Mitochondrial genomes from all three individuals belong to the U5a2d haplogroup. (…) The mitochondrial U5a2d haplogroup is consistent with earlier published results for ancient individuals from Scandinavia, U5a being the most common within SHG. Of the 16 Mesolithic individuals from Scandinavia published prior to our study, seven belong to the U5a haplogroup, nine share the U2 and U4 haplogroups
We divided the SHG group into two groups: SHGa and SHGb (ancient individuals found in contemporary Norway and Sweden, respectively). We based this on both the geographical distribution and the previous studies demonstrating the close relation of SHGa to EHG group and SHGb to WHG group. To further explore the demography within the SHG group, we compared the ancestry of BLE individuals within SHGa and SHGb groups. This comparison revealed a high relative shared drift between BLE individuals and the SHGb group
The results from Huseby Kiev allow us to finally connect the SHG group with the eastern pressure blade technology. However, the higher genetic affinity between Huseby Kiev individuals and the WHG group challenges the earlier suggested tie between eastern technology and EHG genetics. Our results suggest either early cultural transmission, or a more complex course of events involving both non- and co-dependent cultural and genetic admixture.
Seeing how culture is indeed usually associated with the expansion of a certain population, especially at such an early date, I guess this similarity with WHG of incoming eastern peoples comes from an originally EHG population expanding into a mainly WHG area in the west (similar to what happens e.g. with Bell Beakers), or being replaced later by a WHG population which adopted the culture (similar to what happened with late Corded Ware populations in central-east Europe after the expansion of Bell Beakers).
Unlike later periods, it will always be difficult to judge such ancient population movements with few samples covering thousands of years… Probably specific Y-DNA haplogroups would help differentiate between both expanding populations from east and west.
An interesting aspect of the paper, hidden among so many relevant details, is a clearer picture of how the so-called Yamnaya or steppe ancestry evolved from Samara hunter-gatherers to Yamna nomadic pastoralists, and how this ancestry appeared among Proto-Corded Ware populations.
Please note: arrows of “ancestry movement” in the following PCAs do not necessarily represent physical population movements, or even ethnolinguistic change. To avoid misinterpretations, I have depicted arrows with Y-DNA haplogroup migrations to represent the most likely true ethnolinguistic movements. Admixture graphics shown are from Wang et al. (2018), and also (the K12) from Mathieson et al. (2018).
1. Samara to Early Khvalynsk
The so-called steppe ancestry was born during the Khvalynsk expansion through the steppes, probably through exogamy of expanding elite clans (eventually all R1b-M269 lineages) originally of Samara_HG ancestry. The nearest group to the ANE-like ghost population with which Samara hunter-gatherers admixed is represented by the Steppe_Eneolithic / Steppe_Maykop cluster (from the Northern Caucasus Piedmont).
Steppe_Eneolithic samples, of R1b1 lineages, are probably expanded Khvalynsk peoples, showing thus a proximate ancestry of an Early Eneolithic ghost population of the Northern Caucasus. Steppe_Maykop samples represent a later replacement of this Steppe_Eneolithic population – and/or a similar population with further contribution of ANE-like ancestry – in the area some 1,000 years later.
This is what Steppe_Maykop looks like, different from Steppe_Eneolithic:
NOTE. This admixture shows how different Steppe_Maykop is from Steppe_Eneolithic, but in the different supervised ADMIXTURE graphics below Maykop_Eneolithic is roughly equivalent to Eneolithic_Steppe (see orange arrow in ADMIXTURE graphic above). This is useful for a simplified analysis, but actual differences between Khvalynsk, Sredni Stog, Afanasevo, Yamna and Corded Ware are probably underestimated in the analyses below, and will become clearer in the future when more ancestral hunter-gatherer populations are added to the analysis.
2. Early Khvalynsk expansion
We have direct data of Khvalynsk-Novodanilovka-like populations thanks to Khvalynsk and Steppe_Eneolithic samples (although I’ve used the latter above to represent the ghost Caucasus population with which Samara_HG admixed).
We also have indirect data. First, there is the PCA with outliers:
Second, we have data from north Pontic Ukraine_Eneolithic samples (see next section).
Third, there is the continuity of late Repin / Afanasevo with Steppe_Eneolithic (see below).
3. Proto-Corded Ware expansion
It is unclear if R1a-M459 subclades were continuously in the steppe and resurged after the Khvalynsk expansion, or (the most likely option) they came from the forested region of the Upper Dnieper area, possibly from previous expansions there with hunter-gatherer pottery.
Supporting the latter is the millennia-long continuity of R1b-V88 and I2a2 subclades in the north Pontic Mesolithic, Neolithic, and Early Eneolithic Sredni Stog culture, until ca. 4500 BC (and even later, during the second half).
Only at the end of the Early Eneolithic with the disappearance of Novodanilovka (and beginning of the steppe ‘hiatus’ of Rassamakin) is R1a to be found in Ukraine again (after disappearing from the record some 2,000 years earlier), related to complex population movements in the north Pontic area.
NOTE. In the PCA, a tentative position of Novodanilovka closer to Anatolia_Neolithic / Dzudzuana ancestry is selected, based on the apparent cline formed by Ukraine_Eneolithic samples, and on the position and ancestry of Sredni Stog, Yamna, and Corded Ware later. A good alternative would be to place Novodanilovka still closer to the Balkan outliers (i.e. Suvorovo), and a source closer to EHG as the ancestry driven by the migration of R1a-M417.
The first sample with steppe ancestry appears only after 4250 BC in the forest-steppe, centuries after the samples with steppe ancestry from the Northern Caucasus and the Balkans, which points to exogamy of expanding R1a-M417 lineages with the remnants of the Novodanilovka population.
4. Repin / Early Yamna expansion
We don’t have direct data on early Repin settlers. But we do have a very close representative: Afanasevo, a population we know comes directly from the Repin/late Khvalynsk expansion ca. 3500/3300 BC (just before the emergence of Early Yamna), and which shows fully Steppe_Eneolithic-like ancestry.
Compared to this eastern Repin expansion that gave Afanasevo, the late Repin expansion to the west ca. 3300 BC that gave rise to the Yamna culture was one of colonization, evidenced by the admixture with north Pontic (Sredni Stog-like) populations, no doubt through exogamy:
This admixture is also found (in lesser proportion) in east Yamna groups, which supports the high mobility and exogamy practices among western and eastern Yamna clans, not only with locals:
We don’t have a comparison with Ukraine_Eneolithic or Corded Ware samples in Wang et al. (2018), but we do have proximate sources for Abashevo, when compared to the Poltavka population (with which it admixed in the Volga-Ural steppes): Sintashta, Potapovka, Srubna (with further Abashevo contribution), and Andronovo:
The two CWC outliers from the Baltic show what I thought was an admixture with Yamna. However, given the previous mixture of Eneolithic_Steppe in north Pontic steppe-forest populations, this elevated “steppe ancestry” found in Baltic_LN (similar to west Yamna) seems rather an admixture of Baltic sub-Neolithic peoples with a north Pontic Eneolithic_Steppe-like population. Late Repin settlers also admixed with a similar population during its colonization of the north Pontic area, hence the Baltic_LN – west Yamna similarities.
NOTE. A direct admixture with west Yamna populations through exogamy by the ancestors of this Baltic population cannot be ruled out yet (without direct access to more samples), though, because of the contacts of Corded Ware with west Yamna settlers in the forest-steppe regions.
