The genetic complexity of prehistoric Sweden

By Razib Khan | May 17, 2011 12:20 am

Thanks to the fact that northern Europe is cool and archaeological research is rather well developed in the region due to quirks of history, there are lots of findings from ancient DNA which are answering long-standing questions. In particular Scandinavia is of special interest in regards to the transition of Europeans from a hunter-gatherer lifestyle to an agricultural one. We know that hunting and gathering as dominant modes of economic production persisted relatively late in European history in this region, up to ~5,000 years before the present. From my cursory reading of the material on the spread of agriculture in northern Europe one dynamic which seems clear is that the rate of expansion was not always constant, and that at the northern fringes in particular social or ecological frontiers served to demarcate the limits to the expansion of farming groups, which often originated from the south and east. Additionally, on the maritime fringes of the North Sea and Baltic there seem to have been relatively dense agglomerations of hunter-gatherers which resisted or coexisted with farming populations for long periods of time (perhaps they were more accurately termed fisher-gatherers!).

This is where Anna Linderholm’s research comes into the picture. I’ve blogged some of her work before. Linderholm’s goal seems to be to synthesize a range of results from disparate fields in understanding how two partially contemporaneous prehistoric Scandinavian cultures related to each other: the Pitted Ware Culture (PWC) and the Funnelbeaker Culture (TRB, which is an acronym for the German name for the culture). The former were hunter-gatherers who tended to rely upon marine resources, while the latter were agriculturalists who engaged in a great deal of animal husbandry.

You can find her contribution to the book Human Bioarchaeology of the Transition to Agriculture online. It’s pretty accessible for an ignorant lay person, and in the chapter she outlines some really interesting detail about the relationship between the PCW, TRB, modern northern European populations, and the functional genetic characteristics of these ancient groups.


The basic chronological outline seems to be that around 3000 BCE there was a period of hundreds of years of contemporaneous habitation of southern Sweden by the TRB and PCW cultures, though they were spatially segregated. TRB finds seem to be concentrated in inland regions, while PCW were found on the maritime fringe. Additionally, the island of Oland in the Baltic exhibited nearly 1,000 years of coexistence of the two cultures. After 2000 BCE these cultures eventually disappeared and gave way to a homogeneous agricultural Bronze Age society.

Were the two cultures of southern Sweden during the Neolithic simply two modes of production of the same people? And were these the ancestors of modern day Swedes? And what can biological anthropology tell us about what they ate and how they ate?

Let’s start with mtDNA relationships:

The mtDNA results point to the fact that the TRB and PWC are two genetically distinct populations (p G 0.001) (Linderholm, 2008; Linderholm, unpublished). Furthermore, the PWC population appears to be more closely related to the modern Latvian population than the contemporaneous TRB population or any of the other modern population examined. This could imply that the PWC population may have had an eastern/central European origin, whereas the TRB population may have had a continental European origin (Linderholm, 2008; Malmstr€om et al., 2009; Linderholm unpublished). These results could also imply that in Sweden the PWC were part of a large hunter-gatherer complex that spanned vast areas of the central and eastern parts of Europe…

Neither of the two populations (PWC and TRB) shows any genetic affinities to the Sami population when compared on the basis of mtDNA sequence. It therefore appears that the ancient mtDNA analyses provide genetic evidence that the TRB and the PWC cultures were Neither of the two populations (PWC and TRB) shows any genetic affinities to the Sami population when compared on the basis of mtDNA sequence….

There’s a little ambiguity here about the TRB’s relationship to modern Swedes, so let me jump to the discussion where this is clarified:

Over 50 individuals were successfully typed and based on these results it has been shown that the PWC and TRB cultures were genetically distinct populations. The genetic legacy of the TRB continued into the Bronze Age and further in time, suggesting that they made a genetic contribution to modern Swedish populations. In contrast, the PWC seem to have left no genetic legacy to the modern Swedish population, but their genetic signals can be detected in modern north-eastern European populations…..

Modern Sweden was settled ~10,000 years ago after the retreat of the Ice Age. The TRB culture first arrived in Sweden ~6,000 years ago. From these results one might infer then that modern Swedes derive predominantly from a migration of farmers from the north European plain ~6,000 years before the present. In these analyses the identity of the hunter-gatherers resident in region (e.g., Skane) for the 4,000 years after the Ice Age and before the arrival of farming is left unclear. The PWC itself may have been intrusive, and closely related to a broader group of northeast European hunter-gatherer societies. Do recall that this study focused on southern Sweden. We know that the Sami were indigenous to northern Sweden relatively late into the historic period. Whether they too were exogenous on this sort of time scale, we don’t know yet.

