Improved distribution map of Denisova alleles, at SNPs where Denisova is different from chimpanzee and Neandertal. There is an appreciable match with A*1101 HLA alleles

I bet that a few years from now, we will look with amazement at the naivete of the passing Out of Africa orthodoxy that bundled all Africans into an amorphous category of “our ancestors in Africa”. It is also becoming clearer that increased African genetic variation is, at least in part, due to the continent being home to multiple deeply divergent populations that persisted, in various admixtures down to the present. (Dienekes’ blog, 2011-9-16)

The amount of new scientific publications able to shed more light on the issue is already dazzling and for sure preludes an inevitable change of paradigm:

We are only now beginning to harness the power of full human genomes for evolutionary inferences, but it is inevitable that a new theory of human origins will appear that will reconcile the different and conflicting lines of evidence. That theory must take into account latent admixture as a cause of African genetic diversity, and it must also harmonize with the paleoanthropological record. (Dienekes’ blog, 2011-9-19)

Scientists already speak out openly on how the ‘vast amount of recombination information in the human genome has long been ignored or deliberately avoided in studies on human population genetic relationships’ (Xu et al., 2011). The latter now introduced a method of ‘chromosome-wide haplotype sharing’ (CHS) to reconstruct reliable phylogenies of human populations and state that instead of variation being an unbiased measure of population age, ‘the majority of the variation in CHS matrix can be attributed to recombination.’
Unfortunately, time and origin of the admixtures remain hard to quantify. Sampled fossil hominines so far didn’t correspond to much closer modern matches in the wider geographic region. Denisova admixtures have top levels in SE Asia and Australia while traces are virtually absent in South Siberia, where of both Denisova fossils were found. Likewise, Neanderthal admixtures have top levels in East Asia and the Americas, far removed from the European habitat of Neanderthal, where levels are lower. In his blog John Hawks suggests population history and natural selection may have caused this distorted picture, while much of the purported evidence for local continuity of Denisova genes may be of a quite different magnitude. Certain HLA genes, part of the modern human immune system also found in the genomes of Denisova and Neanderthal hominines, would have been straightforward evidence for introgression and local continuity – if only those genes didn’t derive from much older primate genes. Balanced natural selection could have been an alternative to account for the survival of ancient genes, supplying false positives for the survival of a select set of Denisova genes. Still, the current geographical distribution of the HLA-A*11 variety found in Denisova, by Parham, concurred remarkably with that of other derived Denisova alleles, published afterwards:

[...] we found that East Asian populations, particularly Southeast Asian populations, had, on average, a greater frequency of the derived Denisova allele compared with other populations (except for Oceanians) [...]. (Skoglund & Jakobsson, 2011)

Since this particular HLA-A*11 variety is mostly found in Asians and never in Africans this indeed suggests introgression by interbreeding outside of Africa. True, such alleles existed already at the same position in the genomes of apes, but now with supporting evidence this detail should supply an entirely new dimension to the issue: introgression of DNA previously conserved merely on evolutionary sidelines should imply temporal extinction of the gene on the main hominine lineage!
At least, extinction is exactly what happened at a much larger scale with primate ABO blood group genes: chimpanzees so far attested for blood type A and some O, but not for B, and gorillas attested for blood type B and some O, but not for A. The ABO blood type is controlled by a single ‘ABO’ gene, for humans on the long arm of the ninth chromosome (9q34).

In the case of human ABO blood group genes, three alleles (i.e., A, B, and O alleles) have been observed, but it is known that chimpanzees have two alleles (i.e., A and O alleles). (Kitano et al., 2000)

Blood groups are inherited from both parents, that may be homozygous for one or heterozygous for two classes. This means that despite being ancient, any of the groups could become extinct once an epidemic demands a strong response that involves unbalanced selection of another allele. Eg., blood group O was observed to have an advantage against malaria, leading to the hypothesis that ‘selective pressure imposed by malaria may contribute to the variable global distribution of ABO blood groups in the human population.’ (Rowe et al., 2007). Since malaria is strongly related to warm wetland habitats, this may be an example of an ancient immune system whose usefulness and survival may be a matter of habitat. If so, then speciation – that implies a formative stage of isolation within a new habitat – also implies an increased risk. Among the great apes only orangutan attested all three types, suggesting that extinction of immune system-related alleles may indeed have happened more often. Reintroduction of ancient genes that became lost may be an almost impossible affair without subsequent introgression by hybridization. Nowadays such potential hybridization partners may be hard to find for chimps or gorillas, probably characterizing the evolutionary costs of their speciation. However, for the human lineage hybridization remained a possibility with species or subspecies that co-evolved within a wide range of different habitats, where different survival patterns and demands had their unique effect on local adaptations, ie. including the immune system! In summary, balancing selection may play a trick on us here, but the evidence is highly suggestive of the deep time depths involved for the genetic divergence achieved before introgression.
Equally remarkable was the absence of Denisova genes being confirmed for Central and South Asia, closer to the fossil origin. The route of Denisova genes to SE Asia must have led through East Asia, but the replacement in Central Asia by other rare varieties of HLA raises even more questions. Such as the origin of the Central Asiatic allele HLA-B*73, not attested in Denisovans, but in modern humans always associated with the ‘Denisovan’ HLA-C*15 antigen, thus suggesting either an hitherto unknown presence in other Denisovans, or a hybrid neighbor.
A recently deciphered genome “from a 100-year-old lock of hair donated by an Aboriginal man” confirmed that despite long time isolation, their proportion of Neanderthal segments match that observed in European and Asian sequences. This Neanderthal component may have been available when the ‘aboriginal Australians split from the ancestral Eurasian population 62,000 to 75,000 years B.P.’ (Rasmussen et al., 2011). However, even though according to the latest insights East Asians are considered significantly closer to Denisova relative to Neandertal, the morphology of aboriginals is rather related to archaic populations and fossils in the wider region of South and South East Asia:

This study has shown that Southeast Asia was settled by modern humans in multiple waves: One wave contributed the ancestors of present-day Onge [Andaman Islands], Jehai [Malaysia], Mamanwa [Philippines], New Guineans, and Australians (some of whom admixed with Denisovans)
[...]
we considered the possibility that the secondary gene-flow event into the ancestors of Australians and New Guineans came from relatives of Chinese (CHB) rather than western Negritos such as the Onge. (Reich et al., 2011)

Indeed the Denisova admixtures follow a rather geographic pattern, being present in eastern Indonesia and among the Mamanwa ‘Negritos’, but virtually absent in the ‘Negritos’ of Malaysia and the Andaman Islands. Most probably the aboriginals, where tests revealed ‘slightly less allele sharing than observed for Papuans’ – got the component from their closest geographic neighbors, thus flouting the Out of Africa model and increasing the mystery of the timing and origin of Denisova intrusion.

[...] it is becoming increasingly difficult to imagine a structure model that can fully explain the complex pattern of archaic ancestry in non-Africans without invoking any restricted admixture events with archaic humans. Instead, we suggest that direct gene flow from archaic populations is the most likely explanation for the shared genetic ancestry between East Asian populations and the Denisova genome (Skoglund & Jakobsson, 2011)

Likewise, low values of the Neanderthal component in their previous European heartlands and unpublished reports of unrelated archaic admixtures both in Europe and South Asia should make us weary about the significance of current geography for the origin question at all. If dislocation of genetic affinity with fossil samples defines a tendency, we can’t even be sure the African archaic admixtures mentioned above indeed derive from an African core region. However, Wall et al. (HGV2011) reported that part of the genetic regions on the human genome ‘were found in the genomes of both sub-Saharan African and non-African populations’. At least some of the archaic admixture became international, not unlike we already known of the V and M haplotypes of gene ASAH1, coalescent-time depth 2.4 million years ago, that ended up evenly dispersed around the world. The mere age of the African archaic introgressive DNA currently under investigation vastly exceeds the age of Neanderthal and Denisova, even that of their (and our) common ancestor Homo heidelbergensis, an extinct species of the genus Homo that lived between 600 and 400 thousand years ago. This seems to be ever less in agreement with the traditional biological species concept (BSC) that tends to relate speciation with highly effective reproductive isolation, and questions the very nature of the human speciation process over time.
Actually, current biology textbooks don’t hesitate anymore to supply ‘prohibitive’ answers, that simply urge to be applied to the human species:

Typically, gene flow occurs between the different populations of a species. This ongoing exchange of alleles tend to hold the population together genetically. (Campbell Biology, ninth edition, 2011)

Speciation still occurs by the evolution of reproductive isolation, but ‘there are many pairs of species that are morphologically and ecologically distinct, and yet gene flow occurs between them.’ An example is the grizzly bear and the polar bear, that despite occasional natural hybridization remain distinct. ‘This observation has led some researchers to argue that the biological species concept overemphasizes gene flow and downplays the role of natural selection.’ (Campbell Biology, 2011). Some other species appear to be actually fully hybrid, like the Tiger Swallowtail Butterfly (appalachiensis): ‘Inter-specific hybridization is widespread in nature and may have important consequences in evolution, from the transfer of adaptive alleles between species to the formation of hybrid species’ (Kunte et al., 2011). According to the investigators the ‘evolution and persistence of appalachiensis in contact with its parental species suggests that hybridization among animals may result in selectively favored hybrid species that contribute to biodiversity.’ Hence, this example exposes a ‘potential role for natural selection in the origin and maintenance of hybrid species.’
Indeed, ‘hybridization is not always a dead end, as the BSC might suggest, but a potential source of new array of hybrids (hybrid swarms) that may establish themselves, eventually in new ecotone habitats, and evolve as new species. One common feature of the process that accompanies hybridization is the rapidity of genome repatterning which is hardly explained by the conventional mutation and recombination rates. Rather, transposition bursts ensuing hybridization suggest their involvement in these rapid genome reorganizations.’ (Fontdevila, 2005)
The risk of speciation may be genetic isolation and inbreeding, the risk of illimited hybridization of subspecies is the potential loss of local environmental adaptations, like the white color of polar bears following the example above. Over time, reproductive barriers may be either reinforced or weakened. In the latter case, hybridization may eventually lead to ‘the fusion of the parent species’ gene pool and a loss of species.’ (Campbell Biology, ninth edition, 2011)
As such, there is no specific reason why mere biology wouldn’t supply answers for the apparently torturous evolution of the human species, or for the sudden appearance of African apes:

In a punctuated pattern, new species change most as they branch from a parent species and then change little for the rest of their existence. (Campbell Biology, 2011)

Both patterns of how species evolve have been observed for the radiation of the mammals over the last 165-million-year. The majority of mammal species, including even the most speciose orders (Rodentia and Chiroptera), experienced an explosive 10-52 fold increase in the rate of evolution only during their initial formation up to the common ancestor. Stable rates of evolution of species, however diverse, were recorded almost everywhere else.

These results necessarily decouple morphological diversification from speciation and suggest that the processes that give rise to the morphological diversity of a class of animals are far more free to vary than previously considered. Niches do not seem to fill up, and diversity seems to arise whenever, wherever and at whatever rate it is advantageous. (Venditti et al., 2011)

The sudden appearance of chimps and also gorilla in the fossil record may indicate their speciation occurred relatively rapidly, when they occupied and then filled their ecological niches. Over the wider humanoid lineage things may be different:

For species whose fossils changed more gradually [...] it is likely that speciation in such groups occurred relatively slowly, perhaps taking millions of years. (Campbell Biology, 2011)

How literally we could take these ‘millions of years’ for the lineage leading to great apes and modern man? Decades of behavioral studies resulted in the insight that apes are far more hominized than was once believed. According to Krützen et al. (2011) this already applies to the cultural plasticity of orangutan. Since this tendency to share particular behaviors in a group was also demonstrated in wild populations of chimpanzees this suggests evolutionary roots in the ancestors that all great apes share with humans. How realistic this would be for the biological species concept, assuming orangutan diverged from our common ancestor 15 mya, or more? Many hominizing tendencies, like the slowing down of juvenile growth, could develop for all humanoid lineages well after such divergence dates. Or would hominized behavior invoke exactly the opposite of BSC with the first onset of organizing a beneficial habitat to the temporarily impaired, like hybrids? Hominized groups could offer better opportunities to overcome some hybrid-related barriers such as lower fitness and fertility. Better survival opportunities would enhance the possibility that hybrid-related genome repatterning could stabilize recombination by endogenous purifying selection over a longer period, maybe several generations, to the effect that both fitness and fertility could be restored over time by the natural resilience of genetic processes. Indeed, if hybridization may explain parallel developments among humans and great apes, we could postulate genetic exchange between diverging ape lineages spanning millions of years.
The laboratory or Reich in Boston has a record in the study of hybrid-driven evolution, and substantiated hybridization already in 2006 for the hominine lineage. What the team called a ‘realistic upper bound of < 17 Mya for human-orangutan genome divergence' resulted in a common ancestor with chimpanzee less than 5.4 mya, including a considerable period of mutual hybridization:

Most strikingly, chromosome X shows an extremely young genetic divergence time, close to the genome minimum along nearly its entire length. These unexpected features would be explained if the human and chimpanzee lineages initially diverged, then later exchanged genes before separating permanently. (Patterson et al., 2006)

The study concluded that introgression of chimp-like DNA in the human lineage, or the other way round, happened after about 1,2 million years of divergence. This year’s ICHG meeting a team of accredited scientists led by J. X. Sun presented a tight refinement of previous date estimates:

Human-chimpanzee speciation is estimated to be 3.92-5.91 Mya, challenging views of the Toumaï fossil [Sahelanthropus] as dating to >6.8 Mya and being on the hominin lineage since the final separation of humans and chimpanzees. (Sun et al., 2011)

Indeed, despite initial efforts to link this transitional period of hybridization with older hominines, the time span of Ardipithecus would make a much better match, and more so due to the immediate succession of Australopithecus: this hominine ancestor complied even less to the more chimpanzee-like anatomy commonly expected with the genetic similarity of humans and chimps:

In an assessment of fossils from Kanapoi (3.9-4.2 Myr ago), the anagenetic series Ar. ramidus, Au. anamensis and Au. afarensis has been hypothesized. The evidence reported here from the Afar Rift constitutes a strong test from a single stratigraphic succession that fails to falsify this hypothesis (White et al., 2006)

The cited evidence of introgression revolted against traditional phylogeny, but the proposal adhered to the established view: one single lineage up to a common ancestor of humans and chimps. Supportive molecular evidence and dating is commonly cited, but the devil is in the details:

Although chimpanzees are our closest relatives, there are many loci at which humans and gorillas (or chimpanzees and gorillas) are the most closely related; we estimate that this is the case over about 18–29% of the genome (Patterson et all, 2006 Supp.)

Likewise, ‘data based on morphological analyses and with the data based on mitochondrial ribosomal genes [...] suggest a closer relationship between gorilla and chimpanzee’ (Rasheed et al., 1991).
A much longer history of shared ‘hominizing’ evolution that encompass all great apes may be implied, and more so by additional inconsistencies in the order African apes are supposed to have diverged from the lineage leading to modern humans:

The late-divergence hypothesis [...] specifically focuses on the divergence between humans and the ape, emphasizing that there were two different divergence points in the evolution of recent hominoids. (Wolpoff, 1982)

Thin enamel, knuckle walking and specialized feet for grasping are just a few shared features of African apes that are difficult to reconcile with two separate speciation events from an otherwise more hominine fossil record. For instance, despite Begun’s certainty on the knuckle-walking habits of our earliest common hominid ancestors, this remains utterly hypothetical – and utterly unsuported by more complete and recent fossils like A. sediba, and even Ardipithecus had not evolved the hands and wrists of a knuckle-walker. Homoplasty among different African apes that feature knuckle-walking is equally unsupported:

[This study] does not support the hypothesis of a knuckle-walking complex, nor does it support the contention that knuckle-walking could have been easily evolved independently in chimpanzees and gorillas. (Williams, 2010)

Instead, Grehan and Schwartz (2010) make a case for grouping ‘the monophyly of hominids and various Miocene–Pliocene fossil apes and orangutans into a ‘dental-hominoid clade’, with the African apes as a sister clade along with the putative [hominines] Ardipithecus and Sahelanthropus.’ In this view only the early hominine Orrorin could link the ‘dental-hominoid-clade’ to humans, thus suggesting a smooth transition along this lineage to hominids, but leaving the close family relationship with African apes unresolved.

Orrorin is [...] already quite distinct morphologically from the African Great Apes (Gorillidae). This indicates a divergence between Hominidae and Gorillidae that dates back to a substantial period prior to 6 Ma, and we estimate about 8-7 Ma for this event. If so, then the discovery of Orrorin refutes all hypotheses in which humans diverged from apes later than 7 Ma, including most of the recent estimates by molecular biologists who tend to think of the divergence as having taken place later than 5 Ma, and even as recently as 2.5 Ma. (Martin Pickford, 2001)

Molecular evidence favors a close relation with African apes, but a much tighter morphological affiliation of the human lineage with orangutan can be observed:

[...] in addition to the development of low-cusped cheek teeth and thick molar enamel, humans shared a significant number of derived features uniquely with the orangutan (e.g. in reproductive physiology (gestation length, estriol levels, absence of estrus), degree of cerebral asymmetries, fetal adrenal zone size, lack of keratinized ischial callosities, mammary gland separation, hair length, incisive foramen number] (Schultz, 2004)

Indeed, the duration of the menstrual cycle varies with species; about 29 days in orangutans, about 30 days in gorillas and about 37 days in chimpanzees. Only the genus Hoolock (previously Bunopithecus), probably the most basal member of the lesser apes (gibbons), match human females ‘precisely’ – except for a considerable variability – in having an average menstrual cycle of 28 days (Geissmann et al., 2009). Cytogenetic evidence pleads for a shared development that considerably exceeds a basal relationship with great apes but unfortunately, so far gibbons are generally dismissed as even more distant from humans than orangutan.
Grehan (2006) expanded on this paradox by noting that ‘the orangutan relationship is supported by about 28 well-supported characters, and it is also corroborated by the presence of orangutan-related features in early hominids’ such as a thickened posterior palate and anterior zygomatic roots. We could add characteristics of their close fossil relatives such as dental structure, thick enamel, shoulder blade structure, thick posterior palate, single incisive foramen.

Comparative morphology supports a unique common ancestry for humans and orangutans as the only phylogenetic theory with substantial corroborated evidence. Even supporters of a unique common ancestry for humans and chimpanzees collectively support more (26) orangutan-related human characters than they do for chimpanzee-related human characters (Grehan, 2006)

Despite the impressive list of unique correlations shared between the human lineage and orangutans, the vast overall genetic distance between the species is often cited as evidence to a deep and rather straightforward phylogenetic affiliation. Genetic isolation of the orangutan lineage – according to molecular evidence between 14-17 mya (million years ago)- doesn’t help in understanding the last recorded interspecies hybridization event between ancestral humans and chimps from the ‘dental-hominoid’ point of view. In the words of Schwartz such an extended period of geneflow between widely divergent ape lineages certainly “pushes the limits of credulity” regarding the hybridization hypothesis.
For the moment I could suffice to notice that morphological and molecular evidence, that challenges the joint divergence of hominines and chimp ancestors from the ancestors of gorilla, may be explained by introgression: as in the example of hybrid polar bears, hybridization doesn’t necessarily imply the fusion of divergent evolutionary adaptive lineages. But hybridization could have complicated the hominid phylogeny beyond limits over a very long time, and even more so as the common source of adaptive radiation events towards speciation was rooted in a common ancestry that started long ago as a balanced process between evolving species.
Actually, human curiosity supplied comparable results for other species, suggesting relative time depths that correspond to what could be expected for the hybridization of diverging ape lineages:

Recent interspecific hybridizations have been well documented for the Bos and Bison species [...] : zebu and ox in several tropical regions; zebu and banteng in Indonesia; taurine cattle and yak in China, Mongolia and Siberia, etc. Ox-zebu hybrids are completely fertile, while male progeny of other hybridizations are sterile. Earlier introgression events may be indicated by the anomalies in the mitochondrial phylogeny [...] that are incongruent with trees of nuclear genes, AFLP fingerprints (these studies) and Y chromosomal sequence variation
[...]
Since exchange of genetic material depends on the geographical overlap of the regions inhabited by the species and their ancestors, this is consistent with the hypothesis that reticulation influenced the phylogeny of the Bovini. (Buntjer et al., 2001)

Cross-breeding of ox (genus Bos) and bison yielded fertile females and (mostly) infertile males. The same was implied by Reich, Pattison and colleges for hominine interbreeding with chimps. The fossil record already distinguishes bison in the range late Pliocene-Early Pleistocene up to 2 million years ago, while the origin of Bos (ie. cows) is contested. Their divergence is unlikely to be less than 2 million years, and possibly considerably longer:

The oldest clear evidence of Bos is the skull fragment ASB-198-1 from the middle Pleistocene (~ 0.6 – 0.8 Ma) site of Asbole (Lower Awash Valley, Ethiopia).
[...]
Although the origin of Bos has traditionally been connected with Leptobos and Bison [...] we propose here a different origin, connecting the middle Pleistocene Eurasian forms of B. primigenius with the African Late Pliocene and early Pleistocene large size member of the tribe Bovini Pelorovis sensu stricto.
[...]
The Bison lineage [...], based on skull anatomy, can be interpreted as resulting from anagenetic evolution of the Late Pliocene forms of Leptobos across the Plio-Pleistocene transition (~2.0 – 1.7 Ma)
[...]
The cranial anatomy of Bos, however, is highly derived as to be considered the result of a direct anagenetic evolution from any form of Leptobos. Martinez-Navarro et al., 2007)

Bison cows ordinarily conceive for the first time at 2 years of age while chimps reach their reproductive age only at 13.5 years, almost 7 times later. Accordingly, this translates to the feasibility of an extended period of geneflow for the ape lineage that may even exceed the lapse of 10 million years to bridge all genetic divergence between the common ancestor of all great apes and the reported chimp-hominine hybridization event.
As a caveat it should be noticed that fertile first generation (F-1) half-bison bulls are also registered (Wyoming Thunder being one example). This possibility increases when the bull parent already had some cattle blood, though in some cases this could not be confirmed. This observation typifies hybrid infertility as an essentially temporal evolutionary problem.

The grolar bear is a natural hybrid of grizzly bear and the polar bear. To the right the beefalo, fertile offspring of bison and cattle. Below Wyoming Thunder, a F-1 fertile bull from a male bison.

Cladistic analysis of dental, cranial, and postcranial characters separate Hippidion and Equus into two different clades, which share the North American late Miocene Pliohippus as a common ancestor around 10 MA (Prado and Alberdi 1996).
[...]
Alternatively, MacFadden (1997) has suggested that Equus is derived from Dinohippus, and Hippidion from Pliohippus sensu lato (including Astrohippus), implying that the divergence between Dinohippus and Pliohippus occurred prior to 10 MA.
(Orlando et al., 2007)

Molecular evidence proved that the Hippidion, an extinct south american horse with some Pliohippus characteristics, was most similar with moders equids.

Hippidion is considered to be a descendant of the pliohippines, a primitive group of Miocene horses that diverged from the ancestral lineage of equines (a category including all living and extinct members of the genus Equus, such as caballine horses, hemiones, asses, and zebras) prior to 10 Ma ago [...]. A recent study presented genetic data from three southern Patagonian specimens morphologically identified as Hippidion. Unexpectedly, the sequences clustered inside the genus Equus (Weinstock et al., 2005)

Hippidion differed less from caballine equids than all extant non-caballine equids (including donkeys, zebras etc.). This result inspired the team to question the deep morphological split:

The close phylogenetic relationship between Hippidion and caballine horses is in direct contrast to current paleontological models of hippidiform origins. Nevertheless, we are confident that these sequences are those of Hippidion rather than the South American caballine form E. (Amerhippus), which dispersed into South America about 1 to 1.5 Ma later than Hippidion. (Weinstock et al., 2005)

On morphological grounds, the phylogenetic separation between the lineages of Pliohippus/Hippidion and Dinohippus/Equus (modern equines) was previously estimated at 10 mya, and tentatively associated with the merychippine radiation of 15 mya. but the close genetic relationship now attested for Hippidion and caballine equids shatters such deep divergence. Equine evolution now faces the same dilemma as the human origin question, where morphology and molecular evidence are equally at odds. History repeated itself and like what happened before in paleoanthropology in the face of molecular evidence, scientific consensus rather conformed to the biological species concept and concluded that morphology was deceiving:

These data suggest that temporal and regional variation in body size and morphological and anatomic features should be considered a sign of extraordinary plasticity within each of these lineages. Such environment-driven adaptative changes would explain why the taxonomic diversity of equids has been overestimated on morphoanatomical grounds. (Orlando et al., 2007)

Another phylogeny isn’t hard to make, but even the divergence date of extant equines becomes questionable now the DNA of all three extinct American horse species was revealed to cluster specifically with cabelline horses. Genetically, the most recent common ancestor of all modern equids should have lived ‘merely’ ~5.6 mya, and still this appears inconsistent with proposals towards a single Out of America exodus event through Beringia during the glacials, at most 3 mya. Perhaps the easiest way to deal with evolutionary inconsistencies is to just forget about the need of a strict phylogenetic tree at all. For being a viable alternative to morphological continuity, hybridization processes must have been an important element in equine evolution for millions of years. Naturally, divergence dates would easily be overestimated with a last common hybrid ancestor, and easily underestimated for the species – or subspecies – that were the constituent components of the hybridization. For sure this would urge for another evolutionary model, and perhaps the identification of very different genetic signals.
It is suggested that among mammals, equines exhibit the highest rate of chromosomal evolution. Commonly considered Old World immigrants from North America, they feature a progressive ‘decrease’ of their chromosome count moving further away from Beringia: Prezewalski still has 66 chromosomes, 2 more than the domestic horse that is commonly believed to belong to the same species. Of their closest neighbors, the Asian asses, the count ‘drops’ to 56, while the African ass could preserve 62 chromosomes. Extant zebras count 46 chromosomes for the northern Grévy-zebra, 44 chromosomes for the common plains zebra and 32 for the southernmost mountain zebra. Before, maybe inspired by equivocal ideas concerning equine evolution, scientists were inclined to assume the primacy of chromosome fusion in karyotypic evolution, but:

Recent advances in cell-cycle regulation, chromosome behavior, fossil record, and phylogenetic inferences dispute that the primary direction of karyotypic evolution by sequential fusion of chromosomes is toward an arbitrary reduction in diploid number. (Kolnicki et al., 2000)

Lower chromosome counts of equids the further away from the American origin of Equus may now be attributed to conservatism, and the higher counts to ongoing change and evolution.

A key postulate of Todd’s karyotypic fission theory is the idea that in a postfissioned karyotype with a high number of acrocentric chromosomes, a trend for acrocentrics to revert to smaller mediocentrics by pericentric inversion (or centric fusion) repotentiates the karyotype for further fissions correlated with episodes of adaptive radiation directly inferable in the fossil record (Fontdevila, 2005)

For instance, the reduction-argument has been ‘turned on its phylogenetic head’ with investigations on cycads – often mistaken for palms or ferns, but only distantly related to either. The cycad fossil record dates to the early Permian, 280 mya.

Zamia is unique among cycads in that both inter- and intraspecific chromosome numbers range from 16 to 28, excluding 20 [...], with varied karyotype composition
[...]
The large size of chromosomes in all cycad taxa excluding Zamiaceae [...] indicates that chromosomal fission has been rare or absent in these taxa.
[...]
Molecular analysis has generated limited evidence for any chromosomal rearrangements, including chromosomal fission and pericentric inversion, as the main mechanism driving karyotypic evolution in Zamia.
(Olspon et al., 2011)

In chromosomal fission the total number of chromosome arms remains the same, ‘whereas pericentric inversion and/or hybridization of different karyotypes may either add or subtract from the total arm count’:

Pericentric inversions of the different combinations of telocentric chromosomes and/or hybridization were almost certainly involved in generating the chromosome arm numbers observed, a conclusion based on the size and number of chromosomes. Chromosome arm counts for all [Zamia] taxa in this study [...] suggest that chromosomal fission, pericentric inversion, hybridization of different karyotypes and a combination of these mechanisms occurred in the evolutionary history of this genus (Olspon et al., 2011)

All corroborate to the notion that – under certain circumstances – adaptive radiation and speciation are symptomatic to chromosomal change.
The significance is obvious: ‘Chromosomal fission decreases genetic hitchhiking by severing transcentromeric linkages, allowing for direct selection on newly unlinked genes’ (Olspon). However, there must also be a drawback since in the course of evolution the number of chromosomes didn’t increase without limit. As for cycads in general, and most zamia in particular, the preferred count seems to stabilize at a low number chromosomes: ‘most early diverging cycad taxa have 2n= 16 or 18, with mostly metacentric and submetacentric chromosomes’.
Despite the exceptional features of some members of the clade, chromosomal stability and equilibrium is suggested for most cycad species. A straightforward correlation with morphological variability and stressful or widely variable habitats, that would require high levels of genetic adaptation, is falsified by the stable habitat, primitive morphology and high chromosome number found at the species Z. roezlii that inhabits the Colombian rainforest. The high age of the clade may be important evidence that fission may also be reminiscent of adaptive radiation events in the past. Over time this may be compensated by a corresponding reversal of the chromosome count by fusion. At least this is indicated in basal vertebrates, where ‘some ancestral segments were fused prior to the divergence of salamanders and anurans’ (Voss et al, 2011).
A pattern of chromosomal equilibrium may be observed in cichlid fishes:

Perciformes represents the largest order of vertebrates with approximately 9.300 species. It includes more than 3.000 species of the family Cichlidae [1,2] that is one of the most species-rich families of vertebrates
[...]
Phylogenetic reconstructions are consistent with cichlid origins prior to Gondwanan landmass fragmentation 121-165 MYA
[...]
The karyotype formula 2n = 48 st/a [subtelo/acrocentric] elements is characteristic of Perciformes
[...]
Although there is extensive variation in the karyotypes of cichlid fishes (from 2n = 32 to 2n = 60 chromosomes), the modal chromosome number for South American species was 2n = 48 and the modal number for the African ones was 2n = 44.
[...]
Pericentric inversions are thought to be the main mechanism contributing to changes in the basal chromosome arm size of Perciformes. Other mechanisms of
chromosomal rearrangement and translocation probably have contributed to the karyotypic diversification of South American cichlids. The chromosome number variation observed in some species suggests that events of chromosomal translocation followed by chromosome fission and fusion were also involved.
(Poletto et al., 2010)

Like already observed with horses, increased chromosome counts rather represent an evolutionary stage of change rather than a new equilibrium. Kolnicki’s karyotypic fission theory that ‘ posits that all mediocentric chromosomes simultaneously fission’ is just a variation of the theme:

That chromosomal diversity of such distinct taxa is explicable by fission implies this mode of animal evolution is important.
[...]
Increases up to nearly doubling of smaller derivative chromosomes throughout the Cenozoic underlie adaptive radiations, at least in artiodactyls, carnivores, lemurs, Old World monkeys, and apes. (Kolnicki, 2000)

This is not the place to discuss the general applicability of Kolnicki-type processes, except that Müller’s hypothesized ancestor of the lesser apes (Hylobatidae or gibbons) may now be attributed too lightly to an amazing diploid chromosome number of 2n = 64. For clarity I simply discard any existing intention to derive great ape chromosomes from a similar hypothetic ancestor, confident that in the same effort nobody would try seriously to derive monkeys from great apes. All gibbons share the same three chromosomal fission events with other apes that compare with macaque chromosome numbers 2, 7 and 13 and the corresponding fissioned pairs: for humans chromosomes 7 and 21, 14 and 15, 20 and 22 respectively. This detail strongly pleads for an ultimate derivation of gibbon chromosomes from the same macaque-like ancestor having a reduced set of chromosomes (2n = 42), as well as from the same ancestral ape having 2n = 48 chromosomes like great apes. Then subsequent changes should have reduced the count for Hoolock genus to 2n = 38, an even lower diploid chromosome number than macaques, while for other gibbon genera the number is higher: 44 (Hylobates), 50 (Symphalangus) and 52 (Nomascus), respectively. This implies that all additional ape-related fusion events on chromosomal level, as observed for humans and gibbons, were preceded by these three fission events that are shared by all apes.
Fission is one way to trigger accelerated recombination, and to increase plasticity. Hybrid-driven recombination is another, whether or not in combination with the mechanism of fission. Reversely, biological justification of chromosome fusion may rather be found among the advantages of synteny, ie. the physical co-localization of genetic loci on the same chromosome. It is suggested that genetic change by hybridization is not an overnight process, and may create instability on chromosome level for at least a few generations. Hybridization of chromosomes and homogenization processes, per definition occurring after considerable divergence, opens up a whole new array of possible combinations, but also new risks of meiotic nondisposition. Natural selection could become a cumbersome, even impossible strain to reproduction if also the whole array of possible deleterious combinations would eventually result in low-fitness birth. Reproductive consequence would be less for miscarriages, or just lower rates of viable gametes. Still, even the fecundity of plants would be harmed by the possibility of unlimited recombination, not in the least for the reproductory investments of cycads that also have the record for the world’s largest sperm. Encapsulation of heterozygous content in fused chromosomes could reduce this risk to future generations. It goes without saying that fusion of chromosomes also offers a better protection against cytogenetic instability of hybrids. As such, fusion of chromosomes emerges as a potential strategy towards endogenous purifying selection and increased fitness of the hybrid lineage.
An example of chromosomal instability is trisomy, that typically results in miscarriage, or low-fitness offspring. Human low-fitness births related to trisomy are the Patau syndrome (trisomy 13), that affects somewhere between 1 in 10,000 and 1 in 21,700 live births, the Edwards syndrome (trisomy 18) that occurs in approximately 1 in 3,000 conceptions and half this rate for live births, with a median lifespan of 5–15 days, and the Down syndrome, whose incidence is due to trisomy 21 for 95% of the cases, and estimated at one per 800 – 1000 births.