A similar case is found in the Yamna outlier from Mednikarovo south of the Danube. It would be absurd to think that Yamna from the Balkans comes from Corded Ware (or vice versa), just because the former is closer in the PCA to the latter than other Yamna samples. The same error is also found e.g. in the Corded Ware → Bell Beaker theory, because of their proximity in the PCA and their shared “steppe ancestry”. All those theories have been proven already wrong.
NOTE. A similar fallacy is found in potential Sintashta→Mycenaean connections, where we should distinguish statistically that result from an East/West Yamna + Balkans_BA admixture. In fact, genetic links of Mycenaeans with west Yamna settlers prove this (there are some related analyses in Anthrogenica, but the site is down at this moment). To try to relate these two populations (separated more than 1,000 years before Sintashta) is like comparing ancient populations to modern ones, without the intermediate samples to trace the real anthropological trail of what is found…Pure numbers and wishful thinking.
It has been known for a long time that the Caucasus must have hosted many (at least partially) isolated populations, probably helped by geographical boundaries, setting it apart from open Eurasian areas.
David Reich writes in his book the following about India:
The genetic data told a clear story. Around a third of Indian groups experienced population bottlenecks as strong or stronger than the ones that occurred among Finns or Ashkenazi Jews. We later confirmed this finding in an even larger dataset that we collected working with Thangaraj: genetic data from more than 250 jati groups spread throughout India (…)
Rather than an invention of colonialism as Dirks suggested, long-term endogamy as embodied in India today in the institution of caste has been overwhelmingly important for millennia. (…)
The Han Chinese are truly a large population. They have been mixing freely for thousands of years. In contrast, there are few if any Indian groups that are demographically very large, and the degree of genetic differentiation among Indian jati groups living side by side in the same village is typically two to three times higher than the genetic differentiation between northern and southern Europeans. The truth is that India is composed of a large number of small populations.
There is little doubt now, based on findings spanning thousands of years, that the Mesolithic and Neolithic Caucasus hosted various very small populations, even if the ancestral components may be reduced to the few known to date (such as ANE, EHG, AME*, ENA, CHG, and other “deep” ancestral components).
NOTE. I will call the ancestral component of Dzudzuana/Anatolian hunter-gatherers Ancient Middle Easterner (AME), to give a clear idea of its likely extension during the Late Upper Palaeolithic, and to avoid using the more simplistic Dzudzuana, unless it is useful to mention these specific local samples.
Genetic labs have a strong fixation with ancestry. I guess the use of complex statistical methods gives professionals and laymen alike the feeling of dealing with “Science”, as opposed to academic fields where you have to interpret data. I think language reveals a lot about the way people think, and the fact that ancestral components are called ‘lineages’ – while not wrong per se – is a clear symptom of the lack of interest in the true lineages: Y-DNA haplogroups.
It has become quite clear that male-biased migrations are often the ones which can be confidently followed for actual population movements and ethnolinguistic identification, at least until the Iron Age. The frequently used Palaeolithic clusters offer a clear example of why ancestry does not represent what some people believe: They merely give a basic idea of sizeable population replacements by distant peoples.
Both concepts are important: sizeable and distant peoples. For example, during the Upper Palaeolithic in Europe there was a sizeable population replacement of the Aurignacian Goyet cluster by the Gravettian Vestonice cluster (probably from populations of far eastern Russia) coupled with the arrival of haplogroup I, although during the thousands of years that this material culture lasted, the previously expanded C1a2 lineages did not disappear, and there were probably different resurgence and admixture events.
Haplogroup I certainly expanded with the Gravettian culture to Iberia, where the Goyet ancestry did not change much – probably because of male-driven migrations -, to the extent that during the Magdalenian expansions haplogroup I expanded with an ancestry closer to Goyet, in what is called a ‘resurge’ of the Goyet cluster – even though there is a clear replacement of male lines.
The Villabruna (WHG) cluster is another good example. It probably spread with haplogroup R1b-L754, which – based on the extra ‘East Asian’ affinity of some samples and on modern samples from the Middle East – came probably from the east through a southern route, and not too long before the expansion of WHG likely from around the Black Sea, although this is still unclear. The finding of haplogroup I in samples of mostly WHG ancestry could confuse people that do not care about timing, sub-structured populations, and gene flow.
NOTE. If you don’t understand why ‘clusters’ that span thousands of years don’t really matter for the many Palaeolithic population expansions that certainly happened among hunter-gatherers in Europe, just take a look at what happened with Bell Beakers expanding from Yamna into western Europe within 500 years.
If we don’t thread carefully when talking about population migrations, these terms are bound to confuse people. Just as the fixation on “steppe ancestry” – which marks the arrival in Chalcolithic Europe of peoples from the Pontic-Caspian region – has confused a lot of researchers to this day.
When I began to write about the Indo-European demic diffusion model, my concern was to find a single spot where a North-West Indo-European proto-language could have expanded from ca. 2000 BC (our most common guesstimate). Based on the 2015 papers, and in spite of their conclusions, I thought it had become clear that Corded Ware was not it, and it was rather Bell Beakers. I assumed that Uralic was spoken to the north (as was the traditional belief), and thus Corded Ware expanded from the forest zone, hence steppe ancestry would also be found there with other R1a lineages.
With the publication of Mathieson et al. (2017) and Olalde et al. (2017), I changed my mind, seeing how “steppe ancestry” did in fact appear quite late, hence it was likely to be the result of very specific population movements, probably directly from the Caucasus. Later, Mathieson published in a revision the sample from Alexandria of hg R1a-M417 (probably R1a-Z645, possibly Z93+), which further supported the idea that the migration of Corded Ware peoples started near the North Pontic forest-steppe (as I included in a the next revision).
The question remains the same I repeated recently, though: where do the extra Caucasus components (i.e. beyond EHG) of Eneolithic Ukraine/Corded Ware and Khvalynsk/Yamna come from?
Considering 2-way mixtures, we can model Karelia_HG as deriving 34 ± 2.8% of its ancestry from a Villabruna-related source, with the remainder mainly from ANE represented by the AfontovaGora3 (AG3) sample from Lake Baikal ~17kya.
AG3 was likely of haplogroup Q1a (as reported by YFull, see Genetiker), and probably the ANE ancestry found in Eastern Europe accompanied a Palaeolithic migration of Q1a2-M25 (formed ca. 22600 BC, TMRCA ca. 14300 BC).
Combined with what we know about the Eneolithic Steppe and Caucasus populations – it is likely that ANE ancestry remained the most important component of some of the small ghost populations of the Caucasus until their emergence with the Lola culture.
The first sample we have now attributed to the EHG cluster is Sidelkino, from the Samara region (ca. 9300 BC), mtDNA U5a2. In Damgaard et al. (Science 2018), Yamnaya could be modelled as a CHG population related to Kotias Klde (54%) and the remaining from ANE population related to Sidelkino (>46%), with the following split events:
A split event, where the CHG component of Yamnaya splits from KK1. The model inferred this time at 27 kya (though we note the larger models in Sections S2.12.4 and S2.12.5 inferred a more recent split time).
A split event, where the ANE component of Yamnaya splits from Sidelkino. This was inferred at about about 11 kya.
A split event, where the ANE component of Yamnaya splits from Botai. We inferred this to occur 17 kya. Note that this is above the Sidelkino split time, so our model infers Yamnaya to be more closely related to the EHG Sidelkino, as expected.
An ancestral split event between the CHG and ANE ancestral populations. This was inferred to occur around 40 kya.
