Technological revolution in African Acheulean some 800,000 years ago

Well, maybe the title is a bit of a hype but something like that seems to be the most relevant finding on the Acheulean of Konso (SNPP region, Ethiopia): that the technique stood the same for a million years and then, some 800,000 years ago, became more refined in which was apparently one of the first technological leaps of archaic Humankind. Specifically it is the edges of the handaxes (the archetypal Acheulean finding, which may have been more a knife of sorts than a true axe) which became more refined and apt for its cutting purpose.

Yoyas Beyene et al., The characteristics and chronology of the earliest Acheulean at Konso, Ethiopia. PNAS 2013. Open access → LINK [doi:10.1073/pnas.1221285110]

Abstract

The Acheulean technological tradition, characterized by a large (>10 cm) flake-based component, represents a significant technological advance over the Oldowan. Although stone tool assemblages attributed to the Acheulean have been reported from as early as circa 1.6–1.75 Ma, the characteristics of these earliest occurrences and comparisons with later assemblages have not been reported in detail. Here, we provide a newly established chronometric calibration for the Acheulean assemblages of the Konso Formation, southern Ethiopia, which span the time period ∼1.75 to <1.0 Ma. The earliest Konso Acheulean is chronologically indistinguishable from the assemblage recently published as the world’s earliest with an age of ∼1.75 Ma at Kokiselei, west of Lake Turkana, Kenya. This Konso assemblage is characterized by a combination of large picks and crude bifaces/unifaces made predominantly on large flake blanks. An increase in the number of flake scars was observed within the Konso Formation handaxe assemblages through time, but this was less so with picks. The Konso evidence suggests that both picks and handaxes were essential components of the Acheulean from its initial stages and that the two probably differed in function. The temporal refinement seen, especially in the handaxe forms at Konso, implies enhanced function through time, perhaps in processing carcasses with long and stable cutting edges. The documentation of the earliest Acheulean at ∼1.75 Ma in both northern Kenya and southern Ethiopia suggests that behavioral novelties were being established in a regional scale at that time, paralleling the emergence of Homo erectus-like hominid morphology.

Fig. 4. Handaxe refinement through time. Upper, dorsal; Lower, ventral.
From left to right, two each are shown from KGA6-A1 (∼1.75 Ma), KGA4-A2 (∼1.6 Ma), KGA12-A1 (∼1.25 Ma), and KGA20 (∼0.85 Ma). In each pair of handaxes from the respective sites, near-unifacial (left) and more extensively bifacial (right) examples are shown (except with the KGA20 handaxes, which are both well worked bifacially).

I am a bit intrigued by the all-covering work style of the last handaxes, which remind somewhat to the later MSA technology, which belongs already to Homo sapiens. Our species may have also evolved in that very area of the Nile Basin, with the oldest specimen known being from nearby Omo River.

Of course that there are hundreds of thousands of years in between and of course that the peculiar orography of the Rift Valley is susceptible of offering archaeological findings from old much more easily than other areas but still...

Other sources: Pileta, NBC News.



Update: much more than just the edges but a whole technological paradigm change:

I was not really appreciating the whole extent of the technological revolution implicit in these changes. I just took note (reading too fast, too many things to do) of the edge refinement but Va_Highlander has correctly called my attention on that it was a much more ample and complex change in the whole technology of stone flaking and not just the edges, maybe even a whole jump in our mental capacities:

In contradistinction to the >1.2-Ma assemblages, the younger ∼0.85-Ma Konso Acheulean is characterized by considerably refined handaxes. Some of these handaxes are refined to the extent that they would qualify as approaching “three-dimensional symmetry” (i.e., symmetric not only in plan view but also in cross-section form) (Fig. 4 and Fig. S2). Some suggest that manufacturing 3D symmetric tools is possible only with advanced mental imaging capacities and that such tools might have emerged in association with advanced spatial and navigational cognition, perhaps related to an enhanced mode of hunting adaptation. It has been pointed out that purposeful thinning of large bifacial tools is technologically difficult, even in modern human ethnographic settings. In modern humans, acquisition and transmission of such skills occur within a complex social context that enables sustained motivation during long-term (>5 y) practice and learning.
 