Trisomies of all chromosomes with the exception of chromosome 1 have been reported in spontaneous abortions in humans; however, the only numerical autosomal anomalies surviving to birth are trisomies 13, 18, and 21. There are only six reported cases of autosomal trisomies in live horses ([...] 23, 26, 27, 28, 30, and 31) [...]. Similar to that observed in humans, trisomies in horses predominantly involve small chromosomes. (Brito et al., 2008)

Human trisomy indeed involves chromosomes that are either small (18, 21), or with low gene density (13), apparently important preconditions against a spontaneous abortion. Despite the deleterious nature of the three trisomy-ridden human chromosomes mentioned above, these already existed before the lesser apes split off from great ape ancestors. Chromosomes 13 and 18 must have been present even in the common ancestor of apes and macaques. Only different gibbon genera managed to neutralize these potential dangers by fusion: in Hylobates the equivalents of human chromosomes 13, 18 and 21 are all fused (respectively with chromosomes equivalent to 3, fissioned remnants of 1 and 15).

Even though Hominoidea chromosomes 13, 14, 15, 18, 20, 21, and 22 constitute a single, uninterrupted chromosomal block in the lar gibbon, most of them are part of larger chromosomes and/or show internal rearrangements. (Misceo et al., 2008)

Such evidence may indicate that chromosome fusion is neither coincidental nor imperative per definition. This is especially interesting for the unique fusion of two ancestral great ape chromosomes into human chromosome 2.

The human chromosome number 2 is a fusion of two great ape chromosomes. Displayed are respectively the versions of humans, chimps, gorillas and orangutan. The banding pattern of chimps equals the human version most.

This fusion was evidenced by similarities in chromosome banding patterns as well as homologies in DNA sequences, where chimps make the best match. Normally, the extreme ends of chromosomes (telomeres) form a dynamic buffer against loss of internal sequences and prevent chromosomes from fusing, but apparently here telomeric DNA was involved in the fusion, to the extend that some telomeric DNA was preserved at location 2q13 near the new centromere.

Comparative cytogenetic studies in mammalian species indicate that Robertsonian changes have played a major role in karyotype evolution [..]. This study demonstrates that telomere-telomere fusion, rather than translocation after chromosome breakage, is responsible for the evolution of human chromosome 2 from ancestral ape chromosomes. (IJdo et al., 1991)

Somehow this event set us apart from great apes. As the assigned number indicates, chromosome 2 is our second largest chromosome. Commonly described as important for cognitive capacities, this particular fusion implies the importance of synteny for imposing genetic stability on cytogenetic level, that exceeds protection against semi-viable trisomy.
Evolving species may be expected to rely on an evolutionary boost to give them an edge. Since non-deleterious point mutations are rare, much depends on a quick and adequate mechanism to experiment with new complex DNA combinations. Species having low effective population sizes may have a problem if biological divergence is insufficient for regular recombination to produce successful genetic results. At major adaptive radiation events a species could recur to fission as a strategy to compensate for an initially low biological divergence. Next, increased divergence by the process of adaptive radiation may be expected to eventually reduce the need for fission – unless effective population sizes remain too low for harboring major biological variety. Lesser apes may have continued on the strategy of fission where great apes didn’t, implying their divergence probably occurred right here. Instead the lineage of great apes, including hominines, could take advantage of hybridization to incorporate successful mutations in the genome, no matter where these originated. This mechanism must have been a possibility within a certain time window, somewhere between population divergence and irreversible speciation. An extended continuation of this process may have relied on sub-species cross-breeding and hybridization – ie. contrasting with recurrent vicariance events as cited in the case of the gibbon genera, where low effective population sizes urged for continuous cytogenetic change in order to cope with the needs of environmental adaptation. Either way such periods of adaptive radiation should have left traces in the human chromosomes, not in the least if true hybridization was involved:

Contrary to the view that hybrids are lineages devoid of evolutionary value, a number of case studies are given that show how hybrids are responsible for reticulate evolution that may lead to the origin of new species. Hybrid evolution is mediated by extensive genome repatterning followed by rapid stabilization and fixation of highly adapted genotypes. Some well-documented cases demonstrate that bursts of transposition follow hybridization and may contribute to the genetic instability observed after hybridization. The mechanism that triggers transposition in hybrids is largely unknown. (Fontdevila, 2005)

Geneflow within the genetic continuum of a species happens all along, but let’s ‘[...] define hybridization as gene flow between two populations after an isolation barrier has formed between them.’ (Patterson et al., 2006 Supp.). The latter is most likely a major element in evolution, still waiting for recognition. And a potential cul de sac since, contrary to equids, zamia and gibbons, chromosomal changes in great apes and humans are far from impressive. No chromosomal fission event postdated the divergence from lesser apes, not even in hominin-specific evolution. Genetic recombination was significant enough for rapid change and plasticity, but apparently without the need for massive reorganization on chromosomal level, or even significant mutational activity on genetic level. Segmental duplications make a remarkable exception:

Although [the genomes of] terminal hominid lineages show an excess of [segmental] duplications, the most significant burst of activity (4–10-fold [...]) occurs in the common ancestor of human/chimpanzee and gorilla and after divergence of gorilla from the human–chimpanzee lineage [...] We note that this burst of duplication activity corresponds to a time when other mutational processes, such as point substitutions and retrotransposon activity, were slowing along the hominoid lineage. (Marques-Bonet et al., 2009)

Duplication indeed contributes to diversity, though indicative of actual genetic activity rather than anything else:

Gene models associated with signal transduction, neuronal activities (for example, neurotransmitter release, synaptic transmission) and muscle contraction are significantly enriched in human, chimpanzee and orang-utan lineage-specific duplications [...]. Human and great-ape shared duplications or those shared with macaque are, in contrast, enriched for biological processes associated with amino acid metabolism [...] or oncogenesis (Marques-Bonet et al., 2009)

Some sort of genomic destabilization is implied ‘at a period before and perhaps during hominid speciation’. But why and how this process could have been so different for humans, compared with an evolutionary strategy towards increased chromosomal change for lesser apes?
Exchange of alelles between hominine subspecies is nowadays sufficiently attested in modern genome research, but cross-breeding of diverging subspecies – or species! – should suggest chromosomal change of a higher order. Indeed, at least something has happened on chromosome level since the human divergence from apes, most notably the reduction of the chromosome count from 48 to 46, or the unique X-transposed region of the human Y-chromosome, dated right after chimp divergence and a virtually unprecedented event all by itself.

A third sequence class in the human MSY euchromatin — the X-transposed sequences — has no counterpart in the chimpanzee MSY. The presence of these sequences in the human MSY is the result of an X-to-Y transposition that occurred in the human lineage after its divergence from the chimpanzee lineage (Hughes et al., 2011)

Also chromosomal changes in chromosome 7, one of the new ape chromosomes that originated by fission, could be mentioned: the chromosome was subject to a pericentric inversion (including the centromere) after the divergence of orangutan, followed by a paracentric inversion after the divergence of gorilla. Another translocation of genetic information from chromosome 15 to chromosome 4 has been documented only for African apes: gorilla suffered deletions that preclude proper interpretation, but all derived basepairs of the 4q copy in chimps indicate this to be the result of hybrid recombination that happened before the divergence of gorilla:

If the duplication was followed by speciation and independent accrual of mutations, we would expect to find the human 15q and chimpanzee 15q copies to show higher identity to each other than either does to the copy found on chimpanzee 4q. Instead, the two chimpanzee copies are the most closely related pair of this trio.
[...]
By comparing ~19.1 kb of hand-curated, well-aligned block-5 sequences, we find that the chimpanzee 4q and 15q copies are only 1.43% diverged (Jukes-Cantor adjusted). They also share 43 derived mutations, including a 4-bp deletion that disrupts the ORF of gene H, not seen in the macaque or human 15q copies.
[...]
In contrast, the human 15q and chimpanzee 15q copies are 1.65% diverged and share 22 derived mutations not seen in chimpanzee 4q; and the human 15q and chimpanzee 4q copies are 1.64% diverged and share 13 derived mutations not seen in chimpanzee 15q.
(Rudd et al., 2008)

Such macro genetic events may be of an entirely different category than the published evidence that involve autosomal admixture of genetic regions from different hominine subspecies (Neanderthal, Denisova), or even of genetic harmonizing on the X-chromosome, already quoted as valid evidence for true cross-species genetic exchange with chimpanzee ancestors. Still, they are only in modest agreement with the prospected results of radical hybridization.
Notwithstanding an extended period of great ape evolution, lasting millions of years since the divergence of lesser apes, it has all appearance that over time hybridization remained a gradual process and rarely exceeded the level of subspecies crossbreeding.
Ever since this last major hybridization event(s) between chimp and human ancestors the hominine lineage apparently entered an extended period of rapid development. A full discussion is not the context of this article, but we can be sure nature did its utmost to exploit all available intra-species diversity to experiment with genetic recombination. There is an increasing awareness of natural interbreeding and admixture on subspecies level, whose impact on human evolution genetic science only started to disentangle. Another challenge still awaits us in the preposition that hominines could be essentially more related to orangutan than chimpanzees. Genetically this doesn’t seem right, but a successful hybrid is also a collation of DNA and morphology that rather define a new composition than an average of parental features. Current evidence corroborates to the implication that the genetic result of hybridization isn’t even random:

[...] endogenous selection is acting against intermediate hybrid individuals, that is, those that contain the highest number of alien genetic elements (Arnold, 1997). In a similar way, Rieseberg et al. (1996), working with H. anomalus, found that similar linkage groups of genes exist in several artificial hybrid lines with high fertility. (Fontdevila et al., 2005)

All we could propose is a complicated pattern of cross-breeding events to close the evolutionary gap of up to 16 million years since common Griphopiths precursors of great apes started to diverge.

Griphopithecins are the first cosmopolitan hominoid taxon, probably as a result of their powerful jaws and teeth that allowed them to exploit a wide variety of resources.
[...]
I see the entire region from Germany and Turkey in the north to Kenya in the south as a potential core area in which early hominids could have evolved. But there are major gaps in the record. For example, one species of Kenyapithecus is known from 16–16.5 Ma in Turkey and another from Kenya at about 13.5 Ma [...]. It was probably present elsewhere in the intervening interval of time but we have not yet found the fossils. From this core area these stem hominids (not specifically related to either living group of hominids, pongines, or hominines) eventually split, with one segment of the distribution of species dispersing to the north and east and another to the north and west. The causes of this dispersal are unknown, but griphopithecins are the most primitive hominids we know. The later-occurring sivapithecins of Asia and dryopithecins of Europe are more modern, and strong cases can be made that they are related to living orangutans and African apes and humans, respectively. (Begun, 2010)

If both components indeed represent sister lineages of apes rooted in the Miocene, it would be taunting to bolster this almost impossible preposition with genetic evidence.
I suggest the fuss about our purported lack of relatedness with the apparently quite basal gibbon genera might give us an important clue. If both great ape genera acquired their humanizing features by an extended period of hybridization, their common ancestor may indeed remind us to primitive small apes:

[...] Griphopithecus and its relatives retain primitive postcrania. They are more monkey-like than ape-like [...] without any indications of the suspensory capabilities of all later fossil and living great apes (Begun, 2010)

Cytogenetic evidence cited above reveals that chromosomes of small apes share the basic pattern that set great apes apart from macaques and thus may derive from an ancestor having eg. orangutan-like chromosomes. From this point onwards small apes can be defined as monophyletic, what may be confirmed by genetic evidence:

Among hominoid primates, gibbons alone contained Alu elements in their EIF1AY gene of the Y chromosome. (Kyung-Won Hong et al., 2007)

There have been different interpretations of the hylobatid evolutionary history regarding the four different gibbon genera, but: ‘Maximum likelihood and Bayesian analyses support Hoolock as the most basal, and both molecular and karyological studies have supported this alternative’ (Israfil et al., 2010).

Of all gibbon genera the genetic distances of Hoolocks (Bunopithecus), the most basal group, are closest to the human and chimp outgroups.

[Griphopiths] are more monkey-like than ape-like, as is Proconsul, in having fore and hind limbs of roughly equal length, without any indications of the suspensory capabilities of all later fossil and living great apes (Begun, 2010)

The divergence of the lineage leading to small apes must have happened long before at least two parallel lineages of great apes ‘humanized’ together with hominines. In the fossil record brachiating of great apes seems to develop first within the Dryopith clade, and at this stage one might wonder what could be the involvement here of a gibbon-type introgression event. Contrary to our Griphopith ancestors, modern humans and great apes retain many physical characteristics that suggest a brachiator ancestor, including flexible shoulder joints and fingers well-suited for grasping.

Dryopithecus [...] is known from postcranial remains, which are dramatically different from those of the griphopithecins and Proconsul. They show unambiguous indications of the importance of highly mobile limbs and suspensory positional behavior
[...]
I interpret this change to be extremely important in the evolution of the African and human clade. It allowed Dryopithecus to remain relatively large and yet retain the capacity to exploit terminal branch resources, by spreading its weight among the branches and by hanging below them to conserve energy [...]. It also represents the evolutionary origins of human mobile and highly dexterous upper limbs.
In addition to being relatively primitive compared to later species, the teeth of Dryopithecus differ from those of the griphopithecins in having a thin layer of enamel and less rounded cusps. They more closely resemble the teeth of chimpanzees and have been interpreted as adaptations to a soft fruit diet, as in modern chimpanzees [...]. The later occurring dryopithecins Hispanopithecus, Rudapithecus, and Ouranopithecus share even more postcranial derived characters with living great apes (Begun, 2010)

However, again the molecular dates remain difficult to reconcile with the fossil record:

The appearance of Dryopithecus at about 12 Ma parallels the first appearance of Sivapithecus at nearly the same time, suggesting that they diverged from a common ancestor possibly 13 to 16 Ma. (Begun, 2010)

The great ape-gibbon split simply doesn’t concur with the gibbon radiation dates:

Based on a consensus estimate of 15 Ma for the great ape-gibbon split, Chatterjee (2006) undertook molecular clock analyses using cytochrome b gene data and suggests the gibbon radiation dates to approximately 10.5 Ma. (Chatterjee, 2009)

This is still considerably less than the Raaum et al. (2005) estimate of a ‘divergence date of 15.0–18.5 Ma based on the entire mitochondrial genome’ (Chatterjee, 2009), thus apparently contradicting the possible hybridization signal mentioned above.
Molecular clock analysis is often used to estimate the date of speciation events in evolutionary history, but what can be said about its reliability? It didn’t come as a big surprise that paleoanthropologists were reluctant to abandon all the views they still cherished immediately for the sake of biochemistry:

[Wolpoff] admits [...] that ramapithecines are also a stem group for the African apes, whose tooth enamel is very thin. This would be a full reverse evolution towards a dentition very similar to that of Dryopythecus. Is this a parsimonious hypothesis? (Bonis, review on Wolpoff, 1982)

Schwartz criticized newly adapted scenarios ‘defending presumed phylogenetic hypotheses rather than rigorous presentations of such hypotheses’, that essentially left the purported ‘reversal’ in dental characteristics of chimps and gorilla back to the primitive conditions of Dryopithecines, unanswered.

I am not swayed by blanket statements of how similar Pan and [hominines] are because most of the similarities appear to be primitive retentions, and I am so far unpersuaded by karyological and biochemical studies for similar reasons as well as others.(Schwartz, review on Wolpoff, 1982)

Even Wolpoff, trying very hard to conciliate paleoanthropology with molecular evidence, couldn’t help to incite Sarich by stating:

Probably the best way to summarize the very disparate points raised is that the “clock” simply should not work [...] Consequently, although biochemical evidence seems to support a late Homo-Pan divergence, I believe this is a red herring, and that the molecular “clock”does not support any divergence time (Wolpoff, 1982)

One main issue that remains to be answered is how selective processes may have biased the genetic distances between species, and their biochemistry that, as a matter of fact, directly reflects evolutionary changes of their DNA.
Indeed, the molecular “clock” turned out unreliable because of variation ’caused, in part, by uncertainty or assumptions in key parameters, such as divergence times between species, generation times and ancestral population sizes’ (Conrad et al., 2011). But new results on the very mutation rates themselves indicate that apparently there is more internal logic to mutations on chromosome level than mathematicians were able to assume or deal with. Mutation rate differences exist and already turned out to be essentially individual:

[...] future studies promise to revolutionize our understanding of mutation processes and how they vary between individuals and between families as a result of age, genetic background and environmental exposures (Conrad et al., 2011).

This insight came too late for the immediate resurrection of our theorized ‘dental-hominoid’ ancestor, though in some act of posthumous generosity variable mutation processes should certainly provide for more nuance. How some basal species – humans, chimps and … hoolocks? – could remain genetically more ‘related’ on a molecular level than warranted by their morphological distance? Molecular constraints dictated that African apes and the hominine lineage should share a recent common ancestor, but the underpinning assumptions are still full of inconsistencies. ‘Single recent origin’ may be an interesting null-hypothesis to geneticists, but to paleoanthropology this increasingly reverts into the same unmanageable preposition as before. Molecular evidence is still a pretext to challenge common sense where it may oblige human minds to search for answers about themselves within a limited time window, that in turn implies a limited scope of geography. Adding here the constraints of emotional inquisition and lurking nationalism inherent to any limited region of choice, we could conclude ironically that paleoanthropology still finds itself right in the middle of all the fuss where Darwin’s theory of evolution once started.

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• Kivell et al. – Australopithecus sediba Hand Demonstrates Mosaic Evolution of Locomotor and Manipulative Abilities, 2011, link
• Conrad et al. – Variation in genome-wide mutation rates within and between human families, 2011, link, or try here. Supplement
• Chatterjee – Evolutionary relationships among the gibbons: a biogeographic perspective, 2009, link
• Scott A. Williams – Morphological integration and the evolution of knuckle-walking, 2010, link

Recommended:

• Reich laboratory – Population Genetics & Medical Genetics, link
• Anthony J. Olejniczak – Paleoanthropology & Dental Morphology, link
• Overview Human Leukocyte Antigen: HLA – Wikiversity
• John Hawks – Malapa and the “problem” skull KNM-ER 1813, 2010
• John Hawks – The trouble about Kenyanthropus and Ardi, 2009
• John Hawks – What, if anything, is Australopithecus sediba? 2010
• John Hawks – The immune systems of archaic humans, 2011-06-17
• John Hawks – How widespread is Denisovan ancestry today? 2011-11-01
• John Hawks – HLA class-I loci in Neandertals and Denisova, 2011-08-25
• Dienekes Blogspot – 16-12 ka humans with archaic features from Iwo Eleru, Nigeria 2011-09-16
• Dienekes Blogspot – Inference of ancient human demography from individual genomes (Gronau et al. 2011), 2011-09-19
• Verhaegen – Morphological distance between australopithecine, human and ape skulls, 1996, link

The La Tène culture developed in the "warrior fringe" northwest of Hallstatt.

Left: the older futhark. Right, top: Negau helmet; inscription; inscription according to Must; Below: impression of ancient greek alphabet variation

The Harii, besides being superior in strength to the tribes just enumerated, savage as they are, make the most of their natural ferocity by the help of art and opportunity. Their shields are black, their bodies dyed. They choose dark nights for battle, and, by the dread and gloomy aspect of their death-like host, strike terror into the foe, who can never confront their strange and almost infernal appearance. For in all battles it is the eye which is first vanquished.

On the way back through the desolate heath, Odin came upon a leafless tree. Suddenly, his coat was caught in the branches of the tree. Odin hung between heaven and earth. In vain, he tried to free himself. Odin struggled with himself for the ultimate wisdom. Nine nights he hung on the windswept tree. His inner being gradually grew clearer and more luminous. Now he finally found the symbols of life’s noblest values. He bent down deeply from the tree. Groaning with extreme exertion, he took up the signs and cut them into the trunk with his sword (Rúnatal or Óðins Rune Song in the Poetic Edda in the Icelandic Konungsbók)

The earliest runes resemble scrambled messages similar to those attested in Graeco-Roman curse tablets, essentially private in the act of devine communication and hardly fit for developing a public tradition of writing in the runic script. Accordingly, the 4th century Gothic Alphabet created by Ulfilas (or Wulfila) for the purpose of translating the Christian Bible, is thought to have been intended to avoid the use of the older runic alphabet, as it was heavily connected with heathen beliefs and customs. Ulfilas didn’t avoid runic names for his letters, though, and while most of the letters were taken over directly from the Greek alphabet, a few were derived from runic letters to express some unique phonological features of Gothic. Possibly it was just the cryptic nature of the runic tradition that inhibited a mundane use. This detail may be important to discover the timespan and ultimate whereabouts of the runic script and its predecessors.
The first Germanic tribes entered the full light of history in their conflict with the Romans during the Cimbrian War (113-101 BC). The supposed Northsea origin of the participating tribes (Cimbri, Teutons, Ambrones) was a recurrent argument in favour of the original confinement of Germanic people in the northernmost parts of Europe, far away from the purported North Italian source of the runes. Still, it remains difficult to distinguish some of the traces left in the mythology that surrounds the La Tène expansions of protohistory, from a linguistic source akin to Germanic, and rule out Germanic participation in alliance with their Celtic neighbours. Belovesus, the legendary king associated with the earliest Gallic expansions into Italy, was an exponent of the Biturges, well south of the Belgic origin of La Tène between Marne and Moselle (and south of the Belgic Belovaci). His tribe was closely associated with the Druid center in the territory of the Carnute, a Celtic tribe that also travelled with him. His cousin Segovesos took a more northern route along the Alps. A mythological reference of two different ethnic components? The prefix Bel- is widespread in Celtic, but Sego- is ambiguous: this same IE forms have only been encountered in Germanic and Celtic vocabularies. Indeed, northern contigents of La Tène crossed Slovenia when they continued to the Balcans. The name of one of their leaders appears an indication that Belgae were still involved in La Tène expansions as late as 279, when a Bolgius led the invasion in Macedon and Illyria. It is tempting to imagine the different impact of northern and central Italic alphabets on the parties involved, except for the inconvenient circumstance that the Negau B helmet – like the obviously Celtic inscriptions of the Negau A helmet – attest a predominantly central Italic “Etruscan” influence. Strikingly, the development of some letters used at Negau B already diverged where runes rather tend to represent more archaic versions of the alphabet. However, what matters most in this scenario is that La Tène representatives already appear receptive to the use alphabetic scripts. This makes the transmission of writing, from any source that La Tène came in contact with (including North Italic and later Koine Greek), feasible from 400 BC onwards. This Negau helmet decorated with a purported religious “Germanic” inscription, apparently mixed with Italic and associated with finds having Celtic inscriptions, already appears indicative of a melting pot all by itself rather than the result of successive occupation.
The supposedly Germanic inscriptions of the Negau helmet were considered La Tène evidence of Germanic expansions before, especially since the helmet alone was dated to the 5th-4th century. Why presume (too) old fashioned helmets to the authors of the inscriptions? A quantum leap of some interpretors to the 2nd century for dating the actual inscription appears most of all a courtesy to Roman references pointing at a Germanic-Cimbrian presence in Noricum at 113 BC. Even benign dating of the inscription in the 3rd century would have required the helmet to be at least one century old at the moment when it inspired an occasional priest of an ambulant Germanic warrior tribe (!) to carve the text. The priest, according to this interpretation, would have been obliged to continue an older tradition of carving similar inscriptions in another language, and for this purpose he had to find an obsolete helmet without a scratch. How likely this would be?
Slovenia was the stage of La Tène expansions as late as 250 BC. Note the Gallic invasions in the Balcans were dated slightly earlier in 279 BC, while La Tène Celts already crossed the Alps about 400 BC to enter Italy. Names like Brennus are trivial indications that both invasions were indeed carried out by the same people, most probably still at the same stage of development. A later date of La Tène in Slovenia than elsewhere suggests a nucleus of local culture that survived the first onslaughts of the Celts in the Alpine region, that nevertheless must have come in close contact with Gallic culture almost inmediately after the supply of Etruscan helmets was interrupted. Possibly this implies favorable conditions for the survival of North Italic scripts. In this sense, an attractive possibility is that La Tène innovations arrived in Slovenia only after a detour in the south.
There are several North Italic scripts, and by location the North Etruscan script could be considered one of them. Other North Italic scripts are the alphabets of Lugano, the Rhaetic alphabets of Sanzeno, Sondrio (including Camunic) and Magrè, and the Venetic alphabet.
Looijenga rather forgets about Negau at all and comes to a completely different route of North Italian scripts into the Germanic hemisphere:

An archaic North Italic alphabet may have been the precursor of the runes. Borrowing this alphabet may have taken place in North Italy or Raetia, where e.g. the Chauci, Batavi and other Germani served as Cohortes Germanorum in Germanicus’ army in 15, 16 and 69 AD. But, theoretically, Germanic mercenaries may have learned to write anywhere during their tour of duty.
Veterans from the Roman army, for the greater part originating from the mountanous parts of Piemonte and Lombardy (e.g. North Italy) settled in the region near Cologne in the first century AD. Soon they became integrated in the local population. Ubian and Italic elements were intermingled in the common cult of matres and matronae (Derks 1996:104).The indigenous matres cult of the Rhineland knew no votive inscriptions; this custom of writing dedications was introduced by soldiers of Italic and Germanic origin (Derks 1996:75). Here we may find a clue as to how an archaic North Italic alphabet came to the Rhineland. In the first century AD, several letters, known from North Italic archaic alphabets, are still in use in the Rhine area. (Looijenga, 1997)

Strange enough Looijenga can’t decide between a single archaic precursor, or a North Italic mixture that all came together in the Roman army. Here ends all logic, since the Roman veterans in the Rhine area comprised many more nations that could have contributed to the development of a new script. It doesn’t make sense this privilege to contribute was selectively restricted to North Italian veterans, all provenient of such a wide variety of North Italian cultural areas – not even when this selectivity was dictated by the shared cult of matres and matronae. The other way round, why Germanic cohorts would shop around in northern Italy and make a mix rather than retrieving their inspiration from a single source? Needles to comment that Northern Italy wasn’t the only region within the Roman Empire where people had already developed their own peculiar version of the alphabet: where the skilled input of Egyptians, Etruscans or Paleohispanics go, to name just a few?
The presence of an Etruscan Chi (Psi-sign Ψ, derived from a Western Greek/Euboean script, pronounced /Kh/ or /g/) and, strikingly, the very rare Eastern Greek Xi (///) in the Negau B inscription make it very unlikely that Negau B is a “recent” precursor of the runic script. The latter may insinuate direct contacts with Koine Greek at the time of inscription, where the runic script employed the sign *gebo, rather a derivation of pre-Koine “Eastern” Greek “Chi” (Chi-sign X, pronounced /g/).
The origin of the runic *gebo is a problem all by itself. The shape resembles the Latin X, but phonetically the letter suggests another origin. The Latin X has a southern Etruscan precursor of an ultimate Western Greek origin, originally pronounced “Ks”. However, already in early Etruscan this was a sibilant and finally it disappeared altogether in this language, thus implying the intermediate Etruscan role in the development of the Latin X (and indeed of Latin in general, together with other Italic equivalents) must have been ancient and short-lived indeed. Equivalent X-shapes in Rhaetic most likely developed from Northern Etruscan T, pronounced /t/, though opinions may be divided here. Only the Messapic alphabet, much to the south, is agreed upon to have been derived differently and features the non-western Greek X pronounced /Kh/ up to the first century BC. This implies the enigmatic situation that any other forerunner of runic “Chi” (X, /g/) or *gebo than Attic Greek could be problematic. Moreover, even a Greek derivation would be problematic if it happened later than 300 BC, when the original pronunciation /Kh/, necessary to yield the runic sound /g/ of *gebo, disappeared altogether in Greek Koine. Even 3rd century La Tène expeditions as far as Greece are anachronistic for such an early stage of borrowing. The Negau B Ksi (///) might have been the hypothetic consequence of late Classical Eastern Greek or Koine contact during La Tène, but not the runic X. The Western Greek/Euboian Chi (X as /Ks/) disappeared from the Etruscan alphabet, but was continued in Latin as X, while the Western Greek/Euboian Psi (Ψ as /Kh/) never made it to the Latin alphabet. Why should a purported Germanic forerunner of the runic script have incorporated a new /k/ sound that derives from Greek Ksi (///), but keep the Psi-sign Ψ for sound /g/ of Etruscan/Euboean derivation where the runic script would rather plead for a simultaneous acceptance of a new proto-runic Chi (X /g/) from the same source?
Of the runic script, the Euboean/Etruscan(/Rhaetic) trident Psi-Ψ /Kh/ apparently developed further to /-z/ *algiz that has the same shape – a development that for sure postdates Negau B. Looijenga’s assertion this already happened in (Neo-)Etruscan is by no means widely acknowledged. A position dependent softening of Ψ towards /z/ may eventually have made the new loan X /Kh~g/ necessary, but the purported Rhaetic or other North Italic ancestral alphabets don’t give a clear insight for such a process. Anyway, the disappearance of X in early Etruscan and its re-appearance in the runic script with a different sound value and without clear North Italic logic to back this up is striking enough.
The runic re-introduction of the sign X *gebo is even less natural than it may appear. Apart from the above mentioned aspirates, where we might add an otherwise irrelevant Western Greek variety of Ksi that was shaped like the standerd Greek Phi, the Etruscan alphabet already offered three other ‘K’ sounds. Latin accepted those sounds as “C” (like “G” derived from greek Gamma), K (derived from Kappa, very rare in Latin) and Q (derived from Phoenician Koppa, only present in the earliest Greek alphabets). Of this originally Phoenician collection the runic script only could have hooked on to the Latin C, that – like R – indeed appears to be a Latin derivation. Looijenga’s assertion that also the 3rd century Latin derivation G of this C is “clearly the base” of rune *jera /j/, requires some imagination, but it would boost her claim that Latin indeed had a huge influence on the development of the runic script. The proposed subsequent development of *eiwaz as a bindrune melting this transformed Latin letter G (*jera /j/) and I (*isaz /i/) into half a swastika is ingenuous, but raises additional questions about what must have guided so much creativity in such a short time.
Her proposal that *gebo /G/ derives from Latin X /Ks/ seems predominantly guided by a lack of alternative viable scenarios rather than scribe imagination or options. The proposed development of P (*perþ- /p/) out of B (*berkanan /b/) by means of what even she calls “quite a creative variation”, appears inconceivable for ancient runographers who “knew how to spell” and still didn’t know about the considerable variety of existing P-shapes readily available in all the alphabets they allegedly came in contact with. Looijenga’s creative solution to this problem merely indicates how she rejects out of hand any formative stage of the runic script that involved the earliest versions of North Italic scripts, or contact with Greece as a source of inspiration.
Actually, the runic P is most similar to the late Classical Greek or Koine “square” version of Pi, that only about 400 BC was on it’s way te become more popular: Camunic was the only North Italic script whose P resembled this Greek version (coincidence? Shared origin?), but it barely survived until Roman times. Another example where the runic script may be more similar to late Classical Greek or Koine is *tiwaz /T/ that is like a “curved” Tau whereas the Italic versions are either angular or in the shape of a cross.
We may conclude that in the North Italic hemisphere Negau B could emerge as a close relative of the runic script’s forerunner, on the condition that forthcoming influences from late Classical Greek or Koine related to La Tène expeditions deep into Greece are taken into consideration, as well as subsequent Latin modifications at an early stage (eg. the C, the R, possibly the G), thus leaving lots of time for an independent development that allowed the runic script to diverge so much from any other European alphabet-based script we know of. A Germanic presence in Central Europe during La Tène would explain the Germanic runic script well enough, except for the mysterious reappearance of X /g/ in the runic sign *gebo.

