The people who ranged northwards as the ice sheets finally melted around 10,000 years ago were still using stone tools, but their style had changed sufficiently for archaeologists to recognise Mesolithic sites. These were seldom permanent. Foragers need wide hunting grounds to support each band, and might move with the seasons to take advantage of different food sources. The boat would provide the easiest way to travel and these bold colonisers knew how to build and use it. The seas, and the big rivers that drained into them, provided a transport system through Europe. Rich with fish and shellfish they also provided a large part of the Mesolithic diet. So it's not surprising that many Mesolithic sites hugged the coast or riverside. The fisher-folk of Lepenski Vir, on the banks of the Danube in the Iron Gates gorge, took advantage of the plentiful fish supply to build a permanent village. Their enigmatic sculptures seem to combine man and fish. Yet Mesolithic people also adapted to the advancing forest, while some climbed the greening slopes of the Alps, where they might use caves as dwellings or camp beside lakes.1B. Cunliffe, Europe Between the Oceans: Themes and variations: 9000 BC - AD 1000 (2008), chap.3; G. Bailey and P. Spikins (eds.), Mesolithic Europe (2008); C. Bonsall, V. Boroneant and I. Radovanovic (eds.), The Iron Gates in Prehistory (2008). Comparison with modern hunter-gatherers suggests that once Mesolithic people had fanned out to re-colonise the north, their population would be maintained at replacement level. Fertility levels are low among nomadic hunters; late weaning spaces out births.2M.Livi-Bacci, A Concise History of World Population, 3rd edn. (2001), pp. 35-6. Our modern overcrowded planet makes it hard to imagine how few people there were in Europe in those days. Population density would vary according to the terrain and climate, but has been estimated to between 0.04 and 0.1 persons per square kilometre.3F. Reide, Climate and Demography in Early Prehistory: Using Calibrated 14C Dates as Population Proxies, Human Biology, vol. 81, nos. 2–3, (April–June 2009), pp. 309–337.
The ancestors of the Saami probably followed herds of reindeer and other cold-adapted animals as they moved northwards on the shifting steppe and tundra, ending up in the Nordic lands. The dominant Saami mtDNA haplogroup is U5b1b1a, surprisingly closely related to the U5b1b found among Berbers. We can envision their joint ancestors sheltering in Iberia, before going their separate ways, the ancestors of the Saami to trek north-east across Europe. Yet the paternally-inherited Y-chromosome DNA (Y-DNA) tells a different story. The most common Saami haplogroup is N1c.4K. Tambets et al, The western and eastern roots of the Saami: The story of genetic outliers told by mitochondrial DNA and Y chromosomes, American Journal of Human Genetics, vol. 74, no. 4 (2004), pp. 661-682; A. Achilli et al., Saami and Berbers: an unexpected mitochondrial DNA link, American Journal of Human Genetics, vol.76, no. 5 (May 2005), pp. 883-886.