Other samples classified as of the EHG cluster:
Popovo2 (ca. 6250 BC) of hg J1, mtDNA U4d – Po2 and Po4 from the same site (ca. 6550 BC) show continuity of mtDNA.
Karelia_HG, from Juzhnii Oleni Ostrov (ca. 6300 BC): I0211/UzOO40 (ca. 6300 BC) of hg J1(xJ1a), mtDNA U4a; and I0061/UzOO74 of hg R1a1(xR1a1a), mtDNA C1
UzOO77 and UzOO76 from Juzhnii Oleni Ostrov (ca. 5250 BC) of mtDNA R1b.
Samara_HG from Lebyanzhinka (ca. 5600 BC) of hg R1b1a, mtDNA U5a1d.
About the enigmatic Anatolia_Neolithic-related ancestry found in Pontic-Caspian steppe samples, this is what Wang et al. (2018) had to say:
We focused on model of mixture of proximal sources such as CHG and Anatolian Chalcolithic for all six groups of the Caucasus cluster (Eneolithic Caucasus, Maykop and Late Makyop, Maykop-Novosvobodnaya, Kura-Araxes, and Dolmen LBA), with admixture proportions on a genetic cline of 40-72% Anatolian Chalcolithic related and 28-60% CHG related (Supplementary Table 7). When we explored Romania_EN and Greece_Neolithic individuals as alternative southeast European sources (30-46% and 36-49%), the CHG proportions increased to 54-70% and 51-64%, respectively. We hypothesize that alternative models, replacing the Anatolian Chalcolithic individual with yet unsampled populations from eastern Anatolia, South Caucasus or northern Mesopotamia, would probably also provide a fit to the data from some of the tested Caucasus groups.
The first appearance of ‘Near Eastern farmer related ancestry’ in the steppe zone is evident in Steppe Maykop outliers. However, PCA results also suggest that Yamnaya and later groups of the West Eurasian steppe carry some farmer related ancestry as they are slightly shifted towards ‘European Neolithic groups’ in PC2 (Fig. 2D) compared to Eneolithic steppe. This is not the case for the preceding Eneolithic steppe individuals. The tilting cline is also confirmed by admixture f3-statistics, which provide statistically negative values for AG3 as one source and any Anatolian Neolithic related group as a second source
Detailed exploration via D-statistics in the form of D(EHG, steppe group; X, Mbuti) and D(Samara_Eneolithic, steppe group; X, Mbuti) show significantly negative D values for most of the steppe groups when X is a member of the Caucasus cluster or one of the Levant/Anatolia farmer-related groups (Supplementary Figs. 5 and 6). In addition, we used f- and D-statistics to explore the shared ancestry with Anatolian Neolithic as well as the reciprocal relationship between Anatolian- and Iranian farmer-related ancestry for all groups of our two main clusters and relevant adjacent regions (Supplementary Fig. 4). Here, we observe an increase in farmer-related ancestry (both Anatolian and Iranian) in our Steppe cluster, ranging from Eneolithic steppe to later groups. In Middle/Late Bronze Age groups especially to the north and east we observe a further increase of Anatolian farmer related ancestry consistent with previous studies of the Poltavka, Andronovo, Srubnaya and Sintashta groups and reflecting a different process not especially related to events in the Caucasus.
(…) Surprisingly, we found that a minimum of four streams of ancestry is needed to explain all eleven steppe ancestry groups tested, including previously published ones (Fig. 2; Supplementary Table 12). Importantly, our results show a subtle contribution of both Anatolian farmer-related ancestry and WHG-related ancestry (Fig.4; Supplementary Tables 13 and 14), which was likely contributed through Middle and Late Neolithic farming groups from adjacent regions in the West. The discovery of a quite old AME ancestry has rendered this probably unnecessary, because this admixture from an Anatolian-like ghost population could be driven even by small populations from the Caucasus.
While it is not yet fully clear, the increased Anatolian_Neolithic-like ancestry in Ukraine_Eneolithic samples (see below) makes it unlikely that all such ancestry in Corded Ware groups comes from a GAC-related contribution. It is likely that at least part of it represents contributions from populations of the Caucasus, based on the mostly westward population movements in the steppe from ca. 4600 BC on, including the Suvorovo-Novodanilovka expansion, and especially the Kuban-Maykop expansion during the final Eneolithic into the North Pontic area.
NOTE. Since CHG-like groups from the Caucasus may have combinations of AME and ANE ancestry similar to Yamna (which may thus appear as ‘steppe ancestry’ in the North Pontic area), it is impossible to interpret with precision the following ADMIXTURE graphic:
The East Asian contribution to samples from the WHG samples (like Loschbour or La Braña), as specified in Fu et al. (2016), does not seem to be related to Baikal_EN, and appears possibly (in the ADMIXTURE analysis) integrated into he Villabruna component. I guess this implies that the shared alleles with East Asians are quite early, and potentially due to the expansion of R1b-L754 from the East.
It would be interesting to know the specific material culture Sidelkino belonged to – i.e. if it was related to the expansion of the North-Eastern Technocomplex – , and its Y-DNA. The Post-Swiderian expansion into eastern Europe, probably associated with the expansion of R1b-P297 lineages (including R1b-M73, found later in Botai and in Baltic HG) is supposed to have begun during the 11th millennium BC, but migrations to the Urals and beyond are probably concentrated in the 9th millennium, so this sample is possibly slightly early for R1b.
NOTE. User Rozenfeld at Anthrogenica posted this, which I think is interesting (in case anyone wants to try a Y-SNP call):
there is something strange with Sidelkino EHG: first, its archaeological context is not described in the supplementary. Second, its sex is not listed in the supplementary tables. Third, after looking for info about this sample, I found that: “Сиделькино-3. Для снятия вопроса о половой принадлежности индивида была проведена генетическая экспертиза, выявившая принадлежность останков мужчине.”(translation: Sidelkino-3. To resolve the question about sex of the remains, the genetic analysis was conducted, which showed that remains belonged to male), source: http://static.iea.ras.ru/books/7487_Traditsii.pdf
So either they haven’t mentioned his Y-DNA in the paper for some reason, or there are more than one Sidelkino sample and the male one has not yet been published. The coverage of the Sidelkino sample from the paper is 2.9, more than enough to tell Y-DNA haplogroup.
My speculative guess right now about specific population movements in far eastern Europe, based on the few data we have:
The expansion of the North-Eastern Technocomplex first around the 9th millennium BC, most likely expanded R1b-P279 ca. 11300 BC, judging by its TMRCA, with both R1b-M73 (TMRCA 5300) and R1b-M269 (TMRCA 4400 BC) info (with extra El Mirón ancestry) back, and thus Eurasiatic.
The expansion of haplogroup J1 to the north may have happened before or after the R1b-P279 expansion. Judging by the increase in AG3-related ancestry near Karelia compared to Baltic_HG, it is possible that it expanded just after R1b-P279 (hence possibly J1-Y6304? TMRCA 9700 BC). Its long-lasting presence in the Caucasus is supported by the Satsurblia (ca. 11300 BC) and the Dolmen BA (ca. 1300 BC) samples.
The expansion of R1a-M17 ca. 6600 BC is still likely to have happened from the east, based on the R1a-M17 samples found in Baikalic cultures slightly later (ca. 5300 BC). The presence of elevated Baikal_EN ancestry in Karelia HG and in Samara HG, and the finding of R1a-M417 samples in the Forest Zone after the Mesolithic suggests a connection with the expansion of Hunter-Gatherer pottery, from the Elshanka culture in the Samara region northward into the Forset Zone and westward into the North Pontic area.
