First of all, for the lack of a better summary of what these dwarves were all about, here are a couple of quotes from Wikipedia. I checked the original sources and they look legit, so this ought to be accurate:
Dvergar or Norse dwarves (Old Norse dvergar, sing. dvergr) are entities in Norse mythology associated with rocks, the earth, deathliness, luck, technology, craft, metal work, wisdom, and greed. They are sometimes identified with Svartálfar ('black elves'), and Dökkálfar ('dark elves'),[1] due to their apparently interchangeable use in early texts such as the Eddas.
While the word "Dvergar" is related etymologically to "dwarves", the early Norse concept of Dvergar is unlike the concept of "dwarves" in other cultures. For instance, Norse dwarves may originally have been envisaged as being of human size.
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The Dvergar are often called 'black', especially as the 'black elves' (svartálfar). In Old Norse, this byname 'black' (svartr) refers to hair color or eye color.
The illustration above is of Reginn the Dvergr, again courtesy of Wikipedia. Now here's an abstract from a recent open access paper on maritime contacts between the East Mediterranean and Scandinavia during the Bronze Age. Note the references to rocks, technology, craft, metal work and trade (and thus greed, I suppose).
The Bronze Age of Scandinavia (1750-500 BC) is characterized by the sudden appearance of bronze objects in Scandinavia, the sudden mass appearance of amber in Mycenaean graves, and the beginning of bedrock carvings of huge ships. We take this to indicate that people from the east Mediterranean arrived to Sweden on big ships over the Atlantic, carrying bronze objects from the south, which they traded for amber occurring in SE Sweden in the Ravlunda-Vitemölla–Kivik area. Those visitors left strong cultural imprints as recorded by pictures and objects found in SE Sweden. This seems to indicate that the visits had grown to the establishment of a trading centre. The Bronze Age of Österlen (the SE part of Sweden) is also characterized by a strong Sun cult recorded by stone monuments built to record the annual motions of the Sun, and rock carvings that exhibit strict alignments to the annual motions of the Sun. Ales Stones, dated at about 800 BC, is a remarkable monument in the form of a 67 m long stone-ship. It records the four main solar turning points of the year, the 12 months of the year, each month covering 30 days, except for month 7 which had 35 days (making a full year of 365 days), and the time of the day at 16 points representing 1.5 hour. Ales Stones are built after the same basic geometry as Stonehenge in England.
Interesting stuff. The only thing I'd add is that these contacts between the Mediterranean and Scandinavia most likely stretched back to the Neolithic, when Megalithic cultures dominated Southern and Western Europe. Indeed, the remains from a TRB (Funnelbeaker) Culture burial in western Sweden were recently genotype for autosmal DNA and they came out surprisingly Mediterranean (see here).
Nils-Axel Mörner, Bob G. Lind, The Bronze Age in SE Sweden Evidence of Long-Distance Travel and Advanced Sun Cult, Journal of Geography and Geology, Vol 5, No 1 (2013), DOI: 10.5539/jgg.v5n1p78
A study on the ancient DNA of Minoans has shown that these enigmatic Bronze Age inhabitants of Crete were very similar in terms of mtDNA to present inhabitants of Crete, especially those of the Lasithi Plateau where the Minoan sample came from. But here's the really interesting part: as per the gradient maps below, the Minoan mtDNA sequences also show high affinity to those of modern English (a) and Bronze Age Sardinians and Iberians (b). See also Table 1 from the paper, where the top ten "nearest neighbors" to the Minoan sample are ancient and extant Western European populations.
So the results imply genetic links between Bronze Age Crete and Western Europe. Now, Martinez et al. 2007 found that 36.6% of Cretans from the Lasithi Plateau belonged to Y-chromosome haplogroup R1b. They only tested 41 individuals, but that's still an interesting result for Southeastern Europe, where R1b is generally uncommon. Indeed, perhaps the Minoans carried a much higher frequency of R1b, and they (or a related seafaring culture) spread this marker to Western Europe via maritime routes, where it has since become the most important Y-chromosome haplogroup? It's a valid question considering the ancient mtDNA data. The pics of Minoan bull leaping and Spanish bullfighting below are courtesy of Wikipedia (see here).
Update 16/05/2013: To add to my comments above about the Minoans, or a related group, being potentially responsible for the introduction of Y-DNA R1b to Western Europe, it's interesting to note that one of the Minoan mtDNA sequences belonged to the rare H13a1a haplogroup.
Both H13a1a and R1b were recently found in late Neolithic Bell Beaker remains from Germany (see here and here). Moreover, today H13a1a shows a peak in frequency and diversity in the Caucasus, particularly in Dagestan, but also occurs at low frequencies in Italy, Sardinia and Iberia. Interestingly, R1b is found at fairly high frequencies among some ethnic groups in and around Dagestan, like the Lezgins, and obviously also common in Italy and Iberia.
So what am I getting at? Well, it looks like a group with loads of R1b from what is now Dagestan or surrounds - perhaps the deep ancestors of Bell Beakers and Minoans - learned to sail, crossed the Mediterranean Sea from east to west, settled a few islands along the way, and eventually their descendants conquered much of Western and Central Europe. This is certainly not the most parsimonious theory of how R1b might have appeared on the scene in Western Europe during the late Neolithic, but it does make sense considering all the data.
But what might have caused this purported population movement from the Caucasus, and is it a coincidence that both R1a and R1b only appear in European ancient DNA data from the late Neolithic onwards? It's unlikely that the Minoans and Bell Beakers were part of the Indo-European expansion, but perhaps their ancestors in the Caucasus felt the pressure of this expansion from the steppe in the north, which was at that time most likely dominated by Kurgan groups high in R1a?