In light of the above information, it is of interest that our metric analysis shows that there may be a fundamental difference between the handaxe technologies of >1.2 and ∼0.85 Ma. Whereas refinement of handaxe shape did occur from ∼1.6 to ∼1.2 Ma, this refinement did not result in tool thinning and advanced 3D symmetry.

Whale remains reinforce the notion of Magdalenian being linked to sea mammal predation

First it was the whale bone spear point of Isturitz (Basque Country), then the isotope evidence of sea mammal based diet of a Magdalenian individual from Kendric Cave (Wales) and now direct evidence of whale remains in the cave of Nerja (Andalusia). The evidence mounts up for a quasi-Inuit lifestyle of at least some people of the Magdalenian culture of late Upper Paleolithic Europe.

Esteban Álvarez Fernández et al., Occurrence of whale barnacles in Nerja Cave (Málaga, Southern Spain): indirect evidence of whale consumption by humans in the Upper Magdalenian. Quaternary International 2013. Pay per view → LINK [doi:10.1016/j.quaint.2013.01.014]

Abstract

A total of 167 plates of two whale barnacle species (Tubicinella majorLamarck, 1802 and Cetopirus complanatus (Mörch, 1853)) have been found in the Upper Magdalenian layers of Nerja Cave, Mina Chamber (Maro, Málaga, southern Spain). This is the first occurrence of these species in a prehistoric site. Both species are specific to the southern right whale Eubalena australis, today endemic in the Southern Hemisphere. Because of Antarctic sea-ice expansion during the Last Glacial Period, these whales could have migrated to the Northern Hemisphere, and reached southern Spain. Whale barnacles indicate that maritime-oriented forager human groups found stranded whales on the coast and, because of the size and weight of the large bones, transported only certain pieces (skin, blubber and meat) to the caves where they were consumed.

The barnacles

According to the authors, this is the first case of consumption of whale meat and blubber ever documented in Europe. 

The hearth where the remains were found is dated to c. 14,000 years ago. 

Previous evidence from this prolific Andalusian cave have previously informed of consumption of seafood and fish, along with rabbits and the occasional goat, a tradition that dates to Neanderthal times in that region. 

A perplexing curiosity is that one of the whale kinds identified is the southern right whale, which was not known to have lived so far north at all (its main habitat is the Antarctic seas with some extension towards Brazil and the Mozambique Strait). I wonder if it is a case of misidentification and the species is either the North Atlantic right whale or an extinct relative of both.

Sources: Materia[es], Pileta[es].


PS- And what was the blubber used for (besides eating)? Our friend David Sánchez coincidentally just published two successive and quite interesting articles (in Spanish) at his blog on the lamps of the Upper Paleolithic: 1st part, 2nd part.

A particularly beautiful lamp from Lascaux (Dordogne)

Update (Jan 29): another finding of whale consumption in Magdalenian contexts unknown to me until now (h/t David) is from Las Caldas (Asturias). One of the two co-researchers is the same as the lead author of the Nerja paper → direct PDF link.


Update (Feb 22): David again added more interesting information on the matter of possible whaling in the Magdalenian period by pointing us to Colchón Rodríguez & Álvarez Fernández 2008, where they discuss (in Spanish) the presence of sea mammal remains in the cave of Las Caldas (Asturias): a seal tooth (pierced as to be part of a necklace or similar decoration), a pilot whale tooth (only initially worked), a sperm whale tooth (fully sculpted into low reliefs of whale and bison) and also several whale and other sea mammal bones used for tool-making (they made spear points on whale bone, as was documented years ago for Isturitz in the same period) and some mollusks, notably the shell of a whale barnacle (Coronula diadema).

Las Caldas (locator map) is some 20 Km. inland nowadays, in the Magdalenian period maybe 30 Km. or so. The whale barnacle suggests that whale meat was moved all that distance from the coast.