Haplogroup N ranges from Siberia to Norway and south to China. Its distribution suggests that it spread northwards from South-East Asia, probably following herds of game during the Mesolithic. It is common today among widely separated peoples who turned from hunting to herding reindeer in historic times, such as the Saami at the western end of the range, and the Yakuts towards the eastern end.5M. Derenko et al, Y-chromosome haplogroup N dispersals from south Siberia to Europe, Journal of Human Genetics, vol. 52, no. 9 (2007), pp.763-770; A.O. Karlsson, Y-chromosome diversity in Sweden: a long-time perspective, European Journal of Human Genetics vol. 14 (2006), pp. 963–970. (Note that N3 is the old name for N1c); S. Rootsi, A counter-clockwise northern route of the Y-chromosome haplogroup N from Southeast Asia towards Europe, European Journal of Human Genetics,vol. 15 (2007), pp. 204–211. Within Europe subclade N1c1 is strong among peoples speaking Uralic languages, such as Finnish, Saami and Estonian. These languages sprang from a parent spoken, it seems, near the Ural Mountains, probably among the people of the Lyolovo Culure (5000-3650 BC). The western branch probably arrived in Finland with the Comb Ceramic Culture between 4,000 and 3,000 BC. There Saami seems to have developed as a distinct language in the Iron Age. So picture a mingling of reindeer herds and their hunters coming from different directions at different dates.6T. Lappalainen, Regional differences among the Finns: a Y-chromosomal perspective, Gene, vol. 376, no. 2 (Jul 2006), pp. 207-215; C. Carpelan and A. Parpola (eds), Early Contacts between Uralic and Indo-European: Linguistic and archaeological considerations (2001); A. Aikio, On Germanic-Saami contacts and Saami prehistory, Journal de la Société Finno-Ougrienne (Suomalais-Ugrilaisen Seuran Aikakauskirja), vol. 91 (2006), pp.9-55. A genome-wide study of Finnish Saami showed an average 6% East Asian ancestry today.7J.R. Huyghe, A genome-wide analysis of population structure in the Finnish Saami with implications for genetic association studies, European Journal of Human Genetics advance online publication 8 December 2010.
The Saami spread into Scandinavia about 650 BC. They may have been reinforced at around that time by a new influx from the Volga-Ural region. Some Saami carry the Asian mtDNA haplogroup Z1a. This could have arrived in Finland earlier, except for one factor. The common ancestor of Z1a in Finns, Sami, and folk of the Volga-Ural area has been calculated at just 700 BC.8M. Ingman and U. Gyllensten, A recent genetic link between Sami and the Volga-Ural region of Russia, European Journal of Human Genetics, vol. 15 (2007), pp. 115–120.
The Saami are famed for their constancy to the reindeer. Until well into the historic period they continued to hunt these deer. The Viking Ottar (Ohthere) told King Alfred that he loaned six tame deer to the Saami as decoys, enabling them to catch wild deer.9R. Kerr (ed.), A General History and Collection of Voyages and Travels (1824), vol. 1, p.10. (Finnar was the Norse word for Saami.) By the 11th century AD, the Saami were beginning to turn from hunting to herding.10O. Andersen, Reindeer-herding cultures in northern Nordland, Norway: Methods for documenting traces of reindeer herders in the landscape and for dating reindeer-herding activities, Quaternary International, vol. 238 (2011), pp. 63-75. It was a slow process, and incomplete to this day, for wild herds still exist in Norway and Finland. The Saami domesticated the local reindeer, rather than importing domestic animals from Russia, as is clear from a genetic study. But they were selective. It seems that they found the large, gregarious tundra herds more amenable to domestication than the forest animals. This domestication in historical times has much to teach us about the way other species were coralled by man in the more distant past.11K.H. Røed et al, Genetic analyses reveal independent domestication origins of Eurasian reindeer, Proceedings of the Royal Society: Biological Sciences, vol. 275, no. 1645 (2008), pp. 1849-55; K.H. Røed et al., Elucidating the ancestry of domestic reindeer from ancient DNA approaches, Quaternary International, vol. 238, nos. 1-2 (1 June 2011), pp. 83-88.