Next Linderholm reviews previous results on the prevalence of lactose tolerance among these two societies. The results are pretty straightforward: the PWC generally lack the ability to digest lactose. This makes sense, why would they need this mutation if they weren’t engaged in animal husbandry? There’s an allele in northern Europeans which is strongly correlated with lactose tolerance. It operates in a dominant fashion, so the fact that the frequency of the allele in modern Swedes is 75% means that lactose tolerance is nearly at 95%. But the question left unanswered is the state of the situation in the TRB. It seems that this is work that is proceeding now (from what I can tell there was less success amplifying the TRB remains for whatever reason), but the author politely hinted strongly: “Analysis has also been performed on individuals representing the TRB population; the outcome of these analyses has not yet been totally verified. In the dataset we can detect a large rise in T-allele frequency in Sweden amongst the farming community (unpublished data).” This is the reasonable expectation, as the TRB kept cattle. It seems plausible that the rise of farming on the northern European frontier was in large part a function of the innovation of dairying, as in these regions the primary productivity of Middle Eastern cereal crops was far less. Rather, less palatable crops needed to be cultivated, and it may be that it was more efficient to use this as feed for cattle which then produced milk (e.g., oats).

Finally, they assayed the CCR5-D32 variant in their PWC and TRB samples. This is the “HIV resistance” allele found in many Europeans, whose origin has been hypothesized to be 1400 CE or 3000 BCE, depending on which coalescence model you trust. Interestingly Linderholm found that both of these ancient groups carried the resistant allele, likely pushing back the origin of the variant.

So what does this tell us in the end? We need to keep in mind other facts, and not lose the big picture. The populations of Norden, whether Finnic or Scandinavian, are genetically similar. But clearly there is a distinction as one moves from Sweden to Finland. The relationship between the Baltic populations and the Nordic ones is somewhat confused. What I can say from ADMIXTURE runs performed by genome bloggers is that it seems that Scandinavian, as opposed to Finnic, populations seem to have an ancestral component at low, but consistent, levels which is modal in the Middle East. The lack of this distant Middle Eastern affinity among the Finns, and to a lesser extent the Lithuanians, is to me telling. I suspect what you’re seeing is an ancient admixture event, greatly diluted by the time one approaches the far north, which was carried from the southwest by the ancestors of modern Scandinavian populations. The Finns were not impacted by this migration, and so do not have any imprint of that distant Middle Eastern ancestral component.

Second, the relationship of the PWC, hunter-gatherers, and TRB, farmers, needs to be teased apart. It does not seem this was a classic replacement of hunter-gatherers by farmers. Both these groups were intrusive to southern Sweden in relation to the Mesolithic hunter-gatherers who originally repopulated the peninsula. It could be that there was a symbiosis between the PCW and TRB, but it is notable that the genetic distinction between these groups is very strong. Additionally, the authors note that there is no connection between the PCW and modern Swedes, though there is one between the TRB and modern Swedes (in the late Neolithic and early Bronze Age the TRB were replaced by the “Battle-Axe” culture, which may or may not have been an organic outgrowth of the TRB). It doesn’t take rocket science to make an inference of “who won.” All that said, I wonder if one could make the case for the PWC being highly efficient superior hunter-gatherers, the “last stand” so to speak of a non-marginal manifestation of this lifestyle in Europe. Their concentration in coastal fringes suggests that they were focusing on regions where their per unit labor had the greatest returns. It may be that for a period they were at demographic parity with the TRB because the farmers had simply not perfect their cultural toolkit. An analysis of isotope ratios in the bones of the remains points to the possibility that the TRB took a very long time to acclimate to the north: “The results show a gradual transition from a hunter-gatherer diet to a diet based on agriculture took place by the TRB people (which is thought to be associated with the introduction of farming on the island) only by the end of the Neolithic and hence much later than had been previously thought.” If the TRB were facultatively engaging in behavior more typical of foragers then it seems understandable that they didn’t push aside the PWC as easily as they might have, because they lacked the traditional farmer demographic advantage.