Update 18/05/2013: Maju isn't convinced that the gradient maps and "nearest neighbor" analysis show explicit links between the Minoan and post-Neolithic Western European mtDNA gene pools. He calls it a "pseudo-affinity" which should be taken with a pinch of salt (see here). Moreover, he suggests the Minoan mtDNA shows closest links to early European Neolithic mtDNA because of four HVS-1 sequence matches.
But the high affinity between the Minoan and post-Neolithic Western European mtDNA can be seen clearly in two different analyses, so it's real, even if mostly indirect. Therefore, there's no need to take the results with a pinch of salt, they should just be viewed in their proper context. In other words, this affinity is certainly not due to a massive invasion of Western Europe by Minoan women, but the result of the same processes acting on the post-Neolithic Western European and Minoan mtDNA gene pools, which probably included some direct gene flow from the Eastern Mediterranean to Europe during the Bronze Age.
References...
Hughey et al., A European population in Minoan Bronze Age Crete, Nature Communications 4, Article number: 1861, doi:10.1038/ncomms2871, Published 14 May 2013
Martinez et al., Paleolithic Y-haplogroup heritage predominates in a Cretan highland plateau, European Journal of Human Genetics (2007) 15, 485–493. doi:10.1038/sj.ejhg.5201769; published online 31 January 2007
After a recent update the 1000 Genomes sample now includes 62 individuals of South Asian origin belonging to Y-DNA haplogroup R1a-M17. Their full Y-chromosome sequences have been analyzed by Semargl and Maximus (aka. YFull project), with some interesting but not unexpected results:
- All individuals belong to R1a-Z93, which appears to totally dominate South Asian R1a-M17.
- A single Punjabi from Lahore, northeastern Pakistan, is ancestral for the Z94 mutation, which is just below Z93. All the other individuals are derived for Z94.
- Six individuals - of Punjabi, Bangladeshi and Gujarati origin - are ancestral for L657 and Z2124, the two main mutations immediately below Z94.
- All individuals of South Indian and Sri Lankan origin are derived for L657 or Z2124.
- Based on this sample, there appears to be no substructure along ethnic or geographic lines within South Asian R1a-M17 derived for L657 and Z2124.
Thus, it seems the SNP diversity of South Asian R1a-M17 is low, and decreases from Pakistan, North India and Bangladesh to South India and Sri Lanka. In comparison, there are only 12 European R1a individuals in the 1000 Genomes sample, and they represent all the major subclades of this haplogroup: R1a-Z283, R1a-Z93 and R1a-L664. Therefore, sampling bias can't be used as an argument for the more diverse result from Europe.
The lack of substructure along ethnic and geographic lines within South Asian R1a-L657 and R1a-Z2124 looks unusual, especially considering the caste system in India, and needs to be verified with more extensive sampling. However, if this outcome holds up, it'll suggest that paternal gene flow across South Asia has not been restricted by the caste system or geography. Then again, it could mean the caste system appeared after R1a-L657 and R1a-Z2124 arrived in South India via massive population movements from the north.
Below are all the results in as much detail as the current R1a SNP tree allows. Key: BEB - Bengali from Bangladesh; GIH - Gujaratai from Houston, Texas; ITU - Indian Telugu from the UK; PJL - Punjabi from Lahore, Pakistan; STU - Sri Lankan Tamil from the UK.
Z93+ Z94-
PJL - 1
Z94+ L657- Z2124- Z96-
BEB - 2
PJL - 3
GIH - 1
L657+,Y2+ etc.
1) Y9 (inc. Y7)
GIH - 7
STU - 4
ITU - 4
PJL - 8
BEB - 2
2) Y4+, Y8+, Y28+ (inc. Y6+)
GIH - 6
ITU - 6
PJL - 2
STU - 6
BEB - 5
Z2125+ (Z2124+ Z2122- Z2123-)
PJL - 1
Z2123+ (Z2124+ Z2122-, Z2125-)
PJL - 2
STU - 3
BEB - 1
ITU - 6
GIH - 2
One of the striking patterns we see is the relatively high level of sharing of IBD between pairs of individuals across eastern Europe, as high or higher than that observed within other, much smaller populations. This is consistent with these individuals having a comparatively large proportion of ancestry drawn from a relatively small population that expanded over a large geographic area.
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The eastern populations with high rates of IBD are highly coincident with the modern distribution of Slavic languages, so it is natural to speculate that much of the higher rates were due to this expansion. The inclusion of (non-Slavic speaking) Hungary and Romania in the group of eastern populations sharing high IBD could indicate the effect of other groups (e.g., the Huns) on ancestry in these regions, or because some of the same group of people who elsewhere are known as Slavs adopted different local cultures in those regions. Greece and Albania are also part of this putative signal of expansion, which could be because the Slavs settled in part of these areas (with unknown demographic effect), or because of subsequent population exchange. However, additional work and methods would be needed to verify this hypothesis.
