Archive for December, 2010

The European Mesolithisation of a Caucasian Neolithic, or the Origin of the Indo European Language family

December 28, 2010 7 comments

Without any doubt the transition from “Mesolithic” hunting and gathering to a “Neolithic” agricultural way of life was a demographic event of utmost importance, but to what extend? This year (2010) the debate is on again about the Neolithic advance in Europe. New paleogenetic results attested “Caucasian” patrilineal YDNA G2a3 and matrilineal mtDNA N1a in the Neolithic LBK culture north of the Alps, and more mtDNA N1a in Megalithic France. Along the male lineage this particular type of Neolithic YDNA survived with moderate frequencies until today, but the demise of the typical Neolithic matrilineal counterpart was almost complete. Even if we take into consideration an appreciable influence of post-Neolithic selective processes, we can’t ignore a certain sex-biased discrepancy in Neolithic survival rates. Possibly local “European” contributions to the gene-pool may be correlated to “Mesolithisation” processes essentially congruent to Zvelebil’s Neolithic Creolisation Hypothesis, supplying a potential candidate solution to the long standing issue of the Indo European origin.

Genetic distance maps presented by Haak et al. (2010) that show affinities between modern populations and Neolithic LBK samples. Peak values in the Caucasian region are notable and indicative of a Caucasian origin.

Apparently intrusive Neolithic cultures spread from the Near East to Europe along at least two fronts. Despite marked differences in their development and assimilation of local cultures, both groupings were subject to similar processes:

Archaeologically, two main cultural traditions, marked by two different potteries, can be distinguished in the Early Neolithic: the linear pottery culture (or LBK) that runs along the Danubian route and the impressed ware pottery (also called cardial) that spreads along the Mediterranean. This is not just a question on ceramic decoration. The diffusion of the new economy took two main routes after the colonizations of the Balkans that implied different necessities of adaptation of the agriculture and the farming to specific climatic and ecological conditions.(Sampietro et al.,2007)

The Cardial culture advanced west hopping islands and Mediterranean shores, while the Linearbandkeramik Culture (LBK) first developed along the Danube in the Balkans out of other Neolithic cultures before bursting into the North European Plain. The Neolithic stock at both Neolithic fronts could have been local in case Neolithisation was nothing but a cultural process, or otherwise – through a process of “demic diffusion” – closely related to the Anatolian or Levantine people whose ancestors “invented” this new way of life (Ex Oriente Lux!).
The genetic contribution of each of these groupings to the modern Europe population is still a matter of scientific debate. Purported Neolithic intruders must have come in close contact with native cultures wherever they passed, or otherwise acculturated Neolithic populations could have been continuous to at least some of these native cultures.

Two opposing scenarios have been invoked to account for the spread of agriculture in Europe. The demic diffusion (DD) model assumes that the Neolithic transition diffused in Europe from the Middle East by an important movement of population (Ammerman & Cavalli-Sforza 1984; pp. 78–80), without substantial contact with local Palaeolithic populations. On the contrary, the cultural diffusion (CD) model assumes that the Neolithic transition occurred mainly through the transmission of agricultural techniques (Zvelebil & Zvelebil 1988) without large movements of populations. (Currat et al.”, 2005)

Especially the Neolithic “Megalithic” cultures along the Atlantic, according to archeologists, are hard to identify with external Neolithic influences in the area:

The transition to the Neolithic in Atlantic Europe can be viewed as a relatively late phenomenon, with several interesting particularities. Among those, we point out the fundamentally indigenous character of the processes; the existence of a long availability phase, in which hunter-gatherer groups maintained contact with neighboring agriculturalists and probably were familiar with farming and animal husbandry without applying them in a systematic way; and the later development of megalithic monumental funerary architecture.(Pablo Arias,1999)

Cardial culture spread rapidly west along the Mediterranean and reached as far as Portugal about 5400 cal BC. Paleogenetic investigation of mitochondrial DNA (mtDNA, that inherit mother to daughter) revealed absence of ‘LEvantine’ mtDNA haplogroup J in those Neolithic population samples, suggesting that upon arrival the Neolithic farmers didn’t descend anymore from the Levant on the matrilineal side.

The Portuguese Neolithic sample, containing no J haplotypes in 23 samples, indicates that agriculture in Portugal was not brought directly by migrating farmers from the Near East. (Chandler et al., 2005)

At higher genetic resolution these investigated Portuguese Mesolithic and Neolithic groups still show genetic discontinuity, despite sharing mitochondrial mtDNA H, U* and U5, thus also implying important population shifts at the Neolithic transition. Possibly the Neolithic farmers draw from just a subset of local, or nearby Mesolithic mtDNA.