The expansion of R1b-M73 ca. 5300 BC is likely to be associated with the emergence of a group east of the Urals (related to the later Botai culture, and potentially Pre-Yukaghir). Its presence in a Narva sample from Donkalnis (ca. 5200 BC) suggest either an early split and spread of both R1b-P297 lineages (M73 and M269) through Eastern Europe, or maybe a back-migration with hunter-gatherer pottery.
R1b-M269 spread successfully ca. 4400 BC (and R1b-L23 ca. 4100 BC, both based on TMRCA), and this successful expansion is probably to be associated with the Khvalynsk-Novodanilovka expansion. We already know that Samara_HG ca. 5600 was R1b1a, so it is likely that R1b-M269 appeared (or ‘resurged’) in the Volga-Ural region shortly after the expansion of R1a-M17, whose expansion through the region may be inferred by the additional AG3 and Baikal_EN ancestry. Interesting from Samara_HG compared to the previous Sidelkino sample is the introduction of more El Mirón-related ancestry, typical of WHG populations (and thus proper of Baltic groups).
NOTE. The TMRCA dates are obviously gross approximations, because a) the actual rate of mutation is unknown and b) TMRCA estimates are based on the convergence of lineages that survived. The potential finding of R1a-Z645 (possibly Z93+) in Ukraine Eneolithic (ca. 4000 BC), and the potential finding of R1b-L23 in Khvalynsk ca. 4250 BC complicates things further, in terms of dates and origins of any subclade.
The question thus remains as it was long ago: did R1b-M269 lineages expand (‘return’) from the east, near the Urals, or directly from the north? Were they already near Samara at the same time as the expansion of hunter-gatherer pottery, and were not much affected by it? Or did they ‘resurge’ from populations admixed with Caucasus-related ancestry after the expansion of R1a-M17 with this pottery (since there are different stepped expansions from the Samara region)? We could even ask, did R1a-M17 really expand from the east, i.e. are the dates on Baikalic subclades from Moussa et al. (2016) reliable? Or did R1a-M17 expand from some pockets in the Pontic-Caspian steppe, taking over the expansion of HG pottery at some point?
The most interesting aspect from the new paper (regarding Indo-Uralic migrations) is that Ancestral Middle Easterner ancestry will probably be a better proxy for the Anatolia_Neolithic component found in Ukraine Mesolithic to Eneolithic, and possibly also for some of the “more CHG-like” component found among Pontic-Caspian steppe populations, all likely derived from different admixture events with groups from the Caucasus.
NOTE. Even the supposed gene flow of Neolithic Iranian ancestry into the Caucasus can be put into question, since that means possibly a Dzudzuana-like population with greater “deep ancestry” proportion than the one found in CHG, which may still be found within the Caucasus.
If it was not clear already that following ‘steppe ancestry’ wherever it appears is a rather lame way of following Indo-European migrations, every single sample from the Caucasus and their admixture with Pontic-Caspian steppe populations will probably show that “steppe ancestry” is in fact formed by a variety of steppe-related ancestral components, impossible to follow coherently with a single population. Exactly what is happening already with the Siberian ancestry.
If the paper on the Dzudzuana samples has shown something, is that the expansion of an ANE-like population shook the entire Caucasus area up to the Zagros Mountains, creating this ANE – AME cline that are CHG and Iran_N, with further contributions of “deep ancestries” (probably from the south) complicating the picture further.
If this happens with few known samples, and we know of an ANE-like ghost population in the Caucasus (appearing later in the Lola culture), we can already guess that the often repeated “CHG component” found in Ukraine_Eneolithic and Khvalynsk will not be the same (except the part mediated by the Novodanilovka expansion).
This ANE-like expansion happened probably in the Late Upper Palaeolithic, and reached Northern Europe probably after the expansion of the Villabruna cluster (ca. 12000 BC), judging by the advance of AG3-like and ENA-like ancestry in later WHG samples.
The population movements during the Mesolithic and Early Neolithic in the North Pontic area are quite complicated: the extra AME ancestry is probably connected to the admixture with populations from the Caucasus, while the close similarity of Ukraine populations with Scandinavian ones (with an increase in Villabruna ancestry from Mesolithic to Neolithic samples), probably reveal population movements related to the expansion of Maglemose-related groups.
These Maglemose-related groups were probably migrants from the north-west, originally from the Northern European Plains, who occupied the previous Swiderian territory, and then expanded into the North Pontic area. The overwhelming presence of I2a (likely all I2a2a1b1b) lineages in Ukraine Neolithic supports this migration.
The likely picture of Mesolithic-Neolithic migrations in the North Pontic area right now is then:
Expansion of R1a-M459 from the east ca. 12000 BC – probably coupled with AG3 and also some Baikal_EN ancestry. First sample is I1819 from Vasilievka (ca. 8700 BC), another is from Dereivka ca. 6900 BC.
Expansion of R1b-V88 from the Balkans in the west ca. 9700 BC, based on its TMRCA and also the Balkan hunter-gatherer population overwhemingly of this haplogroup from the 10th millennium until the Neolithic. First sample is I1734 from Vasilievka (ca. 7252 BC), which suggests that it replaced the male population there, based on their similar EHG-like adxmixture (and lack of sizeable WHG increase), and shared mtDNA U5b2, U5a2.
Expansion of I2a-Y5606 probably ca. 6800 based on its TMRCA with Janislawice culture. Supporting this is the increase in WHG contribution to Neolithic samples, including the spread of U4 subclades compared to the previous period.
Expansion of R1a-M17 starting probably ca. 6600 BC in the east (see above).
NOTE. The first sample of haplogroup I appears in the Mesolithic: I1763 (ca. 8100 BC) of haplogroup I2a1, probably related to an older Upper Palaeolithic expansion.
It is becoming more and more clear with each new paper that – unless the number of very ancient samples increases – the use of Y-chromosome haplogroups remains one of the most important tools for academics; this is especially so in the steppes, in light of the diversity found in populations from the Caucasus. A clear example comes from the Yamna – Corded Ware similarities:
The presence of haplogroups Q and R1a-M459 (xM17) in Khvalynsk along with a R1b1a sample, which some interpreted as being akin to modern ‘mixed’ populations in the past, is likely to point instead to a period of Khvalynsk-Novodanilovka expansion with R1b-M269, where different small populations from the steppe were being integrated into the common Khvalynsk stock, but where differences are seen in material culture surrounding their burials, as supported by the finding of R1b1 in the Kuban area already in the first half of the 5th millennium. The case would be similar to the early ‘mixed’ Icelandic population.
Only after the emergence of the Samara culture (in the second half of the 6th millennium BC), with a sample of haplogroup R1b1a, starts then the obvious connection with Early Proto-Indo-Europeans; and only after the appearance of late Sredni Stog and haplogroup R1a-M417 (ca. 4000 BC) is its connection with Uralic also clear. In previous population movements, I think more haplogroups were involved in migrations of small groups, and only some communities among them were eventually successful, expanding to be dominant, creating ever growing cultures during their expansions.
Indeed, if you think in terms of Uralic and Indo-European just as converging languages, and forget their potential genetic connection, then the genetic + linguistic picture becomes simplified, and the upper frontier of the 6th millennium BC with a division North Pontic (Mariupol) vs. Volga-Ural (Samara) is enough. However, tracing their movements backwards – with cultural expansions from west to east (with the expansion of farming), and earlier east to west (with hunter-gatherer pottery), and still earlier west to east (with the north-eastern technocomplex), offers an interesting way to prove their potential connection to macrofamilies, at least in terms of population movements.