Ralph P, Coop G (2013) The Geography of Recent Genetic Ancestry across Europe. PLoS Biol 11(5): e1001555. doi:10.1371/journal.pbio.1001555
Peter Ralph and Graham Coop, Exploring recent relatedness - IBD and biparental ancestry, Department of Evolution and Ecology
UC Davis, June 11th, 2012, CIRM Probability, Population Genetics and Evolution
Our genetic distance data also indicate minimal local genetic continuity between the Early Neolithic (ENE) and the Middle/Late Neolithic (MNE/LNE) in Central Europe (Fig. 1; Table 1), again suggesting that ENE lineages were largely superseded during the MNE/LNE (B4100–2200 BC) in a previously unrecognised major genetic transition. This pronounced genetic changeover between ENE and MNE/LNE cultures is comparable to other known major genetic transition, thus far revealed by ancient DNA and coalescent simulations (between indigenous European hunter-gatherers and incoming early farmers from the Near East during the initial Meso-Neolithic transition fromB7500 BC in Central Europe)6,7. When compared with hg H diversity in present-day Central Europe14,15,18,36, the network in Fig. 1 suggests that much of the present-day diversity can be attributed to the incorporation of new lineages in the MNE/LNE and emerging Bronze Age (from 2200 BC). The LNE in particular is known to have been a period of profound cultural and economic change37, with newly emerging pan-European cultures such as the Bell Beaker phenomenon in Western Europe and the Corded Ware culture in north-eastern Europe. It therefore seems likely that these pan-European cultures were associated with the introduction of lineages from outside Central Europe.
Brotherton et al., Neolithic mitochondrial haplogroup H genomes and the genetic origins of Europeans, Nature Communications 4, Article number: 1764, Published 23 April 2013, doi:10.1038/ncomms2656
The consensus haplotype of the three [Corded Ware] individuals (based on most complete profile) gave two exact matches in in an European population sample of 11,213 haplotypes in a set of 100 populations (as of July 2008, Release ‘‘23’’ from 2008–01-15 14:44:25): one individual from Poland (1/939 from Gdansk) and one from Russia (1/48 from Tambov).
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The Y haplotype was predicted using the Web-based program Haplotype Predictor (9). The three individuals of grave 99 belong to
haplotype R1a, with a probability of 100% based on the Y-STR profile of individual 3 (10). To confirm haplogroup status, we
further amplified an 85-bp fragment covering the Y-SNP marker SRY10831.2 characteristic for R1a (11). Primer sequences are
given in Table S6. Sequences and sequenced clones from independent extract of all three individuals show the specific G A transition identifying R1a (Fig. S5).
Wolfgang Haak et al, Ancient DNA, Strontium isotopes, and osteological analyses shed light on social and kinship organization of the Later Stone Age, PNAS, Published online before print November 17, 2008, doi:10.1073/pnas.0807592105
While the U5 mtDNA lineage was virtually absent amongst the Early Neolithic, LBK and Rossen cultures (recorded at a frequency of only 1.7%), it was observed in the Late Neolithic/Early Bronze Age cultures, Corded Ware and Unetice, at a frequency of 18% and 29%, respectively (Table 2). The re-emergence of subhaplogroup U5 during the Late Neolithic/Early Bronze Age in Central Europe, which is particularly associated with the Corded Ware and Unetice cultures, may reflect the Mesolithic origins of these cultures. However, determining where the Mesolithic ancestors of these cultures arose is difficult. The Corded Ware and Unetice cultures may be the descendants of Mesolithic hunter-gatherers from Eastern Europe and are hypothesised to be descendents of the Kurgan culture which was present in the Eastern Steppe. The Corded Ware and Unetice cultures display similarities to the Kurgan culture in pottery styles (eg. impressed ceramics) (Gimbutas 1963) and burial types (i.e. pit graves) (Bogucki, Grygiel 1993; Gallagher, Gunther, Bruchhaus 2009).
Christina Jane Adler, Ancient DNA studies of human evolution, Thesis, 2012, The University of Adelaide, South Australia, School of Earth and Environmental Sciences
The human samples studied here show evidence for consumption of millet, a uniquely Slavic cultigen in Europe that may be useful in studying Slavic migrations. My stable isotope data track millet consumption in Poland back to the Neolithic period (approximately 2,000 BC).
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Millet was cultivated in Poland since the Neolithic, but increased significantly in the Roman Era (Wasylikowka et al. 1991: 227). Millet is common among Slavic populations and has been documented in Southern and Eastern Europe by numerous paleobotanical (Polcyn 2002; Pyrgała 1970; Rösch et al. 1992; Wasylikowa et al. 1991; Zohary and Hopf 1988) and stable isotope investigations (Le Huray and Schutkowski 2005; Murray and Schoeninger 1988; Reitsema et al. 2010). Stable isotope studies in Western Europe (e.g.: Netherlands and Britain) indicate an absence of millet in human diet during and preceding the medieval period (Randsborg 1985; Richards et al. 2006; Schutkowski et al. 1999). This East-West dichotomy suggests Slavs had a cultural preference for millet (Dembińska 1999).
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The first Neolithic skeleton (Neo-1) dates to the Globular Amphora culture (approximately 4,100 years ago) and is from Kowal in the Kujawia region of Central Poland. The other Neolithic skeleton (Neo-2) is of unknown age but likely represents the Corded Ware Pottery culture which occupied much of Northern Europe between approximately 5200 and 4300 years ago (Mallory 1997). It comes from Niedrzwica in Northeast Poland and is thought to represent a more pastoral population, as was typical in Northeast Poland.
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Both Neolithic samples plot far to the left of the C3 protein line, indicating a diet entirely based on C3 protein sources (Fig. 6.6). Δ13C(ap-coll) values of the Neolithic samples are high (9.0‰ and 7.7‰), which along with δ15N values suggests either a diet low in meat or fish and high in plant materials, or including millet as an energy source (Fig. 6.7). A high δ13Cap value from the Kowal individual reinforces the possibility of millet consumption (Fig. 6.6).
