a1a1, dating to ~7 ka
ago, is very unusual among whole-genome mtDNA types
in that it is shared between multiple geographically
distant individuals from Scandinavia, the Baltic, the North
Caucasus, Anatolia, and Morocco. The distribution of
T1a is both widespread and patchy, although at low
frequencies overall, the values rise to ~5% in the South
Caucasus, ~6% in northeastern Iran, ~8% in Tunisia, and
almost 9% in Romania (Table S3). Curiously, despite the
age of T1a1a1, it has not been seen in any Neolithic
remains to date.

valeryz2001 said...

>I think this lineage may have been one of those carried by the Indo-European culture bearers

despite of its rarity in modern Europe?

pconroy said...

@Valery,

The Pala paper doesn't mention that it T1a1 is found at something about 7-8% in Pathans and Baluchis from Pakistan.

So it is found at highest frequencies in an area stretching from Pakistan to Northern Iran to the Caucasus and on to Romania. if the Indo-European incursions were primarily mediated by male invading groups, then we would expect lower percentages in more peripheral areas.

BTW, I'm not suggesting that it is the ONLY mtDNA carried by invading Indo-Europeans - just that it has a very broad range, a very young coalescence time (7,000 yo) and is found wherever there were Indo-European invaders.

Aaron said...

@terryt

Sorry, I misspoke when I said Southern component was lacking in Europeans 4-5k years ago.

What I meant was the Southwest Asian component. So in k7b, only the Southwest Asian component is missing, and in k12b, both the Geodressian and Southwest Asian components are missing. My theory then is that the Geodressian component is Assyrian/Persian, and the Southwest Asian component is the Northern Tribes.

This Southwest Asian component is very high in groups that were known to have left around the time of the Assyrian deportation of the Northern Tribes, like for example the Yemenese Jews, who have remained fairly isolated since their migration south.

You are absolutely right though that our history is based more off of what was written than what actually happened. I am impressed by the level of sophistication of the Cimmarians and the Scythians. They obviously did not arrive ex nihlo, but the question is did they originate from an advanced deported civilization or were they already advanced, but there was just no written record.

andrew said...

* "An early mixture zone between Caucasoids and Mongoloids" so proto-Uralic hunter-gatherers.

* "The Bronze Age arrival of mtDNA-T bearing Andronovo groups, the first pastoralists entering the zone of U+East Eurasian boreal hunter-gatherers; these Caucasoid peoples admixed with the natives of the mixture zone."

Andronovo is conventionally viewed as Indo-European, as are (with strong cultural and late Tarim basin written document support) apparently mtDNA T rich Tarim Basin peoples. The notion of these being pastoralists is also notable - it suggests the possibility of a two track Neolithic with SW Asian pastoralists (mtDNA T) and SW Asian farmers (mtDNA G, I) who were culturally and perhaps linguistically distinct continuing their symbiosis into a mutually interacting migration into hunter-gatherer Europe. Perhaps a farmer-herder population structure made it possible for the pastoralists who assimilated the Uralic populations from having a full range of mtDNA hgs. It also suggests that herder with mtDNA T may have been a highland population in SW Asia and that other non-U West Eurasian mtDNA hgs may have been lowland valley and plain populations in SW Asia.

Being older, one would expect the Androvo to be more similar to the PIE genetics than later populations.

* "The early Iron Age arrival of a full-blown set of Caucasoid mtDNA lineages from the south paving the way for the Iranian Scytho-Sarmatian period"

The implication would seem to be that Bronze Age proto-Indo-Europeans were heavily mtDNA T, but that they assimilating varying populations (Uralic, pre-IE Iranian) in different places and that populations such as the Iron Age Sytho-Sarmatians (presumably themselves the product of Bronze Age PIE population expansions into Iran) may have been, as a result, very different in population genetics from proto-Indo-Europeans. Thus, the original associations of H and J and K and W might be pre-Indo-European lowland farmer Neolithic populations who were militarized and repastoralized by PIE peoples out of Bronze Age collapse chaos.

I think it is fair to question whether the mtDNA T population had an Indo-European identity at first, however. If, for example, PIE is a creole/fusion of Uralic and an early Neolithic mtDNA T rich population that spoke some SW Asian language that is now lost, and that the fusion of two neighboring cultures gave rise to PIE ethnogenesis somewhere in the steppe, then very soon after and directly, the Androvo population, rather having PIE imposed on a Uralic substrate. Androvo may be PIE or something damn close, itself, giving the timing. While Mallory doesn't use the term "creole" the notion that PIE emerged in a cultural boundary zone quite possible in an ethnically mixed population is there.

Thom said...

Hungary Szabadkigyos-Palliget [anc28] Rich grave goods M c. 950 AD N1c [= old N3] Tat C Csányi 2008
Hungary Szabadkigyos-Palliget 7.tomb [anc4] Rich grave goods M c. 950 AD U4

In this village in Hungary were found both Tat C and U4 genetic marker. It is dated 950 AD and Hungarian tribes centrum were in this time in Bekes country(Szabadkigyos-Palliget). If these Hungarian tribes arrived in 895 AD I suppose that it's an Siberian Tat c and U4.