Since it is far more difficult to extract Y-DNA from ancient remains than mtDNA, we still have no Y-DNA earlier than 2600 BC. So no clear picture has emerged of which Y lineages dominated the European Mesolithic. Probably the direct lines of many Mesolithic men died out long ago, leaving a jigsaw puzzle for geneticists with most of the pieces missing. A hesitant finger points towards haplogroup I, which is concentrated in Europe. It represents nearly one-fifth of the present European population. Even so we should not imagine that each of its subclades sprang up in the Stone Age wherever it is now found. The subclades of I carried by living men today are relatively young. Some of the most common seem to have travelled widely in the great wanderings after the breakdown of the Roman Empire. The slender clues to the original source of haplogroup I are its diversity in south-eastern Europe and its correlation with known or suspected migrations from that region, mostly long after farming had taken over from fishing and hunting. An expanding population can leave its genetic mark in a burst of new branch-lines within a haplogoup. We see such a burst in haplogroup I2 at c. 8,000 years ago (6000 BC), as farming reached the Balkans. 12Dating used in the map shown here is taken from Ken Nordtvedt personal communications, rather than Campbell and Tiskoff (2010), whose Y-DNA tree has been used here. S. Rootsi et al, Phylogeography of Y-chromosome haplogroup I reveals distinct domains of prehistoric gene flow in Europe, American Journal of Human Genetics, vol. 75 (2004), pp.128–137; M. Pericic et al, High-resolution phylogenetic analysis of Southeastern Europe traces major episodes of paternal gene flow among Slavic populations, Molecular Biology and Evolution, vol. 22 (2005), no.10, pp. 1964-1975; P.A. Underhill et al, New phylogenetic relationships for Y-chromosome haplogroup I: reappraising its phylogeography and prehistory, in P. Mellars, K. Boyle, O. Bar-Yosef and C. Stringer (eds.), Rethinking the Human Revolution (2007); V. Battaglia et al, Y-chromosomal evidence of the cultural diffusion of agriculture in southeast Europe, European Journal of Human Genetics, vol. 17, no 6. (June 2009), pp. 820-30.
Yet in the South-East we can conjecture that Haplogroup I was carried for thousands of years by men who never cultivated a crop or tended a herd. So did the fisher-folk of Lepenski Vir have I-men among them? They were a settled and successful people when farmers arrived in their district, so they were able to adapt to farming on equal terms, making it more likely that whatever Y-DNA they carried would survive.
Another notably successful culture of hunter-gatherers was the Ertebølle of Southern Scandinavia. They had south-eastern links. The earliest pottery in Europe appeared in the Samara region of south-eastern Russia about 7000 BC.13D. W. Anthony, The Horse, The Wheel and Language (2007), pp.148-9 and p. 480, note 19; D. W. Anthony, Pontic-Caspian Mesolithic and Early Neolithic societies at the time of the Black Sea Flood: a small audience and small effects, in V. Yanko-Hombach, A.A. Gilbert, N. Panin and P. M. Dolukhanov (eds.), The Black Sea Flood Question: changes in coastline, climate and human settlement (2007), pp. 245-370 (361). This is a spillover into Europe of the East Asian tradition of ceramic-making foragers.14B. Fagan (ed.), The Seventy Great Inventions of the Ancient World (2004), chapter 8: pottery. From Samara a distinctive type of pottery with pointed bases and flared rims spread up the Volga to the Baltic and appears in the Ertebølle and as far west as the the Low Countries about 5000 BC.15P. Jordan and M. Zvelebil (eds.), Ceramics Before Farming: The Dispersal of Pottery Among Prehistoric Eurasian Hunter-gatherers (2009); D. Gronenborn, Beyond the models: Neolithisation in Central Europe, Proceedings of the British Academy, vol. 144 (2007), pp.73-98 (87); P. Dolukhanov et al., The chronology of Neolithic dispersal in Central and Eastern Europe, Journal of Archaeological Science, vol. 32, no. 10 (October 2005), pp. 1441-1458.
Were men of haplogroup I involved in this dispersal? The subclade I1 (M253) is most diverse in Denmark, suggesting that it arose from I* there. Unlike I2, it shows no indication of expansion with farming. It has its densest distribution in Fenno-Scandia, but spreads into the regions settled by Vikings, Anglo-Saxons, Franks, Goths and other Germanic peoples.16P.A.Underhill et al, New phylogenetic relationships for Y-chromosome haplogroup I:reappraising its phylogeography and prehistory, in P. Mellars, K. Boyle, O.Bar-Yosef and C. Stringer (eds.), Rethinking the Human Revolution (2007); J. Chiaroni, P. Underhill and L.L. Cavalli-Sforza, Y chromosome diversity, human expansion, drift and cultural evolution, Proceedings of the National Academy of Sciences of the United States of America, vol.106, no. 48 (December 2009), pp. 20174-79, fig. 53b.