Modern proto-writing, resembling runes on the walls of farms and churches in Westhoek, French Flanders.

In the middleages almost all coins depicted a cross on one side. This penny of King Offa dates from c. 787 to 792 AD

Obviously a distinction was made between the mnemonical use of runenames, being a tool that enabled carvers to determine which sound a runic symbol had, and the meaning and use of symbolic runes, used as pars pro toto for some special purpose (Looijenga 1997, p.105)

In the hypothetical evolution of proto-writing towards the rune *gebo we can’t neglect intermediate stages to account for the symbolic decomposition of the Sun cross into “sun” and “cross”. One context where both symbols come together, or get separated, is “money”. Let’s investigate the feasibility of this scenario before recursing to the huge implications this would have for the location of the runic origin.
In the middleages almost all coins depicted a cross on one side, or when it didn’t the “cross” side was represented by the image of a king. The other side was “pila”, what still survives in english as “pile” (of money?), “wealth reckoned in terms of money”. Hence also expressions like croix ou pile (french); Kopf oder Schrift (german); cross or pile, heads or tails (english); kruis of munt (dutch). “Heads or tails” is a game with money, at which it is put to chance whether a coin shall fall with that side up which bears the cross, or the other called pile. The french expression “n’avoir ni croix ni pile”, “having cross nor pile” means to be broke.
In playing cards, Chinese decks used coins and different piles of coins as suits. When playing cards first entered Europe in the late 14th century the suits were very similar to those still used in traditional Italian, Spanish and Portuguese decks, that feature Swords, Staves, Cups and Coins. The shape of suit “diamonds” developed smoothly from the round “coin” shape, and the runic sign *ingwaz makes a perfect match with the diamond. Could this transformation have happened before?
Even nowadays the diamond shape is appreciated for its capacity to envelope other symbols, just like the Sun cross consists of the cross enveloped by a container symbol:

The red cross and the red crescent have been at the service of humanity for more than a century – affording protection to those affected by conflict and to those assisting them. In December 2005, an additional emblem – the red crystal – was created alongside the red cross and the red crescent. (International Commitee of the Red Cross, 2007)
–o–
Protocol Additional to the Geneva Conventions of 12 August 1949 [...], 2006:
Article 2 – Distinctive emblems
1. This Protocol recognizes an additional distinctive emblem [...]
2. This additional distinctive emblem, composed of a red frame in the shape of a
square on edge on a white ground, shall [...]
Article 3 – Indicative use of the third Protocol emblem
1. [Those] which decide to use the third Protocol emblem may [...] choose to incorporate within it [...]:
a) a distinctive emblem recognized by the Geneva Conventions or a
combination of these emblems; or
b) another emblem [...]

One peculiar detail about the *ingwaz rune was, naturally, its associated to the god Ingwaz, whose cult (according to Pliny the Elder, IV-28) was primarily located along the “Ingvaeonic” Northsea coast, still littered by eng- toponyms as far south as the Saxon Shore. His name was attested in several Germanic tribes, in mythical Germanic ancestors and also survived in larger regions like “England” and “Angria”. A connotation of the god Ingwaz (or Ingve in Norse cosmology) with wealth and money is hardly far fetched:

Frey took the kingdom after Njord, and was called drot by the Swedes, and they paid taxes to him.
[...]
Frey was called by another name, Yngve; and this name Yngve was considered long after in his race as a name of honour, so that his descendants have since been called Ynglinger. Frey fell into a sickness; [...] his men [...] raised a great mound, in which they placed a door with three holes in it. Now when Frey died they bore him secretly into the mound, but told the Swedes he was alive; and they kept watch over him for three years. They brought all the taxes into the mound, and through the one hole they put in the gold, through the other the silver, and through the third the copper money that was paid. Peace and good seasons continued. (Ynglinga saga)

Still, the very name of this rune is enigmatic since the cult of Ingwaz was already in decline long before the Migration Period. Indeed, the rune was already pretty rare among the earliest runes and was utterly absent in the Younger Futharc. In it’s original square form it was only found in two futhark inscriptions, suggested to be made for the sole purpose of practice or instruction in Scandinavian carvings of the elder futhark: the Vadstena bracteate and the Kylver stone, a Swedish runestone which dates from about 400 AD. How come that *ingwaz was teached but never applied? According to Looijenga its presence is uncertain the Opedal inscription, the only other possible source, but: “In semantically intelligible texts, it always appears with a headstaff, representing a bindrune.
The name of Ingwaz hardly survived in Germanic cosmology, but may have found an unexpected equivalent in Irish lore, where the cult of namesake Oengus is known in more detail: Oengus Mac IND Óc, a god excelling in youth and beauty (Koch 2006, p553). The possibility of early contact between Celtic and Old Germanic cultures emerges.

[Oengus] was born as a result of a liaison between Bóand and Dagda that occurred when Dagda sent Nechtan away. To hide their infidelity, they asked Elcmar – possibly an alter ego for Nechtan and for Nuadu Argatlám (Nuadu of the solver hand), both of whom may reflect aspects of the god Nodons – to become Oengus’s foster-father. (Koch 2006, p218)

Nodons was a god of healing, though his association with seafaring suggests a different origin, maybe not unlike the Germanic Njord, father of Freyr/Ingwaz; but so does the loss of his arm, that like the Germanic god Tyr may even suggest a more bellic “Mars” origin. This association could be expanded with the notion that the Celtic god of healing Lanus – commonly considered equivalent to Belenos, was venerated in Trier as “god of war” Lenus Mars (Derks, 2009). This rather unexpected qualities of Mars, himself being part of the Roman Archaic Triad along with Jupiter and Quirinus, have the potency to shed light on ancient religious traditions and practices, and possibly make the link between widely different kinds of devine qualities, behaviour and personifications in all related Indo European religions of Celts, Germans, Romans and even Greeks through the use of Gaulish belenuntia, Spanish beleno, the hallucinogenic henbane, whose stems and leaves are covered in fine white hear and whose Latin name is Apollinaris (Koch 2006, p.195).
It is widely accepted that most mythological parallels in the cultures of ancient European people are ultimately due to a common origin, whether Indo-European and linguistic or otherwise. Association of the Oengus cult with the Welsh Mabon / Gallic Apollo Maponos (< mapos ~ young boy) also implies an etymological/genetic link with the Batavian cult of their Hercules Magusanus ("The young-old one"), part of which name appears to be closer to Old-Germanic (< Germanic *magadi- young person) than to truly continental P-Celtic. Others (Toorians, Schrijver – Koch p.1194) would rather call Magusanus a Germanized version of a Celtic tradition. This association also implies a possible genetic link between Maponos and Apollo. Now Apollo was the Greek god of healing "par excellence", at least before this devine task passed over to his son Asclepius. Indeed Apollo remained depicted as a handsome beardless young man, like Maponos, but this didn't apply to what is arguably the most well-known Celtic god of healing "par excellence", that is Belenos. While clearly depicted as a solar god, like Apollo, his image was invariably an old man having a beard. Moreover, rather in agreement with the ambiguous character of Magusanus, Oengus wasn't even specialized in healing:

Oengus has less power than savage medicinemen or gods in myths, who bring the the dead back to life, or than Demeter, who gave life to Dionysos after he was dismembered by the Titans. But the story is an almost unparalleled example of a god’s love for a mortal. (MacCulloch, p67)

Perhaps it would not be too far fetched to see this as quite a different parallel to Apollo as “the god of Good”, maybe even love. A quality that in Germanic cosmology is only paralleled by Baldr, whose dead caused much grief. In Norse mythology his own brother Höðr shot the mistletoe missile which was to slay the otherwise invulnerable Baldr, only because he was misled by the evil Loki that knew his only weakness.
Notwithstanding some pan-Celtic parallels, there is something about the story of Oengus that raises the suspiction of being a later addition to the Celtic cosmology. Not only that he was raised by foster-parents, but he also arrived late when Dagda had already shared out his land among his children: there was nothing left for Oengus until he could recover everything by trickery. However, the example of Apollo shows that a cult can be very old and still foreign to the native cosmology. This would be true as much for the Germanic cosmology that involves Freyr/Ingwaz, and if Baldr belongs to this same category it would be because notwithstanding his popularity he failed to gain the power he might have been entitled to. Apollo, Oengus, Freyr, Baldr: they all remained the young promise they ever were.
The La Tène horizon was previously interpreted as almost exclusively “Celtic”, but the evidence of a smooth exchange of cultural elements demands a re-assessment of the Germanic identity. Part of this exchange may have been ancestral: Magusanus indeed seems to unite the young and the old, the son and the father. The reference of this Germanic cult to Hercules, not Mars, was unique in Roman times, and this hardly derives from a Baldr/Oengus Apollo. This association finds a parallel in Oengus’ devine father Dagda.

The Dagda has two fabulous material attributes, a cauldron of plenty, and a club that can kill the living and raise the dead. The latter has led the comparisons of the Dagda with Heracles and the Gaulish figure Sucellos. The club is so huge that it has to be dragged on wheels. (Koch 2006, p.554)

It becomes untenable to maintain that Germanic beliefs are irreconcilable to Celtic culture and that “true” Germanic territories “thus” should have remained isolated from La Tène influences. All the contrary, most likely such influence was much more important to the Germanic ethnogenesis than traditionally thought. Moreover, even Tacitus already mentioned Germanic tribes in Central Europea that roamed beyond agreed “Celticized” Chatti-like tribes in Central Germany: the Hari and Naharvali being the most peculiar groupings. None of these tribes are easily derived from fresh northern arrivals and in view of their habitat and peculiarities the mutual cultural contact zone is likely to have been much bigger.
We don’t get a straight Classical account on Celtic influences on the Germanic ethnicity, but at least Ceasar was already aware of the profound influences that Germanic tribes had in northern Gallia. Nevertheless, his geographic distinction between Celts and Germani was still a mystery to Strabo:

[...] the Germans, who, though they vary slightly from the Celtic stock in that they are wilder, taller, and have yellower hair, are in all other respects similar, for in build, habits, and modes of life they are such as I have said the Celti are. And I also think that it was for this reason that the Romans assigned to them the name “Germani,” as though they wished to indicate thereby that they were “genuine” Galatae [...] (Strabo, 7.1.2)

Romans knew how to speak their own language, and “germanus” is a Latin word that means “brother”. Romance languages like Spanish still preserve this word (hermano). The Germanic people explicitly received their name to indicate their relationship with their neighbours on the western side of the Rhine, though by doing so Ceasar managed to set them apart from the Gallic lands he managed to conquered. Later, Tacitus expanded on this and set out to exaggerate their “wilder” qualities to accomodate his ideological motive to pinpoint a selective group of people with the noble qualities the Romans could learn from:

The tribes of the interior use the simpler and more ancient practice of the barter of commodities.
[...]
The border population, however, value gold and silver for their commercial utility, and are familiar with, and show preference for, some of our coins. (Tacitus)

This account of Tacitus evolved into the modern myth that all Germanic ancestors of modern Germanic people ancestors could be easily distinguished from their “Celtic” neighbours by their relative lack of culture, while this was only partially true.

Left: early Germanic coins spread from Chatti territory to the west. Right: the coins are usually typified by a triskelion at one side.

The widespread distribution of sunken-featured buildings suggests varied and wide-ranging affinities and origins [like] in northern France, where they may derive from an indigenous, late Antique tradition, rather than representing an ‘intrusive’ Germanic element. Yet, although sunken-floored structures are known from the La Tene period in France, they did not appear in significant numbers until the Migration Period. The debate concerning their ethnic affinities is far from decided, although socio-economic developments seem more likely than Germanic expansionism to explain their appearance in northwest Gaul (H. Hamerow, 2002)

When in 1996 a gilt-silver scabbard mount with a runic inscription was found in Bergakker near Tiel in the Betuwe, well inside supposedly Romanized territory, the apparent knowledge of runic writing in this area came quite as a surprise.

At the same site a Roman altarstone was found [...] The stone, from the second half of the second or first half of the third century AD, was dedicated to the indigenous (Batavian) goddess Hurstrga. The toponym ‘Bergakker’ suggests that the site is higher than its surroundings. [...] The site may have functioned as a ritual centre during the Roman period. A parallel can be found at the temple site Empel (province Noord Brabant), which was dedicated to the Batavian god Hercules Magusanus.
[...]
The interesting thing of Empel was the occurrence of oaks, whereas elsewhere the area was dominated by a vegetation of willow. Together with the runic scabbard mount, a great number of metal objects were found, among which were many coins, fibulae, all sorts of bronze fragments and two objects that may be characteristic for cult-places, namely a small silver votive plate showing three matrones and a silver box for a stamp. (Looijenga, 1997)

The temple at Empel was in use since about 100 BC, when the difference between the strongly related Batavi and Eburones was still difficult to draw. Bergakker was dated 400 AD, when the Batavians were most probably already absorbed by early Frankish (or Salian) tribes. Never before runologists had considered the existence of a Frankish (Merovingan) runic tradition, though evidence from Charnay, Arlon, Amay, Chéhéry and maybe Kent indicated otherwise. Looijenga: “The Frankish king Chilperic (584) proposed the addition of four letters to the Roman alphabet, thus showing his knowledge of runes, since one of the four new letters, described: uui, was shaped after the runic w.” Gregory of Tours despised Chilperic, while his contemporary Venantius Fortunatus (6th century) defined the runic tradition as “foreign”: unfortunately both represented just the Gallo-Roman element of the Frankish empire.
The Merovingians, however, do not seem to have not developed an indigenous runic tradition, after they settled in former Gallia [...] The real powers of those days apparently did not use runes, but the Roman script. (Looijenga)
We still don’t know how or from where the runic tradition arrived in the Germanic world, but the early association to West Germanic, a tradition of coins, triskelions that are another development of the Sun cross and also an otherwise rich tradition of proto-writing, paves the way for a rejection of Looijenga’s stance that runes are due to Roman period parallels between the cults of the matronae in North Italy and the cult of the matres in the Rhineland. Actually, local triple goddesses like Nehalennia represent a very old local tradition that could do very well without an external source of inspiration. Rather, a La Tène setting of the earliest contacts in the wider neigbourhood of Northern Italy could point to exactly the reverse: a thorough early influence of Celto-Germanic influences in Northern Italy tentatively related to an introduction of certain Old Germanic people into the runic art, much earlier than ever imagined.

This Mercury of Soissons represents the old Celto-Germanic tradition of an overall triune god, representing the unity of almost all masculine divine powers. This religious design must have facilitated the intense cultural interaction reflected in Celtic and Germanic cosmology.

Indo-European languages have a long history in the neighborhood. At this stage it is important to be aware that at least the Celtic branch in the area, according to Cunliffe and Koch, was a product of Atlantic processes in the Later Bronze Age, ie. well after the Bell Beaker period that is commonly associated with early Indo-European presence. This as a caution against equating Bell Beaker to proto Celto-Italic and excluding more Germanic-like branches. Eg. according to toponym investigation, especially in Northern France, the glottal stop, an archaic IE feature, seems to have had a much wider distribution than warranted by the spread of IE languages actually associated with the glottal stop, that exclude the Celtic, Italic and Greek branches but include Germanic (Kortlandt, 1989).
As far can be deduced by this evidence, Northern France was rather inhabited originally by people whose language was reminiscent of Old Germanic than Celtic. However, since both Northern France and the Low Countries definitely belong to ancient Beaker territory, Roman accounts of Celtic presence in the north of Gallia, and possibly beyond, would find a logical explanation in an expansion of the Celtic influence towards the north that happened only after the Beaker period. This would be in agreement with Koch and Cunliffe, that date the origin of Celtic only in the Later Bronze Age. They make a case for maritime trade and contact of communities along the Atlantic facade to have culminated into a shared Celtic development, while the underpinning Bell Beaker period can’t be attributed anything else than an (important) contribution to a “Centum” dialectal continuum in western Europe, lasting well into the Bronze Age. For that matter, linguists identified an ancient Celtic north-south cline or continuum that apparently also extended to the Italic branch in the southeast, thus being older than the very division between Celtic and Italic. Eg. Portuguese Celtic features characteristics that apparently relate to Italic languages, while Peter Schrijver attributes the Celtic dialects north of the Alps to substratum rather than any other North European influence. In a Bronze Age proto Celto-Italic stage those mentioned Atlantic contacts could have made the difference in forging northern and southern dialects of the continuum together into one separate “Celtic” branch, diverging from “Italic”. Correspondingly, more to the north something similar could have happened within a “Northern Bronze Age” group of Germanic-like languages that not necessarily already featured all the known Germanic shifts.
The Celtic branch should not be considered the hierarchic offspring of a single standard language, once spoken by some kind of “true” original Celts. In general, the very concept of a single parent language for each Indo European branch is ever more controversial. Not even Latin (nor hypothetical “Vulgar Latin”) makes a perfect ancestor to the Romance languages, that rather originated from shared developments over at least half a millenium that predominantly postdate the Roman Era and even included Germanic influences. Contrary to what hypothetical assumptions into this direction may imply, there was NO single vulgar latin parent language within the Roman Empire. Vulgar latin is nothing more than a collection of texts that don’t follow the strict rules of classical latin. The relation of this collection with romance is still utterly incomplete, inconsistent and hypothetical. Its lack of unity starts in the Classical age, when the difference between literary usage and everyday speech in Rome was merely stylistic, not based on any difference between linguistic systems. Also much later, there is no indication that Latin had split up in two different languages, a spoken proto-romance vernacular next to a written language. Moreover, Vulgar Latin doesn’t even reveal a systematic tendency towards Romance, however deformed it may have been. There is nothing “Romance” about The Satyricon of Petronius, the only “vulgar latin” title of the only author mentioned by contemporaries to have used this kind of “incorrect Latin”. By then there was no trace of the Romance use of definite and indefinite articles. Without written evidence it is already hard to make the difference eg. with the Latin demonstrative, but this Romance feature certainly didn’t happen up to the very last century of the Roman Empire. Not even in case of doubt the position of the article/demonstrative was clear, so in Rumania the development could still be influenced by the Slavic substratum to put the article after the noun – thus suggesting a strong non-Roman convergence even here.
To expand on this example: neither the fate of the neuter follows a strict pattern in Romance that could be derived from a Vulgar Latin parent: neuter forms remain in a few of the Romance Languages, such as Italian (seen in the singular and plural of parts of the body, such as il braccio “the arm”, but le braccia “the arms”, taking the feminine plural definite article, but, at the same time, resembling a feminine singular noun). This is most apparent, in fact, in Romanian, where nouns classified as “neuter” resemble certain classes of masculine nouns in the singular and feminine in the plural. In French, the neuter forms were assigned to the feminine and masculine categories on quite different criteria. Even the case system can’t have been lost in the hypothetical Romance parent language since the plural of Romance north and west of the La Spezia-Rimini Line developed from the accusative case, while in romance south and east of the Line plural developed from the nominative case.
A less antagonized case than Romance against the dogma of a hypothetized single parent language was supplied by Garrett, in his rejection of the existence of proto-Greek as a parent language of Greek.

It is [...] well established that there are linguistic changes found in all first millennium Greek dialects, including Arcado Cyprian, that are not found in Mycenaean. Before the decipherment of Linear B such changes were assumed to be Proto Greek, but now it is clear that they reflect areal diffusion across the Greek speaking area.
[...]if we allow that at least a few post-Proto-Greek changes must already have affected Mycenaean before its attestation (it is after all a Greek dialect), detailed analysis reduces the dossier of demonstrable and uniquely Proto Greek innovations in phonology and inflectional morphology to nearly zero.
[...] it hardly makes sense to reconstruct Proto Greek as such: a coherent IE dialect, spoken by some IE speech community, ancestral to all the later Greek dialects. It is just as likely that Greek was formed by the coalescence of dialects that originally formed part of a continuum with other NIE (Nuclear Indo European, R.) dialects, including some that went on to participate in the formation of other IE branches. With this in mind it is possible to see external links for some Greek dialect patterns. For example, the first-person plural endings -mes and -men are distributed such that -mes occurs in West Greek, across the Adriatic from Italic (with s in Latin -mus), while -men occurs elsewhere, across the Aegean from Anatolian (with n in Hittite -wen).
[...]
I suggested that Greek may be typical of IE subgroups, and that the reason we see the pattern clearly in Greek is that we have Mycenaean. For no other IE branch do we have comparable data — an Italic dialect of 1000 JK, or an Indo Iranian variety documented early in the second millennium. But the coherence of other IE branches can be doubted too. The question of Italic unity has been debated by linguists for at least 75 years. Even for Indo Iranian [...]
I conclude section 2 by noting a pattern in need of an explanation [...] Speaking in the broadest terms, early IE language spread was thus a two phase process [...] in the second phase, late in the second millennium in some cases, changes that gave dialect areas their characteristic phonology and morphology swept across those areas. (Garrett, 2006)

Likewise, Koch does not suggest anything close to a specific “proto-Celtic” parent language, nor a limited point of origin for the Celtic branch. Instead, he holds the Celts to be the ultimate products of processes along the Atlantic coasts.
In this same vein, the Great Barbarian Conspiracy of 367, or the expulsion of Rome from Britain in a coordinated effort from Scottis in Ireland, Picts and Caledonians in Scotland next to Angles and Jutes, is exactly the kind of contact that would be key to a shared “Gaelic” development from Old Irish to Middle Irish, however radical the changes:

I have presented a relative chronology of 22 stages for the phonological developments which characterize the formation of Old Irish (1979). All of these developments are posterior to the Ogam inscriptions, which lack the characteristic features of the Old Irish language. If we use the term “Primitive Irish” for the period before the apocope (my stage 15) and the term “Archaic Irish” for the period between the apocope and the syncope (my stage 19), we may wonder about the applicability of the term “Irish” to the Ogam inscriptions; it may be more appropriate to speak of the variety of Insular Celtic spoken by the ancestors of the Irish. In any case, no reconstruction of Proto-Irish on the basis of Old Irish and later materials comes close to anything resembling the language of the Ogam inscriptions (Kortlandt, 1989)

Kortlandt interprets this rapid evolution as evidence for a younger age of Indo European. It is a pity it did not occur to him to relate this also to a much slower geographic evolution within the branches already in place.
Another example, albeit of another linguistic branch, would be the Frisian trade contacts that kept the Frisian hemisphere knit together with the Anglosaxons, what without doubt was key to a shared Anglo-Frisian linguistic development.
This shared development over this larger (already IE-ed and interrelated) area towards Celtic is “convergence”, made possible by a combination of shared origin and intensive contacts – that may or may not be largely confined to the Atlantic rim. This latter part is less clear at the moment, especially for what happened at the eastern “Hallstatt” borders. Similar processes towards convergence may have caused different linguistic configurations in the North sea region, or even within the La Tene “warrior fringe” between Marne and Moselle that is commonly regarded a key element to the historic La Tene identity. Such impulses deriving from the Celtic fringes thus also demand a new assessment.
How come it took so long for linguists to recognize this simple mechanism of convergence, still so important for understanding the origin of linguistic branches? Only nowadays the acceptance of the universal linguistic models of Chomsky and Greenbergis is in decline. This year investigators from Auckland and Nijmegen may have sealed their demise definitively with results that show languages evolve in their own idiosyncratic ways, and don’t have “simply” a natural tendency to “look the same”. There is no reason to suppose that grammatical structure and rules are governed by universal cognitive factors or “hardwired” by innate linguistic capacities. Instead of the cognitive explanations and universals supplied by Chomsky’s and Greenberg’s theories, cultural evolution is at play.

The central goal of linguistics is to describe the diversity of human languages and explain the constraints on that diversity. Generative linguists following Chomsky have claimed that linguistic diversity must be constrained by innate parameters that are set as a child learns a language [...]contrary to the generative account of parameter setting, we show that the evolution of only a few word-order features of languages are strongly correlated[...]
cultural evolution is the primary factor that determines linguistic structure, with the current state of a linguistic system shaping and constraining future states (M. Dunn et al., 2011)

Actually, Chomsky underestimated the processes of convergence since true “linguistic evolution” is incompatible with the rather creationist-like view of guided internal causes towards change. Parallel evolution never occurs because of internal processes, only as a response to external impulses. Other stale “collectivity theories”, like those that deny convergence in mythology and symbolism, may be equally flawed.
Now what about Old Germanic? Convergence makes the hypothetized Nordwestblock language feasible as a kind of Old Germanic that didn’t participate yet in some “key” Germanic sound shifts – even though the onset of each may have been elsewhere, not unlike the High German consonant shift possibly in another Celto-Germanic contact zone. Common conservative features like the glottal stop tend to set assessments of “Belgic” and “proto-Germanic” apart from “Celtic”, and the mechanism of linguistic convergence virtually dissipates the need for a compulsory set of shared “Germanic” features for Belgae with their eastern brothers (L. germanum). Some common features may have been the result of more recent convergence, and regional features may be retentions of regional variability of a wider “Old Germanic” branch. Even English retains sufficient traces of a Nordwestblock vocabulary, whether of not partially “in situ” as a development among Belgae immigrants, to suggest at least West Germanic has a much longer history than the limited concept of a Migration Period ethnogenesis ever intended to permit.

• Michael Dunn, Simon J. Greenhill, Stephen C. Levinson & Russell D. Gray – Evolved structure of language shows lineage-specific trends in word-order universals, 2011, link, naturenews
• Kurt Braunmüller et al. – Convergence and divergence in language contact situations, 2009, link
• Andrew Garrett – Convergence in the formation of Indo-European subgroups: Phylogeny and chronology, in Phylogenetic methods and the prehistory of languages, ed. by Peter Forster and Colin Renfrew, 2006, pp. 139-151, link
• J.H. Looijenga – Runes around the North Sea and on the Continent AD 150-700; Texts & Contents, 1997, link, or get version 2003
• J.T. Koch – Celtic culture: a historical encyclopedia, Volumes 1-5, 2006, link
• Kortlandt – The Spread of the Indo-Europeans, 1989, link
• Gustav Must – The Problem of the Inscription on Helmet B of Negau, Harvard Studies in Classical Philology Vol. 62, 1957, link
• Frans Plank – From Early Germanic Towards Early English, Evidence for very early Germanic, link
• Helena Hamerow – Early Medieval Settlements: The Archeaology of Rural Communities in North-West Europe, 400–900, 2002, link
• Strabo, Geographica 7.1.2., try here
• Julius Caesar – The Gallic Wars (D.B.G.), link
• Tacitus – Germania, Translated with Introduction and Commentary by J.B. Rives, 1999, link
• Gaius Petronius – The Satyricon, link
• Pliny the Elder – The Natural History, Germania: Book IV-28, John Bostock and H. T. Riley (1855), link
• Snorri Sturluson, Ynglinga saga, ~1225, link
• John Lindow – Norse mythology: a guide to the Gods, heroes, rituals, and beliefs, 2001. ISBN 0-19-515382-0
• Rudolf Simek – Dictionary of Northern Mythology, 2007. ISBN 0-85991-513-1, link
• Ross G. H. Shott – The Dark Arts of Immortality, 2004, link
• Bernard Mees – “Runic erilaR.” NOWELE: North-Western European Language Evolution 42 (March 2003), 41–68.
• John Arnott MacCulloch – Celtic Mythology (1918)), ISBN 0-486-43656-X, 2004
• Simon James – Exploring The World Of The Celts, 1993, ISBN-13 978-0-500-27998-4
• Ton Derks – Van toga tot terracotta: het veelkleurige palet van volwassenwordingsrituelen in het Romeinse Rijk, 2009, link

Useful links:
• Poinikastas: Epigraphic Sources For Early Greek Writing
• Les symboles dits runiques en Flandre
• International Commitee of the Red Cross – The history of the emblems
• Protocol Additional to the Geneva Conventions of 12 August 1949, and relating to the Adoption of an Additional Distinctive Emblem (Protocol III)
• F.A. Stoett, Nederlandse spreekwoorden, spreekwijzen, uitdrukkingen en gezegden (4e druk, 1923-1925), link
• kruimel, krümel, krumel; eng. crumble – dimunitive or frequentative, link
• Carl-Gustav Werner – The allrunes Font and Package (Version 2.1), 2004, link

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)

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 diachtomy 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.

Referenced:

• W. Haak et al. – Ancient DNA from European Early Neolithic Farmers Reveals Their Near Eastern Affinities, 2010, link
• B. Bramanti et al. – Genetic Discontinuity Between Local Hunter-Gatherers and Central Europe’s First Farmers, 2009, link; Supporting Online Material, link
• W. Haak et al. – Ancient DNA from the First European Farmers in 7500-Year-Old Neolithic Sites, 2005, link
• Joachim Burger et al. – Response to Comment on ‘‘Ancient DNA from the First European Farmers in 7500-Year-Old Neolithic Sites’’, 2006, link
• Deguilloux et al. – News from the west: Ancient DNA from a French megalithic burial chamber, 2010, link or try here
• Marc Vander Linden – What linked the Bell Beakers in third millennium BC Europe, 2007, link
• Chandler et al. – Using ancient DNA to examine genetic continuity at the Mesolithic-Neolithic transition in Portugal, 2005, link
• Sampietro et al. – Palaeogenetic evidence supports a dual model of Neolithic spreading into Europe, 2007, link
• João Zilhão – Radiocarbon evidence for maritime pioneer colonization at the origins of farming in west Mediterranean Europe, 2001, link
• Currat et al. – The effect of the Neolithic expansion on European molecular diversity, 2005, link
• Edmonds et al. – Mutations arising in the wave front of an expanding population, 2003, link
• Klopfstein et al. – The Fate of Mutations Surfing on the Wave of a Range Expansion, 2006, link
• Pablo Arias – The Origins of the Neolithic Along the Atlantic Coast of Continental Europe: A Survey, 1999, link
• Hans Peeters, Willem-Jan Hogestijn, Theo Holleman – De Swifterbant Cultuur, een nieuwe kijk op de aanloop naar voedselproductie, 2004. ISBN 90 6825 279 8
• L.P. Louwe Kooijmans – Trijntje van de Betuweroute. Jachtkampen uit de Steentijd te Hardinxveld-Giessendam, 1998, link
• Dirk Raetzel-Fabian – Absolute Chronology and Cultural Development of the Neolithic Wartberg Culture in Germany, 2002, link
• R. T. J. Cappers, D. C. M. Raemaekers – Cereal Cultivation at Swifterbant? Neolithic Wetland Farming on the North European Plain, 2008, link
• Palaeohistoria 41/42 (1999-2000): Institute of Archaeology, Groningen. J.N.Lanting & Van der Plicht – De 14-Chronologie van de Nederlandse pre- en protohistorie, III: Neolithicum, link
• The reseach programme ‘From Hardinxveld to Noordhoorn – From Forager to Farmer’, 2010, link
• Per Johansson – The Lure of Origins, An Inquiry into Human-Environmental Relations, Focused on the “Neolithization” of Sweden,2003, link
• Marek Zvelebil – Indo-European origins and the agricultural transition in Europe, 1995
• Marek Zvelebil – What’s in a name: the Mesolithic, the Neolithic, and social change at the Mesolithic-Neolithic transition, 1996
• Revedin et al. – Thirty thousand-year-old evidence of plant food processing, 2010, link
• Christian Jeunesse – Pratiques funéraires et sociétés Danubiennes au Neolithique ancien, 1998, ISBN: 2 87772 150 7 pp.41-58
• Milenko Bogdanovic – Particularism in the Proto-Starcevo Culture, 2009, link
• Kortlandt – The Spread of the Indo-Europeans, 1989, link
• John D. Bengtson – Materials for a Comparative Grammar of the Dene-Caucasian (Sino-Caucasian) Languages, 2008, link
• Arnaud Etchamendy – Thesis on the Baque language as an Indo-European language, 2007, link or try here.
• Scarre et al. – Long Mounds and Megalithic Origins in Western France: Recent Excavations at Prissé-la-Charrière, 2003, link.
• P. Schrijver – Keltisch en de buren – 9000 jaar taalcontact (Oratie), 2007, link.