There’s a lot more to come I assume. Though there was some analysis of nuclear ancient DNA the focus here was on mtDNA. That’s because mtDNA is copious. But as the techniques get better I assume museums might be more willing to part with parts of their specimens. Overall these results make me more cautious of maximalist clean models of simple replacements. Here’s a speculative scenario which I think might have been:

– TRB emerges out of a hybridization of native north European plain marine foragers and farmers from central Europe, who are themselves a hybridization of southern migrants and indigenes.

– Once TRB crystallizes with an agricultural toolkit more appropriate to the far north it quickly sweeps back the older farming frontier. The old-stock hunter-gatherers of the north melt away. But the TRB then are challenged by the “climax” culture of the northern hunter-gatherers, the very efficient lifestyle of which the PWC were partaking of, focusing on marine resources which were easy to obtain in large quantities. The rise of these hunter-gatherers may partly have been competition, but their lifestyle may also have been dependent upon trade with the dense agricultural societies to their south and west.

CATEGORIZED UNDER: Anthroplogy, Genomics
  • Domino

    Is there any truth in the theory that blondeness of Scandinavia occurred due to the dietary changes involved with farming? Simply put:
    The Gulf Stream allows the Scandinavian/Baltic region to grow grain crops. Nowhere else in the world were these kind of crops so far north. This change in diet meant that they were unable to get enough Vitamin D, which they had previously received from a heavy meat-based hunter-gather lifestyle. Natural selection’s answer was to reduce melanin so that Vitamin D could be absorbed through the skin.

    Other northern peoples, like Eskimos, are still quite dark since they get huge amounts of Vitamin D from fish and meat. Also, the Sami are slightly darker than other Scandinavians.

    The mythology kind of fits this: http://en.wikipedia.org/wiki/R%C3%ADg_(Norse_god)#Synopsis
    ..where the skin coloring goes from dark to light via farmers.

  • Eurologist

    Much of what you write makes sense to me – especially your last two conclusions – I would have expressed it similarly. PWC supposedly also kept pigs and perhaps originated as a subset of a somewhat agricultural-oriented culture. But yeah, for millennia the fishing part all around the Baltics and Scandinavia surely was productive and sustainable.

    “What I can say from ADMIXTURE runs performed by genome bloggers is that it seems that Scandinavian, as opposed to Finnic, populations seem to have an ancestral component at low, but consistent, levels which is modal in the Middle East.”

    Not sure about that, or how important that is. It is already very low in Germans. In my view, Anatolian or West Asian contribution was low to start with in the middle Danube region (NW Balkans), and became almost irrelevant after fringe people largely took over the Danubian culture.

  • http://blogs.discovermagazine.com/gnxp Razib Khan

    Not sure about that, or how important that is. It is already very low in Germans.

    its not noise. it’s consistent.

  • http://dioegenesartemis.blogspot.com/ Diogenes

    Different Mitochondrial DNA haplogroups found in different groups don’t exclude some genetic affinities at the autossomal level. And it is expected that agricultural groups which “abandoned agriculture” in favour of pig-farming and fishing (and with pottery) to present different and much more forager admixed mtDNA haplogroup profiles.

    To compare current Near Eastern genetics with current European ones and upon finding only “limited portions” to be very highly similar (most of the rest being simply closely related), and thus exclude major ancestry from the ancient Near East in such populations is probably quite simplistic, considering the necessarily multi-layered complexity of Neolithic Revolutions.
    Postulating a first “critical point” enabling spread of Neolithic peoples; and secondary points, corresponding to major secondary advances in food production, allowing for new waves with partial replacement of less advanced farming peoples, would explain why the descendants of the periphery today may be moderatelly genetically different compared to the descendants of the core areas, since core innovations would tend to have a major local impact, but decreasing genetic impact with distance. That is, the “wheat agriculture invention” and all necessary adaptations leading to it, may be hard to replicate and pratically impossible for foragers to “copy” in useful time; better seeds, irrigation techniques/social organization may appear in more places and are easier for more primitive farmers (with adequate “mindset”) to eventually adapt to/copy. So an “increasing hybridization with distance” model is very well suited to first farmer/second farmer waves as well (probably better). Secondary Neolithic waves would have “higher drag”.
    First waves would on the other hand be possibly quite “fluid” in forager frontiers.
    So in a first approach, first wave elements could be mistaken for foragers and secondary ones for a single Neolithic wave.