Epipaleolithic finds near Oxford, England

One of the arrowheads found

Archaeologists have found the remains of people living at Didcot (Oxfordshire, England) some 9000 years ago. Previous research had found Neolithic materials almost only but now a clear layer of hundreds of flint tools and weaponry and hearth remains has been unearthed and dated. 

More details at the Archaeology News Network.

Tianyuan, mtDNA B and the formation of Far Eastern peoples

The genetic study of the ancient man of Tianyuan is already online, as I commented yesterday in a quick update.

Qiaomei Fu et al., DNA analysis of an early modern human from Tianyuan Cave, China. PNAS 2013. Open access → LINK [doi: 10.1073/pnas.1221359110]

Abstract

Hominins with morphology similar to present-day humans appear in the fossil record across Eurasia between 40,000 and 50,000 y ago. The genetic relationships between these early modern humans and present-day human populations have not been established. We have extracted DNA from a 40,000-y-old anatomically modern human from Tianyuan Cave outside Beijing, China. Using a highly scalable hybridization enrichment strategy, we determined the DNA sequences of the mitochondrial genome, the entire nonrepetitive portion of chromosome 21 (∼30 Mbp), and over 3,000 polymorphic sites across the nuclear genome of this individual. The nuclear DNA sequences determined from this early modern human reveal that the Tianyuan individual derived from a population that was ancestral to many present-day Asians and Native Americans but postdated the divergence of Asians from Europeans. They also show that this individual carried proportions of DNA variants derived from archaic humans similar to present-day people in mainland Asia.

Mitochondrial DNA

Part of fig. 1

And the old guy (or is it a woman?) happened to carry the matrilineage (mtDNA) B, more specifically B4'5, defined by a relatively long deleted block at positions 8281-8289 (this excludes B6 now linked with R11 and also the other relative of all them R24, see PhyloTree for details). However within B4'5 the lineage could not further be resolved within the modern haplogroups, so it is neither B4 nor B5 but a third branch of the same haplogroup. 

The authors actually talk of "haplogroup B" but they explicitly mention a deletion of a 9-bp motif (5′-CCCCCTCTA-3′, revised Cambridge reference sequence positions 8,281–8,289) as well as a substitution at position 16,189, what makes it unmistakable B4'5 per the current PhyloTree build.

The tree to the right illustrates this fact, placing Tianyuan man's lineage hanging directly from the root of this haplogroup that, beyond reasonable doubt, coalesced somewhere in East Asia (probably SE Asia, with Laos and Hainan being good references judging on diversity) some time before this person lived and died near what today is Beijing. 

What does it tell us? Really nothing new, at least within the parameters I have been managing: it confirms that the expansion of mtDNA B4'5 was already happening back in that time and that it had reached more or less its current area of expansion in East Asia (American and Oceanian B variants expanded later, of course). It also implies that its ancestors R and N, which experienced important successive expansions in the course of the colonization of Eurasia by our species had expanded  at an even earlier date (again nothing new to me but a nice confirmation anyhow).

On the other hand, this person's particular matrilineage went eventually extinct later on. This again does not tell us too much because it is something to expect with the course of time, especially at low population densities, as was the case in the Paleolithic. He can still be ancestral to modern peoples in the area and elsewhere but not by a purely mother-to-daughter line - at least not that we know. 

Chromosome 21 autosomal DNA

Because of the poor state of the DNA, the researchers had a difficult time sequencing it (technical details in the paper), however they managed to reconstruct a good deal of chromosome 21, which they used to compare with modern humans and also with Neanderthals and the so-called Denisovans. 

The result places Tianyuan closer to modern Far Eastern populations than to the rest of modern humans. This clearly indicates that the process of division in various more or less homogeneous subcontinental-sized populations was already somewhat advanced. 

Fig. 2. Maximum-likelihood tree relating the chromosome 21 sequences of the Tianyuan individual, 11 present-day humans, and the Denisovan genome. The most strongly supported gene-flow event is shown in yellow. Bootstrap support for all internal edges is 100% except for the edge putting Tianyuan outside the four Asians, which is 31%. The scale bar shows 10 times the average standard error of the entries in the covariance matrix.