A discontinuity at the Neolithic transition is consistent with the Maritime Pioneer Colonisation model for the arrival of farming in Portugal (Zilhão 1993, Zilhão 2001). In this model, agricultural enclaves were formed by groups of leap-frogging sea-faring colonists who moved around the Mediterranean coast. The source population however is not Near Eastern, as demonstrated both by the absence of haplogroup J in the Portuguese Neolithic population and by the genetic distance observed between the Neolithic Portuguese and Near Eastern populations. More likely, a Mediterranean group which itself had adopted farming through exchange or only limited migration moved into the uninhabited parts of Portugal’s coastal regions to pursue an agricultural subsistence strategy.(Chandler et al., 2005)

Relative frequencies of Mesolithic and Neolithic mtDNA in Portugal, compared to current frequencies in various modern regions (Chandler et al., 2005)

Thus, in Portugal local Mesolithic influences appear to underpin the Neolithic transition on the matrilineal side. Not even subsequent continuity up to modern times poses any problem:

Haplogroup frequencies and genetic distances show that the ancient Portuguese populations studied here, both Mesolithic and Neolithic, are most closely related to the modern Basque and Galician populations of the Iberian Peninsula.(Chandler et al., 2005)

A subsequent investigation in Eastern Iberia confirmed a strong Neolithic continuity here with modern Europeans (“New” Iberian groups printed bold):

The site ‘Camí de Can Grau’ (Granollers, Barcelona, Spain) is a necropolis excavated in 1994, which comprised 23 tombs dated by C14 between 3500 and 3000cal years BC.
[…] 11 sequences were considered to be endogenous and included in the posterior population analysis.
[…] The general [mtDNA] haplogroup composition of the Neolithic sample is: H (36.4%); T2 (18.2%); J1c (18.2%); I1 (9.1%); U4 (9.1%); and W1 (9.1%)
[…]the general composition is not significantly different from that obtained from the current Iberian Peninsula dataset when random resamplings of 11 sequences are made (Sampietro et al.,2007)

Here Near Eastern haplotypes must have trickled in up to a degree of mixture that can already be considered “modern”. However, note this results are substantially younger than the Portuguese finds mentioned above. Whatever happened after the first Neolithic arrivals that changed this Western Mediterranean genetic composition, this must have happened mainly before “Camí de Can Grau”.

Unfortunately, the nature and demic impact of the Neolithic advance north of the Alps proved much more difficult to retrieve and the discussion remains inconclusive:

Archaeological cultures such as the Linear pottery culture (Linearbandkeramik or LBK) and [AVK, the Eastern Linear Pottery Culture] mark the onset of farming in temperate regions of Europe 7500 years ago. These early farming cultures originated in Hungary and Slovakia, and the LBK then spread rapidly as far as the Paris Basin and the Ukraine. The remarkable speed of the LBK expansion within a period of about 500 years, and the general uniformity of this archaeological unit across a territory of nearly a million square kilometers, might indicate that the spread was fueled to a considerable degree by a migration of people (Haak et al., 2005)

The ultimate origin of this Neolithic people was assumed to be Anatolia and recently Myres et al. (2010) published a quite credible reconstruction of their advance based on the genetic structure of the very prolific Eurasian YDNA marker Hg R1b1b2. However, other preliminary assessments led to conflicting results: […]with estimates of Neolithic input into the present population ranging from 20 to 100%. (Haak et al., 2005). A theoretical simulation study by Currat and Excoffier even suggested a minor contribution:

The proportion of Europeans who are descendant from the first farmers from the Levant decreases very quickly with distance from the Neolithic source, as the lineages of Neolithic origin are rapidly diluted along the axis of colonization. Under our simulation conditions, an average local Palaeolithic contribution larger than 0.375% will indeed be enough to prevent Neolithic lineages to diffuse over the whole Europe.
These results imply that, under our model of a progressive range expansion of Neolithic farmers with possible genetic exchange and competition with local Palaeolithic hunter–gatherers, it is very unlikely that the Palaeolithic contribution be globally smaller than 50%. (Currat et al., 2005

Haak et al. were the first to actually verify this. In 2005 they investigated paleogenetic mtDNA of Neolithic samples. From a total of 57 investigated human remains of the prehistoric LBK/AVK cultures, the mtDNA of only 24 individuals could be determined:

Eighteen of the sequences belonged to typical western Eurasian mtDNA branches; there were seven H or V sequences, five T sequences, four K sequences, one J sequence, and one U3 sequence. These 18 sequences are common and widespread in modern Europeans, Near Easterners, and Central Asians (Haak et al., 2005)

Those 18 samples were considered too generic to reveal either a native or a foreign origin. Some may be suggestive of a genetic link to the Near East, that however could well exceed the Neolithic timeframe. Therefore the investigation concentrated on the mtDNA types identified in the other six individuals:

The most striking result is that 6 of the 24 Neolithic skeletons are of the distinctive and rare N1a branch.(Haak et al., 2005)

It could be shown this N1a lineage was universal to both LBK and AVK, but quite rare in modern European populations. A tentative link of the AVK sample to the Central Asian type of N1a is ambiguous. All six LBK/AVK samples can be grouped as pertaining to the “European type” – that despite the name occurs slightly more often in the Near East. Initially this was considered hot evidence against a significant genetic input of Neolithic immigrants in Europe:

The results from the Neolithic sample show that other mtDNA lineages considerably diluted the mtDNA pool of these early Neolithic populations, so that the frequency of N1a in modern Europeans is 150 times lower than in our sample of the first Central European farmers. This is incompatible with the idea that modern Central Europeans—and by implication other Europeans beyond the LBK/AVK area—derive their maternal lineages purely from the earliest farmers of that region. (Haak et al., 2005)

However, this ambiguous picture of Neolithic discontinuity remained confined to Central Europe, as the findings of Haak et al. sharply contrasted with the situation in the Iberian Peninsula that attested a more gradual development towards modern values.

Subsequent investigations on the Y chromosome restored confidence again in an overwhelming Neolithic contribution, albeit along the male lineages. The method heavily depended on the specific Neolithic identity of the European version of YDNA marker Hg R1b, and didn’t consider any possible selective forces on genetic level, but the method was solid, based on the mathematical observation that frequencies of new mutations in an expanding population (congruent to the Neolithic advance) tend to show a wave pattern. Even more noticeable distribution and frequency effects, ie. Allele frequency clines (AFC), were already predicted if a new mutation could manage to surf on the expanding wave front:

The AFCs can be generated by a succession of founder effects along the axis of diffusion of an expansion wave (Barbujani et al. 1995; Fix 1997; Austerlitz et al. 2000).
Our simulations suggest that AFC from the Middle East to north western Europe can be generated equally well by the Neolithic expansion process that occurred 8000 to 3000 BC or by the expansion of the first modern human in Europe ~45000 to 30000 BP. (Currat et al., 2005)

An important milestone was the identification of Europe’s main Y-DNA marker, haplogroup R1b, as “recent” and Neolithic:

Haplogroup R1b frequency in Europe is clinal with increasing frequencies observed in Northwest Europe, a pattern that has been ascribed to the persistance of Palaeolithic Y chromosomes in Europe after a Neolithic demic diffusion from the Near East. Interestingly, attempts to date the Y-STR-based diversity of R1b-M269 chromosomes in populations from Europe and Turkey have yielded Holocene expansion times in both regions. These findings have led to the reappraisal that R1b-M269 in Europe is young and likely associated with a Neolithic demic expansion from the Near East through Anatolia. (Myres et al., 2005)

I dedicated my last blog entry to this latter investigation, and explained how the LBK culture is now credited for being the European intermediary of this Y-DNA haplogroup R1b, that their Anatolian connections then must have brought in from the east. However, the small time window between Neolithic pioneers and Mesolithic populations that may have entered Europe slightly earlier, combined with the success of closely related R1b clades and other Hg R haplogroups that are unrelated to LBK boundaries, suggested more complexity in the events before and after the Neolithic advance.

Thus, invoking the pronounced transformation of the pre-Neolithic European gene pool by intrusive pioneer farmers from the Near East must be viewed cautiously especially when such an argument is based on just a single incompletely resolved haplogroup. Although the transition to agriculture was a pivotal event in human history, the spread of specific haplogroups can occur in more than one migration event. (Myres et al., 2005)

The Neolithic reconstruction became badly in need of a much more benign paleogenetic verification on genetic level. Haak et al. extended their previous investigation of Neolithic DNA, and conceded to less extreme differences regarding the maternal mtDNA composition of LBK compared to current populations, even though their attributed DNA still appears to be pretty unique:

Most importantly, PC correlates of the second component showed that elevated or high frequencies of hgs T, N1a, K, and W were unique to LBK populations, making them appear different from both Europe and Near East. The considerable within-hg diversity of all four of these hgs (especially T and N1a; Table 1) suggests that this observation is unlikely to be an artifact of random genetic drift leading to elevated frequencies in small, isolated populations. (Haak et al., 2010)