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The Roman-era cemetery at Rogowo was excavated in 1999 and 2000. The cemetery includes 137 inhumation and 151 cremation burials (Fig. 4.2). Biritual cemeteries such as this are characteristic of the Wielbark culture, which occupied Poland during the 2nd and 3rd centuries AD. It is possible that Wielbark populations were one of several referred to by the Romans as “Goths” (Heather 1996). It is also possible that the Wielbark are early Slavs from which modern Slavs are descended (for discussion, see Piontek 2006).
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The δ13C values from Rogowo indicate millet contributed between 0% and 35% to overall diet, depending on the individual. The fact that δ13C and δ15N values are not positively correlated also supports an interpretation of direct millet consumption, rather than marine fish, anadromous fish or C4-foddered animal consumption.
Laurie Jean Reitsema, Stable Carbon and Nitrogen Isotope Analysis of Human Diet Change in Prehistoric and Historic Poland, Dissertation, The Ohio State University, 2012.
A Tocharian B feminine gender noun form of the Plural proks-a is attested in MQ 159,3 in comnination with the two attributes derived from names of cereals identified as "milet" (yp-iyana from yap-'millet', yp-iye 'peprtaining to millet', Adams 1999: 518) and "wheat". Klaus T. Schmidt (1999: 3-4), who has recognized the meaning of the word proks-a as 'grain', has also noticed its obvious etymological link to Slavic *proso 'millet'.
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According to Schmidt two ways of semantic reconstruction of the periphery of the term are possible: either the Slavic word (as well as probably, one might add, its possible Old Prussian correspondence if it was not borrowed from Slavic) has retained the original meaning of a special kind of cereal later on generalized in Tocharian or on the opposite the general meaning attested in Tocharian is an ancient one and the development of a special use in Slavic is a product of a following evolution.
Werner Winter, Brigitte L. M. Bauer, Georges Pinault, Language in time and space: a Festschrift for Werner Winter on the occasion of his 80th birthday, Walter de Gruyter, 2003
Our autosomal, Y-chromosomal and mitochondrial DNA analyses reveal that whereas few specimens seem to be related matrilineally or patrilineally, nearly all subjects belong to haplogroup R1a1-M17 which is thought to mark the eastward migration of the early Indo-Europeans. Our results also confirm that at the Bronze and Iron Ages, south Siberia was a region of overwhelmingly predominant European settlement, suggesting an eastward migration of Kurgan people across the Russo-Kazakh steppe. Finally, our data indicate that at the Bronze and Iron Age timeframe, south Siberians were blue (or green)-eyed, fair-skinned and light-haired people and that they might have played a role in the early development of the Tarim Basin civilization.
Christine Keyser et al., Ancient DNA provides new insights into the history of south Siberian Kurgan people, Human Genetics, Saturday, May 16, 2009, doi: 10.1007/s00439-009-0683-0
Regarding the Y chromosomal DNA analyses, the seven males identified all belonged to haplogroup R1a1a. It is most frequently found in Eastern Europe, South Asia and Siberia. In contrast, it is relatively uncommon in Middle Easterners and rare in East Asian [22-24]. It is thought to be a trace of the migration events of early Indo-European [38-39]. The presence of haplogroup R1a1a in the ancient Xiaohe people implies that the parental ancestry of the Xiaohe people originated from somewhere in Siberia or Europe, which is consistent with the origin of maternal ancestry.
C Li, Evidence that a West-East admixed population lived in the Tarim Basin as early as the early Bronze Age, BMC Biology 2010, 8:15, doi:10.1186/1741-7007-8-15
I discussed this topic in an update to my last post, but thought it deserved a more detailed entry of its own. First of all, here's a map of two Neolithic ecological zones (along with some temperature and precipitation stats) and a quote from Banks et al. 2013.
In order to investigate whether the differences between the adaptations and geographic distributions of three major Early Neolithic archaeological cultures are related to the exploitation of different suites of environmental conditions, we apply genetic algorithm and maximum entropy ecological niche modeling techniques to reconstruct and compare the ecological niches within which three principal Neolithic cultures (Impressed Ware, Cardial Ware, and Linearbandkeramik) spread across Europe between ca. 8000 and 7000 cal yr BP. Results show that these cultures occupied mutually exclusive suites of environmental conditions and, thus, were adapted to distinct and essentially non-overlapping ecological niches. We argue that the historical processes behind the Neolithization of Europe were influenced by environmental factors predisposing occupation of regions most suited to specific cultural adaptations.
Map Key: light brown = Impressed and Cardial Ware Mediterranean Neolithic ecological zone, green = Linearbandkeramik (LBK) ecological zone. Another version of the map, marked with archeological sites belonging to the aforementioned Neolithic cultures, can be seen here.
Anyone who has a keen interest in European genetic structure will know by now that modern Europeans carry three main genetic components when analyzed with the popular ADMIXTURE software at optimal K. These vary somewhat in character depending on how each analysis is set up, but they can generally be referred to as North European, Mediterranean and West Asian. The gradient maps below, based on data from an old Eurogenes ADMIXTURE analysis, show their distribution across West Eurasia and North Africa.
The correlation between the Impressed Ware/Cardial Ware ecological zone and the geographic range of the Mediterranean component in Europe is obvious. There's also a close correlation between the North European component and the grey area in Eastern Europe not assigned to any Neolithic ecological zone. On the other hand, the green LBK zone shows weak correlation with the spread of the West Asian component. However, there's a remarkable correlation between the West Asian component and the geographic affinity of ancient mtDNA from LBK burial sites in Western and Central Europe, as per the maps from Haak et al. 2010 below. This makes sense because LBK farmers are thought to have migrated to Central Europe directly from the Near East.