The Neolithic advance was gradual but irreversible, creating a bond with land and property without precedence.

Of all paternal lineages that have been identified in Europe by studying the distinct Y-chromosomal haplogroups, probably the group commonly referred to as R1b has caused most confusion. The occurrence at high frequencies in western Europe and the estimated 110 million of European males currently sharing this R1b lineage, caused the initial consensus to be in favour of a very old, even paleolithic presence. This did not prevent early speculation about more recent connotations, but even the the most hilarious novel “genealogic” age estimates couldn’t reconcile the genetic closeness to its eastern sisterclade R1a – prematurely dubbed the prime “indo-european” marker already in the earliest Y-chromosome publications – with the recent spread of the Indo-European languages: According to the linguists’ consensus this group of languages could never have originated much more than 5000 years ago. The predominance of R1a from Poland to India and R2 from Kurdistan to Bangladesh vaguely suggested some cultural link, but the high genetic age and pretty sharp delimitations between the different genetic groups – including R1b – was hard to reconcile with contemporary archeological evidence.
The human species appears to be quite homogeneous genetically and this applies especially to the male Y-chromosome. The importance of Y-DNA expression is typically played down, and only a few investigations indicate some vague link between Y-DNA haplogroups and a direct genetic advantage. So far, the results tend to be hard to evaluate. Eg. S. Shoaib Shah et al., 2008:

effect-size comparisons allowed us to detect an association of the haplogroups R2 [...] and R1a1 [...] with lower self-reported aggression mean scores in this population

In another study, the European Society of Cardiology links male Y chromosome variants with the risk of heart disease, indicating that the overwhelming Britsh R1b majority population must be better equiped against coronary diseases than the “Native European” haplogroup I population.
All together, low aggression and genetic protection against age- or lifestyle related diseases can hardly be taken serious as an evolutionary advantage for an “Indo-European” society traditionally imagined as violent and dominated by young warriors on horseback. So even if true that the male carriers of haplogroup R had a genetic advantage that facilitated its success, for sure this advantage – just like with the successful gene for lactose persistency – must have been accompanied by a new cultural level.

Like most majority haplogroups in the world, R1b (or more especifically for Europe: subclade R1b1b2) is now incorporated in the select body of major haplogroups that has been attributed a Neolithic age.

Neolithic advance

Frequency of R1b

Variance of R1b

Balareque et al. proposed a relation between the gradual advance of Neolithic farming land and the introduction of male DNA marker Haplogroup R1b. The increased momentum of this change at greater distances from the center resulted in higher frequencies in the west, while diversity (not shown) was assumed to follow a star-structure. Variance was interpreted as an indication of age.

In their 2010 article Balaresque et al. gave the Neolithic kick-off:

…our evidence supports a different interpretation: that R1b1b2 was carried as a rapidly expanding lineage from the Near East via Anatolia to the western fringe of Europe during the Neolithic.

Their dating was in the Neolithic range, having eastern R1b slightly older than western R1b, but their prime argument was the star-like structure they detected in the STR values:

The two hypotheses also make different predictions for the number of sources of diversity within hg R1b1b2: under the postglacial recolonization model, we expect multiple sources, whereas under the Neolithic expansion model, we expect only one.We can test this by examining the phylogenetic relationships among microsatellite haplotypes. A reduced median network of 859 haplotypes (Figure 3) shows a simple star-like structure indicative of expansion from one source [...]. This pattern seems incompatible with recolonization from differentiated refugial populations [...]. The core of the network also contains haplotypes from Turkey (Anatolia), which is compatible with a subpopulation from this region acting as a source for the westwards-expanding lineage.

Almost inmediately this star-like features (and the recent dating) were contested by Morelli et al., who detected a rather bipolar structure due to the inclusion of a single marker (DYSA7.2 or DYS461), BTW to the effect that European R1b was again interpreted as pre-Neolithic:

“The Sardinian haplotypes belong to the Atlantic Modal Haplotype variability, with an interesting internal differentiation shown by the completely Sardinian branch off-shoot (figure 2A). In contrast, the majority of Anatolian samples belong to the DYS39312/DYSA7.2-11 subtype. Interestingly, the bridge between the two main forms, is represented by the intermediate step of a haplotype common in the Balkan region, DYS393-13/DYSA7.2-11.”

Morelli et al. showed how the expected star-structure of European R1b haplotypes turns into a diachtomy, that makes a simple Neolithic scenario an illusion.

The subsequent “Neolithic R1b” article of Myres et al. (2010) conforms to the notion of an important diachtomy dividing R1b in an eastern and western grouping, this time based on SNP M412. This SNP based dichtomy is more or less equivalent to the much older ht15 denomination against ht35 (~ M412-) elsewhere. The investigators disclose a minor indication that R1b (cq. M412-/ht35) might have been present in SE Europe already before the Neolithic:

Although this frequency pattern closely approximates the spread of the Linearbandkeramik (LBK), Neolithic culture, an advent leading to a number of pre-historic cultural developments during the pastr 10 thousand years, more complex pre-Neolithic scenarios remain possible for the L23(xM412) components in Southeast Europe and elsewhere.”

However, Myres et al. gives a different twist to the bipolarity and make a distinction between early and late Neolithic LBK-related expansions:

Archeologically, there are two attested phases regarding the geographic spread of the Linearbandkeramik (LBK). The first phase extended to the upper Danube river near Munich. The second phase extended further to the Paris basin. Furthermore, there is evidence of several post-LBK Neolithic expansions, ca 6000 years BP from the Paris basin region toward Northern Italy, Southern France and Iberia, characterized by the Chasseen horizon, as well as to England.”

These dates are well within the range of the earliest Neolithic activity in Britain and Ireland (Sheridan, 2007):

[...] Neolithic activity, reliably dated to between c.3950/3900 and 3700 cal BC in northern Britain (especially Scotland), that is associated with the use of pottery in the ‘Carinated Bowl’ ceramic tradition [...]. The distribution of this type of pottery extends far beyond the area under review, to encompass much of Britain and much of Ireland. The Carinated Bowl-associated Neolithic is one of at least three distinct strands of the earliest Neolithic activity in Britain and Ireland, the others being i) a strand linking north-west France (probably Normandy) with southwest England during the first quarter of the fourth millennium cal BC [...] and ii) a Breton strand, which is found along the Atlantic/Irish Sea façade and seems to have appeared marginally earlier than the Carinated Bowl tradition, between c. 4200 and 3900 cal BC.

[...] it is clear that the end of the fifth millennium cal BC was a time of agricultural expansion after a period of ‘stasis’, when huntergatherer communities in the Netherlands finally switched to farming (Louwe Kooijmans 2005)—most likely thanks to influences from the south-west— and when farming groups, ultimately deriving from the north-east of the Paris Basin, are suspected to have moved north and eastwards

Balaresque et al. already predicted some exceptions to the star-structure, but these rather applied to the linear expansion of hg E-81 along the mediterranean coast, and less to the semi-circular wave front implied by the continental Neolithic wave of advance in Europe:

Successive founder effects at the edge of the expansion wave can lead to a reduction in microsatellite diversity, even as the lineage increases in frequency.

The innovations in the Near East also spread along the southern shore of the Mediterranean, reflected in the expansion of hg E1b1b1b (E-M81), which increases in frequency and reduces in diversity from east to west.

Only Myres comes up with a possible explanation of the bipolarity, in their results implied by the existence of a M412 hemisphere in western Europe:

Our high resolution SNP genotype results show that the majority of Central and Western European haplogroups relate to common M412 founders whose sub-clades display phylogeographic and temporal patterns consistent with allele surfing at the periphery of expansions.

Where did this M412 surfing start, that must have initiated the European diachtomy for haplogroup R1b in a western and southeastern group? The alternative brought forward by Myres et al. would possibly result in the conclusion that M412 was already in Central Europe (or maybe even further!) at the time that the Early Neolithic arrived there.
Or could it be rather congruent and contemporary to the Late Neolithic expansion?
As far the distribution of R1b is concerned, the Early Neolithic was limited basically to Central and SE Europe. Here is a gap of 1500 km where early occurrence of M412 is problematic because of low frequency, variance and availability of almost all key M412 subclades. This is why the earliest advance of M412+ correlates best to the late Neolithic expansions, that – to the exception of Iberia – covers most of western Europe.

It is a pity the Myres article does not arrive at a deeper analysis here and only seems to clear the field for an answer in the pre-Neolithic, not unlike Morelli et al.
My alternative is that the dates of Myres et al. may need a slight correction for M412 for being partly congruent to the late-neolithic expansion, that started about 4000 BC in western Europe.

How come genetic dating is still causing so much trouble and contradictory results? Contrary to the inflated ages typically in vogue among population genetists (commonly considered 3x too high by genealogic opponents), there is increasing support for genealogic dates that on the other hand are probably sometimes up to 3x too low. Nowadays everybody can order their own marker sets of 12, 37, 67 or more STR for trying to make sense out of a multitude of dating methods, often for genalogic reasons or just amateur interest. Mathematicians, capitalizing on the delight of amateur genealogists, often managed to reverse the paleolithic dates into historical dates, but most of the time the private dating results are achieved without genome-level insight or calibration, even without any sense of historical or archeological context whatsoever. Only this year the dating methods and correlation models that concern the paternal R1b haplogroup lineages are hard on their way to become valuable tools to untangle the European prehistory. Microsatellite diversity at picked locations at the Y-chromosome are considered, where short base-pair sequences (STR) are tandemly repeated by a semi-characteristic value. Unlike the rare base-pair alterations (SNP) that define the different Y-chromosome haplogroups, this microsatellite diversity is commonly interpreted as “datable” for being the result of spontaneous mutation processes from within, that albeit unpredictable are thought to occur at a certain “biological” rate.

More generally, knowledge of the rates and processes of mutation at different classes of marker is fundamental to evolutionary interpretations of diversity data [...]. In this broader context, studies on the Y chromosome are of particular interest because the mutations observed here are the result of exclusively intra-allelic processes. (Jobling et al., 2004)

The strict assumption with STR-age calculations is predictability of spontaneous mutations, to the exclusion of external factors that make mutations inpredictable. The mutation rate at each STR locus may vary significantly, but by taking several loci together the rates could be averaged. Spontaneous mutations are the result of normal DNA metabolism, while induced mutations are the result of agents that are not part of normal DNA metabolism. For instance, agents that cause induced mutations are radiation of various types, highly reactive metabolic byproducts that are produced within a cell, toxins that enter the body from direct contact, inhalation, eating or drinking. Take away all this “open system” factors, and you’ll get the “predictability” you need for making TMRCA calculations.
Some haplotypes are very “average”, while others are very different. How such differences could serve any purpose at all for estimating an age? All haplotypes have the same age counted from any common SNP mutation, so the assumption that mutational processes affect most people more or less in the same way does not hold. Other mechanisms must be at work that especially concern marked outliers.
It should be clear that no assumptions to the effect of spontaneous mutations exist that relate to SNP’s, thus making all speculations about a purported SNP mutation rate much more than precarious, and comparative at most. Moreover, it can’t be excluded that induced mutations causing new SNP’s also excite the STR to new values that may be instable or just very unlikely – thus creating a STR-SNP correlation that does not reflect the STR age, being based on spontaneous mutations.

The genealogic dating issue appears to be almost resolved since the investigation of Zhivotovsky et al., 2006:

Mutation rates observed in pedigree and family studies [...] are, on average, 2.1–2.6 x 10–3 per generation. By studying short tandem repeat (STR, or microsatellite) variation within haplogroups C2 and H1 in populations with known short-term foundation events (Bulgarian Gypsy and New Zealand Maori) and comparing autosomal and Y-chromosomal microsatellite variation, Zhivotovsky et al. (2004) suggested that an effective mutation rate of 0.69 x 10–3 per 25 years could serve as the rate of evolution at Y-chromosomal STR loci (see further discussion in Di Giacomo et al. 2004; Zhivotovsky and Underhill 2005). A mechanism that might explain this 3- to 4-fold discrepancy is that [...]“

Zhivotovsky sought the explanation of his reduced mutation rates in effective population sizes, that should be assumed much lower than the normal population size. however, on haplotype level there was also the issue of outliers. Myres et al. (2010):

The ages of various haplogroups in populations were estimated using the methodology described by Zhivotovsky et al., modified according to Sengupta et al, using the evolutionary effective mutation rate of 6.9 * 10 -4 per 25 years. The accuracy and appropriateness of this mutation rate has been independently confirmed in several deep-rooted pedigrees of the Hutterites. Important caveats to consider include the fact that coalescent times (Td) is sensitive to authentic rare outlier alleles and that multiple founders during population formation will inflate the age estimate of the event.

Most likely those “outliers” are caused by induced mutations, that thus would mess up the observed rates. The matter would become even more complicated when such outliers also become the founders of new clusters. Calculating interclade TMRCA’s would reduce to mere wishful thinking if typically the lifecycle of a subclade started indeed with a mutational STR boost being triggered by an induced mutation.

An example of unstable STR values due to a SNP mutation could be L148, a small subclade of U106+L48+. Comparing Eldon (y-search-id SFVPS), Cripps (BPBJQ) and Mitchell (YNWJ4) using the traditional STR interpretation would indicate Eldon’s haplotype closest to the ancestral haplotype for being closer to both Cripps and Mitchell. Still only Cripps has the normal (U106 modal) value DYS448=19, against the very rare value DYS448=15 for both other haplotyps. A genetic distance jump of GD=4! This giant backmutation to modal values in Cripps lineage suggests that the L148 mutation, when it happened, also involved at least one unstable STR mutation DYS448=15. If we could generalize on this we should be very careful indeed with deriving an age for a subclade using interclade STR calculations.

Statistic results of samples that include outliers would require higher-than-Zhivotovsky mutation rates to compensate for induced mutations. The investigation on Hutterites proved no calibation on mutation rates is necessary as long the outliers are discounted. Since STR dating assumes a predictable rate for spontaneous mutations, it is plain wrong to apply higher mutation rates for the sake of extreme STR fluctuations, even when observed in some documented, genealogic lineages.

More dating dangers are lurking. Multiple founders, cq. potential migrations instead of the gradual geographic expansion of a population, were mentioned by Myres et al. as another source of bias having TMRCA’s getting too old compared to the first occurrence of a subclade in a region. This notion may be important to Iberia, where the proposed LBK related Neolithic advance seems to collapse against unrelated Neolithic traditions at the time R1b is supposed to arrive.

One way to verify the true nature of haplogroup R1b’s advance through Europe is variance. Interpretation is hazardous, especially in relation to origin. This parameter measures the probability distribution of genetic distances compared to the mean. Unfortunately, variance alone won’t be enough to tell the difference between an increased effective population in the process of expansion, or an accumulated distribution of variability near its point of origin.
While often associated primarily with the potential origin of a haplogroup, the truth is that variance is highly sensitive to recent population history in particular, like processes that cause unequal growth and that tend to move away from a normal distribution. Isolated areas often show up with high variances and one-sided interpretation could make us believe that all humanity came from mountainous places like Scotland and Caucasus. Cosmopolitan areas on the other hand almost invariably show low variances. All depends on the success of certain haplotypes within a community, and the facility that successful lineages can also penetrate into the perifery and remote areas.
There is much more to variance than a tenuous indication of age. A subclade in region A having effective population size Na will develop a lower variance than in region B that has an effective population size Nb>Na, due to inbreeding and natural loss.
When the subclade expands from region A to region B, then it will be logical that B will soon develop a higher effective population, ie. a decreased loss via stochastic processes and inbreeding. The result will be that the variance increases most in the expansion areas. The corresponding diversity of haplotypes would be a star-structure, towards an increasing effective population size, thus higher variance at the rims.
Settled Neolithic populations in location A must have had a quite small effective population, almost per definition. This effective population basically consisted of those having the ownership of a small piece of arable land, and was typically stuck in small inbreeding communities.
On the other hand, the expanding part of the same population must have had almost unlimited growth possibilities for a while, thus creating a huge increase of the effective population in location B. At STR level this general situation for an expanding wavefront should comply to a star-structure and a corresponding increment of diversity and variance.

Also, a straightforward interpretation may be hampered by poor sampling that would make the prime assumption of Myres et al. tenuous, ie. that outliers should be excluded from the equation when calculating the age of a population.
This inadequacy of variance as a reliable and unequivocal measure of geographic age becomes dramatically clear when the data of Myres et al.(from table S2) is mapped into separate variance graphs for mutations U106 and P312, as has been done by Vincent Vizachero (acknowledged by Myres et al. “for alerting us to the potential of M520, M529, S116, L11, L23 SNPs and for insight regarding the DYS390, 19 repeat allele and M73″)

Variance of R1b-L23

Variance of R1b-U106

Variance of R1b-P312

The R1b data of Myres et al. (table S2) becomes complicated and ambiguous once mapped separately into variance graphs for the components M412-, U106 and P312.

These varance graphs magnify inconsistencies that are difficult to reconcile with the Neolithic wave of advance. Indeed, the inconsistencies are so pronounced that it is hard to believe that the Neolithic advance is the only process that determined the current M412 distribution. It shouldn’t be surprising that now the variance of P312 and U106 (see above) could be interpreted as an indication of origin in western Europe.The absolute dates are debatable, but on a relative basis the variance graphs of P312 and U106 would indicated two separate, even contrary movements west to east against east to west that globally met each other at the axis East-Germany-Czechia-Croatia. Such results are at odds with the Neolithic advance that is the focus of current discussion, and no issue at all in the article of Myres et al. This doesn’t imply that variance is completely useless, but for sure this situation appeals to creativity.

Superficially the variance graph of P312 is more or less in agreement with the expected state of R1b at its arrival in western Europe, and – using one eye – can even be conciliated with the neolithic advance. Closer examination reveals that the peak values of P312 variance can be contributed almost exclusively to the U152 subclade. Even undifferentiated P312* seems to have attributed a lower variance. I figure the explanation could be that some relevant P312 subclades already existed at the eve of new expansions that eminated from central western Europe, and could have been post-Neolithic. This expansion inevitably involved a reduction of SNP diversity the further P312 moved away from such a purported expansion center in western Europe. This scenario for P312 would be in agreement with the predominance of subclade U152 towards the east and the predominance of subclade L21 towards the west.

The graph of U106 shows an increasing variance towards the eastern Donau region and beyond that superficially appears to be absolutely incongruent to the Neolithic advance. The graph could be reconciled with the western European focus of the P312 graph only if we just try to grasp the basics, that are quite ambiguous as far variance is concerned. Two mutually exclusive possibilities apply to U106:

• Western U106 is younger and didn’t accumulate as much variance, or:
• Western U106 suffered the limitations of a smaller effective population size compared to the expanding eastern part, that thus could build up an increasing variance and variability in a star-like fashion as a function of geographic distance from the source (Belgium?).

If we now just reverse the observations for P312 above then it looks likes U106 already started to expand before it was significantly differentiated. Expansion then involved an inevitable incremental increase of variability and variance. In the expansion areas formerly obscure lineages continued to expand in a star-like fashion. Indeed, I could show a remarkable relatedness of western U106 haplotypes “in the centre of the star” with respect to eastern haplotypes that exceed the internal relatedness of eastern haplotypes, insinuating that the center of expansion of U106 is indeed Western Europe.

BTW., the same ambiguity about variance as an indicator of population history could be emphasized for Iberian P312*:

• At the end of the wave of advance, Iberian P312* could be younger when it didn’t accumulate as much variance over time, or:
• Iberian P312* is older in NW Iberia and once settled, suffered the limitations of a smaller effective population size compared to an expanding U152 subclade, also essentially associated to expanding Bronze Age cultures (Iberian Bell Beaker). Only U152 could thus build up an increasing variance as a function of geographic distance from the (Iberian) source, at increasing variability in a star-like structure.

At this stage we should go back to the Myres et al. investigation and ask why M412 doesn’t comply to the star-structure one would expect with a truly Neolithic wave of advance.

• first there is the transition to M412 itself, that doesn’t really comply to the star-structure of a wave front that propagates in a semi-circular way. Myres et al. explains this bipolarity or diachtomy in European R1b1b2 as a surfing-effect along the wave.
• second there is a circular cline of variance for P312 that seems to emanate from the western Alps. Half of this cline runs in eastern direction, what would also violate the Neolithic star structure that is expected to run east to west, either way: taking variance as an unambiguous indicator of age or taking variance as an unambiguous indicator of increasing effective population size along the direction of the wave of advance.
• third there is the semi-circular cline of increasing variance of U106 in eastern direction. Since obviously this cline is unambiguous variance can be taken here as both supportive to the Neolithic advance (continuous surfing along the direction of the wave of advance, what is anyway a violation of the expected star-structure); or a violation of the Neolithic wave of advance by focussing on the star structure that is now contrary to the wave of advance.
• fourth there is a maximum variance in NW Iberia, that is predominantly P312*. There is no way that this could have reached Iberia through the Neolithic wave of advance, so the inflated TMRCA must have been caused by the multiple founders typical of migration. IMO this proves the oversea origin of Iberian Beaker from NW Europe, where in Belgium there is about the same situation of high variance P312*.
• fifth there is an untangible mix of P312* and large public P312+ subclades in the peak region of P312 variance nearing the western Alps. If this peak values are due to U152 then probably we could identify two different movements here:
  • A Late-Neolithic movement of P312* emanating from the Paris Basin, conform the proposal of Myres et al. that links the Neolithic advance to LBK-derived cultures. This wave reaches a frequency peak in the Basque country, or rather Aquitaine where the Basques are supposed to originate, where such the peak would be expected. Also, since in Iberia the Neolithic influence was determined by the mediterranean Cardial Culture complex (often associated with a westward spread of haplogroup E), this continental R1b wave should be expected to stop here.
  • Another movement in opposite direction bringing P312+ subclades having a star-structure in eastern direction, emanating from the west. This wave must have been younger and IMO originates ultimately in NW Iberia (not the Basque country).

R1b variance graph of Balaresque et al.indicates a higher variance in northwestern Iberia comparable to the values in northwestern Europe, what indeed would be in agreement with archeological indications towards the oversea, post-Neolithic link indicated by the Corded Beaker remains in Castelo Velho, Northern Protugal, at rock bottom (Susana Oliveira Jorge, 2001) and the Beaker remains in La Soria that according to Rojo-Guerra et al. (2005) has its best parallels in French Brittany, and ultimately in the Corded Ware complex (“Su decoración de líneas horizontales y paralelas a lo largo del vaso y su peculiar forma tienen sus mejores paralelos en la Bretaña francesa, y en última instancia en la Cerámica cordada”)

Are we here on the verge of a breakthrough that links the Neolithic advance to a common European Beaker culture, tied together by a single SNP of R1b – ie. M412?

Desde nuestro punto de vista esta tendencia tipológica tiene su origen en última instancia en los perfiles característicos de los vasos de la Cultura de la Cerámica Cordada, en los que esas características se muestran de forma mucho más marcada (…)

In our point of view, this typological tendency [i.e.narrow neck and low belly] has its origen ultimately in the characteristic profiles of the beakers of the Corded Ware culture, where these characteristics are much more noticeable.

En lo que respecta a la decoración, y más allá de la obvia derivación del patrón lineal de los esquemas cordados, es posible encontrar los paralelos más cercanos de la ornamentación de nuestro vaso en una curiosa variante regional del Campaniforme típica de la Bretaña francesa, con derivaciones hacia las regiones aledañas, que se caracteriza precisamente por el empleo de líneas horizontales y paralelas impresas, de forma corrida por toda la superficie externa del recipiente, como única decoración (…)

Regarding the decoration, beyond the obvious derivation of the linear patterns from the Corded schemes, it is possible to find parallels that are closer to the ornaments of our beaker [in central Spain] in a curious regional variety typical to the Bell Beaker style of French Brittany and its derivatives in the neighbourhood, that are characterized exactly by this application of horizontal lines and parallel imprintings, that run through the whole external surface of the beaker, being the only decoration.

The only difference is the tool employed for making the decoration:

todas las decoraciones campaniformes son impresas, lo único que varía es el instrumento empleado para ello: una cuerda en los tipos cordados, un peine de púas en los marítimos, y un peine liso en los hasta ahora llamados tipos incisos.

All bell beaker decorations are imprinted, the only difference is made by the employed tool: a cord for the corded prototypes, a dented comb for the maritime prototypes and a smooth tool for those prototypes that so far are known as incised.

The subsequent success of the Iberian Bell Beaker tradition and its hypothetical association to Atlantic Celts and Celto-Italic expansions deep into Central Europe, or possibly even much further, would for sure move us too far away from the Neolithic scope of this article. Still, it must be clear that unfortunately the Neolithic is not the only answer to all our questions. Actually, paleogenetic investigations (most notably the Lichtenstein Caves) didn’t confirm at all the predominance of R1b in prehistory. At the end the “R1b takeover” may have been a gradual, albeit inevitable process over thousands of years that was rather due to the disproportionate genetic contribution of the farming component to society. Even today in genealogy it can be observed that the most successful surnames derive from rural areas, having the owner of a farm as a founder – my own lineage started with a single obscure farmer and expanded to 2000 households worldwide within barely 500 years.

Often obscure and of humble status, the economical niche and attachment to the land of farming families offered continuity and survival, even under the hardest circumstances.

Referenced:

• Myres et al. – A major Y-chromosome haplogroup R1b Holocene era founder effect in Central and Western Europe, 2010, link or try here.
• Alison Sheridan – From Picardie to Pickering and Pencraig Hill? New information on the ‘Carinated Bowl Neolithic’ in northern Britain, 2007, link
• Pierre Allard – The Mesolithic-Neolithic transition in the Paris Basin : a review, 2007, link
• P. Balaresque et al. – A Predominantly Neolithic Origin for European Paternal Lineages, 2010, link
• L. Morelli et al. – A Comparison of Y-Chromosome Variation in Sardinia and Anatolia Is More Consistent with Cultural Rather than Demic Diffusion of Agriculture, 2010, link
• Cruciani et al. – Strong intra and inter-continental differentiation revealed by Y chromosome SNPs M269, U106 and U152, 2010, link or try here.
• European Society of Cardiology – Study links male Y chromosome variants with the risk of coronary heart disease, 2010, link
• S. Shoaib Shah et al. – Y haplogroups and aggressive behavior in a Pakistani ethnic group, 2008, link
• Mark A. Jobling and Chris Tyler-Smith – The Human Y Chromosome: An Evolutionary Marker Comes Of Age (2004), link
• Zhivotovsky et al. – Difference between Evolutionarily Effective and Germ line Mutation Rate Due to Stochastically Varying Haplogroup Size, 2006, link
• Rojo-Guerra et al. – Un peculiar vaso Campaniforme de estilo Marítimo del túmulo de La Sima, Miño de Medinaceli (Soria, España): Refleciones en turno de las técnicas decorativas Campaniformes y los sistemas de intercambios de larga distancia, 2005, link
• Susana Oliveira Jorge – An All-Over-Corded Bell Beake in Northern Portgal: CASTELO VELHO DE FREIXO DE NUMÃO (VILA NOVA DE FOZ CÔA): Some remarks, 2001, link
• Schilz – Molekulargenetische Verwandtschaftsanalysen am prähistorischen Skelettkollektiv der Lichtensteinhöhle, 2006, link

Rapid genetic change affected the global population once the genes were available. Next to evolution came the Magic Stick.

Green et al. investigated the differences between Africans and non-Africans in relation with the (draft) Neanderthal genome and revealed 1-4% of the modern human genepool could be accounted for by Neanderthal only. Green et al., 2010:

“Under the assumption that there was no gene flow from Neandertals to the ancestors of modern Africans, the proportion of Neandertal ancestry of non-Africans, f, can be estimated by…”
“Assuming that gene flow from Neandertals occurred between 50,000 and 80,000 years ago, this method estimates f to be between 1 and 4%“
“We note that a previous study found a pattern of genetic variation in present-day humans that was hypothesized to be due to gene flow from Neandertals or other archaic hominins into modern humans (81). The authors of this study estimated the fraction of non-African genomes affected by “archaic” gene flow to be 14%, almost an order of magnitude greater than our estimates, suggesting that their observations may not be entirely explained by gene flow from Neandertals.”
“We expect that further analyses of the Neandertal genome as well as the genomes of other archaic hominins will generate additional hypotheses and provide further insights into the origins and early history of present-day humans.”

But let us define “gene flow”: the study explicitly excluded shared ancestral genes and gene flow from Neanderthals to the ancestors of modern Africans. This 1-4% only reflects the measurable genetic differences between African and non-African haplotypes explained by Neanderthal, and thus is just a minimum amount. What we miss is an assessment of the maximum amount, that would instead reveal the percentage of early African genes that exclude any Neanderthal input in the genome of non-African modern humans. This could prove to be a much more cumbersome excercise, not in the least because we don’t know yet how to tell apart all truly African from other non-Neanderthal hominin genes.

Only the genetic deviations in modern humans with respect to the Neanderthal genome were investigated. None were found or mentioned in genes that nowadays harbour truly cross-hominin haplotypes. Previously the “European” H2 haplotype of the gene MAPT and the “Eurasian” haplotype D of the Microcephalin-1 gene were recognized as definitely different from the haplotypes associated to Homo Sapiens, and consequently dubbed (prematurely) “Neanderthal” by some. The sheer differences between the various haplotypes of such genes already suggested cross-breeding of hominins that share a common origin with modern humans that coalescence to dates that vastly exceed the AMH-Neanderthal split under investigation. Indeed, neither haplotype was found on Neanderthal and by default the Neanderthal version of MAPT and Microcephalin-1 must be considered compatible with the haplotypes commonly associated with modern humans. Green et al., 2010:

This analysis shows that some old haplotypes most likely owe their presence in present-day non-Africans to gene flow from Neandertals. However, not all old haplotypes in non-Africans may have such an origin. For example, it has been suggested that the H2 haplotype on chromosome 17 and the D haplotype of the microcephalin gene were contributed by Neandertals to present-day non-Africans (12,79, 80). This is not supported by the current data because the Neandertals analyzed do not carry these haplotypes.