However, the previous stance that LBK is genetically extinct has now been considerably nuanced, apparently even abandoned. Important post-Neolithic events are still suggested, but there is no further denial that paleogenetic LBK survived somehow in both European and Near Eastern populations:

[The LBK dataset was] grouping with Europeans because of a lack of mitochondrial African hgs (L and M1) and preHV, and elevated frequencies of hg V. In contrast, low frequencies of hg H and higher frequencies for HV, J, and U3 promoted Near Eastern resemblances. (Haak et al., 2010)

The widely divergent results of LBK samples compared to current populations possibly found a different explanation:

The pooled European and Near Eastern meta-populations are necessarily overgeneralizations, and there are likely to be subsets of Near Eastern populations that are more similar to the Neolithic population. Interestingly, both the PCA [i.e. based on mtDNA haplogroup frequencies] and the MDS plots [Multidimensional scaling plot of genetic distances based on haplogroup frequencies] identified Georgians, Ossetians, and Armenians as candidate populations (Figures 2 and S1). (Haak et al., 2010)

Still, extinction and survival of LBK related genes appear to have gone hand in hand. This is true for mitochondrial DNA, where the correlation of some unique haplotypes with modern populations still poses a problem, but that for now can’t be correlated to contemporary Mesolithic populations either:

The frequency of N1a was 13.6% for Derenburg samples (3/22) and 14.3% for all LBK samples published to date (6/42). Notably, N1a has not yet been observed in the neighboring hunter–gatherer populations of Central Europe before, during, or after the Early Neolithic nor in the early Neolithic Cardial Ware Culture from Spain. (Haak et al., 2010)

There is a vague indication those mitochondrial haplogroups indeed could derive from a subset of Near Eastern populations:

The only relatively close neighbor of haplotype 16319-16343 is found in Iraq (16129-16189-16319-16343), in agreement with the Near Eastern affinities of the informative LBK haplotypes. (Haak et al., 2010)

Recent investigation of Megalithic mtDNA suggests this haplogroup apparently arrived as far as western France, thus locally congruent with the expansion of post-LBK cultures as suggested by the wave front spread of YDNA R1b according to Myres et al. (2010). From an archeological point of view such evidence was always so difficult to discern:

To extend existing knowledge of the mitochondrial European Neolithic gene pool, we examined six samples of human skeletal material from a French megalithic long mound (c.4200 cal BC). We retrieved HVR-I sequences from three individuals and demonstrated that in the Neolithic period the mtDNA haplogroup N1a, previously only known in central Europe, was as widely distributed as western France. Alternative scenarios are discussed in seeking to explain this result, including Mesolithic ancestry, Neolithic demic diffusion, and long-distance matrimonial exchanges. In light of the limited Neolithic ancient DNA (aDNA) data currently available, we observe that all three scenarios appear equally consistent with paleogenetic and archaeological data. In consequence, we advocate caution in interpreting aDNA in the context of the Neolithic transition in Europe. Nevertheless, our results strengthen conclusions demonstrating genetic discontinuity between modern and ancient Europeans whether through migration, demographic or selection processes, or social practices. (Deguilloux et al., 2010)

Matrilineally, this particular component of LBK diluted in the European genepool, and not only there. Even the Near Eastern origin of N1a became tenuous because of low frequencies, but the virtual extintion of N1a would be even more difficult to explain if it were also firmly rooted in the European Mesolithic. Tentatively, its virtual extinction appears to post-date the Late Neolithic wave of advance, and the resulting discrepancies with modern European mtDNA suggests important post-Neolithic matrilineal population shifts, not unlike the processes already specifically associated with the 3rd millenium advance of Beaker cultures:

The distribution of Bell Beakers could thus reflect the movement of marriage partners. (Marc vander Linden, 2007)

The post-LBK decrease of Near Eastern mtDNA components in Europe could have been less dramatic for mtDNA T – if indeed this type could be confirmed at all as fully oriental rather than Central/East European additions.
New paleogenetic information that concerns patrilineal DNA (YDNA) tends to confirm this picture of post-Neolithic decline.