Now, the LBK culture flourished for several thousand years, but collapsed around 4,000 BC. Latest studies show that climate fluctuations, overpopulation and resulting conflicts were responsible for this outcome. The vacuum was initially filled by a variety of Middle Neolithic cultures, but the replacement process culminated during the late Neolithic/early Bronze Age with major invasions of Central Europe by groups from the eastern and western peripheries of Neolithic Europe. Interestingly, the cultures spreading from the east, such as the Corded Ware, Unetice, Single Grave and Battle Axe, brought with them new social organization and customs. On the other hand, from the west came the Bell Beaker culture, which has been described by archeologists as a copy of the Corded Ware Culture, but also a reaction to it. See, for instance, Jan Turek 2012.
So while the Maritime Beakers are clearly a south-western element (Case 2004, 14, with further references), the symbolic system of the burial rites is based on the eastern Corded Ware and even earlier Yamnaya tradition (Harrison and Heyd 2007, 193-199, fig. 45).
As already mentioned, ancient DNA suggests that LBK farmers were genetically West Asian-like, and it's improbable that all traces of them were completely wiped out from Europe since the Neolithic. Indeed, the West Asian ADMIXTURE component is often found deep in Europe at non-trivial frequencies, and this is probably in large part due to lingering genetic influence from LBK farmers.
The other thing we're now learning from ancient DNA is that, compared to the LBK farmers, Corded Ware and Unetice groups were closely related to North and East European Mesolithic hunter-gatherers (see here). What all of this suggests is that during the early Neolithic much of Europe basically became a colony of West Asia, but the process was reversed to some degree thanks to the expansion of late Neolithic/early Bronze Age groups from Eastern Europe harboring significant pre-Neolithic ancestry. I have little doubt that this expansion - in effect, a revival of hunter-gatherer ancestry in Central Europe - is in large part responsible for the modern North European ADMIXTURE component.
Of course, interpreting the results of ADMIXTURE analyses should always be done with extreme caution and a lot of skepticism. However, I think in this case the proof is in the pudding, and the correlations I pointed out above are difficult to dismiss as coincidental.
Indeed, those familiar with ADMIXTURE will know that the components it produces are usually the result of relative isolation and/or rapid population expansions. Well, we now know that early Neolithic European cultures occupied "mutually exclusive ecological niches", which also means they probably didn't readily mix with each other for thousands of years. Moreover, the late Neolithic/early Bronze Age expansions of pan-European cultures like the Corded Ware and Bell Beaker appear to have been massive, with profound effects on the genetic structure of Europe at the time. As far as I can see, there's no reason why these events wouldn't register in our genomes today.
Update 05/05/2013: I was about to make a GEDmatch ancestry test based on the three Neolithic components described in my post, but then I realized that such a test was already available there, and it's called the Eurogenes K9. All you have to do is re-name several of the components in the following way...
North European > Corded Ware, Single Grave, Battle Axe, Unetice and/or Bohemian Bell Beaker
Mediterranean > Impressed Ware, Cardial Ware, Maritime Bell Beaker and/or Funnelbeaker (TRB)
Caucasus & Southwest Asian > Linearbandkeramik (LBK)
Ancient DNA will tell us one day whether these assumptions are correct. However, please note that the new component names won't be relevant for everyone. For instance, if you're from East Africa or Turkey, then the Caucasus-like admixture you score won't be due to LBK farmers, because they were based in Western and Central Europe.
References...
Banks et al., Ecological constraints on the first prehistoric farmers in Europe, Journal of Archaeological Science, Volume 40, Issue 6, June 2013, http://dx.doi.org/10.1016/j.jas.2013.02.013
Haak W, Balanovsky O, Sanchez JJ, Koshel S, Zaporozhchenko V, et al. (2010) Ancient DNA from European Early Neolithic farmers reveals their Near Eastern affinities. PLoS Biol 8(11): e1000536. doi:10.1371/journal.pbio.1000536
Christina Jane Adler, Ancient DNA studies of human evolution, Thesis, 2012, The University of Adelaide, South Australia, School of Earth and Environmental Sciences
Brotherton et al., Neolithic mitochondrial haplogroup H genomes and the genetic origins of Europeans, Nature Communications 4, Article number: 1764, Published 23 April 2013, doi:10.1038/ncomms2656
Fokkens, H. & F. Nicolis (eds) 2012: Background to Beakers. Inquiries into regional cultural backgrounds of the Bell Beaker complex. Chapter 8, Origin of the Bell Beaker Phenomenon: The Moroccan Connection. Leiden: Sidestone Press
Jocelyne Desideri & Marie Besse, Swiss Bell Beaker population dynamics: eastern or southern influences?, Archaeol Anthropol Sci (2010) 2:157–173, DOI: 10.1007/s12520-010-0037-9
This Banks et al. paper argues that the two main early Neolithic groups in Europe - Mediterranean and Linearbandkeramik (LBK) - were adapted to very specific and mutually-exclusive ecological zones. This would certainly help to explain why the LBK disappeared from Central Europe just before the Middle Neolithic, because this was a time of severe climate fluctuations in the region. Eventually it was replaced by late Neolithic/early Bronze Age cultures from the peripheries of Neolithic Europe, like the Bell Beaker, Corded Ware and Unetice.
The genetically Western European-like Bell Beaker folk expanded from Iberia, which suggests they might have sprung, at least in part, from remnants of the Mediterranean Cardial Ware culture. It's still not clear where the genetically much more Northeast European and Mesolithic-like Corded Ware and Unetice groups came from exactly, but looking at the maps below, particularly the last one (F), it's very likely it was from somewhere within the vast grey area east of the green LBK zone.