    Genetic correspondences between the current Near East and Europe likely correspond to such secondary waves, since the Near East may have suffered major internal change due to such secondary “critical points” rendering the “homelands” of European frontier Neolithic waves difficult to recognize today. Another point is the case for pig-farming, pottery-making (possibly occasional oats and rye planters?) being mistaken for original paleolithic foragers just because in a difficult climate they (at first) adapt by fishing and hunting readily available resources.
    High density forager environments (rich coasts) likely presented unique niches for Neolithic organization expertise. Innovations related to high density in farming societies would possibly allow for major competitive advantages of hybridized populations versus “pure” foragers in such rich foraging enviroments. One would expect such hybridization to occur, with high levels of original forager mitDNA in the resulting population. These populations/cultures might even be expected to spread to similar niches further “out” as a “developed forager” wave (a “preshock” to what was coming). Nonetheless they would still have unique disadvantages versus exclusive farmers once appropriate farming technology for such colder coastal regions appeared: they would then likely then be swamped out in a “secondary wave” fashion. So late highly developed settled foragers communities may also be explained in an opposite model: as a first stage of Near Easterner Neolithic colonization with at first only partial replacement of natives and not as aboriginal adaptation.

    So such patterns in Scandinavia can be explained by alternative scenarios, for which some support I think can be found in ADMIXTURE runs. A first Ukraine river valley Neolithic sweep from the Near East; a second related but separate advanced pastoralist incursion into the area; the expansion of the farmers in conflict with pastoralists to coastal marginal lands for agriculture, partial reversion into foraging in rich coastal environments and consequent hybridization with foragers, followed by their substitution by more advanced/adapted farmers later on (from the Corded Ware region where advanced farming with winter-rye/barley may have developed).
    In colder areas such as Lithuania, forager mitDNA would persist since the first hybridizing impact of foragers was great. In richer areas Neolithic mitDNA carried by exclusive farmers may be more important. In both, autossomes likely speak of complete replacement.
    I think it’s possible Uralic tongue expansion may correspond to such a hybrid population with high density/cooperation adaptations.

  • pconroy

    Razib,

    I presume you’re talking about the putative “West Asian” (aka Georgian usually) component that many genome bloggers have found, that is usually associated with a wave of farming out of the Middle East, right?

    In terms of Northern Europe, you make the case – quite rightly – that this element is lacking in Finnic populations, but present in Scandinavia at a low consistent rate.

    Not surprisingly, this West Asian component is also present in Britain and Ireland, other Northern European countries.

    However, based on the Dienekes’ Dodecad Project, there are 17 Irish 23AndMe samples currently, and of these 11 have publicly identified themselves. Based on this Irish K=12 data, I have put together an Irish K=12 chart.

    The only sample that I know of that has no non-Irish ancestry is my father DOD098, and interestingly, he has NO West Asian element, while also having both the highest NE European element and the highest Basque element. On 23AndMe he appears as the most NW European in the European view.

    What does this mean? Initially I thought it probably meant Viking ancestry, but maybe it just means that he has relic North European ancestry – which is more like NE European ancestry, while also have relic Atlantic ancestry, which is more like Basque??

  • Robert

    “I think it’s possible Uralic tongue expansion may correspond to such a hybrid population with high density/cooperation adaptations.”

    :clap: Very tightly argued. I knew there had to be an explanation for the existence of pickled herring in sour cream and sweet onions!

  • http://washparkprophet.blogspot.com ohwilleke

    Interesting stuff.

    * “it seems that Scandinavian, as opposed to Finnic, populations seem to have an ancestral component at low, but consistent, levels which is modal in the Middle East. The lack of this distant Middle Eastern affinity among the Finns, and to a lesser extent the Lithuanians, is to me telling. I suspect what you’re seeing is an ancient admixture event, greatly diluted by the time one approaches the far north, which was carried from the southwest by the ancestors of modern Scandinavian populations. The Finns were not impacted by this migration, and so do not have any imprint of that distant Middle Eastern ancestral component. . . . the authors note that there is no connection between the PCW and modern Swedes, though there is one between the TRB and modern Swedes (in the late Neolithic and early Bronze Age the TRB were replaced by the “Battle-Axe” culture, which may or may not have been an organic outgrowth of the TRB).”