While it is generally acknowledged that Papuans cluster at very deep level with East Asians, the authors are not fully persuaded of the exactitude of this tree, particularly in this aspect. They declare:

We note, however, that the relationship of the Tianyuan and Papuan individuals is not resolved (bootstrap support 31%). Further work is necessary to clarify whether this reflects the age of the Tianyuan individual relative to the divergence between modern human populations.

The caveat is particularly relevant because the colonization of New Guinea is at least as old as 49,000 years ago, some ten millennia before Tianyuan, what does not fit too well with the tree at that level of detail, assuming (as I do) that modern Papuans are direct unmixed descendants of those early settlers. Papuans do carry at high frequencies a related matrilineage (P also basal descendant from R) but that is also true of modern Europeans and they appear more distant in the tree above.

A good contrast to understand better the difficulties in getting a good picture from autosomal DNA, especially one so old, is table 1:

Here we can appreciate the differences and proximities by another measure. The closest compared modern person to Tianyuan man is a Karitiana, followed closely by the Han and, surprisingly, by the Sardinian and the French, and only then the Dai and Papuan. 

The distance of the Karitiana to Tianyuan man is still greater than that with not just the Han or the Dai but also the Europeans. However this can be argued to be because Native Americans must have a deep dual East Asian and West Eurasian origin, the latter via Altai (Y-DNA Q, mtDNA X2).

Let's check the Han then, who are not believed to have any meaningful West Eurasian admixture. Curiously the paradox happens again: the Han is somewhat closer to Europeans by this measure than to Tianyuan, and even their comparison with the Papuan shows up slightly less differentiated. 

This is admittedly harder to explain but we can conclude that either (a) this method can only grasp affinity/divergence to some degree or (b) that the Tianyuan partial genome indicates a very preliminary level of continental differentiation. Or (c) both. Of course time is the main cause of genetic differentiation and by no means we can imagine that such an ancient individual would be too similar to his modern plausible descendants but, on the other hand, all (including Tianyuan mtDNA) indicates that the process of continental differentiation was already well developed 40,000 years ago (most European ancestry must come from people living in Europe or West Asia back then) so we can either blame subtle flows like Siberian migrations that have kept both genetic pools somewhat closer than in pure isolation or we must assume that the measure is not too exact.

Admixture with other human species

The paper also deals with Denisovan and Neanderthal admixture, finding that Tianyuan man was within the modern range for both parameters in East Asia. 

This is very important because it ratifies the mainstream model of two minor admixture episodes: (1) with Neanderthals at the exit from Africa and prior to the Great Eurasian Expansion (so all non-Africans, including Tianyuan man, have very similar levels of Neanderthal admixture today) and (2) with a relative of Denisovans (Homo erectus?) maybe in Indonesia affecting only (or almost only) the aboriginal peoples of Oceania (and Filipino Negritos but not the other so-called Negritos from Malaysia or the Andaman, who are not particularly related anyhow).

(As a side note notice that the Denisovan-like gene flow into Papuans in fig. 2 appears to hang not from the end of the branch but from a very high position, suggesting it was a relative and not the known Denisovans of Altai themselves who became admixed into Papuans and other Oceanian populations, probably a relative living in the route to Australasia).


Update: Marnie just published a mention of a previous work on Tianyuan 1, which focuses on the isotopic evidence for a fish-based diet. 

Ancient North Chinese from 40,000 years ago closely related to modern locals

The information is sketchy as of now but the news in the press indicate that , near Beijing, whose fragmented remains were discovered in 2003, was closely related to modern East Asians and Native Americans.  

The paper is not yet online but the information released to the media strongly suggests that East Asians were already distinct from other populations some 40,000 years ago. This would seem to be based on the sequencing of the mitochondrial DNA and the explicit mention of Native Americans indicates that the lineage must be A, B, C or D (X, the fifth and less common matrilineage of Native Americans, is not found in East Asians, with some exceptions from Siberia, so we can exclude it safely). 