In order to investigate whether the differences between the adaptations and geographic distributions of three major Early Neolithic archaeological cultures are related to the exploitation of different suites of environmental conditions, we apply genetic algorithm and maximum entropy ecological niche modeling techniques to reconstruct and compare the ecological niches within which three principal Neolithic cultures (Impressed Ware, Cardial Ware, and Linearbandkeramik) spread across Europe between ca. 8000 and 7000 cal yr BP. Results show that these cultures occupied mutually exclusive suites of environmental conditions and, thus, were adapted to distinct and essentially non-overlapping ecological niches. We argue that the historical processes behind the Neolithization of Europe were influenced by environmental factors predisposing occupation of regions most suited to specific cultural adaptations.
For background info on the fatal effects of climate fluctuations on LBK communities just prior to the Middle Neolithic refer to Gronenborn et al. 2013.
According to the applied age model, the simple farming societies of the LBK (5600–4900 cal BC) in west-central Europe were not immediately and devastatingly affected by most climate fluctuations. Yet, they might have been one destabilising component within broader processes. However, periods of decreased or irregularly spaced rainfall are contemporaneous to periods of population decline, while periods of increased rainfall may have favoured population growth. Towards the end of the 6th millennium cal BC, the final years of LBK in western Central Europe are contemporaneous to a general trend to less rainfall punctuated by short-term increases in precipitation. During this climatically highly volatile period LBK reaches its highest population rates and at the same time experiences a period of warfare. Thereafter population rates decline and LBK gradually vanishes from the archaeological record, being replaced by Middle Neolithic societies.
Update 01/05/2013: I had an epiphany after writing this entry. The two Neolithic ecological zones, plus the grey eastern area, look very ominous to me. They seem to correlate closely with the three main modern European autosomal components - Mediterranean, West Asian and North European - which show up in many ADMIXTURE analyses. So perhaps Impressed Ware/Cardial Ware = Mediterranean component, LBK = West Asian component, and Corded Ware/Unetice = North European component? Check out the map of the ecological zones again and compare it to three gradient maps from an old Eurogenes ADMIXTURE run which show the aforementioned components.
But before you point out that the LBK territory is located in Central Europe and not West Asia on the Banks et al. map, consider that LBK samples usually come out significantly West Asian in ancient DNA studies (for instance, see the Keerl et al. abstract and Haak et al. maps below). Moreover, as already mentioned, LBK groups were eventually replaced by genetically more European populations.
30 successfully typed mitochondrial haplotypes from the Alföld region were compared with 43 samples from Transdanubia and analysed in the light of previously published Neolithic ancient DNA studies. We found similarity between the ALPC and LBK from Transdanubia as well as the published Northwestern LBK datasets of Vedrovice and Derenburg. The studied Linear Pottery Culture groups are distinct from other known European early and middle Neolithic populations, as well as from the Mesolithic hunter-gatherer data. Comparing the ALPC-LBK mitochondrial DNA datasets with modern Eurasian and North African populations, they showed a Southwest Asian affinity. From an mtDNA-perspective, there is evidence for genetic continuity between the LBK in Hungary and the LBK in Central Europe which would be in accordance with the presumed fast expansion of LBK farmers. Furthermore, our results support the Southwest Asian origin of LBK ancestors and the importance of western Hungary for the formation of the Central European LBK.
References...
Banks et al., Ecological constraints on the first prehistoric farmers in Europe, Journal of Archaeological Science, Volume 40, Issue 6, June 2013, http://dx.doi.org/10.1016/j.jas.2013.02.013
Gronenborn et al., ‘Adaptive cycles’ and climate fluctuations: a case study from Linear Pottery Culture in western Central Europe, Journal of Archaeological Science, In Press, Available online 2 April 2013, http://dx.doi.org/10.1016/j.jas.2013.03.015
Keerl et al., Ancient DNA from middle Neolithic Hungary suggests a common ancestry of the Linear Pottery Culture in Central Europe, Early Farmers: the view from archeology and science, 2012
Haak W, Balanovsky O, Sanchez JJ, Koshel S, Zaporozhchenko V, et al. (2010) Ancient DNA from European Early Neolithic farmers reveals their Near Eastern affinities. PLoS Biol 8(11): e1000536. doi:10.1371/journal.pbio.1000536
See also...
Ancient mtDNA from Unetice Culture + genetic origins of modern Central Europeans
Neolithic mtDNA H genomes + Bell Beaker Culture origins in Iberia
This Brotherton et al. paper confirms latest archeological and anthropological findings that the Bell Beaker Culture expanded from the Iberian Peninsula. However, it seems the authors are somewhat befuddled by these results, and attempt to explain them in the following way:
The genetic affinities between Central Europe’s Bell Beakers and present-day Iberian populations (Fig. 2) is striking and throws fresh light on long-disputed archaeological models3. We suggest these data indicate a considerable genetic influx from the West during the LNE (Late Neolithic). These far-Western genetic affinities of Mittelelbe-Saale’s Bell Beaker folk may also have intriguing linguistic implications, as the archaeologically-identified eastward movement of the Bell Beaker culture has recently been linked to the initial spread of the Celtic language family across Western Europe39. This hypothesis suggests that early members of the Celtic language family (for example, Tartessian)40 initially developed from Indo-European precursors in Iberia and subsequently spread throughout the Atlantic Zone; before a period of rapid mobility, reflected by the Beaker phenomenon, carried Celtic languages across much of Western Europe. This idea not only challenges traditional views of a linguistic spread of Celtic westwards from Central Europe during the Iron Age, but also implies that Indo-European languages arrived in Western Europe substantially earlier, presumably with the arrival of farming from the Near East41.