I am quite convinced right now that it would be possible to connect the expansion of R1b-L754 subclades with a speculative Nostratic (given the R1b-V88 connection with Afroasiatic, and the obvious connection of R1b-L297 with Eurasiatic). Paradoxically, the connection of an Indo-Uralic community in the steppes (after the separation of Yukaghir) with any lineage expansion (R1a-M17, R1b-M269, or even Q, I or J1) seems somehow blurrier than one year ago, possibly just because there are too many open possibilities.
David Reich says about the admixture with Neanderthals, which he helped discover:
At the conclusion of the Neanderthal genome project, I am still amazed by the surprises we encountered. Having found the first evidence of interbreeding between Neanderthals and modern humans, I continue to have nightmares that the finding is some kind of mistake. But the data are sternly consistent: the evidence for Neanderthal interbreeding turns out to be everywhere. As we continue to do genetic work, we keep encountering more and more patterns that reflect the extraordinary impact this interbreeding has had on the genomes of people living today.
I think this is a shared feeling among many of us who have made proposals about anything, to fear that we have made a gross, evident mistake, and constantly look for flaws. However, it seems to me that geneticists are more preoccupied with being wrong in their developed statistical methods, in the theoretical models they are creating, and not so much about errors in the true ancient ethnolinguistic picture human population genetics is (at least in theory) concerned about. Their publications are, after all, constantly associating genetic finds with cultures and (whenever possible) languages, so this aspect of their research should not be taken lightly.
Seeing how David Anthony or Razib Khan (among many others) have changed their previously preferred migration models as new data was published, and they continue to be respected in their own fields, I guess we can be confident that professionals with integrity are going to accept whatever new picture appears. While I don’t think that genetic finds can change what we can reconstruct with comparative grammar, I am also ready to revise guesstimates and routes of expansion of certain dialects if R1a-Z645 is shown to have accompanied Late Proto-Indo-Europeans during their expansion with Yamna, and later integrated somehow with Corded Ware.
However, taking into account the obsession of some with an ancestral, uninterrupted R1a—Indo-European association, and the lack of actual political repercussion of Neanderthal admixture, I think the most common nightmare that all genetic researchers should be worried about is to keep inflating this “Yamnaya ancestry”-based hornet’s nest, which has been constantly stirred up for the past two years, by rejecting it – or, rather, specifying it into its true complex nature.
This succession of corrections and redefinitions, coupled with the distinct Y-DNA bottleneck of each steppe population, will eventually lead to a completely different ethnolinguistic picture of the Pontic-Caspian region during the Eneolithic, which is likely to eventually piss off not only reasonable academics stubbornly attached to the CWC-IE idea, but also a part of those interested in daydreaming about their patrilineal ancestors.
Sometimes it’s better to just rip off the band-aid once and for all…
Most mtDNA lineages found are characteristic of the early Neolithic farmers in south-eastern and central Europe of the Starčevo-Kőrös-Criş and LBK cultures. Haplogroups N1a, T2, J, K, and V, which are found in the Neolithic BKG, TRB, GAC and Early Bronze Age samples, are part of the mitochondrial ‘Neolithic package’ (which also includes haplogroups HV, V, and W) that was introduced to Europe with farmers migrating from Anatolia at the onset of the Neolithic17,31.
A noteworthy proportion of Mesolithic haplogroup U5 is also found among the individuals of the current study. The proportion of haplogroup U5 already present in the earliest of the analysed Neolithic groups from the examined area differs from the expected pattern of diversity of mtDNA lineages based on a previous archaeological view and on the aDNA findings from the neighbouring regions which were settled by post-Linear farmers similar to BKG at that time. A large proportion of Mesolithic haplogroups in late-Danubian farmers in Kuyavia was also shown in previous studies concerning BKG samples based on mtDNA only, although these frequencies were derived on the basis of very small sample sizes.
A significant genetic influence of HG populations persisted in this region at least until the Eneolithic/Early Bronze Age period, when steppe migrants arrived to central Europe. The presence of two outliers from the middle and late phases of the BKG in Kuyavia associated with typical Neolithic burial contexts provides evidence that hunter-farmer contacts were not restricted to the final period of this culture and were marked by various episodes of interaction between two societies with distinct cultural and subsistence differences.
The identification of both mitochondrial and Y-chromosome haplogroup lineages of Mesolithic provenance (U5 and I, respectively) in the BKG support the theory that both male and female hunter-gatherers became part of these Neolithic agricultural societies, as has been reported for similar cases from the Carpathian Basin, and the Balkans. The identification of an individual with WHG affinity, dated to ca. 4300 BCE, in a Middle Neolithic context within a BKG settlement, provides direct evidence for the regional existence of HG enclaves that persisted and coexisted at least for over 1000 years, from the arrival of the LBK farmers ca. 5400 BCE until ca. 4300 BCE, in proximity with Neolithic settlements, but without admixing with their inhabitants.
The analysis of two Late Neolithic cultures, the GAC and CWC, shows that steppe ancestry was present only among the CWC individuals analysed, and that the single GAC individual had more WHG ancestry than previous local Neolithic individuals. (…) The CWC’s affinity to WHG, however, contrasts with results from published CWC individuals that identified steppe ancestry related to Yamnaya as the major contributor to the CWC genomes, while here we report also substantial contributions from WHG that could relate to the late persistence of pockets of WHG populations, as supported by the admixture results of N42 and the finding of the 4300-year-old N22 HG individual. These results agree with archaeological theories that suggest that the CWC interaction with incoming steppe cultures was complex and that it varied by region.
About the analyzed CWC samples, it is remarkable that, even though they are somehow related to each other, they do not form a tight cluster. Also, their Y-DNA (I2a), and this:
When compared to previously published CWC data, our CWC group (not individuals) is genetically significantly closer to WHG than to steppe individuals (Z = −4.898), a result which is in contrast with those for CWC from Germany (Z = 2.336), Estonia (Z = 0.555), and Latvia (Z = 1.553).
Włodarczak (2017) talks about the CWC period in Poland after ca. 2600 BC as a time of emergence of an allochthnous population, marked by the rare graves of this area, showing infiltrations initially mainly from Lesser Poland, and later (after 2500 BC) from the western Baltic zone.
Since forest sub-Neolithic populations would have probably given more EHG to the typical CWC population, these samples support the resurge of ‘local’ pockets of GAC- or TRB-like groups with more WHG (and also Levant_Neolithic) ancestry.
The known presence of I2a2a1b lineages in GAC groups in Poland also supports this interpretation, and the subsistence of such pockets of pre-steppe-like populations is also seen with the same or similar lineages appearing in comparable ‘resurge’ events in Central Europe, e.g. in samples from the Únětice and Tumulus culture.
About the Bronze Age sample, we have at last official confirmation of haplogroup R1a1a (sadly no subclade*) at the very beginning of the Trzciniec period – in a region between western (Iwno) and eastern (Strzyżów) groups related to Mierzanowice – , which has to be put in relation with the samples from the final Trzciniec period in the Baltic published in Mittnik et al. (2018).
EDIT (8 OCT 2018): More specific subclades have been published, including a R1a-Z280 lineage for the Bronze Age sample (see spreadsheet).
This confirms the early resurge of R1a-Z645 (probably R1a-Z282) lineages at the core of the developing East European Bronze Age, a province of the European Bronze Age that emerged from evolving Bell Beaker groups in Poland.