Ancient DNA from cell nuclei and maternally inherited mitochondria indicates that this individual belonged to a population that eventually gave rise to many present-day Asians and Native Americans, says a team led by Qiaomei Fu and Svante Pääbo, evolutionary geneticists at the Max Planck Institute for Evolutionary Anthropology in Leipzig, Germany. 

This would seem to discard some adventurous hypothesis floating around about tremendous demographic changes in the Paleolithic and afterwards, at least for this region. Probably not even when "mode 4" technology arrived to the region (from Altai) c. 30,000 years ago. 

In other words: the seeds of modern populations were already there c. 40,000 years ago in East Asia (and surely also in most other regions) and, even if they may have changed somewhat, they have remained the same at least to some notable degree.

Furthermore, the autosomal DNA also seems to have been sequenced to at least some degree because the researchers state that Denisovan and Neanderthal genetic inputs are at the same levels as modern North Chinese (i.e. some 0% and 2.5% respectively):

The partial skeleton, unearthed in Tianyuan Cave near Beijing in 2003, carries roughly the same small proportions of Neandertal and Denisovan genes as living Asians do (SN: 8/25/12, p. 22), the scientists report online January 21 in the Proceedings of the National Academy of Sciences.

Or in the words of the Max Plank Institute:

The genetic profile reveals that this early modern human was related to the ancestors of many present-day Asians and Native Americans but had already diverged genetically from the ancestors of present-day Europeans. In addition, the Tianyuan individual did not carry a larger proportion of Neanderthal or Denisovan DNA than present-day people in the region.

This also seems to discard models implying Denisovan admixture happening in Siberia or NE Asia and would indirectly support my own hypothesis of admixture with Homo erectus (for which Denisovans, plausibly an Erectus-Neanderthal hybrid, would be just a proxy) in or near Indonesia.

Sources: Science News, Max Plank Institute.


Update (Jan 22): the paper is already online and is open access (cool!)  I don't think I have time to discuss it today but will do tomorrow without doubt (other than the sky falls on my head, you know).

Y-DNA of Moldovans

Moldova is the easternmost generally recognized sovereign state in Europe speaking a Romance language. Achieving its independence from the Soviet Union in 1991, Moldova saw part of its territory segregated in the mostly unrecognized Republic of Transistria (multiethnic, under military control of Russia). There was some talk about joining the related Republic of Romania but this plan seems to have been abandoned for now.

Alexander Varzari et al., Paleo-Balkan and Slavic Contributions to the Genetic Pool of Moldavians: Insights from the Y Chromosome. PLoS ONE 2013. Open access → LINK [doi:10.1371/journal.pone.0053731]

Abstract

Moldova has a rich historical and cultural heritage, which may be reflected in the current genetic makeup of its population. To date, no comprehensive studies exist about the population genetic structure of modern Moldavians. To bridge this gap with respect to paternal lineages, we analyzed 37 binary and 17 multiallelic (STRs) polymorphisms on the non-recombining portion of the Y chromosome in 125 Moldavian males. In addition, 53 Ukrainians from eastern Moldova and 54 Romanians from the neighboring eastern Romania were typed using the same set of markers. In Moldavians, 19 Y chromosome haplogroups were identified, the most common being I-M423 (20.8%), R-M17* (17.6%), R-M458 (12.8%), E-v13 (8.8%), R-M269* and R-M412* (both 7.2%). In Romanians, 14 haplogroups were found including I-M423 (40.7%), R-M17* (16.7%), R-M405 (7.4%), E-v13 and R-M412* (both 5.6%). In Ukrainians, 13 haplogroups were identified including R-M17 (34.0%), I-M423 (20.8%), R-M269* (9.4%), N-M178, R-M458 and R-M73 (each 5.7%). Our results show that a significant majority of the Moldavian paternal gene pool belongs to eastern/central European and Balkan/eastern Mediterranean Y lineages. Phylogenetic and AMOVA analyses based on Y-STR loci also revealed that Moldavians are close to both eastern/central European and Balkan-Carpathian populations. The data correlate well with historical accounts and geographical location of the region and thus allow to hypothesize that extant Moldavian paternal genetic lineages arose from extensive recent admixture between genetically autochthonous populations of the Balkan-Carpathian zone and neighboring Slavic groups.