Likewise, the study doesn’t resolve the precise origin of the other newly identified non-Neanderthal genes. Was the AMH gene pool already amalgamated with a wide variety of exogenetic hominin contributions at the moment African AMH and Neanderthal met? Such contacts should have increased the genetic differences between early African AMH and Neanderthal about the time of ingression. On the other hand, as far they could verify with modern genes, Green et al. were pretty determined that their Neanderthal samples did not feature ingression:

Thus, all or almost all of the gene flow detected was from Neandertals into modern humans.

This emerging picture of the AMH genome being like a sponge to exogenetic influences against isolated Neanderthal communities that preserved their genetic purity until extinction is amazing. Admixtures being confined to quite recent developments confirm the effects of culture in human evolution, that thus must have included increased contact and communicaton between hominin groups. To a certain degree admixtures may have involved even African AMH’s own African neighbours. Green et al.:

[...]old population substructure in Africa has been suggested based on genetic as well as paleontological data.

However, in the article this substructure suggested by Green et al. only concerned a possible African’s AMH or sub-saharan special relationship to Neanderthal up north:

If after the divergence of Neandertals there was incomplete genetic homogenization between what were to become the ancestors of non-Africans and Africans, present-day non-Africans would be more closely related to Neandertals than are Africans.

For sure, recent cross-hominin gene flow of the H2/MAPT or D/Microcephalin-1 type would require an African substructure much deeper than the one implied by this study. Older cross-hominin events may include “African” V and M haplotypes of gene ASAH1, where a coalescent-time depth of 2.4 million years ago tend to marginalize the differences with Neanderthal. The Green paper did not give a clue about the Neanderthal haplotype and thus does not resolve the question of potential post-split date hybridization in Africa. Based on dates and global currency, haplotype M could have been of heidelbergensis and haplotype V a successful African cross-hominin addition. Kim et al. 2007:

it should be noted that the pattern of genetic diversity of ASAH1 and other loci is compatible with the proposal that the human population was once geographically structured and genetically differentiated in Africa

The predominance of the V lineage is observed in both Africans and non-Africans
[...] Furthermore,the number of haplotypes in the V lineage is only 6, yet it is 11 in the M lineage. These hold true in both African and non-African samples

The most parsimonious explanation for the sharing of this pattern across all four [worldwide reference] populations is that the sweep occurred prior to the radiation of modern humans out of Africa.

the TMRCA of the V lineage is estimated as 200 +/-50KY from Genetree analysis [...] and 340 +/- 80 KY on the basis of the average nucleotide diversity [...] On the other hand, the TMRCA of the M lineage is 320 +/- 70KY from the Genetree analysis and 680 +/- 180 KY from the nucleotide diversity. Compared with the M lineage, the relatively recent origin of the predominant V lineage implies that it has been rapidly increasing in frequency.

Haplotype M/ASAH1 would not be surprising for Neanderthal, but haplotype V/ASAH1 would raise questions about the extend of early human hybridization events, and where it all started. Such and other potentially recent African cross-hominin evidence should put the impact of the Neanderthal lineage on modern humans in a completely different perspective. Unfortunately the Green paper didn’t intend to resolve the interbreeding questions already raised in previous publications on the subject. Moreover, single genes are hardly significant to the degree of admixture and the significance of African non-AHM contributions are still a matter of debate. Wall et al. (2009) insist on at least a considerable West African non-AMH hominin component:

We find evidence for this ancient admixture in European, East Asian, and West African samples, suggesting that admixture between diverged hominin groups may be a general feature of recent human evolution.

Instead, the announced results of a New Mexico investigation “didn’t find evidence of interbreeding in the genomes of the modern African people included in the study” (Nature News,20 April 2010). Conventional wisdow still has it that hominin admixture was a recent AMH feature, primarily linked to Out of Africa expansions and post-dating the AMH-Neanderthal split, but at least an African origin of haplotypes H2/MAPT and D/Microcephalin-1 would apply for special pleading. A specifically eastern Asian RRM2P4 haplotype even computes a coalescent-time of 2.3 million years ago and a quite short LD sequence virtually rules out a recent adquisition of this gene – that moreover must have been confined to the people of eastern Asia!

Let us return to the 1-4% measurable admixture. Like John Hawks already put it, this estimate “is so high that this is not just a few genes introgressing in from Neandertals — it is a big fraction of the neutral, non-coding part of the genome.” This portion was deduced from the signal of gene flow retrieved from the segments with the lowest divergence to Neanderthal: [Non-African] “segments, with few differences from the Neandertals, tend to have many differences from other present-day humans, whereas African segments do not“. However, I don’t agree it is just this “visible” part of the Neanderthal genome that survived. This must be just the low-mutation-rate part of Neanderthal that pops up from the graphs, where strongly homogenizing selective forces in Africa about 200 kya must have reduced variability among AMH. Subsequent strong selective forces during AMH expansion kept variablity lower at non-African high mutation rate segments.

Only non-African high variability segments with an unusually low divergence to Neanderthal (box) tend to have an unusually high divergence to other modern humans (here Craig Venter) because the segments are not shared by all, being specifically derived from Neanderthal. One additional reason may be some of these segments are typified by low mutation rates and bottlenecked in Africa, where the registered variability of these segments is low.

The premise of this certainty of dealing with Neandertal admixtures is that other causes of a low divergence to Neanderthal, such as low mutation rates, “would produce monotonic behaviors” on the above diagram. Here it shows that African and non-African segments that most closely resemble the Neanderthal genome may diverge considerably from the reference genome of mister Craig Venter, thus confirming high variability, and 94% of these diverging, high variability genes are of confirmed European ancestry. Moreover, the results of “identified regions in which there is considerably more diversity outside Africa” (Green et al.) show 80% tag SNPs (133 out of 166) and 83% of the regions (10 out of 12) pointing to a specific Neanderthal origin. Note the potential contribution of other hominins to this unexpected variability seems to be rather low, and we can only assume the remaining non-Neanderthal tag SNPs and regions could be African – if this weren’t so contradictory to Out of African type bottleneck scenarios. All these results seem to be in sheer contrast to concurrent genetic and phenotype results that rather indicated a “smooth loss of genetic diversity with increasing distance from Africa” (Manica et al., 2007). The contradictory clines involved inhibit a simple extrapolation of the diagram to the Neanderthal survival of the remaining 96% of their genome, ie. somewhere hidden on the part where diagram behavior is “monotonic” (ie. a straight line). Basically, in line with the comments made by Relethford, 2008, the current data doesn’t exclude this possibility at all:

[...] the observation of higher African diversity supports the other genetic (and fossil) evidence for an African origin for modern humans, but does not distinguish between an African origin with replacement and an African origin with admixture outside of Africa except to say that if there was any admixture it was not of sufficient magnitude to erase the genetic signature of an African origin.

High non-African variability on certain regions is remarkable, but so is the implied low African variability at these same locations. Let us reverse the assessment and consider the potential backmigration or diffusion from a hybrid source in Eurasia into Africa, conform the one already implied by the purported cross-hominin D haplotype of the microcephalin-1 gene found in only 30% of subs-saharan Africans. The Green paper is not conclusive about the origin of African counterparts of the mentioned non-African high variability genetic regions, but for sure the ingression back into Africa of a reduced set of “bottlenecked” non-African haplotypes would have a similar result on the diagram if regions of higher Neanderthal divergence were caused by higher mutation rates of these regions, and thus bottlenecked were to remain within the bandwith of African variability. However, the low world-wide frequency of this off-modal variability would also imply the reduction of high mutation speed variability among non-African populations, possibly due to some kind of post-Neanderthal event. This scenario would be in agreement with a more recent ingression of new genes and the heavy selective pressures implied during this process.

It should be investigated if much of the higher African variability indeed involve segments having a higher mutation rate, since strong natural selection at an early stage, contrary to the Neanderthal situation, could have obliterated much of the deeper African substructure. If this would be the case, there wouldn’t be any argument left against extrapolating the survival of virtually the whole genome of Neanderthal among non-Africans – i.e. except for the few genes and sweep areas associated to AMH related evolutionary changes elsewhere. This result would be irrespective of the African substructure proposed by Green et al. or sheer Neanderthal relatedness of early AMH, and in agreement with the announced New Mexican results, that Africa remained largely exempt of reminiscent cross-hominin interbreeding variability. African hybridization, if any, thus would have been concurrent with strong selective processes.

Long time, bidirectional genetic diffusion should be taken in consideration, and we still don’t know to what extend the advance of AMH into Neanderthal territory also involved hybridization with other hominins. However, having this assumption of 1-4% of the modern human genome just being the detectable part of the surviving Neanderthal genome in mind, it will be much easier to estimate Neanderthal survival. Green et al. (2010) estimate the frequency of candidate Neanderthal regions among non-Africans averaging “13%, and all less than 30%“. Now even the Mesolithic survival of Europeans as a result of the Neolithic wave of advance is still a matter of scientific dispute, this possible 13% Paleolithic result of potential Neanderthal survival may shed more light on this issue.

The real Neanderthal differences were found in regions where “the Neanderthal carries fewer derived alleles than expected relative to the human allelic states. A unique feature of this method is that it has more power to detect older selective sweeps”, ie. neighbouring DNA of a mutation in a region that defines a haplotype where reduced variation is an indication of few chromosome cross-overs, and thus of recent and strong positive natural selection. Such haplotypes are often referred to as a region of LD (linkage desequilibrium) for being characterized by DNA patterns that are overrepresented in the population, thus being not as random as it should be after a certain count of cross-overs. The Green et al. investigation yielded a limited set of such AMH related genetic regions that are low or depleted of Neanderthal derived alleles, and that thus could prove useful in the search for the genome or genomes of African hominins.

We identified a total of 212 regions containing putative selective sweeps [...] We ranked the 212 regions with respect to their genetic width [...] because the size of a region affected by a selective sweep will be larger the fewer generations it took for the sweep to reach fixation [...] Thus, the more intense the selection that drove a putative sweep, the larger the affected region is expected to be. [...] The widest region is located on chromosome 2 and contains the gene THADA,where a region of 336 kb is depleted of derived alleles in Neandertals. [...] Changes in THADA may thus have affected aspects of energy metabolism in early modern humans. (Green et al., 2010)

Mutations in several genes on the 20 widest regions have been associated with diseases affecting cognitive capacities. One single AMH gene at such a location (RUNX2) was suggested to be related to the most striking morphological changes associated with the Neanderthal “extinction”.

The Green paper is not very clear about the reasons why these sweep areas are lower in derived Neanderthal DNA compared to other regions, except for saying that the method to identify regions where the Neandertal carries fewer derived alleles than expected relative to the human allelic states [...] has more power to detect older selective sweeps. This by itself has the potency to even put the truly ‘Sapiens’ origin of this cherrypicked sweep areas in doubt, if the difference with Neanderthal would turn out to be too much. Note it is useless to play down the Neanderthal component based on unrelated selective sweep areas, while even larger sweep areas from a different hominin origin exist that are not even as closely related to AMH as Neanderthal. E.g. Hardy et al. (2005) about the (non-Neanderthal) H2/MAPT haplotype:

Interestingly, a recent assessment of LD (linkage disequilibrium) across the genome in different populations suggested that the MAPT locus was the longest region of LD in Europeans.

A close relationship between AMH and Neanderthal only solves part of the problem of knowing what DNA haplotypes originated where. Indeed, it complicates matters considerably knowing that especially the larger sweep areas of the AMH genome segments found exceptionally low in derived Neanderthal DNA, qualify most for being admixtures from hominins unrelated to AMH. Successful genes are most likely to reach saturation and to become fixed into the whole human population, and also are most likely to reduce overall variability. A deep African substructure as a source for new genes that are low in derived Neanderthal DNA would only make sense if evolution accelerated also among groups whose genes were less related to AMH’s proposed Neanderthal-like kin in the neighbourhood. However, the African origin of AMH is currently sought in the north of sub-saharan Africa, not south. Those AMH thought to have ultimately passed the selective sweep out of Africa thus should have received at least some genetic improvements from further south, what is remarkable considering the attested focus of AMH development around Ethiopia. If the alternative could be sought rather in local hybridization then one might wonder about the fate of southern African hominins. Where did the evidence of interbreeding go in the genomes of the modern African people? Indeed, the team of the University of New Mexico that conducted this analysis, due for publication in the near future, claims no such evidence exists. The gradual increase of variability towards the south is no argument in favour of interbreeding, nor acceptable as evidence of a southern African origin of AMH. It is evidence of continuity.

At the annual meeting of the American Association of Physical Anthropologists in Albuquerque, New Mexico, on 17 April, this team asserted that extinct species interbred with the ancestors of modern humans twice, allegedly first at about 60,000 years ago in the eastern Mediterranean. Nature News: “The researchers suggest that the population from the first interbreeding went on to migrate to Europe, Asia and North America. Then the second interbreeding with an archaic population in eastern Asia further altered the genetic makeup of people in Oceania.” The precise details are still unpublished, but especially the genetic interbreeding model of the second event “at about 45,000 years ago in eastern Asia” raises questions “about the range of species, like H. heidelbergensis”, since “Human skeletons found at Lake Mungo in New South Wales, Australia, have robust features, which may represent the result of interbreeding“.

Homo heidelbergensis is often considered the last direct ancestor of both Neanderthal and modern humans, though others rather consider Homo antecessor in this role. The remains of this hominins center in Spain and are dated between 1.2 million and 800,000 years ago. About 600 kya the first heidelbergenses in Europe already appear to be on the the evolutionary line to Neanderthal. Ever more robust forms of purported heidelbergensis derivation arrived on the scene, but not only in future Neanderthal territory. Robust phenotypes appeared almost everywhere, including southern Africa where apparently closely related forms (Rhodesia Man) became prominent between 600 – 250 kya. However, not even the mtDNA split date between Neanderthal and modern humans exactly indicate Homo antecessor as the most recent common ancestor (MRCA):

mean = 465,700 years ago; 321,200–618,000 years ago, 95% HPD (Krause et al., 2010)

The specimen of Rhodesian Man found at Broken Hill, Zambia, also referred to as Kabwe skull, put the mark of a general development towards maximum robustness before this African lineage gave away to a geographically rather scattered development towards gracile AMH. Unfortunately there is a dating problem here. Rhodesian Man shared many morphological features with rapidly modernizing precursors of AMH in northeast Africa that could have been contemporaneous. Like European Neanderthal, it doesn’t look like Rhodesian Man was much in a hurry to evolve into the “right” direction. Human modernization probably reached southern Africa from the north. It is striking that geographic isolation probably wasn’t a precondition to the development of African AMH, since an Ethiopian origin is rather somewhere in the middle between Neanderthal and Broken Hill. Indeed, rather quite close to the eastern Mediterranean, where according to the New Mexico team, in a Recent Out of Africa (ROA) scenario about 60,000 years ago, the first Neanderthal hybridization event should have taken place.
The current substructure and global distribution of mtDNA is unlikely to resolve the origin, for being too young. The mitochondrial “Eve”, the female MRCA that was the ancestral mother of all modern humans, was born only 200,000 years ago, making it impossible to even define an older African origin. In the vein of recent insights this discrepancy could be interpreted as a strong signal of natural selection involving mitochondria, even within Africa. In his blog article on the new paper of Green et al. John Hawks reproaches everyone that still don’t know, and that take their misconception as an argument in favour of extinction scenarios:

I’ve been saying it for years. I’ve published it. Will you learn to listen to me, already?
The mtDNA of Neandertals is gone because it conferred some disadvantage. There are many reasons to suspect this — the Neandertal variation is itself apparently recently derived; the human variation is clearly in disequilibrium, especially outside Africa; the mtDNA genes affect functions that differ greatly in Neandertal and recent populations, including energetics, longevity, and brain; there are clear signs of mtDNA selection in many recent human populations.

But where did the pre-sapiens mtDNA in Africa go in view of a heidelbergenis takeover, and did current mtDNA really originate in Africa? Neanderthal mtDNA isn’t so very different to take this for granted anymore. The overwhelming success of apparent heidelbergensis mtDNA and the absence of a significant signal of African hybridization may imply the death blow to older African lineages before heidelbergensis. Current African variability can’t even beat the overall post-heidelbergensis variability deduced from a draft of the Neanderthal genome. Extinction scenarios like those that traditionally accompany Out of Africa hypotheses may not be utterly useless after all.
Mitochondrial extinction was not just an issue of Neanderthal: the age of mtDNA recovered from LM6, the early modern human of Lake Mungo, Australia, even exceeds the age of modern mtDNA. Similar DNA at chromosome 11 of especially Eurasian modern humans indicate an inclusion event of a type of mtDNA that may have been common outside Africa once, and that may point to an eastern expansion of heidelbergenses dating back up to even 300,000 years ago. This mtDNA coalescent-time depth suggests that eastern Asia and Oceania may have been overrun by the heidelbergenses that were geographically closer to East Africa. In between, in the Middle East, the type of mtDNA might have changed with the fortunes of Neanderthal, or otherwise early Asiatic Neanderthal may have been the carriers of this type of mtDNA, in which case there should have been a reversed link to eastern Africa. The findings of Xinzhi Wu are in agreement to the latter:

There is a morphological mosaic between H. s. erectus and H. s. sapiens in China. The existence of common features and the morphological mosaic suggest continuity of human evolution in China. That there are a few features which are more commonly seen in the Neanderthal lineage, occurring in a few Chinese fossil skulls, probably suggests gene flow between China and the West. (Xinzhi WU, 2004)

Based on the evidence of continuity and gene flow, a new hypothesis, Continuity with Hybridization, was proposed in 1998 for characterizing human evolution in China. (Xinzhi WU, 2004)

At least across the eastern fringes of generally accepted Heidelbergensis/Neanderthal territory, hybridization was noticed in the fossile record, suggesting this process started indeed long before the purported exodus of AMH out of Africa.

“It is also suggested the Homo heidelbergensis is represented in Asia by the Dali skull and the Jinnishuan skeleton, both from China, and dated at between 200,000 and 300,000 years old. Precise dating of these fossils is important, because they might be contemporaneous with the las Homo erectus fossils in China” (de Arsuaga & Martinez, 1998)

According to the New Mexico announcement, hybridization in East Asia has been confirmed on the genetic level. A possible origin in a hypothetized African substructure in a Recent Out of Africa scenario is no issue here, though the investigators are careful to link a possible pre-Sapiens reminiscence in the East Asiatic (and Oceanic) genes to more or less related heidelbergensis rather than local hominins in the region that derive directly from much older forms. Nature News:

Theodore Schurr, a molecular anthropologist at the University of Pennsylvania in Philadelphia, said the genetic model showing interbreeding raises questions about the range of species, like H. heidelbergensis.

The exotic genetic results of the Denisova hominin, for that matter, proves that we can’t think too light about the close genetic distance of eastern Asian people to the rest of the world. The genetic differences of hominins that developed in isolation for such a long time should have left a mark impossible to miss. The few haplotypes of genes that indeed attest an extremely high age, just don’t add up to appreciable levels of admixture that, like now attested with Neanderthal, represent a certain percentage. Thus all along the bulk of exogenetic influences appears drowned amidst very much related heidelbergensis genes, whose differences to modern genes can be assumed to be more moderate. The coalescent-time depth of heidelbergensis is much less than a million years while the genus Homo might be three times that age.

The early arrival of heidelbergensis as a new hominin quickly replacing older hominins, also created an opportunity to differentiate into geographical phenotypes. Naturally, the degree of differentiation would increase closer to the place of the heidelbergensis origin. The proximity of Europe and West Africa to Iberia, home to the above mentioned ancestral Homo antecessor, would locate potential hotspots of heidelbergensis variability in those same places, thus making heidelbergensis hybridization feasible. Contact zones could be assumed at the fringe of heidelbergensis dispersal, but also close to more archaic forms that may now be assumed in Neanderthal Europe and West Africa. Prospected contact zones for hybridization can thus be hypothetized to include geographic regions where diverged branches of a wider heidelbergensis family (also Neanderthal, early sapiens etc) probably met:

• Middle East, the most likely contact zone between African sapiens and Neanderthal
• West Africa, probably not so far from the oldest heidelbergenses and deepest heidelbergensis substructure in case of an important role of nearby Homo antecessor as an inmediate precursor
• China, the likely scene of prolonged contact that most probably involved much earlier local hominins as well

Without exception, these prospected zones of genetic interaction and the earliest hotspots of modern features in the fossile record turn out to be close together. Indeed, the earliest attestations of AMH also include Morocco at the western side of Africa:

… an early Homo sapiens juvenile from Morocco dated at 160,000 years before present displays an equivalent degree of tooth development to modern European children at the same age. (Smith et al., 2007)

These include China, where “existence of common features and the morphological mosaic suggest continuity of human evolution” (Xinzhi WU, 2004) from about 200 kya, when hominins like Dali start to appear.

Attestations, of course, also include NE Africa, at the perifery of the Middle East, where the first emergence of AMH (Omo I and II) appears to be especially typified by variety, rather than being evidence of a single, isolated lineage. McDougall et al., 2004:

Here we confirm that the Omo I and Omo II hominid fossils are from similar stratigraphic levels in Member I of the Kibish Formation, despite the view that Omo I is more modern in appearance than Omo II

Our preferred estimate of the age of the Kibish hominids is 195 5 kyr, making them the earliest well-dated anatomically modern humans yet described.

The first unequivocal modern human features emerge in Ethiopia. It is difficult to define the differences as extreme variability or progressive hybridization.

Hominin variety has been noticed and studied before, but rarely to this degree at a single site. The evolutionary trend towards the modern features of AMH is not unlikely to have been preluded by inter-hominin contact, cq. starting about 200kya, rather than that inter-hominin contact was the unequivocal result of quite recent AMH expansions “out of Africa”. Like Garrigan et al. already put it in 2005:

Alternatively, the AMH phenotype may be the by-product of such admixture events.

What new clue would this observation give us about the evolution of AMH?

Cosmopolitan behavior and universal physial acceptance might have been of prime importance to those early humans that were located at the inter-hominin contact zones, and whose survival depended on their ability to cope with human differences, both physical and behavioural. The underlying assumption is that communication and language can be properly understood by taking into account their relation with other important behavioural, social, and cognitive processes – and the corresponding genetic modifications to ascertain a selective advantage. About 200 kya the level of human development must have reached a critical point, when the most economic response to first contact with other groups had changed. The nature of selective forces changed as well, thus accelerating cognitive improvements that boosted the evolution of e.g. that select set of genes mentioned by Green et al., 2010. Indeed, this must have happened right in the middle of contact zones.
Note accelerated evolution in contact zones doesn’t strictly imply hybridization, since it basically involves an adaptive response to increased environmental stress due to the demands of frequent “cosmopolitic” contacts.
Physical acceptability might have been another factor. Wearing clothes could have been one strategy to conceal the differences and this custom seems to originate from about the same time. Science News, 8th of May 2010:

Using DNA to trace the evolutionary split between head and body lice, researchers conclude that body lice first came on the scene approximately 190,000 years ago. And that shift, the scientists propose, followed soon after people first began wearing clothing.

Cloths as a social invention would overcome the practical arguments against a correlation with decreasing body hair in hot climates.
The human response to the physical appearance of others might also have accelerated the development of typical “modern” features that accompanied the rise of AMH. A positive response that commonly derives from typical “child-like” features associated with AMH might have invoked another strategy involving physical change. The investigation of Green et al., 2010, teaches us that much of these modernizing changes are possibly regulated (suppressed?) by a single new gene, considered exempt from a Neanderthal origin:

One gene of interest may be RUNX2(CBFA1). It is the only gene in the genome known to cause cleidocranial dysplasia, which is characterized by delayed closure of cranial sutures, hypoplastic or aplastic clavicles, a bell-shaped rib cage, and dental abnormalities (70). Some of these features affect morphological traits for which modern humans differ from Neandertals as well as other earlier hominins. For example, the cranial malformations seen in cleidocranial dysplasia include frontal bossing, i.e., a protruding frontal bone. A more prominent frontal bone is a feature that differs between modern humans and Neandertals as well as other archaic hominins. The clavicle, which is affected in cleidocranial dysplasia, differs in morphology between modern humans and Neandertals (71) and is associated with a different architecture of the shoulder joint. Finally, a bell-shaped rib cage is typical of Neandertals and other archaic hominins. A reasonable hypothesis is thus that an evolutionary change in RUNX2 was of importance in the origin of modern humans and that this change affected aspects of the morphology of the upper body and cranium.

A natural selection-driven advance of a small set of modernizing genes would do the rest of the trick. Neanderthal did not evolve slowly to AMH, but neither did Neanderthal disappear. Neanderthal survived, because they tricked human evolution by swaying the magic stick. The Neanderthal phenotype disappeared as rapidly as it took for a small set of AMH genes to gain prevalence. Cultural changes were the precondition for the success of these new genes, including RUNX2. The physical change could have been a matter of a couple of generations.

Or does all of this mean that Neanderthal disappeared anyway, because Neanderthal hybridization already happened long before AMH entered Europe and the rest of the world? Very unlikely, since on their natural selection-driven way through European Neanderthal territory, the new AMH genes were brought by those that also carried the exclusive haplogroup H2/MAPT gene. The Neanderthal genes may have been the same east and west, so instead we have to focus on the way how AMH genes entered. Now understanding hybridization better as the trigger for AMH related change, we should recognize this cross-hominin genetic MAPT admixture as the genetic marker to be associated with the “modernization” of European Neanderthal. This is the ultimate indication that AMH related hybridization didn’t stop in the Middle East. The magic stick even touched some unknown, unrelated hominin that exclusively roamed European Neanderthal borderland on the eve of modern ingression, but none could override the Neanderthal genes already there.

• Green et al. – A Draft Sequence of the Neandertal Genome, 2010, link
• Xinzhi Wu – On the origin of modern humans in China, 2004, link
• Xinzhi Wu – Fossil Humankind and Other Anthropoid Primates of China, 2004, link
• Garrigan et al. – Evidence for Archaic Asian Ancestry on the Human X Chromosome, 2005, link
• Burbano et al. – Targeted Investigation of the Neandertal Genome by Array-Based Sequence Capture, 2010, link
• John Hawks Weblog – Neanderthals Live! 2010, link
• John Hawks Weblog – Multiregional evolution lives! 2010, link
• Hardy et al. – Evidence suggesting that Homo neanderthalensis contributed the H2 MAPT haplotype to Homo Sapiens, 2005, link
• Neanderthals may have interbred with humans – Nature News 20 April 2010, link
• Wall et al. – Detecting Ancient Admixture and Estimating Demographic Parameters in Multiple Human Populations, 2009, link
• Manica et al. – The effect of ancient population bottlenecks on human phenotypic variation, 2007, link
• The Human Lineage – Matt Cartmill,Fred H. Smith,Kaye B. Brown, 2009, link
• Juan Luis de Arsuaga and Ignacio Martínez – The chosen species: the long march of human evolution, 1998, English translation 2006, link
• Smith et al. – Earliest evidence of modern human life history in North African early Homo sapiens, 2007, link
• McDougall et al. – Stratigraphic placement and age of modern humans from Kibish, Ethiopia, 2004, link
• Science News – Lice hang ancient date on first clothes, May 8th, 2010, link
• Garrigan et al. – Deep haplotype divergence and long-range linkage disequilibrium at xp21.1 provide evidence that humans descend from a structured ancestral population, 2005, link
• Hie Lim Kim and Yoko Satta – Population Genetic Analysis of the N-Acylsphingosine Amidohydrolase Gene Associated With Mental Activity in Humans, 2008, link
• Relethford – Genetic evidence and the modern human origins debate, 2008,link

The Celtic from the West proposal was first presented in Barry Cunliffe’s Facing the Ocean (2001) and has subsequently found resonance amongst geneticists. It provoked controversy on the part of some linguists, though is significantly in accord with John Koch’s findings in Tartessian (2009). The present collection is intended to pursue the question further in order to determine whether this earlier and more westerly starting point might now be developed as a more robust foundation for Celtic studies.

This new approach turns the focus away from a Central European origin of Celts and instead advocate an Atlantic gravity. Also the time-line of Celtic spread is due for a thorough revision, now it becomes increasingly secure to consider the Celtic world to be firmly rooted in the Copper Age (Maritime Bell Beaker) and a result of shared development that culminated in the Late Bronze Age:

“Barry Cunliffe, 2001, 261-310, has proposed the origins of the Celtic languages should be sought in the maritime networks of the Atlantic Zone, which reached their peak of intensity in the Late Bronze Age and then fell off sharply at the Bronze-Iron Transition (IXth-VIIth centuries BC).” (Koch, 2009)

Also Koch (2009), when he identifies the Tartessian language of southern Spain as Celtic, agrees that the Celtic identity was already forged in the Atlantic Late Bronze Age, 13th-8th century BC.

That general conclusion could carry important implications for historians and archaeologists. It reinforces something we have known for some time, namely that the Celtic languages in the Iberian Peninsula—possibly unlike those of Gaul and Britain—cannot be explained as the result of the spread of the La Tène and Hallstatt archaeological cultures of the central European Iron Age. To find Celtic extensively used so far to the south-west at such an early date must also call into question the relevance of Hallstatt’s Late Bronze Age forerunner, the Urnfield cultures, in the Celticization of the Peninsula. (Koch, 2009)

Middle Bronze Age massgrave in Wassenaar. The crimes were most likely committed by Hilversum Culture neighbors, that could make a fit for the people that introduced Atlantic (Celtic?) influences in the north.

Or earlier? There is no agreement yet about the Celtic identity of Tartassian, but an Indo-European language closely affiliated to Italic and Celtic would probably do just as fine in the recognition of an important Copper Age spread of Indo-European emanating from the Atlantic shores. Western continuity all the way back to Maritime Bell Beaker, that established the main fluvial routes in western Europe, suggests scholars are now returning to the idea first proposed in the 1970s that the Celts arrived in the British Isles with Bell Beaker. This steering away from a Celtic origin in Hallstatt and La Tene, even Urnfield, requires a reevaluation of the Celtic nature of the people involved within these cultural horizons. Adherence to the view that Celtic indeed spread along the Atlantic coast would explain the new scrutiny for a Celtic presence in the Low countries. From this viewpoint it would be strange indeed if Celts didn’t reach to the coastal plains of our region, since the North Sea was certainly within Celtic reach.

The linguistic situation in the Low Countries in Roman and pre-Roman times is still far from clear. Obviously, a kind of P-Celtic not unlike Brythonic was held in high esteem, at least among the local elites, and it is likely the Romans sought to tune their local policy accordingly. Local rites and mythology correlate well with the Celtic world and Ingvaeonic language features of Germanic languages along the North Sea suggest a Celtic substrate. However, accepted Celtic toponyms are rare and often associated with Roman military strongholds. Koch is among those that criticize the scientific climate in the Low Countries against clear cut answers about Celtic evidence:

This tendency can be illustrated by the practice of place-name studies in the Netherlands, where names are categorized as either Germanic, Roman, or pre-Roman. However, this last category is rarely specified. (Koch, CCHE, p1192)

However, his criticism is impeded by his own observation that since WW II the same restraints apply for Germanic evidence. He praises Toorians for being one of the few that “dares to say so” (ie. that Celts were omnipresent), but insinuates a lack of linguistic and Celtologic equipment among those that indeed perceived a pre-Germanic substratum but didn’t consider it Celtic and coined the name “Belgic” for it. He did not question their equipment for making a link to Germanic, eg. concerning their observation of glottal stops and Latin transcription issues that may have distorted our interpretation. The political assessment of Koch to address a purported local lack of commitment to the Celtic versus Germanic controversy is probably how foreigners see it: impatient to look into an archeological mirror, they conceive petty Low Countries historical matters as interfering with their demand for the kind of answers that would allow Low Countries to fit in smoothly on their map. Place-name evidence remains controversial and contested in the Low Countries, and evidence that was never filed as Germanic does not imply Celtic by any means. Even Celticist Koch has to content himself with the current status quo: “the conclusion must be that in the Roman period Celtic and Germanic existed side by side“.