Unfortunately, this approach ignores the mainstream view that the Bell Beaker folk only became Indo-European speakers after they settled in Central Europe, and it was their descendents who introduced Celtic languages to Western Europe from the east in a series of population movements that, in some ways, could be described as back-migrations. Indeed, last year Patterson et al. reported a strong signal of gene flow from Northern Europeans to Iberians dating back to the Bronze Age, which seemed to match the "reverse" spread of Bell Beaker derived groups from Central to Western Europe.
We have detected here a signal of gene flow from northern Europe into Spain around 2000 B.C. We discuss a likely interpretation. At this time there was a characteristic pottery termed ‘bell-beakers’ believed to correspond to a population spread across Iberia and northern Europe. We hypothesize that we are seeing here a genetic signal of the ‘Bell-Beaker culture’ (HARRISON, 1980). Initial cultural flow of the Bell-Beakers appears to have been from South to North, but the full story may be complex. Indeed one hypothesis is that after an initial expansion from Iberia there was a reverse flow back to Iberia (CZEBRESZUK, 2003); this ‘reflux’ model is broadly concordant with our genetic results, and if this is the correct explanation it suggests that this reverse flow may have been accompanied by substantial population movement.
So there's no need to propose that the Bell Beaker Phenomenon was an Indo-European expansion from Iberia. It's more plausible that Bell Beaker groups spoke Vasconic languages related to modern Basque. Note that Basques carry high frequencies of mtDNA H and Y-DNA R1b, and the latter was recently found in Bell Beaker remains from Germany (see here).
Interestingly, it seems that the latest genetic results back up Theo Vennemann's controversial Vasconic substratum theory. In other words, perhaps Vasconic languages once dominated most of Western Europe - and this was due to the Bell Beaker expansion from Iberia - but were eventually replaced by Indo-European languages everywhere except Basque country, leaving their influence in some Indo-European toponyms and vocabulary?
Anyway, the Brotherton et al. paper also includes new evidence which suggests that mtDNA H is a marker of the Neolithic expansion into Europe from the Near East, rather than a signal of the pre-Holocene re-population of Europe from various Southern European Ice Age refugia.
The demographic reconstruction, which is based on direct calibration points, has major implications for understanding post-glacial human history in Europe. Our new estimate is incompatible with traditional views that the majority of present-day hg H lineages were carried into Central, Northern and Eastern Europe via a post-glacial human population expansion before the Holocene (12 kya)13. Our data complement a recent study, based on present-day mt genomes, which describes a pronounced population increase at ~7000 BC (interpreted as a Neolithic expansion into Europe), but followed by a slow population growth until the present day26. By including ancient DNA data from across the critical time points in question, our skyride plot corrects for missing temporal data and suggests substantial growth of hg H from the beginning of the Neolithic and continuing throughout the entire Neolithic period.
...
Although an expansion of hg H could in principle be compatible with a post-glacial resettling of Northern and Central Europe from southwestern refugia12,16 (as indicated by our population skyride and PCA plots), we instead propose that the rise of hg H to become the predominant mtDNA branch in Europe was mediated by subsequent demographic events during the Neolithic, as shown by a general increase in hg H frequency and strong population growth during this period (Fig. 1b). Support for this position comes from data suggesting that hg H was virtually absent among Central and Northern European hunter-gatherers6,43 and formed only 19% in LBK individuals, most likely introduced from Southeast Europe and/or the Near East7. In our updated data set from Mittelelbe-Saale, hg H appears to have been established by the LBK period and increased in frequency after 4000 BC (Supplementary Fig. S3).
So as far as I can see, the mystery of the origins of European mtDNA H is solved. But in any case, mtDNA only tells us half the story. What really needs to happen now is a systematic survey of Y-chromosome haplogroups from prehistoric European remains, which will hopefully be possible thanks to more widespread use of next-generation sequencing (NGS) technology since last year.
Update 27/04/2013: I just did a bit of checking as a result of some of the comments below. True, the Hervella et al. 2012 paper shows two Upper Paleolithic Iberian samples belonging to H6 and H (via HVR polymorphisms).
But I'm not sure about these results, especially the H6. Why would H6 be present in Iberia before the Ice Age, when today it's found at decent levels in Central Asia and the Arabian Peninsula? It's not impossible, but seems unlikely. If anything, I'd expect Upper Paleolithic Iberians to belong to H1 or H3, which are the H subclades most characteristic of Western Europeans today (see here).
Well, if the Hervella et al. samples get tested at a higher resolution and they're confirmed to belong to haplogroup H, or someone else proves beyond any doubt that H was indeed present in pre-Ice Age Iberian samples, then I'll accept it. Until then, I remain skeptical.
Update 28/04/2013: Here's an excellent study from 2010 on the dental morphology of Bell Beaker groups by Jocelyne Desideri and Marie Besse, titled Swiss Bell Beaker population dynamics: eastern or southern influences? I already referenced an earlier paper on Bell Beaker teeth by Desideri below, and this one is similar in many respects, but I think it spells out more clearly what the results say about the origins and movements of the Bell Beaker Culture, and its interactions with Corded Ware and Unetice cultures. It's remarkable how closely the dental data match all the latest ancient DNA studies on late Neolithic/early Bronze Age Europe.