Most interesting is without doubt the list of haplogroups:

Table 2 - Kharahasani is located to the South and Sofia to the North, the Romanian and Ukranian samples are both from nearby regions (Romanian Moldavia and Transistrian Ukranians).

My notes (see ISOGG for nomenclature):

• The high diversity of haplogroup I (also in nearby Romania and Ukraine), including I1-M253, I2a1b-M423 and I2a2-M223 is consistent with the wider region being, arguably, ancestral to this lineage. However "Low Germanic" I2b does not show up, as doesn't "West Mediterranean" I2a1a nor Anatolian-Caucasian I2a*.
• Among Neolithic-specific inputs, which are particularly important in the Balcans, Moldovans show notable (13%) presence of E1b1b1a1-M78 variants, especially the well studied E1b1b1a1b-V13, related now even by ancient DNA to European Neolithic flows. They also have some (2%) E1b1b1b2a-M123, an Eastern and NE African lineage found at low frequencies in Southern Europe.
• Another clearly Neolithic lineage is G2a-P15, found among Moldovans only at very low frequencies (more common in the context of the Mediterranean Neolithic it seems).
• Not yet documented by aDNA but also likely Neolithic in Europe is haplogroup J, found in Europe mostly as J2 (originating in Highland West Asia) but also, mostly in the Balcans, as J1 (originating maybe in Palestine). Moldovans show both at low frequencies (4% each).
• Almost all the rest belongs to the largest European clade, R, mostly its Eastern variant R1a1a-M17 (30%). Western R1b1a2-M269 makes up 16%. 
• Minor clades are H (Romani), T (South and West Asian, with extensions into East Africa and, thinly, in Europe), N (NE European and North Asian) and Q (probably from West Asia).

In the PC analysis (fig. 2, not shown) Moldovans appear intermediate between Balcanic and Central-Eastern European populations but rather leaning towards the latter.

In spite of their historical and ethno-linguistic connection Romanians and Moldavians do not appear to be particularly related in the genetic aspect:

The genetic relationship between Moldavians and Romanians deserves special attention, since these two groups speak practically the same language and share many cultural features. It is reasonable to assume that Moldavians and Romanians inherited genetic lineages, shared with other Balkan populations, from Vlachs who, in turn, received them from Paleo-Balkan tribes. However, Moldavians and Romanians do not form a cluster that would have separated them from the neighboring populations. Indeed, in the space of multi-dimensional scaling based on the R_ST distances between STR haplotypes, Romanian populations appeared scattered among the Balkan populations and did not cluster with the Moldavians (Figure 3). According to the AMOVA analysis, the degree of within-group differentiation among Moldavian and Romanian populations was significantly greater than genetic differences between either Romanians or Moldavians and the group comprised of the Balkan populations (Table 3). Moldavians and Romanians also appear dissimilar on the diagram of binary lineages (PC plot, Figure 2). Thus, sharing nearly the same language is not accompanied by specific genetic similarity between Moldavians and Romanians. Furthermore, Italian populations that share the Romance/Latin language with Moldavians and Romanians, show little genetic similarity with them. These results agree with previous genetic studies suggesting that the genetic landscape of southeast Europe had been formed long before the modern linguistic/ethnic landscape was shaped [16], [48].

Instead the genetic affinities of Moldovans lean strongly towards their Slavic neighbors from Eastern and Central Europe:

In contrast to Romanians and most other Balkan populations, Moldavians show a clear genetic similarity to western and eastern Slavs. This is strongly implied by haplogroup R-M17, which dominates the paternal lineages of the Slavs and is broadly represented in Moldavians. (...)
The noteworthy domination of R-M17 chromosomes in Moldavians compared to Romanians is due to the R-M458 subclade. Haplogroup R-M458 likely has its roots in western/northern Poland, where it has its greatest modern concentration and microsatellite diversity [49].