Y chromosome SNPs could be typed for only three out of the eight male individuals (37.5%; Table S2)
[…] individual deb34 was found to belong to hg G (M201)
[…] downstream SNP S126 (L30), placing it into G2a3.
[…] whereas individuals deb20 and deb38 both fall basally on the F branch (derived for M89 but ancestral for markers M201, M170, M304, and M9)
[…] to distinguish between F and H
[…] deb20 and deb38 were shown to be ancestral at M69 and hence basal F (M89), and remained in this position because we did not carry out further internal subtyping within the F clade. (Haak et al., 2010)

Like mtDNA N1a, those two LBK YDNA F* samples could have pertained either to a European grouping that is currently very rare, maybe even merely theoretical (cq. extinct hg IJ*), or otherwise to an equally rare Near Eastern grouping. The third sample, however, survived in appreciable frequencies over a wide Eurasian territory: ‘Caucasian’ Hg G2a3. Since a shared YDNA history with that other F* samples is irreconcilable with the latter’s virtual extinction, both at a hypothetic Anatolian source and at its Central European destination, this F* is most likely to a pre-Neolithic addition to the LBK gene-pool. In this sense the ‘asymmetric’ survival of YDNA Hg G2a3 compared to F* could possibly compare with the survival of mtDNA T2 compared to mtDNA N1a.
Equally noteworthy is the absence of more common modern YDNA:

Interestingly, we do not find the most common Y chromosome hgs in modern Europe (e.g., R1b, R1a, I, and E1b1), which parallels the low frequency of the very common modern European mtDNA hg H (now at 20%–50% across Western Eurasia) in the Neolithic samples. (Haak et al., 2010)

Note the author doesn’t even dare to mention Hg J2, that before was the example ‘par excellence’ of Neolithic YDNA from Anatolia. Is it really? But for sure the absence of all these haplogroups in LBK can’t be taken for granted altogether, even though Corded Ware Y-DNA samples found in Eulau apparently indicated a rather Late Neolithic (post-LBK?) association for Hg R1a:

The few published ancient Y chromosome results from Central Europe come from late Neolithic sites and were exclusively hg R1a [31]. While speculative, we suggest this supports the idea that R1a may have spread with late Neolithic cultures from the east. (Haak et al., 2010)

No Late Neolithic arrivals from the east are known by archeology other than those already associated with European cultures cq. LBK, or at least comparable Balkanic cultures rooted in a Neolithic that is slightly older. How all these explicit and implicit claims of Near Eastern cq. Caucasian haplogroups, all having different occurrences and distributions throughout Europe and elsewhere, could ever be possibly accomodated within a single and progressive Neolithic wave of advance? Even the distribution of Hg G2a3 appears erratic compared to the tidy Wave of Advance model for Hg R1b1b2 claimed by Myres et al. (2010).

Let me explain briefly how these very different scenarios for the distribution of a wider array of Caucasian haplogroups from a single source are indeed possible, before I move on to transgress about the unlikelihood of ALL haplogroups being unequivocally Caucasian, and their possible sex-biased local “European” contribution.
Lots of ink have been spilled on expanding populations to describe the kind of founder effects that would reverse the normal, star-like expansion pattern, to the effect that the importance and behaviour of normal star-structured expansion patterns tends to be forgotten. The normal expansion pattern typically features an increased effective population at the front relative to the expanding population as a whole, what means that into the direction of an expanding wave front a growing portion of the population will be actually involved in reproduction, since better opportunities (especially of the unsettled younger population component) are the raison d’être to the expansion. The expansion is most profitable for those that effectively acquired new opportunities in the expansion areas. Even though founder effects are likely to occur at the front of the expanding population and are normally adverse to variance, ie. when the offspring of one founder tends to outbreed other lineages in the neighbourhood at the cost of diversity; their adverse impact on the overall variance is typically exaggerated when elsewhere this same lineage is less successful or remains absent altogether. Only a succession of founders recurring on the same lineage can make a difference, for instance when founder X1 is the ancestor of founder X2 at T=50 generations, that is the ancestor of founder X3 at T=100 generations and so on. This feature causes the anti-thesis of the star-pattern of an expanding population, ie. the Allele frequency cline (AFC).

Allele frequency clines (AFCs) can result from […] subsequent founder events during a range expansion (Klopfstein et al., 2006)

Klopfstein et al. are careful to explain that AFCs are a rare phenomenon, in population history rather to be expected in paleolithic low-population density scenarios than in Neolithic high-population density scenarios.