    The case that this genetic component and cultural transition represent the appearance of the Germanic languages carried by an immigrant Indo-European superstrate that arrives at this point seems very strong.

    * “there is no connection between the PCW and modern Swedes,”

    So, it would seem that PCW is unlikely to be the source of the blond/blue eyed phenotype which is so common in modern Swedes (selection could do the trick, but one would think that selection in favor of PWC appearance genes would carry at least some PWC mtDNA with it, and the populations that are closest to the PWC genetically are not the strongest in that phenotype). It would also seem odd for a population source that shows up as a Middle Eastern ancestral component (which I suppose to be proto-Germanic and to date roughly to the late Nordic Bronze Age) to have that phenotype. (FWIW, the blond-blue eyed phenotype in modern humans is also known to be rooted in non-Neanderthal genes). By process of elimination it would seem that TRB is the most plausible source of that phenotype, which it might have evolved around the same time that it was becoming lactose tolerant.

    * “Neither of the two populations (PWC and TRB) shows any genetic affinities to the Sami population when compared on the basis of mtDNA sequence.”

    (N.B. there seems to be a glitch in your blockquote at this point.)

    It doesn’t seem unfair to suppose that the PWC was probably linguistically Uralic, and the expanse of the PWC corresponds reasonable well with the historical range of the Uralic languages and the fact that the Uralic urheimat was probably not in Northern Scandinavia. Given the lack of genetic affinity between the Sami and the PWC, and the fact that the Sami now speak a Uralic language, it is fair to suppose that it is more like that the Sami lost their language in favor of a Uralic language from a PWC people, instead being the source of the Uralic languages.

    The Sami uniparental lineages with the Berbers that are almost absent from intermediate populations, and have a high frequency of mtDNA hg U5, one of the modal hgs of the oldest European hunter-gatherer mtDNA. The geographic discontinuity in lineages shared by the Sami and Berbers, along with the fact that nether population was a farming population (at least until very recently), suggests that these populations have roots in a pre-Neolithic migration – one natural path of such a migration would have been along the Atlantic coast since any population that could have crossed the straights of Gibraltar and the Baltic Sea in measurable numbers would have had to have had more than rudimentary maritime capabilities.

    Also, while both Sami and Berbers have had predominantly nomadic pastoralist subsistence patterns in historic eras, it is notable that reindeer are not part of the Fertile Cresent package of domesticates, nor in the conventional account are camels a core part of that package. Thus, there may have been domestications of core animals species by both groups distinct from the Fertile Cresent package. This argues from a transition from hunting and gathering to pastoral nomadic subsistance that was a result of cultural imitation rather than admixture with other herders. The absence of Berber domesticates in the Sami pastoral package (and visa versa), while it could be simply a product of substitution given the necessities of climate, is not inconsistent with a common genetic ancestor of the groups during a hunter-gatherer rather than a pastoral nomadic era (again consistent with a pre-Neolithic link).

    Given the fact that the current territory of the Sami was not inhabitable at the time of the LGM, however, they must have arrived from refugia to the South after the LGM. Thus, the migration that links the Sami and the Berber was probably Epipaleolithic. In contrast, ancient DNA from 12,000 years ago in North Africa shows genetic continuity between that pre-Neolithic population and modern Berbers.

    Given that there is population genetic unity to some extent within Afro-Asiatic language macrofamilies (Semitic, historically Coptic, Berber, Chadic, Cushitic/Omotic), but not between Afro-Asiatic language macrofamilies, it is fair to infer the the Berber’s probably acquired their Afro-Asiatic language as a result of cultural transfer rather than demic replacement (to my knowledge, no one has seriously argued that Berber is the most basal Afro-Asiatic language), most plausibly in connection with the other cultural changes associated with the early Neolithic in North Africa. If that scenario is correct, then the language spoken by the Berbers in the Epipaleolithic was probably not Afro-Asiatic (and the probably related proto-language of the Sami probably wasn’t either). The original languages of the Berbers and Sami have probably now been replaced by Afro-Asiatic Berber and Uralic Sami languages respectively with their original language lost unless it is a relative of Basque (a population that shares some hgs otherwise mostly found in Berbers and Sami, that shows cultural continuity from a fisher-gatherer era rather than merely back to the farmer era, and hence a possibility that is not to be dismissed out of hand, although Basque has also frequently been suggested as related to the megalithic culture, which would suggest a linguistic link to the TRB rather than than to the Sami).