This supports my impression of R1a1a1b1a1-M458 being not spread by Slavic migrations (it is very rare among Balcanic Slavs but has an notable presence in Greek Macedonia instead) but much earlier, plausibly by Indoeuropean migrations which had a major sub-center in Poland in the Chalcolithic period.

Mitochondrial DNA of some Slavic peoples

With emphasis on haplogroup H5 and H6.

Marta Mielnik-Sikorska et al., The History of Slavs Inferred from Complete Mitochondrial Genome Sequences. PLoS ONE 2013. Open access → LINK [doi:10.1371/journal.pone.0054360]

Most interesting is maybe table 1 (right), which lists two Polish populations (Kashubia, at the Baltic coast, and Podhale: the Carpathian piedmont), Ukrainians and Czechs.

We can see here that the most common lineages among these Slavs are not different from other European populations, namely H*, H1, U5a, U5b and also the, arguably Neolithic, lineages J1 and T2. I find relevant in this sense that there is a significant amount of T(xT1,T2) among Kashubian Polish especially.

Another point of interest is the minor presence of North and Central Asian lineages A, C, D and G, for which the authors present an elaborate rationale:

... we were able to pinpoint some lineages which could possibly reflect the relatively recent contacts of Slavs with nomadic Altaic peoples (C4a1a, G2a, D5a2a1a1).

They also suggest that the L2a1l2a, found among the Polish, is of Ashkenazi Jewish origin. L1b1a8 found in Polish and Russians belongs to the wider L1b1a, recently argued to be European-specific.

Another point of notice may be the rare HV0(xV) found at significant frequencies among Ukranians (4.5%).

But the authors make a particular effort to discern within haplogroups H5 and H6, which they find of particular interest. H5 might be (with doubts) of Italian origin and they consider its coalescence age (on the dubious molecular clock estimate methods) as clearly pre-Neolithic.

Based on these speculative methods they argue that several Slavic-specific clades within H5 may be contemporary in origin with U4a2, common in Central and Eastern Europe. They consider both to be roughly from Early Neolithic times.

Figure 1. Complete mtDNA phylogenetic tree of haplogroup H5.

Legend:

→ Green: Polish, Czech, Slovaks, Ukrainians and Russians
→ Red: German, Dutch and Austrians

→ Yellow: Italians and Spaniards

→ Blue: Irish, British, Danes and Finns

→ Magenta: Tunisia

→ Black: Levant

→ Grey: USA

→ White: unknown geography

If they are correct in their interpretation of the tempo of H5, the hypothesis of H sublineages migrating Northwards, from Southern to Central Europe, within Neolithic would seem to gain some support.

However H5 is less common in the Czech Republic and Austria than in Poland or Ukraine and the Neolithic colonization of Poland should have gone via the Czech Republic and, previously, Austria. Of course we cannot reject upfront a founder effect specific to Poland but what about Ukraine, which was almost totally oblivious to the Balcano-Danubian Neolithic phenomenon?

The other focus is H6, which is found almost only among Ukranian and Czechs of the four target populations. Generally speaking H6a and the most rare H6c are European, while H6b is Central and West Asian. In spite of its extreme rarity, the authors detected H6c in three individuals (one Czech, one Pole and one Slovak), all non-Slavic H6c are from Central-West or NW Europe (or from unknown locations). This seems to define H6c as a rare Northern-Central European haplogroup (excluding Eastern Europe apparently).

The remaining H6 samples sequenced in our study belong to different H6a subclusters being identified as singletons (H6a1a*) or as members of subclusters H6a1a4, H6a1a9 and H6a1b3. Subcluster H6a1a9 is novel, comprising of two haplotypes found in Russians and Ukrainians. Subcluster H6a1b3 is also interesting because it contains, except for European individuals of unknown origin, a founder haplotype of Czech origin and two Polish haplotypes.

Figure 2. Complete mtDNA phylogenetic tree of haplogroup H6 (legend as above).