Our study further suggests that mutations having arisen during Paleolithic range expansions should show larger absolute frequency differences than those having occurred during a pure Neolithic expansion […]
Conversely, mutations that are found today at very low frequencies and nevertheless show a clinal pattern […] are much more likely to have been spread during the Neolithic than during the Paleolithic expansion. Finally, we would predict that new mutations being highly localized and at relatively low frequencies are more likely to have spread during the Neolithic expansion. (Klopfstein et al., 2006)

Contrary to popular opinion, the extinction (or low extant frequency) of a mutation does not imply the extinction of the whole expanding population, and neither in the AFC test cases. Actually, an expanding population can do very well without an AFC! Quantified in a straightforward way, for a hunting band of 60 individuals remaining on the narrow edge of the wave of advance and having an effective size of maximum 10:

Of the 64,000 simulations, ~18% were successful […] Altogether, the majority of mutations remained near the origin (~78%) whereas the remainder (~22%) traveled in the direction of the expansion, and on average their centroid can be found about midway between the place of origin of the mutation and the end of the expansion.
The stationary [centroids] have, on average, very few mutants; in the majority of simulations, the number of mutants remains below the level of polymorphism (Edmonds et al., 2003)

In the latter case an average lineage, here marked by a new mutation, doesn’t experience a strong founder effect at all and even tends to be phased out by neighbouring expanding lineages at the front “while the wave rolls on”. Thus, an expanding population doesn’t necessarily feature the AFC of any mutation at all, according to the mathematical investigations involved not even in the majority of cases.
In brief applied to the Neolithic wave of advance: the haplogroups that expanded together with LBK in Europe could result concurrently in vastly diverging expansion patterns, that may vary from a combination of star-like and AFC patterns applicable especially to Hg R1b1b2, to low frequency-long range haplogroups like Hg G2a3, and even to currently extinct haplogroups. The population shifts at the Neolithic transition seem to involve predominantly the arrival and subsequent diverging distributional processes of DNA that potentially derive from the Near East.

The low availability of maternal mtDNA in LBK territory that to a certain degree of confidence could be considered typical European, like mtDNA U5 or H, still suggests that European Neolithic populations experienced a considerable ingression of the local female element in a later period, not unlike the findings in Megalithic Europe. This would also imply that another population was still around. Paleogenetic samples strongly correlated eg. mtDNA U5 to the mesolithic cultures of the north of the Alps, while Mesolithic H appears to be a rather Atlantic phenomenon (west of LBK). Both distributions might have overlapped at the northwestern boundaries of LBK, even though so far it was impossible to retrieve reliable samples from the marshy soil to verify this. The continued expansion of male Neolithic DNA until modern times strongly contradicts the annihilation of Neolithic cultures by a new society of militant intruders, but the apparent extinction of matrilinear DNA supports the arrival of a new component on the scene that most likely was firmly rooted in Mesolithic Europe. In this process LBK mariage partners among the local Mesolithic population may already have preceded the shifts theorized by Marc vander Linden in Beaker times. These sex-biased genetic changes may coincide with a process that in modern archeology was forwarded as “Mesolithisation”.

Archeology traditionally presumed a Neolithic takeover in Europe of agriculturists from the Near East that took advantage of their high cultural level. They were supposed to have squeezed Mesolithic Europe into virtual extinction, without much attention to the possible effects of acculturation. Paleogenetic investigation tends to confirm the Neolithic victory at this “clash of cultures”, even though so far the results are far from unambiguous. At least the female Neolithic component obviously received a blow in more recent times, that could be due to Neolithic (LBK or post-LBK) or post-Neolithic marriage partners among the local Mesolithic population – conform the proposals of Marc vander Linden in 2007.
My previous article gave an overview on the current status of the discussion on the Neolithic advance as a wave that started in Anatolia and propagated with increasing R1b frequencies grosso modo to NW Europe. That article pointed at the possibility of a secondary Late-Neolithic expansion area in the neighborhood of the Paris Basin that could have been responsible for the spread of most of Western European R1b, represented by the especially numerous M420+ subclade. The LBK cultural complex was pivotal to this advance, and the investigation of Haak et al. (2010) suggests YDNA G2a3 could help in defining a specific Caucasian origin. “Caucasian” mtDNA N1a (similar to the “Central Asian” type N1a found in an AVK context) was also found at the Megalith site Prissé-la-Charrière dated 4200 BC, thus supporting the view of Myres et al. that the same LBK-related Neolithic wave of advance continued in the Late Neolithic:

We reproducibly retrieved partial HVR-I sequences (nps 16,165 to 16,390) from three human remains (Prisse´ 1, 2, and 4, Table 1), one adult and two children deposited during different stages of use of the burial chamber. Corresponding sequences could be unambiguously assigned to haplogroups X2, U5b, and N1a (Deguilloux et al., 2010)