Results of multivariate analyses are eloquent. The first analysis testing possible eastern populational influences gives a very clear picture of relationship between Switzerland and eastern Europe at the end of the Neolithic. Swiss populations group together and do not mix with assemblages from the eastern domain. It should be noted that while the Hungarian non-Bell Beaker assemblages are close to their Czech congenerics, the Hungarian Bell Beaker is clearly distinguished and seems much closer to its western contemporaries. The configuration of Bohemian populations corresponds perfectly to the view of the end of the Neolithic in eastern Europe: with populations from the Final Neolithic (Corded Ware culture) intermediate between the Bell Beaker and the Early Bronze Age (Unetice culture), telling elements of the involvement of the Corded Ware in the appearance of successive periods, which in contrast seem to evolve independently of one another. The second analysis testing possible southern influences clarifies relationships between Switzerland and southern Europe. The Swiss samples are grouped again, but this time associated with later Iberian and French periods.
So, we have seen that the Swiss sites do not mix with the eastern domain, but fit well with the southern domain. The axis of external influences is clearly southern, whether this occurred during the Final Neolithic or the Bell Beaker in western Switzerland. Discussing only the Bell Beaker period, these results conflict with interpretations based on earlier anthropological data (Menk 1979, 1981) used to construct scenarios of Bell Beaker settlement in this region with ideological influences from the south and population movements from the east (Gallay 2006).
...
We have seen that a moderate external population contribution present in the Final Neolithic was increased during the Bell Beaker in western Switzerland. Today, we can clarify, based on the analysis of nonmetric dental variability, that these were mainly people coming from the Mediterranean domain, which is in agreement with the archaeological data from the 3rd millennium BC.
However, it's still a total mystery of who the ancestors of the Bell Beaker folk were and when they arrived in Iberia. It's highly unlikely that they were in large part descendents of Upper Paleolithic Iberians. A more plausible scenario is that their (presumably mostly Y-DNA R1b and mtDNA H) ancestors migrated to Iberia from the Near East during the Neolithic or Epipaleolithic. But how? There are only three options: 1) via the Mediterranean Sea; 2) through Southern and/or Central Europe; 3) across North Africa.
I'd say the first option is probably the best one. After all, the early Bell Beaker folk had strong maritime traditions. That's how they initially colonized much of Western Europe, and they might even be responsible for the R1b found in the Canary Island aboriginal remains (see here).
Update 29/04/2013: There's a new website up called Haplogroup R1b. It's raising funds to study ancient DNA remains in the hope of finally solving the mystery of where European R1b came from (see here). I don't know who's behind the effort, and whether it's legitimate, but their articles are informative and well balanced. Here are some quotes from a piece about the Kromsdorf Bell Beaker samples from Lee et al. 2012, titled R1b and the Bell Beaker Phenomenon.
Busby et al. stated that an east to west migration could not be inferred based on STR variance of R1b1a2’s largest European subclade, R-L11 (aka S127) which makes sense in a rapid spread scenario.
...
Unfortunately the distribution of L11(xP312,U106) is too fragmented and samples too few to draw any conclusion about its origin. The lack of a variance cline is probably the result of a rapid expansion and is attested by similar STR modal values for the three major subclades of R-P312 (aka S116), which are U152, DF27 and L21. In fact, it is at this level that a slight difference in variance (Table 1) and GD (Table 2) can be observed. [Walsh, B. Computing Genetic Distances, (24 Nov. 2003), http://nitro.biosci.arizona.edu/ftDNA/Distance.html, (visited 27 Jun 2012)] The increased variance and decreased genetic distance from P312 makes a case for U152 being the oldest subclade of P312. As such, it is also the likeliest to have occurred near the P312 origin point (Figure 2).
Speculative:
DF27 = Maritime Beaker expansion out of Iberia
U152 & L21 = Reflux expansion from the Alps which would give rise to Italo-Celtic
...
If we take radio carbon data into account which tells us the earliest Bell Beakers occurred in Iberia, an out-of-Iberia DF27 expansion becomes even more intriguing. Finally, if we take the definition of the North South cluster as one with a Northern and Southern coastal distribution, it aligns with the expansion of Maritime Bell Beakers along the Atlantic and Mediterranean coasts. Maritime Bell Beaker cluster seems more appropriate at this time than North-South cluster. There has finally been an Iberian L165 sample found, which would mean the 10x Isles sample bias may be in play.
If we look at the variance of DF27′s siblings, we see that they are younger than DF27 and may have been involved in the later Bell Beaker reflux expansions. The ordering of variance (1. DF27 2. U152 3. L21) is also a good match for radio carbon dating that shows Bell Beaker age as oldest in Iberia/S. France/N. Italy and then progressively younger as one goes north and east.
Main reference...
Brotherton et al., Neolithic mitochondrial haplogroup H genomes and the genetic origins of Europeans, Nature Communications 4, Article number: 1764, Published 23 April 2013, doi:10.1038/ncomms2656
Other references...
Patterson et al., Ancient Admixture in Human History, Genetics: Published Articles Ahead of Print, published on September 7, 2012 as 10.1534/genetics.112.145037
Jocelyne Desideri & Marie Besse, Swiss Bell Beaker population dynamics: eastern or southern influences?, Archaeol Anthropol Sci (2010) 2:157–173, DOI: 10.1007/s12520-010-0037-9
Jocelyne Desideri, Europe during the third millennium BC and Bell Beaker culture phenomenon: peopling history though dental non-metric traits study, Prix Latsis
Universitaires 2008: Présentation des travaux des quatre Lauréats, 2008, vol. 29, p.
15-33
Lee et al., Emerging genetic patterns of the European Neolithic: Perspectives from a late Neolithic Bell Beaker burial site in Germany, American Journal of Physical Anthropology, Article first published online: 3 MAY 2012, DOI: 10.1002/ajpa.22074
See also...
Mitochondrial haplogroup H, lactase persistence, and the formation of the modern European "R1 + H" gene pool
Ancient mtDNA from Unetice Culture + genetic origins of modern Central Europeans