  • Lassi Hippeläinen

    Before jumping to conclusions, remember that the blue-eyed blonde phenotype is common also in Finland and Russia. IIRC, there are more blondes east of the Baltic Sea than west.

    Also, the seminomadic reindeer herding of the Sami is a recent development. They probably had some reindeer for their own needs since ancient times – it being a common practise with all Arctic peoples – but it became important only in 15th and 16th centuries with the spread of money economy.

  • http://blogs.discovermagazine.com/gnxp Razib Khan

    Before jumping to conclusions, remember that the blue-eyed blonde phenotype is common also in Finland and Russia. IIRC, there are more blondes east of the Baltic Sea than west.

    also, the site of the original mutation isn’t necessarily where it is most common today. the blue eye variant in the mid east seems to be the same one as in europe (same haplotype), and dates to ~10,000 B.P. one can imagine (this may not be true) that it came north with the farmers, who had it as part of their background variation, and that it was selected upward in the north for whatever reason.

  • http://washparkprophet.blogspot.com ohwilleke

    “Also, the seminomadic reindeer herding of the Sami is a recent development.”

    What was their subsistence economy like before that?

  • http://dioegenesartemis.blogspot.com/ Diogenes

    Metis people are quite numerous for an “aboriginal” people…
    http://en.wikipedia.org/wiki/M%C3%A9tis_people_(Canada)
    As are Griqua for a khoisan-descended group.
    http://en.wikipedia.org/wiki/Griqua_people
    Also lingua geral was a good candidate for national language of colonial Brazil for a while
    http://en.wikipedia.org/wiki/L%C3%ADngua_Geral_Paulista
    As was Guarani in some other parts of South America
    http://en.wikipedia.org/wiki/Guaran%C3%AD_language
    All products of early European colonization of new forager areas.
    I believe there are a few now marginal American examples as well, but can’t remember names.

  • Pingback: The genetic complexity of prehistoric Sweden | Gene Expression … | Today Headlines()

  • Lassi Hippeläinen

    @10: The Sami were hunter-gatherers. Hunting meant mostly fishing and trapping birds or rodents. And the short but furious summer produces lots of berries and mushrooms.

    Since they were mobile people, they had no problems in following reindeer herds. That’s why even the Swedish government pressured them to change from h&g to reindeers.

    They still do h&g, but more as a hobby these days. Reindeering is now an industry. Nomadism is practically gone. They live in villages and visit the herds with snowmobiles or fourwheel bikes. It shows how ready they are to adopt new ideas. That wouldn’t work if they had long traditions to defend.

    Besides, reindeering is not specific the Sami, it is specific to Lapland. Also many Swedes and Finns who moved to the Wild North got into the business.

  • pconroy

    Also interesting in light of the eclipse of the PWC by the TRB is the fact that ancient dog remains from Sweden don’t match current dogs there at all.

    This article finds that:

    More precisely, the data obtained on French Neolithic dogs,
    combined with the analysis of Swedish (Malmstrom et al., 2008)
    and one Italian (Verginelli et al., 2005) Neolithic dogs, confirm that
    Hg C was well represented all over the western part of the European
    continent during this period. As Hg A encompasses the majority of
    extant European dogs (70% for Hg A, 6% for Hg C), we speculate that
    Hg C has been replaced through time by Hg A in Europe. Malm-
    strom et al. (2008) also clearly identified a lineage replacement in
    Scandinavia (with a clear emergence of Hg D in modern dog gene
    pool).

    Ancient DNA supports lineage replacement in European Dog genepool

  • neshumno

    “the blue eye variant in the mid east seems to be the same one as in europe (same haplotype), and dates to ~10,000 B.P.”

    This time estimate, like many others, could be wrong:

    http://alturl.com/u8doi

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About Razib Khan

I have degrees in biology and biochemistry, a passion for genetics, history, and philosophy, and shrimp is my favorite food. In relation to nationality I'm a American Northwesterner, in politics I'm a reactionary, and as for religion I have none (I'm an atheist). If you want to know more, see the links at http://www.razib.com

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