In short, the traditional view on the Neolithic revolution could hardly be illustrated better than by this interpretation of a genetic wave of advance of Y-chromosome marker R1b that originated in a single male somewhere in Anatolia and then – Ex Oriente Lux – propagated with ever increasing frequencies until the darkest outposts of the western world where the sun goes down.
However, this doesn’t solve the marked differences with the distribution patterns of other Neolithic haplogroups, both YDNA like G2a3 and mtDNA. These inconsistencies may be much more than the mathematic phenomena as described above. Actually, modern archeology currently favours an important revision that challenges almost anything Neolithization traditionally stands for. The apparent genetic association with Neolithic culture in Europe is still to ambiguous to demonstrate the Neolithic advance to be exactly what modern archeology doesn’t support anymore: a deterministic transition referred to as Neolithization. This model regarded the introduction of domesticates as inevitable, the Neolithic way of life inescapable, the evolution and progress involved universal and its origin unique and singular.

The main proposition of this view is that the term and the concepts of the Neolithic inhibit clear thinking about what happened. Proponents consider the traditional dichotomy between huntergatherers and agriculturists already debunked as an artificial construct. Instead, the key development of the transition lies within the overall management of the natural environment, including specific methods like the systematic burning of forests all over the world. Features like the domstication of animals and growing crops are secundary to this evolving management of nature. A growing body of archeological evidence shows up to prove this. Eg. in Korea people already grew rice about 13000 BC, 6000 years later this same people also included millet and domesticated pigs (7000 BC). Domestication of pigs probably also happened in Germany in Rottenburg-Siebenlinden and Gottingen. Forest management in pre-Neolithic Northern Europe was directed at harvesting oaks and hazelnut. The list is long and includes many Neolithic features scattered all over the world in pre-Neolithic cultures: like pottery between 11800-8000 BC in the Jomonculture, Japan; in El Adam, Egypt at 9000 BC; Mesolithic La Hoguette pottery already existed before the arrival of LBK. And why the Neolithic Revolution in Europe would be associated to a unique package of “Neolithic” changes that may be rather ethnically defined? Its introduction was apparently associated with new ethnic elements and their culture, but this does not mean that the native population was “evangelized” into a new way of life of Neolithic copycats. Especially in Northern Europe parallel cultural developments can be discerned that apparently evolve from internal impulses. The Neolithic represented by cultures like LBK thus emerges as a shared development that was already initiated in the paleolithic all over the world:

European Paleolithic subsistence is assumed to have been largely based on animal protein and fat, whereas evidence for plant consumption is rare. We present evidence of starch grains from various wild plants on the surfaces of grinding tools at the sites of Bilancino II (Italy), Kostenki 16-Uglyanka (Russia), and Pavlov VI (Czech Republic). The samples originate from a variety of geographical and environmental contexts, ranging from northeastern Europe to the central Mediterranean, and dated to the Mid-Upper Paleolithic (Gravettian and Gorodtsovian). The three sites suggest that vegetal food processing, and possibly the production of flour, was a common practice, widespread across Europe from at least ~30,000 y ago. It is likely that high energy content plant foods were available and were used as components of the food economy of these mobile hunter-gatherers. (Revedin et al., 2010)

The earliest Neolithic was still far away from large scale exploitation and production, that developed only much later and under completely different circumstances. Rather this early Neolithic cultures were an expression of an imported ideology, meeting another completely different Mesolithic ideology. The imported ideology was rigid, restricted to certain soils and techniques and unable to get full profit out of local circumstances, while the Mesolithic ideology was flexible and diverse like expressed in Swifterbant culture:

Our knowledge of Late Mesolithic hunter-gatherer food strategies in the area suggests that they included the exploitation of a wide range of food sources to avoid dependence on a single food source. (Cappers et al., 2008)

The integration of Mesolithic exploitation strategies must have been critical to the survival of Neolithic immigrants. Indeed, Mesolithic interaction can already be discerned at the earliest stages of the Neolithic advent. The Neolithic culture of Starcevo (Serbia) may have inherited from similar Anatolian traditions as LBK, and has even been named as an important precursor to LBK. There is also a strong Mesolithic component. Bogdanovic (2009): “Mesolithic and Neolithic horizons in Lepenski Vir show that in both groups of inhabitants there are only slight differences in what they ate”. Some discussion remains about the anachronism of these Mesolithic influences, since according to the interpretation of Bogdanovic the site Lepenski Vir should be “attributed signs of a conservative variant of the Proto-Starcevo culture”:

At the time Lepenski Vir was discovered and the Proto-Starcevo culture promoted, there was an ideological change in interpreting Early Neolithic of the central Balkan. The Starcevo culture was ultimately shifted as secondary. (Bogdanovic, 2009)