I don’t have any hope that the Balto-Slavic evolution through BBC Poland → Mierzanowice/Iwno → Trzciniec → Lusatian cultures is going to be confirmed any time soon, until we have a complete trail of samples to follow all the way to historic Slavs of the Prague culture. However, I do think that the current data on central-east Europe – and the recent data we are receiving from north-east Europe and the Iranian steppes, at odds with the Indo-Slavonic alternative – supports this model.
I guess that, in the end, similar to how the Yamna vs. Corded Ware question is being solved, the real route of expansion of Proto-Balto-Slavic (supposedly spoken ca. 1500-1000 BC) is probably going to be decided by the expansion of either R1a-M458 (from the west) or R1a-Z280 lineages (from the east), because the limited precision of genetic data and analyses available today are going to show ‘modern Slavic’-like populations from the whole eastern half of Europe for the past 4,000 years…
The Middle Neolithic is known to mark the westward expansion of Comb Ware and related cultures in North-Eastern Europe.
Mathieson et al. (2017 and 2018) had this to say about the Middle Neolithic in the Baltic:
At Zvejnieki in Latvia, using 17 newly reported individuals and additional data for 5 previously reported34 individuals, we observe a transition in hunter-gatherer-related ancestry that is opposite to that seen in Ukraine. We find that Mesolithic and Early Neolithic individuals (labelled ‘Latvia_HG’) associated with the Kunda and Narva cultures have ancestry that is intermediate between WHG (approximately 70%) and EHG (approximately 30%), consistent with previous reports34–36(Supplementary Table 3). We also detect a shift in ancestry between Early Neolithic individuals and those associated with the Middle Neolithic Comb Ware complex (labelled ‘Latvia_MN’), who have more EHG-related ancestry; we estimate that the ancestry of Latvia_MN individuals comprises 65% EHG-related ancestry, but two of the four individuals appear to be 100% EHG in principal component space (Fig. 1b).
Other samples and errors on Y-SNP calls
The truth is, this is another sample (Latvia_MN_dup.I4627.SG) from the same individual ZVEJ26.
There is another sample used for the analysis of ZVEJ26, with the same data as in Mathieson et al. (2018), i.e. better coverage, and Y-DNA R1b1a1a(xR1b1a1a2).
Most samples in the tables from Wang et al. (2018) seem to be classified correctly, as in previous papers, but for:
Blätterhöhle Cave sample from Lipson et al. (2017), wrongly classified (again) as R1b1a1a2a1a2a1b2 (I am surprised no R1b-autochtonous-continuity-fan rushed to proclaim something based on this);
Mal’ta 1 sample from Raghavan et al. (2013) as R1b1a1a2;
Iron Gates HG, Schela Cladovey from Gonzalez Fortes (2017) as R1b1a1a2;
Oase1 from Fu (2015) as N1c1a;
samples from Skoglund et al. (2017) from Africa also wrongly classified as R1b1a1a2 and subclades.
It seems therefore that the poor coverage / SNPs hit on autosomes is the key common factor here for these Y-SNP calls, and so it is in the Zvejnieki MN1 duplicated sample. Anyway, if all Y-SNP calls come from the same software applied to all data, and this is going to be used in future papers, this seems to be a great improvement compared to Narasimhan et al. (2018)…
EDIT (25 JUN 2018): I have been reviewing some more papers apart from Mathieson et al. (2018) and Olalde et al. (2018) to compare the reported haplogroups, and there seems to be many potential errors (or updated data, difficult to say sometimes, especially when the newly reported haplogroup is just one or two subclades below the reported one in ‘old’ papers), not only those listed above.
The sample accession number in the European Nucleotide Archive (ENA) is SAMEA45565168 (Latvia_MN1/ZVEJ26) (see here), in case anyone used to this kind of analysis wishes to repeat the Y-SNP calls on both samples.
EDIT (25 JUN 2018): Added that it is another sample with lesser coverage from the same ZVEJ26 individual.
In 1898, two megalithic tombs containing graves of a local social elite dated to the Early Bronze Age were discovered by N. I. Veselovsky near the village of Tsarskaya (modern Novosvobodnaya, Republic of Adygeya) (Fig. 1 (a)) (Baye 1900, 43–59; IAC 1901, 33–8; Sagona 2018, 281–97).
Radiocarbon dates place both tombs within the Novosvobodnaya phase of the Maykop culture, between c. 3200 and 2900 BC (Trifonov et al. 2017). Along with the human remains (one adult individual was interred in each dolmen), the tombs yielded rich funerary offerings, including artefacts made of gold, silver and semi-precious stones. (…) This paper presents results of a technical analysis of just one type of artefact, from kurgan 2 at Tsarskaya: thin-walled jointless beads made from gold.
Ever since M. I. Rostovtzeff noted a stylistic similarity between Maykop art and Sumerian art (Rostovtzeff 1920) and M. V. Andreeva described this phenomenon within a broad cultural and chronological context (Andreeva 1977), new archaeological studies have only extended this picture of a vast cultural province that appeared between the Caucasus and the northern fringe of Western Asia (Trifonov 1987). The discovery of the Leyla-Tepe culture (Narimanov 1987) and Maykop-type kurgans in Azerbaijan (Lyonnet et al. 2008) and adjacent Iran (Muscarella 1969, 1971, 2003; Trifonov 2000) has confirmed the spatial and temporal unity of this phenomenon as a precondition for free circulation of cultural patterns and technical innovations across vast areas of the Caucasus and Western Asia. Jewellery made of gemstones and precious metals, primarily gold, was probably one such innovation.
Attempts to demarcate the historical region where the Maykop culture emerged and developed have emphasized the role of Upper Mesopotamia in the development of the Sumerian civilization and the definition of a northern centre of urbanization, independent from the centres of the south (Rothman 2002; Oats et al. 2007). The turn of the fourth millennium BC saw the development of various cultural traditions in south-east Anatolia, north-east Syria and north-west Iran; on the northern fringe, these traditions manifested themselves through the Maykop culture. Perhaps it is no coincidence that the first high-status burials containing gold and gemstone jewellery (including carnelian, turquoise and lapis lazuli) appear in these northern, rather than southern, centres in the first quarter of 4000 BC (e.g., Tepe Gawra, graves 109, 110) (Piasnall 2002). With regard to funeral rites and stylistic characteristics of jewellery pieces, these graves have many parallels with early Maykop burials (Munchaev 1975, 329; Trifonov 1987, 20).
It still remains unclear if the goldsmiths of Upper Mesopotamia mastered the technique of making thin-walled jointless beads. The gold beads from Tepe Gawra are described as spherical or ball-shaped, but their maximum diameter (5–8mm) always exceeds the length of the bore (3–4mm) (Tobler 1950, 89, 199, pl. LV, a). On the whole, these measurements are consistent with the proportions and sizes of some Maykop beads.(…)
It is quite possible that a distinctive technique of making thin-walled jointless beads from gold was a regional technological development of Maykop culture goldsmiths, within a wider tradition of Near East metalwork, as a type of production regulated by ritual beliefs (Gell 1992; Benzel 2013).
These deep-rooted Near East traditions of ritualization of the production and use of jewellery pieces made of gold, silver and gemstones in the Maykop culture, on the one hand, maintained familiar canons of ritual behaviour and, on the other, made perception of sophisticated symbolism of gemstones more difficult for neighbouring cultures with different living standards, levels of social development and value systems to understand. The jewellery traditions of the Maykop culture had no successors in the Caucasus or the adjacent steppes. In the third millennium BC, the goldsmiths of Europe and Asia had to reinvent the technique of making thin-walled jointless gold beads from scratch (Born et al. 2009).