The differences between the two linguistic groups, Celtic and Germanic, are significant and difficult to reconcile within one small geographical context, and still the linguistic record in the Low Countries points to all directions at the same time. Maybe there is an argument against impatient bipolar solutions that are ahead of full understanding?

Finds traditionally considered “Celtic” start in the Netherlands at about 825 BC. Now knowing about the direction of Celtic spread between the 13th-8th centuries BC – who can tell us those expansive elements that reached the Low Countries afterward were new groups of ethnical Celts at all? If we follow the logic of Cunliffe and Koch, implying an early Northsea diffusion of the Celtic element, these allegedly Celtic finds could even involve any other foreign element whose ethnic imprint so far couldn’t be conceived.

The Middle Bronze Age Hilversum Culture is the only potential Atlantic intrusion that, in the vein of Cunliffe, could vie as "Celtic".

Cunliffe-style Celtic North Sea diffusion could have been as old as the Hilversum Culture. Indeed, this culture achieved a deep penetration and makes a much better case for a lasting Celtic prehistory in the Low Countries.

Halfway through the Early Bronze Age the practice of cremation and the burial of the remains in Hilversum urns under barrows surrounded by ditches and banks was introduced in the southern part of the Netherlands (especially North Brabant). This clearly indicates close connections with the urn burial traditions of south England (esp. the Wessex biconical urns). The development is now generally interpreted as an evolution, based on regular and intensive contacts, continuing those of Beaker times, and no longer as an indication of the arrival of British immigrants. (L.P. Louwe Kooijmans, 1993)

The culture achieved a strong foothold in the west (Holland, Utrecht) and the south (Brabant and connected Belgian territories), but never succeeded to replace a native element that (also) inherited from the Barbed Wire horizon in the north and, especially, northeast.

This northern Dutch group represents the westernmost manifestation of a tradition that was common all over the North German Plain and the southern part of Scandinavia.
In the late part of the Early Bronze Age and the Middle Bronze Age there was therefore a marked difference in burial practices between the northern and southern parts of the Netherlands.(L.P. Louwe Kooijmans, 1993)

Possibly Hilversum Culture was a culmination of Atlantic interchange dating back to 3rd millennium Maritime Beaker times, but the first attestation of this culture north across the Rhine estuary was a mass grave in Wassenaar, near The Hague. Twelve individuals were slayed and hastily buried, including young children and women.

The use of particular bunal rules and the personal attention paid to the dead suggest that they were buried by captive or escaped kinsmen.
[...]
Very important and most interesting is the aforementioned small flint arrowhead that was found between the ribs of No 10. Its position suggested that it had been shot into the body.
[...]
The arrowhead did not resemble any of the fairly large number of arrowheads found in Bell Beaker graves in the Netherlands (Lanting/Van der Waals 1976). Barbed Wire Beaker graves have yielded hardly any grave goods and no arrowheads whatsoever (Lanting 1973) However, a fairly characteristic, rather sophisticated type of arrowhead with recurved barbs has been found in domestic assemblages. Close parallels have been found in an early Hilversum Culture pit fill at Vogelenzang near Haarlem [...], which are to be dated around 3400 BP
[...]
This evidence soundly dates the grave to c. 3400 BP, 1700 cal. BC, around the transition from the Early to the Middle Bronze Age.(L.P. Louwe Kooijmans, 1993)

Far from being evidence of Atlantic “convergence”, this rather attests post-Beaker divergence. Only the Hilversum culture attest a certain degree of Atlantic convergence. Instead, apparent northern expansion happened at the onset of the Middle Bronze Age and may have been less peaceful than expected, and also less enduring in most places.

The evidence of the Wassenaar grave fits in with Bronze Age burial traditions as far as the extended postures and the custom of inhumation are concerned, although the two were not to become common until a few centuries later.

This pattern of returning native features also emerges in other northern sites:

The practice of cremation was introduced in the northern part of the Netherlands (Drenthe) at the beginning of the Middle Bronze Age. It remained the common form of burial throughout the first half of that penod (MBA-A) and was replaced by inhumation of fully stretched corpses (preserved äs silhouettes) in proper coffins in the second half of the Middle Bronze Age (MBA-B)

Despite the differences both cultural complexes north and south also influenced each other:

There were, however, also similarities between the two areas: in both areas barrows were erected on heavily podzolized soils using sods cut from those soils, in both areas these barrows were surrounded by circles of postholes during the Middle Bronze Age and were used for later secondary burials [...] Moreover, stretched inhumation was occasionally practiced in the south while cremation started to be introduced in the north.

I wonder what Cunliffe and Koch already figured out about the people those purported North Sea Celts met here, since genetic evidence suggests important population changes. Elsewhere along the Atlantic belt, Celts may have spread in the Late Bronze Age, but this approach fails in the Low Countries. Earlier, for sure even the Hilversum Culture didn’t replace all native communities, that kept its most important stronghold in the contemporary Elp Culture just to the northeast. I count on Cunliffe to have noticed this already, since especially in the (Ingvaeonic) northwest a Middle Bronze Age arrival would do much better. Too early?
However, if Celtic expansion to the Low Countries could be traced back to Atlantic influences in the Bronze Age, then the issue of local cohabitation should be addressed as well. The Celtic blueprint of the coastal languages much later (Frisian!) might have been a complicated mixture to start with and only nominally Celtic.

Unfortunately so far we didn’t get a straight genetic picture. Genetic evidence includes the abundance of YDNA R1b-P312 markers within the Celtic hemisphere, but different degrees of marker differentiation into subclades might point at special circumstances in the Netherlands. At first sight an intrusion from the Isles would suggest a profusion of typical British YDNA markers like R1b-L21, but so far such results are far from convincing. The region is littered with R1b-U106 instead, a marker that typically runs high among Germanic Anglosaxons and low among Celts. This R1b-U106 marker clearly diverged from the combined Atlantic “Celtic” association under investigation, that in turn rather points at the very prolific R1b-P312 sisterclade. The vast majority of current YDNA results in the Low Countries thus doesn’t reveal the Celtic affiliation we are looking for: the association between P312 and U106 would confirm Belgae as close relatives to Celts at most. We can’t rule out that some P-Celtic strongholds survived up to Roman times eg. at the coastal regions in the west, then where did the genetic evidence go?

Recent YDNA investigation in Belgium and the southern Netherlands revealed lots of the potentially Celtic P312 marker. However, unlike the British situation this P312 marker is hardly differentiated. Only one subclade, characterized by the U152 mutation, has a clear reading in the south, but nothwithstanding the frequent local association with another L2 mutation this “Celtic” subbranch rather links to the Upper Rhine or (Roman?) Italy than Britain. U152 can’t be the Atlantic marker that accompanied the Celts when they arrived in the coastal marches.

One explanation for the undifferentiated portion of the candidate P312 left may be that Celts arrived in the neighbourhood already before the differentiation of P312 in the main current subclades even occurred. Assumed that local domination of each known subclade of P312 essentially reflects younger events than the initial spread of (undifferentiated) P312*, the current subclade distribution may simply be not decisive in reflecting early Celtic subdivisions. The early age of the Hilversum Culture is in agreement with this results, but there is more in it than a possible earlier onset of Celtic influence: Why did P312 differentiate at the Isles into predominantly L21 subclades, and could differentiation into prevalent subclades happen almost anywhere in Western and Central Europe, but not in the Netherlands? Beware, there still may be some undiscovered Low Countries mutation out there, somewhere downstream P312, that could point at an equivalent local development of the initial Celtic P312 marker, but what does it mean?

Recent YDNA results in the Low Countries reveal substantial portions of potentially Celtic/Atlantic "undifferentiated" P312, but native U106 took the lead. 56% of all R1b in both West- and East-Flanders (WVL / OVL) is U106, compared to 44% in Brabant, and 36% in Limburg. P312* (in blue) shows an opposite gradient: 43% in Limburg, 35% in Brabant, 31% in East-Flanders and 26% in West-Flanders.

The predominant survival of undifferentiated P312 may simply indicate the lack of successful young subclades. Was the development of subclades that inherited from the Celtic element effectively suppressed by others that took the lead? Departing from the abundant availability of undifferentiated P312, the purportedly Celtic element that entered the Low Countries can’t be denied to have been numerous, but apparently their domination was short-lived. The subsequent leading groups inherited from a local element that was characterized by essentially non-Celtic U106.

Schrijver made a case for Celtic substrate influences, ultimately linked to the Neolithic advance. Especially the Celts of NW Europe were affected, and they in turn passed some of their linguistic features over to the Ingvaeonic hemisphere, that includes the West-Germanic coastal regions of the Northsea: West Flanders, Holland, Frisia, England, and to a lesser extend also more inland continental regions. Exaggerated Ingvaeonic use of verbs like to come, to become and to do all seem to compensate for ancestral Celtic restrictions. Still, linguists such as Gysselink and Van Durme identified ancestral linguistic features in the Low Countries that are impossible to derive from Celtic, dubbed Nordwestblock or Belgic. For instance, the Celto-Italian languages miss the glottal stop, probably an important PIE feature, and their IE vocabulary is remarkable distinct. Nordwestblock linguistic features, on the other hand, definitely included the glottal stop, like Germanic. This adds up to the scenario that Celts were immigrants that remained in close contact to surviving native elements. How did this mixture look like, and how “Celtic” this mixture could remain?

Caesar was the first to tell us more about the Belgae, a brave powerful confederation of tribes in the north of Gallia, that was clearly distinct from neighboring tribes. The precise ethnic identity of the Belgae, as well as their language, is still a mystery. Caesar suggested their tribal affiliations were predominantly Germanic, at least historically:

…that the greater part of the Belgae were sprung, from the Germans, and that having crossed the Rhine at an early period, they had settled there, on account of the fertility of the country, and had driven out the Gauls who inhabited those regions; and that they were the only people who, in the memory of our fathers, when all Gaul was overrun, had prevented the Teutones and the Cimbri from entering their territories (Ceasar, D.B.G., 2.4.2)

Still, a lot of reasons were perceived to link Belgae to the Celts, not least of all because the Romans assigned their territory to Gaul. Their status of “having crossed the Rhine at an early stage” thus was readily argued to be part of a premeditated design to claim their “Gaulish” lands. However, bolstered by the Celtic names that prevailed among the Belgae, archeologists could readily reconcile this trans-Rhine origin according to Caesar with a Celtic identity anyway, because this happened to coincide perfectly with their alleged Central European Hallstatt/La Tene origin: Caesar had it all wrong, instead they were proposed as Celtic refugees being pushed over the Rhine by increasing Germanic pressures and expansion, rather than being of Germanic stock themselves. Indeed, unlike the Germanic people described by Tacitus, the Belgae were participants of the La Tene culture and their personal and tribal names were often Celtic. Caesar already distinguished the “Germani cisrhenani” among them: Germanic tribes that according to Caesar crossed the Rhine much earlier. Note that about 150 years later the surviving representatives of Ceasars “Germani cisrhenani” were not mentioned by the Roman historian Tacitus anymore. Instead he claims the Tungri were among the first Germani to cross the Rhine. This appears irreconcilable to Caesar’s account. For really being the first the Tungri should have been the contemporaries of the Eburones, unless Tacitus employed a very different definition of the Germanic identity than Ceasar. Modern interpretation holds them to have crossed the rivers later, to occupy the abandoned wastelands of the Eburones (Germani cisrhenani), once these people were claimed exterminated by Caesar.

The name Germany, on the other hand, they say is modern and newly introduced, from the fact that the tribes which first crossed the Rhine and drove out the Gauls, and are now called Tungrians, were then called Germans.

Caesar didn’t mention the Tungrians at all, but Tacitus didn’t mention his Germani cisrhenani so probably this potential anachronism also reflects changing contemporary ideas about the Germanic ethnicity.

It has all appearance the “Germani” received their Latin name for being the brothers (~ L. germanus) of people that dwelled on the conquered hither, or Roman side of the Rhine. This set them apart as the brothers of the Belgic “intruders”, thus being next of kin to the perceived “aliens” in Gaulish territory that the Caesar claimed for the Romans in his war against the Celtic “foe”. What was the truth behind the Roman assertion that Belgae were not indigenous? Unlike the Aquitani, that harbored another non-Celtic population, the Belgae were not subordinate in any way to the fiefs of Celtica. Neither they were directly involved in the historic Gallic sack of Rome at 390 BC and thus didn’t make a credible natural enemy of Rome. Wouldn’t Caesars claims on territories so far to the north be void if the Belgae were held native to this part of Gaul? And if they were native, what gave the Romans the credence that possibly they weren’t? Maybe the Belgic expansive past had something to do with it, like with Sequani, or with the Suessiones from the neighbourhood of Soissons, possibly an even more important Belgic tribe:

That the Suessiones [...] possessed a very extensive and fertile country; that among them, even in our own memory, Divitiacus, the most powerful man of all Gaul, had been king; who had held the government of a great part of these regions, as well as of Britain (Ceasar, D.B.G., 2.4)

Remarkable that Caesar could relate the veracity of this history to his people’s “own memory”. Does this imply an exception, and utter ignorance of all other previous whereabouts of the Belgae? The Roman picture of Celts being the original inhabitants deserves closer scrutiny. From Cunliffe and Koch we already know that the Celts may have spread along the Atlantic coasts. Their foothold in the northern region does not exclude a mixed population and a strong native element. Indeed, the Belgae, non-Celtic element could have originated in the neighbourhood, beyond the memory of anybody. We should assess their cultural, linguistic and ethnic position to make a comparison with the only other potential natives of the wider region we know of: the Germanic tribes.

Caesar was inclined to assign the Belgae a Germanic origin or identity, but apparently Tacitus narrowed down the criteria substantially. Even though he idealizes the physique “purity” of those he considered truly Germanic, he focused on cultural arguments to set the Germanic world apart, albeit ignoring linguistic evidence. Obviously, in their Celtic association the Belgae didn’t meet his “ideal” of cultural backwardness, at least in his time, long before the ultimate Roman withdrawal that preluded the abysmal cultural decline known as the Migration Period. Belgae claims to any Germanic identity were reduced to an ideal: “The Treverians and Nervians aspire passionately to the reputation of being descended from the Germans“.

The modern Germanic denomination is defined by language and can’t be equated to the Roman definition. Due to the Celtic influence and their cultural achievements it may have been impossible for them to appreciate the ethnic ties between Belgae and Germanic. Until today Tacitus’ approach is influential and archeologists still tend to distinguish ethnicity by a purported ability to tell a metal coin apart from a cow. But all the contrary, the traditional denomination of the Germani may involve just a subset of a single people on both sides of the Rhine.

While Ceasar reckon the Belgae essentially as Germani that adapted to the Gallic way of life, Tacitus obviously disagreed when he wrote in Germania:

“The Germans themselves I should regard as aboriginal, and not mixed at all with other races through immigration or intercourse.”

“For my own part, I agree with those who think that the tribes of Germany are free from all taint of intermarriages with foreign nations, and that they appear as a distinct, unmixed race, like none but themselves.”

Tacitus’ picture of the Germani clearly owes much to the tradition of ‘hard primitivism’ (Rives). His Germania was a political document to contrast Roman decadence with the values of the “noble savage”. The purported “purity” of race and behaviour, however, was contradicted by their strife and tribal heterogeneity. Archeology doesn’t confirm Tacitus’ presentation of a homogeneous Germanic people, nor does YDNA markers reveal genetic unity: YDNA haplogroup R1b-U106, once considered a “Germanic marker”, turned out to be dominant only in the NW (Low Countries, NW France, England): the subclade is low in Scandinavia, while having intermediate frequencies in Germany. Even at low frequencies U106 could be considered widespread at almost all Germanic speaking areas, but the highest frequency zone fits the historic Belgae whereabouts almost exclusively and seems to be there especially to defy the would-be Germanic homogeneity that is commonly upheld.

Closer scrutiny reveals even Tacitus ambiguous about the fabled Germanic homogeneous unity and the following line, when he expressed his serious concerns about the Germanic power, says it all:
“May the Gods continue and perpetuate amongst these nations, if not any love for us, yet by all means this – their animosity and hatred towards each other: since whilst the destiny of the Empire thus urges it, fortune cannot more signally befriend us, than in sowing strife amongst our foes.”

The processes towards heterogeneity are likely to have affected both Celts and the native element. Ingvaeonic features may point at a gradual language shift from Celtic to Frisian, but from a Nordwestblock perspective there might have been an intermediate stage. I could be argued that Frisian didn’t draw directly from a Celtic substratum, but from a heavily Celtified native element that inherited from a linguistic group more closely related to Germanic. Nordwestblock languages are described to feature an abundance of suffices already lost in Germanic, that instead follow the Celtic use of compound words. Phonetic conservatism and features like “nn” gemination , a typical doubling of consonants in toponyms often identified as Nordwestblock, are proposed to point at tendencies that refer to a linguistic situation dating before the Celtification Belgium and Northern Gaul. To a lesser extend West and Central Gaul may be included, until the Garonne. More striking are similarities in Germanic, where gemination accompanied the weakening of the glottal.

Glottalization is found in five out of the ten surviving branches of Indo-European, viz. Indic, Iranian, Armenian, Baltic, and Germanic. (Kortlandt, 1993)

This feature tends to correlate the Nordwestblock substratum geographically to the group of languages in the North European Plain rather than the Atlantic, thus to the archeological horizon that also includes the northern Dutch group, referred to by Louwe Kooijmans.

The emerging Atlantic view and the potential exclusion of the Celtic origin from the North European Plain is screaming for a new assessment that relates Celtic and Germanic from the perspective of a western contact zone. The Germanic vocabulary might owe more from the west than previously conceived. Visualized by some examples, the Cornish/Welsh word lann or llan occurs frequently in place-names. Originally meaning “land”, it gradually came to mean “churchyard” and then “church” and “parish”. But how strong this original meaning ‘land’ an be confirmed to be embedded in the Celtic language? It could have been a Belgic loan, closely related or equal to Germanic “land”. In the Atlantic view the reverse might be true.

The Dutch river mouth mentioned by Pliny the Elder was called Helinium in accusitivus, what would probably indicate a Latin river name Helinius, consistent with the Latin ending -us that generally applies to rivers. Normally, -ius instead of simply -us would imply a derivation of a region called Helinus. In Frisia numerous toponyms feature the hel element, what Clerinx translates into “low lands, marsh” and subsequently connects to Brythonic “marsh” or “estuary”. Other Celtic etymologies have been proposed, like “salt” – probably inspired by now obsolete ideas that involve a Hallstatt origin of Celtic. The Friso-Brythonic etymology does a much better job in addressing reminiscent Celtic features in Frisian, or the Ingvaeonic hemisphere as a whole. The implication would be that the -lin suffix might as well have been Germanic, distorted by Latin transcription issues. This intertwining of ancestral Celtic heritage and west Germanic loans and culture could be extended to the puzzling etymology of local goddess Nehalennia (also transcribed as Neihalennia), that now from a mixed local heritage easily translates to water-ghost (nikker ~IE *neig, to wash) of a region called Halennia – not unlike the latinized form Helinus deduced above. Since the description of the region delimited by Pliny between Helinium ac Flevum neatly corresponds to the historic region of Holland, I wonder if this is mere coincidence or that Holland indeed represent the ultimate indication of a lost Celtic heritage. This mixture could be symptomatic for the almost intangible potpourri that is might be implicated by the Nordwestblock or “Belgae” denomination. This may have been nothing but emerging West Germanic from a shared heritage.

Worse, it might be impossible at all to draw the line between the northern proto-languages of Germanic and Nordwestblock. The Proto-Germanic period is where Grimm’s Law took effect and evolved into Germanic , having time-range 700-100 B.C. generally considered to be the best shot in the dark. The length of this period resembles the High German consonant shift, globally dated 3rd to 5th until 9th century AD, thus having taken 400-600 years to complete. This very recent 100 BC date boundary for Germanic is remarkable and may reveals a huge insecurity about the Germanic origin and expansion. Much more recent split dates between groups make the assumptions involved in the Germanic genesis even more tenuous. IMO, if that 100 BC date boundary is based on the first indications of post proto-Germanic Germanic, then those first Germanic attestations may as well represent the first onset for the spread of this sound shift. Germanic sciences are traditionally hampered by an overkill of migrational mischief, and most of the ideas are 18th-20th century debris. Taken the fragmentary information of frontier tribes in consideration, the Proto-Germanic period could as well have lasted between 100BC – 500AD, or 300 BC-100 AD, and thus in an alternative grouping equally well have included conservative Nordwestblock/Belgae dialects, depending on the degree of Celtic integration already achieved in (pre-)Roman times.

If we take Cunliffe and Koch seriously, the Celtic influence along the North Sea was a lot older than conventional wisdom that stems from migrational La Tene or Hallstatt theories and the Roman interpretation: Late Bronze Age at least, in a Bronze Age Atlantic context. In the Low Countries such an unequivocal Atlantic period is very likely to be of an even older date. Some Celts might have lingered in the swamps for a longer period, but increasing continental contacts from the North German Plain and returning native styles seriously challenge the survival of an unequivocal Celtic ethnicity up to Roman times. The change from Celtic to Germanic, and especially the Ingvaeonic part, could have been much more gradual. A Celtic world strongly suggests the feasibility of an adjacent non-Celtic world, and the Low Countries is where both worlds met. It would be silly to suggest a unified Celtic world that existed since Bronze Age, but deny any consistency of a non-Celtic world in the North German Plains that was attested largely contemporaneous. A shared development at the contact zone for over at least 1000 years opens up the possibility of thorough Celtic influences on Germanic vocabularity and linguistic features in the north that ultimately were be no means confined to the Ingvaeonic hemisphere.

• B. Cunliffe – Iron Age communities in Britain: an account of England, Scotland and Wales from the Seventh Century BC until the Roman Conquest, 2005, link
• J.T. Koch – A Case for Tartessian as a Celtic Language, 2009, link
• J.T. Koch – Celtic culture: a historical encyclopedia, Volumes 1-5, 2006, link
• P. Schrijver – Keltisch en de buren: 9000 jaar taalcontact,2007, link or try translate
• H.Clerinx – Kelten en de Lage Landen, 2005 ISBN: 90-5826-324, p.78, 82
• S. James – Exploring the World of the Celts, 1993, ISBN-13:978-0-500-27998-4, p.103,141,146
• L P Louwe Kooijmans – An Early/Middle Bronze Age multiple burial at Wassenaar, the Netherlands, 1993, link
• J.E. Bogaers – Over de naam van de godin Nehalennia, 1972, link
• F. Kortlandt – General Linguistics and Indo-European Reconstruction,1993, link
• Julius Caesar – The Gallic Wars (D.B.G.), link
• Tacitus Germania – Translated with Introduction and Commentary by J.B. Rives, 1999, link. Try online translation here.
• DNA-project oud hertogdom Brabant, 2009, link
• Larmuseau et al. – Micro-geographic distribution of Y-chromosomal variation in the central-western European region Brabant, 2010

Further reading:

• Celtic from the West: Alternative Perspectives from Archaeology, Genetics, Language and Literature, edited by Barry Cunliffe and John T. Koch (June 2010), announced
• M. Counihan – An Etruscan Solution to a Celtic Problem, 2009, link

The collection of personal adornments and artifacts suggestive of symbolic behavior from the Early Upper Paleolithic deposits of Denisova Cave, Altai, is one of the earliest and the most representative of the Upper Paleolithic assemblages from Northern and Central Asia. Especially important is a fragment of a bracelet of dark-green chloritolite, found near the entrance to the eastern gallery of the cave in the upper part of stratum 11. The estimated age of the associated deposits is ca 30 thousand years. According to use-wear and technological analysis, techniques applied for manufacturing the specimen included grinding on various abrasives, polishing with skin, and technologies that are unique for the Paleolithic – high-speed drilling and rasping. The high technological level evidences developed manual skills and advanced practices of the Upper Paleolithic cave dwellers. (Derevianko et al., 2008)

Humans spend more time gathering around the campfire to celebrate their victory on Nature, only to challenge evolution in an entirely new way.

Neanderthal were readily dismissed as potential authors of local Upper Paleolithic art, due to what boils down to a deep distrust against anything that would deem them capable of such a feat, and they were the only other early hominins around that we knew of – at least culturally speaking, since we don’t have much more than a little pinky after all. And indeed, the first genetic results showed the world was right about one thing: the little finger did not belong to a Neanderthal child. But nobody could have guessed how wrong the usual lot of junk scientists were about almost anything else. This was not the child from the same flesh and blood of modern humans, but a member of a previously unknown ancestral human subgroup.

Dr. Johannes Krause, of the Max Planck Institute in Germany, sequenced the entire mitochondrial DNA (mtDNA) genome and showed almost two times as many differences to modern human mtDNA as does Neanderthal mtDNA. You can find the genome at GenBank or EMBL using record ID FN673705 and check it out by yourself: Even Neanderthal was a close relative to modern humans compared to this hominin!

A phylogenetic analysis similarly shows that the Denisova hominin mtDNA lineage branches off well before the modern human and Neanderthal lineages (Fig. 3). Assuming an average divergence of human and chimpanzee mtDNAs of 6 million years ago, the date of the most recent common mtDNA ancestor shared by the Denisova hominin, Neanderthals and modern humans is approximately one million years ago (mean = 1,040,900 years ago; 779,300–1,313,500 years ago, 95% highest posterior density (HPD)), or twice as deep as the most recent common mtDNA ancestor of modern humans and Neanderthals (Krause, 2010)

Established paleo-anthropology is now faced with the challenge to rewrite the book of human evolution. And of course first things first, the dates were adjusted to make a better fit with pre-AMH cultures:

We note that the stratigraphy and indirect dates indicate that this individual lived between 30,000 and 50,000 years ago. At a similar time individuals carrying Neanderthal mtDNA were present less than 100 km away from Denisova Cave in the Altai Mountains, whereas the presence of an Upper Palaeolithic industry at some sites, such as Kara-Bom and Denisova, has been taken as evidence for the appearance of anatomically modern humans in the Altai before 40,000 years ago. (Krause et al., 2010)

Nobody has ever heard of pre-AMH bracelets, so let’s conveniently forget for a while about that fragment of a polished bracelet with a drilled hole, that was found earlier in the same layer that yielded the bone. Is it possible that here we have evidence that points to a third species, next to Neanderthal and AMH? A species, that might have been as civilized as a AMH, or a beast our ancestors didn’t breed with, or anything else that didn’t involve “us” so we can understand? The publication of Krause carefully omitted this pressing question and the word went out that for sure Krause had already access to autosomal data that could explain why. That Denisova child might have been anything but a Yeti.

Sure, mtDNA doesn’t make a species, no matter how different it may be from modern humans. There was no need for Krause to mention this. But divergence of mtDNA lineages has been taken as an indication of divergent hominin developments before. Explicitly with respect to Neanderthal, whose attested and validated mtDNA lineage was deemed sufficiently homogeneous and different from ours to provoke a definite ordeal. However, now we have the Denisova mtDNA sample to teach us modesty. After all, there are lots of things about mtDNA that need better understanding before we can even attempt to solve the question of how the modern forms spread, and how they evolved.

All conspires against the notion that paleogenetic mtDNA of Neanderthal, and now even more so the mtDNA from Denisova Cave, might be the precursor of modern mtDNA. It couldn’t have evolved so rapidly to modern mtDNA. A study on 44,000-year-old remains of Adelie penguins in Antarctica even confirmed the potential overestimation of the mutational change that is used for dating mtDNA of paleogenetic samples. This stems from a bias that is caused by nonsynonymous mutations, involving notable coding changes that are potentially deleterious and most likely won’t persist very long due to natural selection. Accordingly, only a portion of the mutational changes can be observed over a longer period of time:

Rates of evolution of the mitochondrial genome are two to six times greater than those estimated from phylogenetic comparisons. Subramanian et al., 2009)

The investigation showed that only the effect of synonymous mutations (“silent mutations”) in the mtDNA genome, that involve coding synonyms for the same proteins, remain stable. To retrieve the phylogenetic dates only these “silent mutations” should be measured, ie. changes on coding genes that produce coding synonyms that won’t affect the function of the gene. Mutations that effectively change the functionality of a gene and thus are most likely to be (slightly) harmful, get lost over time, since such mutations would finally bring about the extinction of a lineage and thus shouldn’t count for calculating the age of surviving lineages. The mtDNA “molecular clock” thus should only involve properly identified “silent mutations”.
This results were also important for interpreting the paleogenetic mtDNA samples of hominins.

Mildly deleterious mutations initially contribute to the diversity of a population, but later they are selected against at high frequency and are eliminated eventually. Using over 1,500 complete human mitochondrial genomes along with those of Neanderthal and Chimpanzee, I provide empirical evidence for this prediction by tracing the footprints of natural selection over time. The results show a highly significant inverse relationship between the ratio of nonsynonymous-to-synonymous divergence (d_N/d_S) and the age of human haplogroups. Furthermore, this study suggests that slightly deleterious mutations constitute up to 80% of the mitochondrial amino acid replacement mutations detected in human populations and that over the last 500,000 years these mutations have been gradually removed. (Subramanian, 2009)

Interestingly, this d_N/d_S ratio among Neanderthal was initially reported strikingly high.

These results suggest that slightly deleterious amino acid variants segregate within populations, and that differences in the intensity of purifying selection may affect mtDNA dN/dS ratios. Previous estimates based on mean pairwise differences (MPD) within the mtDNA HVRI suggested that Neandertals (MPD = 5.5) had an effective population size similar to that of modern Europeans (MPD = 4.0) or Asians (MPD = 6.3), but lower than that of modern Africans (MPD = 8.1) (Krause et al., 2007b). Recent population genetic analyses have revealed a higher mtDNA amino acid substitution rate (Elson et al., 2004) and relatively more deleterious autosomal nuclear variants (Lohmueller et al., 2008) in Europeans than in Africans, presumably due to the smaller effective population size of Europeans. Thus, it seems plausible that Neandertals had a long-term effective population size smaller than that of modern humans. (Green et al., 2008)

However, the new information supplied by the Denisova hominin reveals this assumed feature of Neanderthal mtDNA was actually a mistake:

The 12 proteins encoded by the Denisova hominin mtDNA (excluding ND6, Supplementary Information) show low per-site rates of amino acid replacements (dN) when compared to the per-site rates of silent substitutions (dS), consistent with strong purifying selection influencing the evolution of the mitochondrial proteins (dN/dS=0.056). Notably, when the evolution of mitochondrial protein-coding genes in modern humans, Neanderthals, chimpanzees and bonobos is gauged in conjunction with the Denisova hominin mtDNA, a previously described reduction of silent substitutions causing an increased dN/dS in Neanderthals is not observed. This is probably due to a more accurate reconstruction of substitutional events when the long evolutionary lineage leading to modern humans and Neanderthals is subdivided by the Denisova hominin mtDNA (see Supplementary Information) (Krause et al., 2010)

The immediate result of this new finds is that an earlier proposed reduction in length of the Neanderthal mtDNA lineage “about three times as large as would be expected if it was entirely due to the age of the fossil” (Green, 2008), resulting in an earlier common ancestor to modern humans, is wrong. The shrunken phylogenetic tree was accordingly corrected for by Krause: the mean age of the most recent mtDNA ancestor of modern humans and Neanderthal went down from 660.000 t0 465,700 years ago.

(mean = 465,700 years ago; 321,200–618,000 years ago, 95% HPD) (Krause et al., 2010)

The feature that contemporary dN/dS values of modern humans are high, especially among Europeans, also corresponds to current assumptions that concern a younger age or (in the case of Europeans) of a smaller effective population size. May this be another lousy interpretation of results that are barely understood? This could be another example of a solution that supplies an easy way out of a complex issue.

There might be more. COX2 is a coding gene located on mtDNA. According to Green et al.(2008):

COX2 has experienced four amino acid substitutions on the human mtDNA lineage after its divergence from the Neandertal lineage [...]