However, elsewhere archeologists are rather inclined to conclude that Mesolithic influences altered the Neolithic ideology in a later phase. Eg. Christian Jeunesse makes a distinction between different LBK traditions, one of them flexible (tradition II) and the other one rigid (tradition I). Even though he is acquainted with the possible mixed Starcevo-related origin of LBK, he is rather inclined to consider the effect of regional Mesolithic influences. At a LBK site in Vaihingen, garbage pits were found with human remains that were more robust than usual. This was interpreted as reminiscent to native hunter-gatherer funeral traditions, being clearly distinct from typical LBK funeral traditions. However, dumping of the remains of subordinates, slaves or hostages couldn’t be excluded – in which case any Mesolithic integration probably wasn’t accomplished through the male lineage.
The final breakdown of LBK culture has often been associated with internal stress when agricultural traditions proved to be insufficient to compensate growth with further expansion and increase of productivity. LBK lacked the flexibility to take full advantage of local resources like their Mesolithic neighbours. The LBK society had a subsistence economy based on just a few products. Previous growth only complicated the internal problems until the whole system broke down, hence the collapse of food production and social structure. Nobody ever cited clear external causes to the LBK demise: the eventual introduction of new techniques may rather be related to renewed growth. However, the most striking innovation after the collapse of LBK is the return to a wide spectrum economy, where the people rediscovered natural resources that were already employed in the Mesolithic. Hence post-LBK growth may be attributed to the Mesolithic heritage: Mesolithisation.
The Swifterbant culture was one of LBK’s Mesolithic neighbours, that had kept their wide spectrum economy. Their flexibility to use different kinds of habitats and their resources (hunt, fishing, gathering, small scale foodproduction) proved a strong argument against any need of Neolithization, rather the contrary might be true: the Mesolithization of Neolithic societies. The same kind of flexibility developed in post-LBK cultures like Rössen and Michelsberg. According to this view Swifterbant culture, like other contemporaneous Northwestern European groups, continued to develop at their own pace towards a more pronounced management of the natural environment. These groups developed into Funnelbeaker, generally considered ancestral to the vast Corded Ware horizon that ultimately emerged as a society where – next to agriculture – hunting and fishing was the basis of subsistence, with an increasing reliance on the exploitation of marine resources. This transformation could happen without much external influences – paving the way to a whole new period of thoroughly “de-Neolithized” Beaker cultures, that could be considered fully “Mesolithized” if for a moment we would be willing to discount the new dynamism of trade and contact. From that moment on a new unity embraces the people of Europe – that often is referred to as Indo-European.

Bengtson grouped the North Caucasian languages together with Basque and Burushaski in a single Macro-Caucasian family, that apparently was also shared by the LBK culture.

At this point we deduced the development of a Neolithic core population under the influence of Mesolithic neighbouring populations into a new Indo-European entity, not unlike Zvelebil’s Neolithic creolisation hypothesis, first put forward in 1995. This archeological argument was predicted linguistically by Kortlandt when he argued thus, albeit having a completely different location of the Caucasian substrate in mind:

What we do have to take into account is the typological similarity of Proto-Indo-European to the North-West Caucasian languages. If this similarity can be attributed to areal factors, we may think of Indo-European as a branch of Uralo-Altaic which was transformed under the influence of a Caucasian substratum. (Kortlandt, 1989)

According to linguist Peter Schrijver the Neolithic substrate in NW Europe must have spoken a language that feature complex verbs, not unlike current NW Caucassian languages. The results of Haak’s investigation, both in YDNA and mtDNA, allow us now to fully identify the North-West Caucasian-like substrate as LBK, and the Mesolithic influences as the transformed superstratum that gave rise to Indo-European.
However, if this was the case the population shifts that accompanied this transformation can’t have been predominantly male-driven as has been always taken for granted. Instead, the changing composition of mtDNA combined with more or less static YDNA strongly suggests the transformation was rather in line with the exchange of marriage partners described by Marc Vander Linden.
Ultimately, to give this interpretation of the Neolithic Creolisation Hypothesis more substance, I would recommend reading about the progress made on the Basque issue. John D. Bengtson groups Basque, Caucasian and Burushaski together in a single Macro-Caucasian family, thus supplying evidence that a Neolithic precursor of Basque could fit the profile of a Caucasian-like substratum to Indo-European. Arnaud Etchamendy (2007) even defended his thesis that Basque vocabularity should make this language essentially another integrant of the Indo-European family of languages.


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