Related excerpts, about the climate of an adjacent region of the Caucasus before, during, and after the Maykop culture:
The 7th millennium BC featured a warm and arid climate, so that time corresponds to the steppe landscapes in the final stage of the Mesolithic. It is likely that the formation of a producing economy in the mountainous zone of Dagestan gradually emerged against this background. In the Neolithic period, the area remained almost treeless, as it was still warm and quite dry. However, archaeological data indicates that long-term settlements with well-developed farming spread in the mountainous zone around 6200-5500 BC.
The beginning of increasing humidity and the appearance of deciduous forests corresponds to the early Chalcolithic period of the Eastern Caucasus. It is the most poorly studied period in the history of this region. Covering a time span of 2000 years, this period was the least saturated by archaeological sites. At the start of this period, only the stands of herdsman in the mountain zone are known, dating to the second half of the 6th millennium BC (Gadgiev, 1991). It is still not clear whether the mountains were not settled in such a favorable climatic stage. The uncertainty may be due to the fact that people have chosen other ecological niches, or it could be we simply do not have data due to the insufficient archaeological survey of the territory. It is surprising that the turn to drier climate and the reduction of deciduous forests in the inner mountainous part of Dagestan, the large, long-term settlements like Ginchi emerge with pronounced specialization in agriculture (Fig. 7 panel (2)) (Gadgiev, 1991).
After the dry climate, simultaneously with cooling, the subsequent spread of pine forests coincides with the beginning of expansion of Kura-Araxes culture from the territory of Georgia through Chechnya to the mountainous Dagestan. Debates on the impact of past climate on Kura-Araxes societies in Transcaucasus have a long history (for the comprehensive review see, for example, Connor and Kvavadze, 2014 and references therein). In general, it is clear that after 3000 BC, forest cover in most areas of the Kura-Araxes region in the Transcaucasia reached its maximum extent in the Holocene (Connor and Kvavadze, 2014). However, at the same time lakes in Central Anatolia began to dry out and Caspian Sea levels fell (Roberts et al. 2011; Leroy et al. 2013), and arid conditions were identified in mountainous Dagestan in the 4th millennium. Clearly the regional moisture balance shifted in the Eastern Caucasus only in the late 4th to early 3rd millennium BC (this study). The only available radiocarbon dating of Dagestan confirms that the agricultural settlements of the Early Bronze Age appear not in the middle of the 4th millennium BC, but in the early 3rd millennium BC; that is not earlier than the stage of increasing moistening and the appearance of pine forests.
The Turganik settlement in the Orenburg Region constitutes part of the so-called Ivanovo microregion of cultural heritage monuments, along with the Mesolithic Starotokskaya site; an Ivanovskoye multi-layered settlement (Neolithic, Eneolithic [or Chalcolithic], Late Bronze Age); Toksky I and Toksky II settlements attributed to the Late Bronze Age (the Timber-Grave archaeological culture); an Ivanovsky ground burial dated to the Eneolithic; and the Ivanovsky kurgan cemetery of the Early Iron Age (Fig. 1).
The ancient settlements are located at the Turganik River mouth, where the river joins the Tok River (the Samara River drainage basin). The Turganik River enters an old channel of the Tok which continues to flow due to that fact. Both valleys are wide and dissected by multiple river channels. The floodplain landscapes are mostly wet meadows with rich herb and grass vegetation, pastures, and hay fields. On both sides of the Turganik River, and farther along the right side of the Tok valley there are flat-topped elevations, with occasional forests (Chibilev, 1996). The Turganik settlement was positioned on a slightly elevated surface at the confluence of the Turganik and Tok rivers, on the right side of the valley. The settlement was inhabited in the Eneolithic and the Late Bronze Age, the fifth to fourth millennia BC.
Results and discussion
Pollen assemblages of the Atlantic optimum ~ 5500 yr BP indicate some increase in moisture supply and related afforestation of the floodplain (Lavrushin and Spiridonova, 1995). As follows from our data, the site was abandoned at that time and the no-longer-functioning cultural layer VI was gradually buried under deposits of frequent floods. According to the 14C ages obtained on archeological materials, the age of layer VI (or the second stage of the Eneolithic epoch on the Turganik settlement) may be dated to 4237–3790 cal yr BC, that is, somewhat earlier than the Holocene optimum suggested by palynologists.
Layer V shows another interval marked by increasing climate aridity and the dominance of grass steppes. As stated by the above-cited authors, the climate at the time that layer V was functioning was even dryer than during the formation of layer VI. That is confirmed by our data on the layer V composition, was formed during early Pit-Grave culture (the Early Bronze Age), in the range from 3800–3360 BC, according to the dates obtained on archeological materials (Morgunova et al., 2016b). As follows from the above, the maximum of aridity coincided with the Atlantic optimum.
It follows from the above that the Atlantic period of the Holocene was mostly characterized by arid environments; the peak of aridity fell on the early Bronze Age, the time of the early (Repino) stage of the Yamnaya culture in the Cis-Ural steppes. The Subboreal and Subatlantic periods were relatively colder and more humid, though short episodes of aridity could occur and some of them happened to be recorded in the sequence under study.
The reconstructed history of the climate changes in the Cis-Ural steppes during three intervals of the Holocene is in a good agreement with the results obtained in other regions. According to Alexandrovskiy (1996, 2000; Alexandrovskiy et al., 1999, 2004), the Atlantic period was the most arid one in the south of Russia, the subsequent intervals being comparatively wetter and colder. The extreme aridity was recorded on the Ukraine territory at the final Atlantic period, a few less arid chrono-intervals having been identified over the entire period (Kotova, 2009).
There are, however, other schemes of climate fluctuations in the central part of the Russian steppe zone; a few of them consider the Atlantic period to be humid, or even the most humid, as compared with the second half of the Holocene (Ivanov, 1992; Demkin, 1997). Also acceptable is a scenario of climatic fluctuations occurring at different times in different regions (Chendev et al., 2010). Further investigations and accumulation of empirical data would help to gain a better insight into the problem.
The ancient people inhabited the place from 5000 to 4000 BC (actually throughout the Atlantic period), when the place was not subjected to flooding. At the time of human habitation, the climate was mostly arid. Paleosols of that time are attributable to the Kastanozems (Endosalic Protosodic). They developed under grass (or herb and grass) steppes. The peak of aridity falls on the final Atlantic period. At the end of Eneolithic epoch (the fifth millennium BC) and in the Early Bronze Age (the fourth millennium BC) there were short-term but violent floods, which forced people to leave the habitable place.
During the Subboreal and Subatlantic periods of the Holocene, the climate became more humid, the floods became regular, the vegetation was dominated by meadow forbs and herbs growing on meadow-chernozem soils (Luvic Chernozem [Stagnic]), and the settlement was completely abandoned. In general, the studied sedimentary record at the Turganik archeological site reveals traceable climate change towards lower temperatures and increasing humidity in the second part of the Holocene, with occasional episodes of aridity that did not affect the general trend.