Fixed mutations indeed tend to define both human lineages as mono-phyletic blocks. But the paper only mentions COX2 as a potential indication of divergent evolution, and due to the new information revealed by the Denisova hominin nothing remains of Green’s assertions that Neanderthal coding mtDNA is strikingly different from modern human mtDNA. The main argument why this would be irreconcilable with a continuous development can now be rejected:

The observation of four nonsynonymous substitutions on the modern human lineage, and no amino acid changes on the Neandertal lineage, stands in contrast to the overall trend of more nonsynonymous evolution in Neandertal protein-coding genes (Table 1), and deserves consideration. (Green, 2008)

There is NO overall trend among Neanderthal towards a more nonsynonymous evolution, hence the four new proteins that correspond to four nonsynonymous substitutions on the modern human lineage do not indicate a striking new tendency, since this kind of mutations happened all the time, also among Neanderthal.
The age calculations gain in reliability once the synonymous mutations involved are better identified and harbored on the phylogenetic tree, by comparing more hominins and branches. Quite considerable purifying selection has now been identified as applicable to both Denisova and Neanderthal mtDNA. However, the mtDNA of an old skeleton in Australia already showed us that neither of this leads us closer to the mtDNA of modern humans.

Whatever the nuance of details, that scream variety and continuity in human evolutionary development, we can’t deny a striking, almost exclusive unity of AMH mtDNA compared to the different forms that have been recovered from Neanderthal and – even more – Denisova:

The genealogies of mtDNA sequences in most human populations, including Aboriginal Australians, characteristically have very little hierarchical branching structure. This pattern of sequence variation is consistent with a population expansion following a population bottleneck and is generally taken as supporting the recent out of Africa model. Under this model, all contemporary sequences spread globally with an expanding population that replaced all other people and all other lineages. Africa has been postulated as the source of the expansion because some populations in Africa have more sequence diversity than populations anywhere else. (Adcock et al., 2001)

Almost, since the discovery of ancestral mtDNA of the gracile early human, found at Lake Mungo, Australia (code named LM3, age 62 kya), that is unmistakably an AMH, also attests the extinction of quite distinct outliers. There must have been a huge and progressive selective thrust towards modern mtDNA. The mtDNA of LM3 was kind of “modern” alright, but definitely the genetic distance fell outside the range of modern humans. The investigators observed this find poses a serious challenge to the “interpretation of contemporary human mtDNA variation as supporting the recent out of Africa model” (Adcock et al., 2001), effectively reducing Africa as a refuge for outgroups that have accumulated change and drifted apart rather than being a true indication of the source of all AMH related mtDNA. But even more so, the find strongly indicates that the current lack of hierarchical branching structure among humans can’t be understood as the direct result of a succession of AMH migrational waves alone. Some waves phased out and lost their origin from the record. Could it be possible that something about mtDNA triggered the worldwide substitution of extremely divergent older forms by the reduced array of current forms? Then how did this happen?

The mtDNA genome, modelled as a circle.

Let us regard the issue in a wider genetic perspective and forget about cheap scenarios of cannibal hominins exterminating each other, a view that conveniently ignores autosomal evidence of inter-hominin gene flow. One little segment of non-coding mtDNA can be found on the Displacement (D-) loop or control region, that is involved in repair activities. It has an analogy in the telomers of nuclear DNA, that are highly prone to insertion and deletion processes. This little region may be subject to the random change and stochastic speed-density that are necessary to infer a neutral “molecular clock”, but the location of this region on the mitochondria introduces a substantial bias in the basic assumption of overall neutrality. I will return at this issue.
Several studies have demonstrated the ongoing transfer and integration of mitochondrial DNA sequences into nuclear chromosomes. The evolutionary inclination of mtDNA genes to move from the D-loop control zone to the nuclear autosomal part of the DNA could be studied in more detail on the paleogenetic sample of an AMH fossil found near Lake Mungo, Australia, dated 40kya (Bowler et al.,2003):
“His mtDNA belonged to a lineage that only survives as a segment inserted into chromosome 11 of the nuclear genome, which is now widespread among human populations.” (Adcock et al., 2001)
This particular strand of early human (AMH) mtDNA vanished from the mitochondrial record ever since, all over the world, but the insertion in chromosome 11 flourished, especially outside Africa:

Overall, 39% of chromosomes tested carried the insertion. In four African populations, the frequency of chromosomes carrying the insertion ranges between 10 and 25%, whereas it varies between 38% and 78% in populations tested in Europe, Asia, Oceania, and South America.(Zischler et al., 1995)

Assuming a lower evolutionary rate in nuclear DNA, “these mitochondrial integrations might preserve the ancestral state of the mitochondrial sequence that existed at the time of transposition and could therefore be regarded as ‘‘molecular fossils.’’” (Zischler et al., 1998). Previous investigation on a similar, albeit much older Insert on chromosome 9 that “took place on the lineage leading to Hominoidea (gibbon, orangutan, gorilla, chimpanzee, and human) after the Old World monkey–Hominoidea split” (Zischler et al., 1998), that happened in the range of 17–30 MYA in a common ancestor of all hominoids, already established the value of nuclear insertions for reconstructing ancestral mitochondrial sequences of the Most Recent Common Ancestor (MRCA):

Thus, the MRCA sequence deduced from homologous integrations in different species will represent the ancestral mtDNA sequence more reliably and with less sequence ambiguities than an ancestral sequence deduced from the fast-evolving mtDNA sequences. (Zischler et al., 1998)

The remarkable affiliation of the autosomal Insert with both LM3 and hominoid mtDNA. The newly discovered, potentially ancestral affiliation with the Denosova hominin is not drawn.

The Insert on chromosome 11 definitely suggests fossil information of some early AMH individual, or at least of a hominin that interbred with early AMH. The closest match to the mtDNA of this particular individual was indeed an AMH, the gracile LM3 dated 40kya (Bowler et al., 2003) found in Australia at Lake Mungo. However, a simple comparison of the Insert to the current genome of modern human mtDNA reveals that this individual can’t possibly be the direct ancestor of modern human mtDNA. No close mtDNA matches of LM3 nor the Insert survived and the mtDNA of LM3 doesn’t indicate direct matrilinear inheritance of the original mtDNA source of this autosomal Insert either.

The LM3 Sequence Belongs to an Early Diverging mtDNA Lineage. The divergence of the LM3 sequence before the MRCA of contemporary human sequences is indicated by its grouping with the Insert sequence, which other reports have suggested diverged before the MRCA of sequences in living humans.
[...]
Although this analysis did not reliably establish an early divergence of the LM3/Insert lineage, it demonstrated that the lineage is unusually long. (Adcock et al., 2001)

This presentation of the Insert as a member of a single branch together with LM3 may be an oversimplification. The location of the Insert at the mtDNA phylogenetic tree of humans suggest an even more pronounced outlier:

Upon comparing 243 bp of a human-specific integration (Zischler et al. 1995) that corresponds to the conserved part of the mitochondrial D-loop of all available hominoid (n=14) and human (n=261) mtDNA sequences, only two insert-specific substitutions were traced, with both the ape mtDNA sequences and all human mtDNA sequences being identical at these positions. (Zischler et al. 1998)

Salient detail is that the two Insert specific substitutions (A on 16259 and C on 16288) are now covered by the mtDNA of the Denisova hominin. Even though the other differences with Denisova are big enough to exclude a close affiliation, this remarkable detail invites to the tentative proposal that the divergence of the Insert sequence could have happened long before the MRCA of human sequences that also include LM3.

Between 16,259-16,381 the mtDNA variation of the Insert nucleotides is covered by the corresponding nucleotides of apes, Lake Mungo 3, current aboriginal polymorphisms (not drawn) and the Denisova hominin.

This rare scope on a deep Eurasian affiliation, combined with extant aboriginal polymorphisms that echo the survival of Insert and LM3 features in the haplogroup N and M branches of modern mtDNA, suggest a much more complicated phylogenetic tree than the one currently in use. Aboriginal mtDNA polymorphisms drawn in the figure of Adcock et al., 2001 (above) are part of a mixture of the closely related haplogroups N and M that up to now define the earliest Out of Africa scenario, appear to be closer to an extinct group of Eurasian outliers than African branches. Also typical East Asian loci of mtDNA show a remarkable similarity, what might point at a counter-indication of African branches being ancestral to haplogroups N and M. The reverse may be true. African divergence from CRS and current variability doesn’t cover the Insert loci set in any systematic or significant way. The establishment of such a “reversed tree”, however, is hampered by the apparent extinction or extreme “pruning” of what might have been an enormous Eurasian mtDNA variability. Any scenario that reverses the tree should account for this low extant Eurasian variability in comparison with Africa.

Let’s return to the assumed “neutrality” of mitochondrial DNA inheritance. High variability of the control region might suggest otherwise. One of the prerequisites of fast evolution is a fast mechanism underneath genetic change, and the purpose of fast mtDNA mutations could be just that, to put the precondition of rapid evolutionary change. Anyhow, a similar observation was made concerning the massive STR of chimps on the Y-chromosome, that seem to be secondary to the incredible evolutionary changes on the Y-chromosome as observed in the recent study of Hughes et al. (2010) I already wrote about here.

A set of interesting differences of mtDNA between humans is located on the Hypervariable Region (HVR). Most strikingly, HVR is not highly variable per definition. For instance, investigations on the Ayu fish (Takeshima et al., 2005) revealed the Hypervariable region may also turn into a Hypovariable region, what suggests a special functionality of the property defining HVR (or general D-loop) variability. And a substantial susceptibility to damage.

The mitochondria continuously reproduce themselves at intervals averaging about 2 weeks, like bacteria by a process of binary fission. They generate most of the cell’s (chemical) energy supply and because mitochondria use oxygen as an electron acceptor, they produce harmful free radicals that may cause genetic damage, often deletion mutations. This free radical damage to mtDNA cannot be repaired, basically because the regular repair mechanisms of the cells can’t access the mitochondria and the mitochondrion has no repair mechanism of its own. Therefore, mitochondria accumulate damage at each mitochondrial generation, what gradually leads to malfunction and ultimately affects the health of the organism as a whole.

However, this dreary scenario must have some constraints, or else all life on earth would already have ended millions of years ago. Somehow the reproductory system must have been exempted from this process, or at certain circumstances, and also it seems the genetic damage to mitochondria can be slowed down by exercise, both physical and mental, but especially by consuming antioxidants like vitamin C or omega-3 fatty acids. These are abundant in fresh fruit, raw meat and fish, indispensable supplements to the species that lost the functionality of the L-gulonolactone oxidase (GULO) gene – amongst whom one of the two major primate suborders, the Anthropoidea (Haplorrhini), that happens to include human beings, together with tarsiers, monkeys and apes. Originally meant as a genetic “improvement” for getting rid of the old and weak when food shortages occurred, ie. those most badly in need of antioxidants to remain healthy, the loss of this gene also effectively confined this suborder of primates to subsistence in the tropics. Only humans succeeded in finding new habitats in colder climates. They left the hot places where fruits were available all year round and traditionally made up an important addition to the menu, because they could. Only humans evolved into great hunters, and developed the necessary skills to catch fish, in order to compensate for the irregular availability of fresh fruits. Notwithstanding unfavorable climates, they managed to keep their necessary supply of antioxidants at a save level. And they did, for hundreds of thousands of years. Until everything changed at the eve of Upper Paleolithic – when human cultural advance reached a critical level.

What went amiss when humans reached their first cultural highlights? Their success triggered important improvements in their living standards, that moved their prime focus away from the concerns of harsh survival, and towards the community around the fire. They spend more time preparing their meals, started to cook their meat and fish and thus destroyed their main antioxidant food supplies. Degenerative diseases made their introduction and invoked new selective pressures, that caused a steady gene flow from the south to rejuvenate the slowly degenerating mitochondrial lineages in the north. In the mean while females ceased to worry about the survival of the fittest and developed a preference for “feminine looking men over their more rugged counterparts” (DeBruine et al., 2010), triggering the most notorious changes in the human anatomy that resulted in Anatomically Modern Humans as a progressive tendency all over the world.

However, this does not fully explain the current low overall variability of mtDNA even in fruit-rich tropical territories in comparison to the attested mtDNA of early AMH such as Lake Mungo 3. Still, cultural level related natural selection might be a good trail to follow.

Booming AMH culture most probably also entailed a closer contact between different groups within a wider economical areas. For sure this new behavioral patterns would have initiated a catastrophic increase of contagious diseases as soon viruses and bacteria could circulate freely among newly interconnected communities. However, this also implies a strong relation between resistance against (new) infections and mtDNA, that vastly exceed the benign effects of Vitamin C. The relation between mtDNA, antioxidants and the development of new “genetic” cures may reach a lot further. At this point it is tempting to regress to the behaviour of mtDNA and its facility to travel to nuclear DNA, and evaluate the genetic potential of mtDNA as a genetic laboratory against new diseases. Indeed, the immune system is where human DNA might have evolved most and is where most human variability occur.

Despite the high homology between chimpanzee and human genes at the level of amino acid sequences, human genome contains 1418 genes that do not have direct orthologues in chimpanzee, many of which are related to immune defence.
[...]
A number of genome-wide scans for positive selection have recently been performed (Wagner, 2007). They confirm that many immune genes and their regulatory sequences have been the subjects of positive selection in humans.
Population genomics is still in its infancy and the specific predictions may vary among studies but this is where future discoveries are anticipated. (Danilova, 2008)

Then, survival of just one little branch of early human mtDNA must point directly to the main focus of Upper Paleolithic development. Of the early mtDNA strands only those that accumulated in Africa were safeguarded against the effects of progressive damage, due to the continuous availability antioxidants. But in the center of change the preconditions for rapid change were set, including the extinction of mtDNA that did not meet the new standards of natural selection against the inevitable pandemics of cultural cohabitation and coexistence. Relatively low population density prevented the accumulation of high haplotype diversity, and the surviving mtDNA haplogroups in Eurasia obliterated all traces of a long, rich and diverse hominin history. To the effect that the false positives of mtDNA lured the public opinion into thinking that a long list of pre-AMH hominins, that include famous names like Neanderthal, Peking Man, Rhodesian Man, Denisova hominin etc., became extinct.

We can’t solve the origin question with a narrow scope, since the only truth is that we still don’t know. However, the Denisova hominin shows us one important clue: the more we know, the more complicated the solution. And most probably, the more hominins involved.

• Krause et al. – The complete mitochondrial DNA genome of an unknown hominin from southern Siberia, 2010, link (paysite): try here
• Krause et al. – A complete mtDNA genome of an early modern human from Kostenki, Russia; 2010, link
• Derevianko et al. – A Paleolithic Bracelet from Denisova Cave, 2008, link
• Howell et al. – Molecular clock debate: Time dependency of molecular rate estimates for mtDNA: this is not the time for wishful thinking, 2008, link
• Adcock et al. – Mitochondrial DNA sequences in ancient Australians: Implications for modern human origins, 2001, link
• Ovchinnikov et al. – Molecular analysis of Neanderthal DNA from the northern Caucasus, 2000, link
• Orlando et al. – Revisiting Neandertal diversity with a 100,000 year old mtDNA sequence, 2006, link
• Green et al. – A Complete Neandertal Mitochondrial Genome Sequence Determined by High-Throughput Sequencing, 2008, link (paysite): try here
• Takeshima et al. – Unexpected Ceiling of Genetic Differentiation in the Control Region of the Mitochondrial DNA between Different Subspecies of the Ayu Plecoglossus altivelis, 2005, link
• Sankar Subramanian – Temporal Trails of Natural Selection in Human Mitogenomes, 2009, link
• Subramanian et al. – High mitogenomic evolutionary rates and time dependency, 2009, link
• Zischler et al. – A nuclear ‘fossil’ of the mitochondrial D-loop and the origin of modern humans, 1995, link
• Zischler et al. – A Hominoid-Specific Nuclear Insertion of the Mitochondrial D-Loop: Implications for Reconstructing Ancestral Mitochondrial Sequences, 1998, link
• DeBruine et al. – The health of a nation predicts their mate preferences: cross-cultural variation in women’s preferences
  for masculinized male faces, 2010, link
• Nadia Danilova – Evolution of the Human Immune System Evolution of the Human Immune System, 2008, link
• Allard et al. – Control region sequences for East Asian individuals in the Scientific Working Group on DNA Analysis Methods forensic mtDNA data set, 2004, link (paysite), try here
• Bowler et al. – New ages for human occupation and climatic change at Lake Mungo, Australia, 2003, link
• PhyloTree.org – Global human mtDNA phylogenetic tree, 2010, main

Recommended reading:

• Kris’s Archaeology Blog – Possible New Hominid Species Identified
• John Hawks Weblog – Hobbit version 2.0: the undiscovered hominin
• John Hawks Weblog – An earlier initial Upper Paleolithic at Kostenki
• Science News – Ancient Penguin DNA Raises Doubts About Accuracy of Genetic Dating Techniques
• The Scientist – Surprising mtDNA diversity
• Hendrickson et al. – Mitochondrial DNA Haplogroups influence AIDS Progression, 2009, link (declining Mesolithic mtDNA U5 may be a possible clue towards repeated extinction of northern mtDNA haplogroups).

Buffalo Bill meets his Indian brothers.

Once upon a time there were two brothers. Well, actually there were a lot more of them, but these two were very special: they were destined to become the most prolific progenitors of the world. Their offspring divided in two. Those that went east to cross the Bering Strait, to become the most prominent inhabitants of the New World as far as Tierra del Fuego, were the descendants of one brother, while the numerous descendants of the other brother that remained hither caused a shock-wave in the Old World that could be felt as far as West Africa, the Atlantic and India. Like in a fantasy tale, both people founded incredible civilizations, each highlights of human talent and splendor. Their feats were almost unequaled and competing – even though they were separated by two mighty oceans. Each completely forgot about the existence of the others, until the people on the hither side learned how to tame the waves and crossed the oceans. Their final reunion should have been a reason for joy and celebration but when they met again, 500 years ago, oh pitiful wretches! Moans of distress filled the air and blood soaked the earth. The once great people of the New World never recovered the blow of what could probably qualify in history as the most tragic example of fratricide.

Human Y-DNA genetic tree. The yellow square indicates the relationship between the Eurasian R-clade (the world's most prolific clade), the Q-clade (that dominates the Americas), and the ancestral P-clade.

In the genetic tree these two people are nowadays known as members of the R-clade and the Q-clade. The offspring of the multitudes of other brothers and close patrilinear relatives that couldn’t catch up with their success, dwindled, but some of them survived in fringe areas on both sides of the oceans. For convenience sake these people are considered members of the same ancestral clade as their fathers, the P-clade. Geneticists would easily recognize the Y-DNA markers of each clade, and say that their respective haplogroups were R, Q or ancestral P. Together they form a superclade on a single branch of the genetic tree. They make up a substantial part of the human population, but still they represent nothing but a single branch that on a paleolithic scale can’t be considered very ancient.

Global approximation of the pre-colonial distribution of the P-clade, that include the R-clade (red, west) and the Q-clade (yellow, east and Americas). Note that there is a substantial overlap in Central Asia.

Where did those very successful P-derived haplogroups originate? The story of the two brothers may be an undue simplification since the full development of each clade involved nomads that probably roamed wide swaths of territory during many generations. The Old World was subject to an accumulated prehistory of up to a million years in every single corner, what tend to render our information for reconstructing the relevant demographic events almost intangible. Their legacy doesn’t include a unified language, not even overall genetic uniformity. Their genetic “kind” remains essentially restricted to the male lineage, that never became numerous in most of Africa and remained virtually absent in Oceania and the Far East. The Near East traditionally harbored a rich variety of their male genes in all stages of development, but so far nothing points at a historic predominant position in the region in comparison to other clades that eg. include notorious “stayers”, defined by haplogroup markers like Hg J and Hg E. These current distribution issues hardly give any clue about the origin, especially since more often than not the trail leads to desparate isolated and remote areas, or otherwise to historically cosmopolitan areas that received too much foreign visitors to reveal unambiguous answers. The only security we have is that on a genetic scale those three clades were exceptionally close related along the male lineage and thus that the incredible success of two P-derived branches may point to something systematic underneath, that also applies to the virgin soils of the Americas.

In stark contrast to the abysmal disagreements that surrounds our knowledge about the peopling of the Old World, the disagreements that concern the Americas are almost negligible. The Q-clade entered through the Bering Strait in one or two waves and were the first human beings to do so. They are theorized to originate in southern/central Siberia:

Globally, Y-chromosome data therefore emphasize the critical role of southern/central Siberia in the peopling of the Americas, since this region appears to be at the origin of two major male migratory waves of colonization. The data presented here are also consistent with the intriguing possibility of ancient links between proto-Europeans and proto–Native Americans, an idea that has been put forward in previous Y-chromosome studies (Karafet et al. 1999; Santos et al. 1999; Wells et al. 2001; Lell et al. 2002). An ancestral connection between these groups has also been suggested on the basis of morphological (Brace et al. 2001) and mtDNA (Brown et al. 1998) data and could ultimately trace back to ancient east/west human dispersals from a common source in central Asia. In agreement with this scenario, the P-M45 lineage has been found to be oldest in central Asia (Wells et al. 2001; Zerjal et al. 2002), where the Tuvan population includes haplogroups M242, M45, M173, and Tat, which are now dispersed in Europe and/or America. (Bortolini et al., 2004)

More recent migrational events, of new people that never succeeded to be equally successful in entering America, caused their kind to become rare in East Siberia, thus cutting the umbilical cord that once must have interconnected the great P derived clades on both sides of the Bering strait. We can’t easily derive from facts the precise events that should have linked the clades mentioned above with proto-europeans, but the link to proto-Native Americans is virtually straightforward and unequivocal. This security may serve as a sound base to analyze and verify the processes involved in deeper detail. Let us depart from the general agreement, that concern an origin from Central Asia and widespread habitation of the Americas during the end of the last glacial period, known as the late glacial maximum, around 16,000 — 13,000 years before present. How should this affect our assumptions that link medium term migrational processes to the origin of languages and genetic variability, and how the corresponding observations should contribute to our understanding of what happened elsewhere, ie. in the Old World, including Europe?

First of all, let us consider the pre-columbian era. Native Americans feature a seemingly disproportionate amount of linguistic groups and isolates, that at first sight hardly agree with the homogeneous origin implied by one or two migrational events from the same direction. Only the Na-Dené languages of North America, that include speakers of Athabaskan languages like Apache, seem to find a distant relative in the virtually extinct Ket language of Central Asia – that indeed feature the Q-haplogroup in remarkable high proportions. The languages are tentatively grouped together in the Dené-Yeniseian language family. Possibly the proto-Native Americans migrated wholesale to the Americas and didn’t leave much of a trace in Asia behind, or the descendants of their relatives that once spoke related languages in Asia became overwhelmed by groups that entered later into Siberia from the southeast, from the south or from the west across the Ural mountains. Both scenarios may apply, especially considering the advance of additional haplogroups as far as the Pacific coast of East Siberia that remain strikingly rare or absent in the Americas, like Hg N, several ancestral Hg C varieties and even Hg D. We don’t know about American Hg R, since up to now the ample presence of this haplogroup among virtually all Amerindian groups has been interpreted as recent European admixture. As a side-note could be mentioned that claims of Amerindian Hg R are indeed made and may find some confirmation in STR mutations on the Y-chromosome whose statistical properties deviate significantly from European Hg R (eg. less than 10% DYS390=23 among Native-American Hg R individuals, what plainly contradicts a West-European, and especially a Northwest-European origin). However, the general picture is that the migrational events that are responsible for the Amerindian presence were dominated by a small group of Siberian people whose males belonged especially to the Q-clade. Whatever the genetic and linguistic variety of Amerindian people nowadays, somehow this must relate to this potentially quite homogeneous people that arrived from the Arctic.

This purported ancestral homogeneity, barely 15.000 years ago, is confirmed by low intra-population variety in South America, but how this could be reconciled with the maximum genetic distance with respect to the rest of the world?

(2) Consistently with previous studies, South Amerindians and Oceanian populations show the lowest intra-population diversity when compared with autochthonous populations from other continents. (3) Within South America, Western South Amerindian populations show the highest intra-population diversity, consistently with an evolutionary model previously proposed by us, which suggest a higher long term effective population size and levels of gene flow among them. (4) Comparing the different continents, the between-population diversity is clearly the highest for South Amerindians (Fst = 0.19). This value is almost twice the level of differentiation observed for the worldwide population (Fst =0.11).(E. Tarazona-Santos, 2009)

The human world population attests an overwhelming importance of geneflow: Fst increases proportionally with distance rather than anything else. South America is at the end of the line.

Fst (Fixation Index) measures random admixture levels but whatever this may imply, this extreme South American value can’t be due to mere ancientness of the population. On a world-wide scale, the globally linear increase of Fst with geographic distance doesn’t confirm Heyerdahl-style immigration from weird directions and rather indicate other, more sofisticated processes are at work. The geographic position at the end of all migration lines, ie. more so if indeed traversing the Bering Strait, and a corresponding distance-driven isolation against the effects of geneflow may offer a more plausible explanation.

Could we hypothesize similar causes behind the incredible variety of Amerindian languages? Simple language drift may be responsible. This mechanism defines language as vocabulary cast into the mold of a particular syntax, subject to change. Over time the basic structure of the sentence, held together by functional items (~grammar), with the lexical items (~words) filling in the blanks, will thus be deeply modified, together with the “outer appearance” of a particular language, and affect grammar in all its (morphological and syntactic) aspects. The process may be gradual, the product of chain reactions or subject to cyclic drift, and causes the development of completely different linguistic characteristics of originally related languages that can’t be explained by the mere development in isolation of vocabulary.

In the Americas – including the most southern parts – we are thus confronted with a native population that exceeds the arctic origin location in linguistic and genetic diversity. This is completely contradictory to the common wisdom that homogeneous populations are younger and derived from populations that feature more diversity! So let’s return to the Old World and investigate how this insight might shed new light at the origin of the closely related R-clade.

The descendants of that other brother, that was the progenitor of the R-clade, are difficult to define. If we choose to ignore the possibility that some might have followed their kin of the Q-clade to the Americas through Siberian, we can observe that the R-clade essentially has a more western and southern distribution. The clade is subdivided in several groups that rarely escaped the attention of those that sought and Indo-European association. An unambiguous exception to the Indo-European bias was provided by the Westafrican subclade defined by mutation R1b-V88, that instead was related to the Afrosasiatic linguistic group. More specifically, the Chadic subgroup:

The analysis of the distribution of the R-V88 haplogroup in 41800 males from 69 African populations revealed a striking genetic contiguity between the Chadic-speaking peoples from the central Sahel and several other Afroasiatic-speaking groups from North Africa. The R-V88 coalescence time was estimated at 9200–5600 kya, in the early mid Holocene. We suggest that R-V88 is a paternal genetic record of the proposed mid-Holocene migration of proto-Chadic Afroasiatic speakers through the Central Sahara into the Lake Chad Basin, and geomorphological evidence is consistent with this view. (Cruciani et al., 2010)

Since this subgroup of the R-clade doesn’t derive from the subclades that are most frequently indicated as “Indo-European”, the simple conclusion should be that the R-clade is much older than the Indo-European language family and hence that the success of the Hg R-clade can’t be explained by the Indo-European advance, that indeed has been dated a lot later.

The origin of another important subgroup was recently associated with the Neolithic wave of advance, that is theorized to have started in Turkey:

Does the time to the most recent common ancestor (TMRCA) of the hgR1b1b2 chromosomes support a Paleolithic origin? Mean estimates for individual populations vary (Table 2), but the oldest value is in Central Turkey (7,989 y [95% confidence interval (CI): 5,661–11,014]), and the youngest in Cornwall (5,460 y [3,764–7,777]). The mean estimate for the entire dataset is 6,512 y (95% CI: 4,577–9,063 years), with a growth rate of 1.95% (1.02%–3.30%). Thus, we see clear evidence of rapid expansion, which cannot have begun before the Neolithic period. (Balaresque et al., 2010)

The debate on the origin of virtually all subclades of haplogroup R was repeatedly derailed by the discovery of ancestral samples on disparate locations. Especially the Middle East proved itself a valuable source, to the effect that previous assertions that pointed to Central Asia lost much of their attraction. However, drawing our lessons from the Amerindian situation we could wonder if this change of mindset is justified. A Russian publication (excerpts translated by Google) on the genetic variety of the Bashkir people, just south of the Ural mountain, will possibly make the difference. This study confirms that Bashkir R1b1b2 (also written as R1b-M269), otherwise especially associated to the Neolithic and Europe,  might be very ancient in the region, notwithstanding the exceptionally low diversity.

Baskhir R1b1b2 has an intermediate position between European and Asiatic populations. The largest cluster (γ) occurs equally in all referenced regions (Europe, Balkans, Asia and South Urals), while the β cluster haplotypes correspond predominantly to the populations of Europe, and the majority of haplotypes in cluster α are from South and West Asia.

Some researchers believe that haplogroup R-M269 was spread throughout Western Eurasia in the Upper Paleolithic [Semino et al 2000; Al-Zahery et al. 2003]. The high frequency of the paternal line in populations of the Southern Urals is an unexpected finding, as this haplogroup is not typical for either allied areas (Central Asia, Eastern Europe and Siberia). In order to clarify the origin of this haplogroups in on the southern Urals, we analyzed the phylogenetic relationships between microsatellite haplotypes in populations of Bashkir and West Asia, South Asia and Balkans, Europe (Fig. 4). As a result, phylogenetic analysis identified three clusters microsatellite haplotypes, designated as α, β and γ (Table 6, Fig. 4). It was evident that the bulk of the β cluster haplotypes corresponds to the populations of Europe (50 out of 70 haplotypes cluster β), while the majority of haplotypes in the cluster α (65 of 79 haplotypes) are from South and West Asia.

It should be noted that the largest cluster of the phylogenetic tree, cluster γ haplotypes, occurs equally in all regions (Europe, Balkans, Asia and South Urals). Over 70% of the haplotypes among the Bashkir belong to this cluster, which apparently is the result of the earliest stage (Upper Paleolithic) resettlements of mutation M269 carriers, that cover a larger area.

The inevitable conclusion:

Relatively low population density, and hence the effective population of this region compared to the densely populated regions West Asia, Europe, the Balkans and South Asia prevented the accumulation of high haplotype diversity.

This approach is essentially different from deducting the origin of a subclade – or clade – by only taking in consideration the diversity of surviving haplotypes. Naturally, at the benign living conditions of old cultural areas a high variance can be “accumulated”, when “fossile” haplotypes are “collected” over a large period of time. In the harsh climate of Siberia, however, the living conditions are completely different, and also the survival conditions and subsequent “pruning” of haplotypes. Then the alternative approach for retrieving age should thus consider the diversity in the widest sense, ie. of reminiscent clusters that can be compared with extant clusters on a world wide scale. I gather the “collector” function eg. of the middle east cultural area for “old” R-clade haplotypes has a biological analogy in the jungle, where you can find an enormous variety of species, including “living fossils”. I don’t think this should be explained by a theory saying that all living creatures evolved in the tropics. Variance and diversity of Y-DNA genes thus shouldn’t be taken as an unequivocal indication of origin.

Bashkir R also includes the R1b1b1 subclade, whose Central Asiatic antecedents are rarely disputed, and R1a (R – SRY10831.2). Accordingly, the study concludes:

Since there is no trace of mass migrations from Europe and western Asia, the reasons for the predominance of the main haplogroups (R – SRY10831.2 and R-M269) in the southern Urals are to be found in the processes of early settlement this region.

Possible Holocene migrational nodes of the R1b-clade assuming a correlation to the southern Ural and mtDNA U5b.