The central East European plains are famous for their Epigravettian sites that date to around 15–12 14C ka BP (ca. 18.2–13.8 cal ka BP) and display impressive large structures made from the bones of woolly mammoth (Mammuthus primigenius; e.g., Gladkih et al., 1984; Soffer, 1985; Hoffecker, 2002). The origin of the large accumulations of mammoth remains is still a matter of debate, with the main hypotheses being the collection of natural occurrences versus active hunting (e.g., Soffer, 1985; Haynes 1989; Svoboda et al., 2005). In favor of this second scenario, studies of the mammoth remains of Yudinovo (Germonpré et al., 2008) concluded that the mammoths were hunted and, at Mezhyrich, mammoths were obtained by combined procurement via collection of carcasses and active hunting (Péan, 2015). Hunting practices were observed in older sites of the Gravettian culture in the Dnieper and Desna valleys (Demay et al., 2016).
Between ca. 22 and 12 14C ka BP (ca. 26.2–13.8 ka cal BP), the Dnieper and Desna basins correspond to the southern part of the geographical distribution of the woolly mammoth (Markova et al., 2013; Kahlke, 2014). Over time their distribution shifted northwards, while the density of the mammoth population decreased (Markova et al., 2013). According to Markova et al. (2013), the combined effect of gradual warming and growing human pressure is most likely to have had a negative impact on the mammoths, resulting in their local extinction in the Russian and Ukrainian plains around 14–12 14C kaBP (ca. 17.0–13.8 ka cal BP; Stuart et al., 2005).
Discussion and conclusion
Humans could have taken advantage of the mammoth vulnerability, as reflected by lower δ15N values, to access animals that died naturally, collect bones, and hunt the most fragile individuals. This, with the possible assistance of domesticated dogs as hunting partners, could have countered a possible return to more suitable conditions for mammoth (Sablin and Khlopachev, 2002; Shipman, 2015). Our results confirm at least that mammoth specimens from Mezhyrich, Buzhanka 2, and, to a lesser extent, Eliseevichi were part of late mammoth populations surviving in sub-optimal conditions. They were thus most likely vulnerable to any pressure from environmental and/or human origin. Detecting further such cases among the late surviving mammoth populations using stable isotopic tracking may be a way to test if mammoth populations still had an optimal ecology or were metastable and, therefore, vulnerable to extinction. For instance, the insular Holocene population of mammoth on Saint Paul Island exhibited low and variable δ15N values, indicating suboptimal ecological conditions preceding their final disappearance (Graham et al., 2016).
The results of δ34S analyses showed no differences among mammoth according to the site but possibly a forage range partitioning between mammoth and coexisting large ungulates. Thus, variability in the mobility pattern for the mammoth between the high and low δ15N groups, such as migratory versus sedentary individuals, is not supported so far. We consider that rapid environmental modifications over time, probably not detectable through radiocarbon dating, can be a valid alternative explanation. Combined with direct competition with other large herbivores, such as the horse, and hunting of the most vulnerable individuals, the loss of their optimal habitat was likely to be the driving factor behind the local extinction of the mammoth in the central East European plains.
NOTE. I think one of the important changes in this version compared to the preprint is the addition of the recent Iberomaurusian samples.
Abstract (emphasis mine):
The extent to which prehistoric migrations of farmers influenced the genetic pool of western North Africans remains unclear. Archaeological evidence suggests that the Neolithization process may have happened through the adoption of innovations by local Epipaleolithic communities or by demic diffusion from the Eastern Mediterranean shores or Iberia. Here, we present an analysis of individuals’ genome sequences from Early and Late Neolithic sites in Morocco and from Early Neolithic individuals from southern Iberia. We show that Early Neolithic Moroccans (∼5,000 BCE) are similar to Later Stone Age individuals from the same region and possess an endemic element retained in present-day Maghrebi populations, confirming a long-term genetic continuity in the region. This scenario is consistent with Early Neolithic traditions in North Africa deriving from Epipaleolithic communities that adopted certain agricultural techniques from neighboring populations. Among Eurasian ancient populations, Early Neolithic Moroccans are distantly related to Levantine Natufian hunter-gatherers (∼9,000 BCE) and Pre-Pottery Neolithic farmers (∼6,500 BCE). Late Neolithic (∼3,000 BCE) Moroccans, in contrast, share an Iberian component, supporting theories of trans-Gibraltar gene flow and indicating that Neolithization of North Africa involved both the movement of ideas and people. Lastly, the southern Iberian Early Neolithic samples share the same genetic composition as the Cardial Mediterranean Neolithic culture that reached Iberia ∼5,500 BCE. The cultural and genetic similarities between Iberian and North African Neolithic traditions further reinforce the model of an Iberian migration into the Maghreb.
FST and outgroup-f3 distances indicate a high similarity between IAM and Taforalt. As observed for IAM, most Taforalt sample ancestry derives from Epipaleolithic populations from the Levant. However, van de Loosdrecht et al. (17) also reported that one third of Taforalt ancestry was of sub-Saharan African origin. To confirm whether IAM individuals show a sub-Saharan African component, we calculated f4(chimpanzee, African population; Natufian, IAM) in such a way that a positive result for f4 would indicate that IAM is composed both of Levantine and African ancestries. Consistent with the results observed for Taforalt, f4 values are significantly positive for West African populations, with the highest value observed for Gambian and Mandenka (Fig. 3 and SI Appendix, Supplementary Note 10). Together, these results indicate the presence of the same ancestral components in ∼15,000-y old and ∼7,000-y-old populations from Morocco, strongly suggesting a temporal continuity between Later Stone Age and Early Neolithic populations in the Maghreb. However, it is important to take into account that the number of ancient genomes available for comparison is still low and future sampling can provide further refinement in the evolutionary history of North Africa.
Genetic analyses have revealed that the population history of modern North Africans is quite complex (11). Based on our aDNA analysis, we identify an Early Neolithic Moroccan component that is (i) restricted to North Africa in present-day populations (11); (ii) the sole ancestry in IAM samples; and (iii) similar to the one observed in Later Stone Age samples from Morocco (17). We conclude that this component, distantly related to that of Epipaleolithic communities from the Levant, represents the autochthonous Maghrebi ancestry associated with Berber populations. Our data suggests that human populations were isolated in the Maghreb since Upper Paleolithic times. Our hypothesis is in agreement with archaeological research pointing to the first stage of the Neolithic expansion in Morocco as the result of a local population that adopted some technological innovations, such as pottery production or farming, from neighboring areas.
By 3,000 BCE, a continuity in the Neolithic spread brought Mediterranean-like ancestry to the Maghreb, most likely from Iberia. Other archaeological remains, such as African elephant ivory and ostrich eggs found in Iberian sites, confirm the existence of contacts and exchange networks through both sides of the Gibraltar strait at this time. Our analyses strongly support that at least some of the European ancestry observed today in North Africa is related to prehistoric migrations, and local Berber populations were already admixed with Europeans before the Roman conquest. Furthermore, additional European/ Iberian ancestry could have reached the Maghreb after KEB people; this scenario is supported by the presence of Iberian-like Bell-Beaker pottery in more recent stratigraphic layers of IAM and KEB caves. Future paleogenomic efforts in North Africa will further disentangle the complex history of migrations that forged the ancestry of the admixed populations we observe today.
Also, from the main author’s Twitter account:
I just realized that the paragraph with information on data availability is missing! Sequence data in the European Nucleotide Archive (PRJEB22699). Consensus mtDNA sequences are available at the National Center of Biotechnology Information (Accession Numbers MF991431-MF991448).
I find it hard to believe that this genetic continuity from Upper Palaeolithic to Late Neolithic could be representative of an autochthonous development of Afroasiatic. An important population movement – likely more than one – must be found in ancient DNA influencing North-Central and North-East Africa, probably during the time of the Green Sahara corridor.