The question how the R-clade became involved in the Neolithic expansion and how it also reached Africa will be the sequel of this issue. At the moment it will suffice to point at a possible relation of the Bashkir homelands with the archeological Botai culture, the most likely place where the horse was domesticated and where the first consumption of  (mare)milk was attested. I already mentioned the potential association between the R-clade and gene T-13910 for lactase persistency (including West Africa), what might be especially interesting because Enattah et al. (2007) already located the origin of this gene in the neighborhood of the Ural mountains. Consequently, mitochondrial haplogroup mtDNA-U5b and the autosomal gene associated to Bloodtype B may serve as valuable markers to locate Holocene migrations to West Africa, while the Neolithic wave of advance as a medium for the expansion of the R1b-clade into Europe – due to the low occurence of such migrational markers  on the Neolithic route – could thus be confirmed as essentially characterized by geneflow.

How this processes might have affected also the spread of R1a and R2, the other members of the R-clade, is yet another chapter, being most probably intertwined with that Indo-European question. The issue of the Two Brothers, however,  should be considered without linguistic or cultural implications. The issue of this Siberian Brotherhood is far more important, since the success of the arctic brothers can’t be a coincidence. For sure this issue touches the evolutionary pressures that are especially in force at the conditions of a harsh environment. Hence this issue directly touches our existence. We would never have existed without. Everybody already received all the benefits, since powerful geneflow around the globe continuously forges our species into a single, indivisible brotherhood of mankind.

• Lobov Artem – Structure of the Gene Pool of Bashkir Subpopulations, 2009, Russian link, Translated excerpt
• Bolnick et al. – Asymmetric MAle and Female Genetic Histories among Native Americans from Eastern North America, 2006, link
• Lell et al. – The Dual Origin and Siberian Affinities of Native American Y Chromosomes, 2002, link
• Zegura et al. – High-Resolution SNPs and Microsatellite Haplotypes Point to a Single, Recent Entry of Native American Y Chromosomes into the Americas, 2004, link
• Bortolini et al. – Y-Chromosome Evidence for Differing Ancient Demographic Histories in the Americas, 2003, link
• E. Tarazona-Santos et al. – The genetic structure of Native Americans: inferences from SNPs in genes involved in carcinogenesis, immunity and pharmacogenetics, 2009, link
• Balaresque et al. – A Predominantly Neolithic Origin for European Paternal Lineages, 2010, link
• Cruciani et al. – Human Y chromosome haplogroup R-V88: a paternal genetic record of early mid Holocene trans-Saharan connections and the spread of Chadic languages, 2010, link (paysite)
• Enattah et al. – Evidence of Still-Ongoing Convergence Evolution of the Lactase Persistence T-13910 Alleles in Humans, 2007, link

Further reading:

• Ted Goebel et al. – The Late Pleistocene Dispersal of Modern Humans in the Americas, 2008, link

“It is very important not to use the same lab to analyze the DNA of living and dead people as there may be confusion in the results,” Dr Hawass said. “I used to be against the DNA tests for mummies, because it was done by foreigners, and the mix of DNA of the dead and the alive could lead to inaccurate results” (Chinaview).

Nefertiti, the sister of Akhenaten was also his wife.

An interesting comment, that makes me wonder if Tutankhamun was already sampled before by foreigners, either or not within their own compounds when testing started in 2008, and if the first results were rejected for the reason of an alleged contamination by foreign genes – i.e. western genes. Now the first paleogenetic results come out we should expect that any risk of western contamination is properly dealt with, since the tests were done exclusively by their own people. However, certain irregularities in the divulgation of the results are causing the word to be spread that “western contamination” was possibly meant in the broadest sense, including historical facts. Valuable information is left in the drawer, and this time it ain’t extraterrestrial evidence. Come hither and behold an example of successful Internet espionage.

The 17th of February 2010, The Independent was not the only newspaper to come up with an article on the valuable results that were published by the team of Zahi Hawass to reveal ancestry and pathology in King Tutankhamun’s Family:

The study, published in the Journal of the American Medical Association today, was carried out by a team of archaeologists led by Dr Zahi Hawass, one of the world’s leading Egyptologists. Its aim was to determine the relationships between 11 royal mummies of the New Kingdom, looking for common features which might have been caused by inherited disorders or infectious diseases.

The genetic investigation was successful in attesting malaria (plasmodium falciparum) by tests on genes STEVOR, AMA1 and MSP1, and also resulted into rejection of some genetic diseases that were previously suspected. Most importantly, genetic testing resulted in the identification of family members of the young pharaoh:

More than 55 bone biopsies were used to elucidate the individual relationships of 18th-dynasty individuals, with the result that several of the anonymous mummies or those with suspected identities are now able to be addressed by name. These include KV35EL, who is Tiye, mother of Akhenaten and grandmother of Tutankhamun, and the KV55 mummy,who is most probably Akhenaten, father of Tutankhamun (Figure 2, eAppendix, and online interactive kinship analysis and pedigree). The latter kinship is supported in that several unique anthropological features are shared by the 2 mummies and that the blood group of both individuals is identical.

Most probably Akhenaten?? To prove KV55 as Akhenaten, they have to claim a much older age for the remains than has been attested in 3 full body examinations (1931, 1966 and 2000) to make the age be old enough. This Hawass pet theory seems to rely upon the idea that his successor Smenkhkare simply didn’t exist, what few experts are ready to accept. It has all appearance Hawass is eager to have his version in Wikipedia as the Mainstream Absolute Truth for Dummies, since this is what he achieved. However, what everybody in the world of genetic genealogy is eager to know, the way how Tutankhamun and the other pharaohs of the 18th Dynasty fit into the human family-tree as a whole, was not revealed. Even though the publication mentioned the investigation of the necessary genetic markers. A short explanation. One popular way to find out about the ethnic and geographic origin of a person is doing a male specific test. Genetic details of the paternal line may include a panel of Short Tandem Repeat, STR, markers retrieved from the male Y-chromosome. Such results commonly serve as a first approximation to a subset of (apparently) underpinning mutations, better known as single-nucleotide polymorphisms (SNP), that are DNA sequence variations on a certain base pair. Somehow STR testing is considered more accessible, so up to now most genetic comparison involves balanced panels of STR, that produce a genetic signature referred to as “haplotype”. Now, biological clades may be determined by a unique range of SNP or “haplogroup” that is featured by a group of people having at least one common ancestor. It is commonly assumed a haplotype may give a reliable approximation to the haplogroup the tested person belongs to, thus to the genetic clade or even subclade that may reveal the genetic history of the tested person. At least in theory the ethnic and geographic origin of a person may be derived from this information.

The cited paper on Tutankhamun mentioned the investigation of sufficient STR markers to come at a valid approximation of the genetic profile, thus to the genealogic history of his family line. Explicitly mentioned where STR indicated by the following names: DYS456, DYS389I, DYS390, DYS389II, DYS458, DYS19, DYS385, DYS393, DYS391, DYS439, DYS635, DYS392, Y-GATA-H4, DYS437, DYS438 and DYS448.  However, the values of only DYS393 and Y-GATA-H4 were published. This already triggered a lot of irritation. Why objective scientists would withhold the world information like this?

The family united again after years of mummified solitude. Thanks to the genes!

The 18th Dynasty Haplotype corresponds most likely to European R1b-ht15.

How a European haplotype, having possible Jewish connotations, could have entered the Egyptian royal house to begin with? This doesn’t look like an issue an Egyptian team would be eager to investigate:  Starting with the political and religious animosities between Israël and Egypt, being nowadays part of the Arab world. The biblical history of the pharaoh who promoted Joseph’s rise to authority (Genesis 41:39-46) and subsequent claims of an important role of the Jewish people in Egypt until Exodus, already kindled debate on the Egyptian influence in the origin of monotheism. The biblical account comes close to a complete takeover. Genesis chapter 47:

1. Then Joseph went in and told Pharaoh, and said: ‘My father and my brethren, and their flocks, and their herds, and all that they have, are come out of the land of Canaan; and, behold, they are in the land of Goshen.’  2. And from among his brethren he took five men, and presented them unto Pharaoh.  3. And Pharaoh said unto his brethren: ‘What is your occupation?’ And they said unto Pharaoh: ‘Thy servants are shepherds, both we, and our fathers.’  4. And they said unto Pharaoh: ‘To sojourn in the land are we come; for there is no pasture for thy servants’ flocks; for the famine is sore in the land of Canaan. Now therefore, we pray thee, let thy servants dwell in the land of Goshen.’  5. And Pharaoh spoke unto Joseph, saying: ‘Thy father and thy brethren are come unto thee;  6. the land of Egypt is before thee; in the best of the land make thy father and thy brethren to dwell; in the land of Goshen let them dwell. And if thou knowest any able men among them, then make them rulers over my cattle.’

20. So Joseph bought all the land of Egypt for Pharaoh; for the Egyptians sold every man his field, because the famine was sore upon them; and the land became Pharaoh’s.  21. And as for the people, he removed them city by city, from one end of the border of Egypt even to the other end thereof.  22. Only the land of the priests bought he not, for the priests had a portion from Pharaoh, and did eat their portion which Pharaoh gave them; wherefore they sold not their land.

Note the “land of the priests” should here have included Thebe, the prime powercenter of the 18th Dynasty. We don’t know the historical context of Joseph, though all we can say is that even biblical sources would be rather in agreement with an essentially Egyptian character of the 18th Dynasty! The genes of possible European and Levantine origin could have arrived into the Egyptian royal lineage by mere coincidence, but one important indication remains the same: the European-like Y-chromosome was around.

Akhenaten leads his family and nation to the only god Aten. Inherited from the Indo-European "bright one" Apollo/Belinus, having the Levantine Baäl/Adon as an intermediary?

Akhenaten, confirmed by the genetic investigation as the father of Tutankhamun, was probably the most remarkable pharaoh of the most remarkable dynasty. He patronized the highly appreciated art of his time and display a considerable zeal to establish the god Aten as the exclusive, monotheistic god of Egypt. In his book “Moses and Monotheism”, Sigmund Freud already proposed Akhenaten for being the pioneer of a monotheistic religion that later became Judaism. The course of events might have been slightly different now Akhenaten appears to be genetically closer to the Jews than he and the people of Moses (“child” in the Egyptian language) to the Egyptians. However, if Akhenaten indeed has an ultimate Hyksos background then his affinity to Canaan as the ultimate source of inspiration to his “monotheistic” ideas would be evident.

Other barriers to a neutral investigation may be found in animosities between the Jewish people and Euro-centrists, what for sure might initiate a new round of false claims and counterclaims concerning the Indo-European issue, especially in relation to Semites. Bottom line, however, is that oriental R1b-ht15 is most likely of European provenance. Even though it might have been around in the Levant for thousands of years, it rather points to European expansions than to that few attested strains of oriental R1b-ht15 being native to the region. If indeed 18th dynasty R1b genes would be confirmed being of the European type, this is bound to support traditional claims that concern a rather Indo-European character of the Hyksos invasions into Egypt. For sure the Hyksos introduced a Semitic element into Egypt, though the ultimate initiators of the most outstanding  Hyksos innovations, like the chariot, were always understood as Indo-European.
However, if so, we could find another barrier in the general dislike of Egyptology against the Hyksos, that have a terrible reputation as destroyers of Egyptian culture. Especially the 18th Dynasty, even though firmly rooted in the 17th Dynasty that was contemporaneous to the Hyksos period, had a reputation to uphold as liberators against Hyksos occupation since their rise to fame was closely connected to this nationalistic event. Tuthmose III, the great-great grandfather of Akhenaten, may have approached these former oppressors initially as far as Megiddo, where – suggestively – he already managed to force the Indo-European Hittites into paying tribute. The great-great-grandfather of Thutmose III was Ahmose I, who at the onset of the 18th Dynasty (he himself was a product of the 17th Dynasty, Thebe) made a career out of the subordination of the Hyksos pharaohs in the north. The (Hyksos) Rhind papyrus document, however, still refers to Ahmose by the inferior title of ‘Prince of the South’ rather than king or pharaoh. The 17th Dynasty and Ahmose may already have pertained to the Hyksos world, even though the chose to pursue a different policy, thus it isn’t at all that unlikely their royal lineage derived from the Hyksos, that themselves may relate to neighboring Anatolian or Hittite stock.
In a 1993 article, Helck pointed out that the Hyksos could be part of a sea invasion of Indo-European peoples from mainly Anatolia, bastion of Indo-European “Anatolian” languages like Hittite. Unfortunately, we don’t know enough of the Hyksos period nor the Hittites to confirm a close historic tie between both people on historic grounds.

The relevance of a possible Indo-European heritage, written in genes rather than papyrus, is that foreign influences of the kind may have extended up to the end of the 18th Dynasty. We already know that the Egyptian goddess Anat arrived in Egypt together with the Hyksos, and she was worshiped long after. Then why shouldn’t the same apply for Aten, the new sun god of Akhenaten? The Northwest Semitic Adon or Baäl (both “Lord”) portrayed next to Anat in the Ugarit Baäl cycle as her brother/lover.

Strange enough, the Ugarit Adon-Anat cycle finds a parallel in the biblical Adam and Eve: Adam and Aden are namesakes. Equally, Eve appears to have inherited some of the less admirable traits of Anat in her relation to men, and at least the Canaanite Anat had something with snakes. More importantly, another parallel can be found in the twin children of Zeus and Titan daughter Leto: Apollo and Artemis. Both being “powerful archers”, the question arises how close their cult could have been connected to the archeological remains attested in the eastern Mediterranean (Heyd), that were closely related to Bell Beaker – whatever the cultural interpretation. The Baäl-Anat cult was definitely unique in the Semitic world, a strong indication of foreign contact of a kind, that coerced influence on the borderland between Semitic and Anatolian cultures and along the sea. But were it the Phoenicians that accomplished the Mediterranean integration of mythology and culture, or did they build further on the work already done by others, like Maritime Beaker traders? Especially, when these people from the west, that must have carried R1b-ht15 gene markers, could be identified as the Indo-European avant-garde that had already arrived in the Levant.

The introduction of the snake-cult and the custom of deformed skulls in Malta coincided with the collapse of older Megalithic civilizations, from 2500 BC on, i.e. (Maritime) Beaker time. Popular science, like here Adriano Forgione, already noted that the megalithic temple of Hal Saflieni, Malta, featuring deformed doliocephal skulls of a culture that identified with the snake, could supply an ideal link to the east. The genetic investigation of Hawass proved the exaggerated elongated head in the portrayal of Akhenaten in sculptures and carvings, and also his female appearance, to be fully cultural (one reason why the mummy of Akhenaten was never fully recognized before).

In an Indo-European context, snakes typically relate to the underworld. In this specific form this mythological element didn’t make it to Egypt, but it can be recognized in the Levant. Here the Anet cult is closely associated to Astarte, as though the different goddesses personified a host of contradictory female aspects, united into one single divine principle. In Indo-European mythology this multiple identifications are more common, especially in the reconstructed mother-cult of the triple-goddess. Note that the Celtic goddess Brigit has also been identified as a triple goddess and she was associated with the snake. Possibly the underpinning theme was general and wide-spread in Indo-European cultures. In Greek mythology the snake represented the chthonic power connected with the Goddess of Earth, the animal was sacred to Demeter, was depicted on the shield of the goddess Athena, and Artemis send Admitos snakes to his wedding bed. A Mediterranean Diana cult might have traveled together with the Beaker culture, to leave behind a trail of related cults of goddesses that may all derive from the same stem, including names like the Egyptian Neith, Ankh, the Phoenician Tanat and Anat and the Greek goddess Athena. Conform the current version of the Dutch view concerning Beaker stratification, this Mediterranean Beaker agent, despite of having an originally northern (Corded Ware/TRB horizon) connotation,  had direct Iberian (thus most likely R1b-ht15) precedences. Since Beaker cultures in the Mediterranean region seem to correlate in particular to Celto-Italic cultures, this would be the particular Indo-European branch to look for in the Hyksos homelands and the Levant. Also in Greece, where this element could have been part of the substratum.

A Beaker connection would make the association of Baäl to an Apollo like god, i.e. the brother of Artemis/Diana, more likely. Note also that Apollo/Baäl has a Celtic counterpart in Belinus. A semitic etymology of Baäl may ultimately be proven false, or derived. An alternative IE etymology would be “the bright one”. This would be a suitable epitaph to Aten indeed, now this Egyptian god may correlate to the Canaanite god Baäl/Adon.

So now, what we can make of the Indo-European identity of this devine pair Baäl-Anat versus Apollo-Artemis? In an Indo European context, the themes related to the Apollo and Artemis cults are pretty universal. Apollo represents the sun, Artemis the moon. The line between Artemis and the Mother God is often hard to draw, but where the divine female powers of Mother Earth are benign to the snake, the male “Apollo” was not. There is a parallel between snake-mastering myths, like snake Pythus being subdued by Apollo, and dragon slaying myths, like Typhon being battled by Zeus. Those myths seem to derive from the same source and can be recognized as a common Indo European theme in myths like Sigurd killing dragon Fafnir (in the Sigurd stones depicted as a snake), Thor fighting Jormungand, Vritra “the enveloper” or world-snake beaten by Indra (India), In Hittite mythology, Illuyanka was a serpentine dragon slain by Tarhunt. As a salient detail, this kind of mythological snakes or dragons are generally imagined as a source of knowledge and foresight that can be accessed once the monster is slayed. Hence the oracle of Delphi and the revelations to Sigurd. The biblical snake, that thus may be a relict of the Hyksos ancestors that reached the Levant already during an early Indo-European expansion, offered his advice for free, having hostile intentions. The message: divine submission is essential for being trustworthy. The pursuit of Aten was to eradicate all other gods, since none can be trusted. Like JHWH and Allah, Aten was such a god that chose to be intolerant to other gods.
Note that the biblical monster Leviathan was actually a whale, and the Egyptian Seth that killed Osiris was rather a god than a monster and depicted as an animal that seems to unite a host of strange desert creatures (aardvark, donkey, jackal) except snakes. Apparently, the snake-dragon distinguishes the Indo-European mythology from others, and still the biblical theme of Adam-Eve-Snake seems to echo the Indo-European Apollo-Artemis-Pytho theme. The Dutch equivalent of the triple goddess Nehalennia even carried a basket of apples, even though her connotation with the underworld was invariably represented by a dog, no snakes.

Ramsund carving, depicting Sigurds victory over the world-snake Fafnir. Once in command he has access to knowledge and truth, like Apollo that subdued the snake Python and initiated the oracle of Delphi.

The duality between the Indo-European male-female divine powers was clearly mirrored in the brother-sister and lover relation between Baäl and Anat in the Levant, the presumed Hyksos homelands, and possibly even in Genesis’ Adam and Eve. The proposed genetic connection between Hyksos and the 18th Dynasty in Egypt strongly insinuates that much of the Levantine  “Hyksos” influence survived until Tutankhamun. Not just the goddess Anat, but also the “monotheistic” god Aten may link the whole Levantine region together, and thus, albeit secondarily, draw the whole Mediterranean into the Indo-European hemisphere. The consanguinity of the divine couple may have inspired the attested brother-sister marriage of Akhnaten and Nefertiti (mummy KV35YL), his parents.

And yes, a European haplotype of Akhenaten could shed light on all of this questions that concern the integration of people and cultures that apparently happened long before and independently from the Phoenicians. The issue is complicated and in need of new perspectives. Are we ready to deal with all available evidence? So far I can only hear silence. Let’s hope Hawass will come up soon with all genetic results that are relevant for history!

Referenced:

• Zahi Hawass et al.- Ancestry and Pathology in King Tutankhamun’s Family, 2010, link
• Othmar Keel,Christoph Uehlinger – Gods, goddesses, and images of God in ancient Israel, 1992, translated 1998, link
• John T. Koch – Celtic culture: a historical encyclopedia, Volumes 1-5, 2006, link
• Helck, W. – Das Hyksos-Problem, Orientalia 62.2 (1993) p. 60 – 66
• Volker Heyd – When the West meets the East: The Eastern Periphery of the Bell Beaker Phenomenon and its Relation with the Aegean Early Bronze Age, 2007

Recommended reading:

• On the identity of Akhenaten: Kate Phizackerley’s blog

Decorative shell ornament used by Neanderthal 50,000 BP.

In their announcement of a recent study by João Zilhão et al., BBC evaluates the discovery of Neanderthal make-up as modern human behaviour that preceded the arrival of modern humans:
Until now it had been thought by many researchers that only modern humans wore make-up for decoration and ritual purposes.

There was a time in the Upper Palaeolithic period when Neanderthals and humans may have co-existed. But Professor Zilhao explained that the findings were dated at 10,000 years before this “contact.”
“To me, it’s the smoking gun that kills the argument once and for all,” he told BBC News.
“The association of these findings with Neanderthals is rock-solid and people have to draw the associations and bury this view of Neanderthals as half-wits.”

In the original paper of January 19, 2010, João Zilhão et al. conclude:
“The Iberian finds show that European Neandertals were no different from coeval Africans in this regard, countering genetic/cognitive explanations for the emergence of symbolism and strengthening demographic/social ones.”

In other words, in the course of human evolution something altered in the human being that can’t be defined by mere IQ, and that still is important to the way we behave differently from animals – and even from early humans. Both Neanderthal and Anotomically Modern Humans (AMS) crossed this evolutionary boundary.

Note the evolutionary changes that led to the use of make-up among early men involve aesthetics here, not the capacity or just a way to solve a problem in order to survive. Aesthetics, that stem from the Self and that imply self-reflection, an important “human” criteria. Foucault wrote about the Self as a source of aesthetics, albeit his focus was rather on quite less primitive stylistic technologies of the self:
“I am referring to what can be called the “arts of existence.” What I mean by the phrase are those intentional and voluntary actions by which men not only set themselves rules of conduct, but also seek to transform themselves in their singular being, and to make their life into an oeuvre that carries certain aesthetic values and meets certain stylistic criteria. These “arts of existence,” these “techniques of the self,” no doubt lost some of their importance and autonomy [...]  Still, I thought that the long history of these aesthetics of existence and these technologies of the self remained to be done, or resumed (UP, 10-11).“

Why African Homo Sapiens would have reached this same evolutionary level about the same time as Neanderthal, and how come the geographic transition to modern humans (that could have been a matter of domestication processes rather than genetic development) is congruent to this change?

In 2008, the Washington University in St. Louis already summarized the implications of a more precise dating of the late Neandertals and Modern Human contact in Southeastern Iberia, where the Protoaurignacian and the Aurignacian I remain unknown:
“The human fossils from the upper levels of the Sima de las Palomas are anatomically clearly Neanderthals, and they are now securely dated to 40,000 years ago. They therefore establish the late persistence of Neanderthals in this southwestern cul-de-sac of Europe. This reinforces the conclusion that the Neanderthals were not merely swept away by advancing modern humans. The behavioral differences between these human groups must have been more subtle than the Middle-to-Upper Paleolithic technological contrasts might imply.”

The intermediate position of the local Neanderthal were described thus in the paper referred to (Walker et al., coauthored by  Zilhao and Trinkhaus):
“At the same time that the Palomas humans exhibit a suite of derived Neandertal features and archaic Homo configurations long since lost among early modern humans, their morphological variation indicates that they deviate from the expected Neandertal ranges of variation. This pattern may be result of genetic drift in relative isolation, directional change or, perhaps more likely, population contact to the north.”

In a new publication, Zilhao arrives at an extended interpretation of the transition. The time-lines are now defined such that Aurignacien (taken indicative to modern humans) started there only at 37,000 BP. The Neanderthal phenotype that exhibited the above mentioned suite of derived Neanderthal and archaic Homo configurations had already disappeared at the time of the anatomically mixed AMH specimen of Lagar Velho:
“With the last of the region’s Neandertals dating to five millennia before the child was borne, crossbreeding between immediate ancestors (e.g., parents or grandparents) drawn from distinct ‘‘modern’’ and ‘‘Neandertal’’ gene pools is empirically untenable. Therefore, those features must represent evolutionarily significant admixture at the time of contact.”

The Lagar Velho child thus attests Neanderthal admixtures that are considered reminiscent of much earlier contact, since by then the Neanderthal type had already disappeared.

Venus of Hohle Fels, made by the first AMS or the last Neanderthal?

However, the most recent publications of Zilhao don’t answer our curiosity that concern the human phenotype up north across the Pyrenees, that was responsible for the early art at the basal Aurignacian of Hohle Fels Cave in southwestern Germany. How did they look like?
A carving found in six fragments depicts a woman with a swollen belly, wide-set thighs and large, protruding breasts, that carbon dating suggested to be at least 35,000 years ago.
“It’s the oldest known piece of figurative sculpture in the world,” said Jill Cook, a curator of Paleolithic and Mesolithic material at the British Museum in London in the news. By the way, most likely the contemporaneous women looked quite different.

First flute, Hohle Fels. Made by the first AMS or the last Neanderthal?

Another find of the same artistic magnitude, a flute, was found here as well. Conard, 2009:
“Although arguments have been made for Neanderthal musical traditions and the presence of musical instruments in Middle Palaeolithic assemblages, concrete evidence to support these claims is lacking. Here we report the discovery of bone and ivory flutes from the early Aurignacian period of southwestern Germany. These finds demonstrate the presence of a well-established musical tradition at the time when modern humans colonized Europe, more than 35,000 calendar years ago.”

It is assumed that these finds are to be associated to anatomically modern humans. Conard:
“It’s 100 percent certain that, by the time we get to 40,000 years ago in Swabia, we’re dealing with people just like you and me.”
However, this seems to be at odds to the investigation results of Finlayson and Carrion (2007):
“Overall, it is clear that diagnostic and well-dated AMH fossils are not found west of the Iron Gates of the Danube before 32 kya, implying a relatively recent entry into western Europe.”

The first dog was found in Goyet Cave, Belgium (31,700 BP). Dog domestication appear to be older than attested AMH in the region.

This factual evidence of a late arrival date of anatomically modern humans would also leave the domestication of the dog rather in the hands of (pre-AMH) Neanderthal offspring. Remains of the first dog were found in the Goyet Cave in Belgium and was dated 31,700 BP.

The replacement of Neanderthal in western Europe is still an enigma and strictly spoken the first known figurative sculpture and musical instrument might as well have been manufactured by Neanderthal, or people that featured a similar suite of derived Neanderthal and archaic Homo configurations as their contemporary Neanderthal neighbours found in Palomas. Like the people that domesticated the first dog in Belgium and the chiuld of Lagar Velho, they could also be modern humans that featured evolutionary significant Neanderthal admixtures.

Or does all of this indicate continuity of a magnitude that hitherto remained unrecognized?

Let’s take a look again at the Neanderthal Genome project. Green:
“We tested more than 70 Neanderthal bone and tooth samples from different sites in Europe and western Asia for bio-molecular preservation by removing samples of a few milligrams for amino acid analysis. The vast majority of these samples had low overall contents of amino acids and/or high levels of amino acid racemization, a stereoisomeric structural change that affects amino acids in fossils, indicating that they are unlikely to contain retrievable endogenous DNA. However, some of the samples are better preserved in that they contain high levels of amino acids (more than 20,000 p.p.m.), low levels of racemization of amino acids such as aspartate that racemize rapidly, as well as amino acid compositions that suggest that the majority of the preserved protein stems from collagen.“

Next, Green investigated if the sampled mtDNA was Neanderthal-like or not, taking mtDNA that resembles modern mtDNA as contaminated “Assuming that the ratio of Neanderthal to contaminating modern human DNA is the same for mtDNA as it is for nuclear DNA“.

“Whereas only around 1% of the mtDNA present in three samples from France, Russia and Uzbekistan was Neanderthal-like, one sample from Croatia and one from Spain contained around 5% and 75% Neanderthal-like mtDNA, respectively. One bone (Vi-80) from Vindija Cave, Croatia, stood out in that 99% of the 63-base-pair mtDNA segments and 94% of the 119-base pair segments are of Neanderthal origin.“

Thus, having samples rejected when apparent Neanderthal mtDNA resembles modern human mtDNA, then of course the result will be that “all Neanderthals sequenced to date differ from all contemporary humans“. If mtDNA is accepted when different for 99%, then why amino acids having low levels of racemization shouldn’t be investigated too since this is what Green did. However, Wall and Kim suggested a new contamination criteria departing from the positive correlation of nuclear DNA, and took degraded DNA as indicative to the differences between Neanderthal and modern humans. The circular interpretation of the results should be obvious, and may even extend to the current measurements of Neanderthal mtDNA.

In my opinion the apparent lack of evidence for gross admixture in human DNA either indicates the extinction of the Neanderthal offspring, or otherwise – intriguingly -the full scale integration of Neanderthal DNA into the modern human gene-pool.

Assuming the extinction of Neanderthal offspring is not so hard to do, even though this would complicate the previous extinction scenarios that concerned a whole would-be species rather than a hybrid lineage. However thus, talking about circular thought, the assumption of extinction seems to derive from meeting the preposition rather than independent investigation. To say it can’t so it isn’t, rather than it could but it isn’t, invites to the kind of circular thought we should try to avoid.

Moreover, “extinction” is a way to deal with Neanderthal traits of archaic AMH that does not even occur to the writers of a new article about the child of Lagar Velho, Priscilla Bayle et al. (2010). This article reminds us about the evolutionary changes of modern humans since 40 kya, that involve characteristics that can also be found among Neanderthal.

Therefore, a simple Neandertal versus modern human dichotomy is inadequate to accommodate the morphostructural and developmental variation represented by Middle Paleolithic and earlier Upper Paleolithic populations. These data reinforce the complex nature of Neandertal-modern human similarities and differences, and document ongoing human evolution after the global establishment of modern human morphology.

It will be hard to sustain that Neanderthal were not involved as ancestors of the modern humans only because Neanderthal traits were simply lost in the course of evolution. Then, considering full scale integration instead as the only other possible option, how could it be? How could we even imagine the integration of Neanderthal DNA within the modern human species to this extend, in such a way that not any sign of significant hybridization can be discerned within the modern gene-pool?

Genes that for sure crossed the phenotype boundaries have already been identified, and a mixture between archaic Asiatic types (divergence time 2 million years) and African types for certain genes is already generally accepted. There is still discussion on how this mixture stabilized (eg. Diffusion wave hypothesis). An overview:

- Some genes have very deep, non-African branches: the RRM2P4 pseudogene has a MRCA of ~2 MYA in East Asia (Garrigan et al., 2005); PDHA1 locus is another ~2 mya example supporting multiregionality
- Microcephalin entered the modern human genepool at 37 kya, since older type has MRCA at 1.7 mya
- Disequilibrium at locus Xp21.1 (MRCA 1.9 myr old) points to admixture
- Gutiérrez, Sánchez & Marín (2002) show ancient mtDNA is very sensitive to phylogenetic methods, diagenetic modifications have altered the sequences, and conclude that Neanderthal and modern mtDNA may overlap.
- FOXP2 in Neanderthal is a potential argument for being of the same species as modern humans, to the point of being identical.

Moreover, by examining the mutations in DNA near Alu insertions in two completely sequenced modern human genomes, Huff et al.(2010) could calculate a significant bottleneck at 1.2 million years ago. This is in agreement to the assumed split time of African Homo Sapiens and Neanderthal. The average age of human DNA is about one million years, and some genes are even estimated at two million years. From this perspective Neanderthal DNA should indeed be consistent to modern DNA.

Neanderthal (left) and archaic AMH of Predmosti (right) feature the same occipital bun at the back of the skull.

The solution to this problem may be all but genetic. The flexibility of human phenotypes may rather suggest a good deal of non-genetic differences between Neanderthal and modern humans. Still other differences correlate with current typological differences, such as the “occipital bun” that has also been documented among Europeans.

As for now, there is no knowledge of genes that unequivocally account for robust “Neanderthal” morphological features, such as those that involve a lower and larger cranium, a larger browridge, a larger nose, larger shoulder joints, a larger and broader rib cage, larger elbow joints, broader hips, shorter forearms, larger hip joints, larger and thicker patellae, shorter and more flattened tibiae and larger ankle joints. All these features may stem fully or partially from epigenetic, environmental interaction.

There is no reason to assume the disappearance of once outspoken Neanderthal features would to be due to evolutionary developments that are essentially different from domestication-like processes. Anatomically Modern Humans only look different.
The Neanderthals are us.

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