With mtDNA we are on firmer ground. U5 is one of the oldest European mtDNA haplogroups.17P. Soares et al., Correcting for purifying selection: an improved human mitochondrial molecular clock, American Journal of Human Genetics, Vol. 84 (2009), no. 6, pp. 740-759; K. Tambets et al, Complex signals for population expansions in Europe and beyond, in P. Bellwood and C. Renfrew (eds.), Examining the Farming/Language Dispersal Hypothesis (2002), p.451. It seems to have evolved in Europe and spread northward in the Mesolithic as the climate warmed. U5b1 probably spread from Iberia, while U5b3 seems to have expanded along the Mediterranean coasts from a refuge in the Italian Peninsula. U5a is more strongly Eastern European and may have evolved in a refuge in the Balkans or elsewhere in south-eastern Europe.18B. Malyarchuk et al., The Peopling of Europe from the mitochondrial haplogroup U5 perspective, PLoS ONE, vol. 5, no.4 (2010): e10285; M. Pala et al, Mitochondrial haplogroup U5b3: a distant echo of the Epipaleolithic in Italy and the legacy of the early Sardinians, The American Journal of Human Genetics, vol. 84 (12 June 2009), pp.1-8. U5 has actually been found in Mesolithic DNA. Cheddar Man lived about 7,000 BC near Cheddar, England. MtDNA was extracted from a tooth of his by Bryan Sykes, and found to be haplogroup U5. Tests on hunter-gatherers' remains in Central and Northern Europe found that U5 and U4 overwhelmingly predominated. The oldest lived about 13,400 BC at Hohler Fels, in Germany. They carried mtDNA U*. These were early adventurers north, before the last cold attack. U4, U5a, U5a1, U5b1, and U5b2 were found among later hunter-gatherers in Germany, Lithuania, Poland, Russia and Sweden, including some who made pointed-based pottery. 19B. Bramanti et al, Genetic discontinuity between local hunter-gatherers and Central Europe’s first farmers,Science, vol. 326. no. 5949 (October 2009), pp. 137-140 includes samples from the Narva Culture, with which pointed-based pottery is connected; H. Malmstrom et al, Ancient DNA reveals lack of continuity between Neolithic hunter-gatherers and contemporary Scandinavians,Current Biology, vol. 19 (Nov 2009), pp. 1-5. Today mtDNA haplogroup U5 is widely spread over Europe, though comparatively thinly outside the far north-east, which was relatively untouched by the farmers who brought new mtDNA haplogroups to Europe from the Near East.
Haplogroups H1 and H3 have been proposed as other candidates for a spread northwards in the Mesolithic, since they have their greatest frequency in Iberia. It was initially thought that they were also most diverse there. 20L Pereira et al, High-resolution mtDNA evidence for the late-glacial resettlement of Europe from an Iberian refugium, Genome Research, vol. 15 (2005), pp. 19-24. H3 is scarcely found in the Near East, which seemed a persuasive argument against its arrival with farming. Yet increasing data-gathering has shown that the greatest diversity of H3 is in North Africa, and that for H1 in the Near East.21H. Ennafaa et al., Mitochondrial DNA haplogroup H structure in North Africa, BMC Genetics, vol.10, no. 8 (2009). That suggests that both arrived with early farmers, H3 springing from H* sometime along the Mediterranean route. A recent study from Spain trained its guns specifically upon the idea that the Basques are the modern-day representatives of the hunter-gatherers of the Franco-Cantabrian refuge. Its authors point out that H1 and H3 show a low diversity in Cantabria and particularly among the Basques. They found sub-clade H1t specific to Iberia and calculated a coalescence age for it at 5,800 years. This fitted neatly with their dates for two other mtDNA haplogroups in Iberia to suggest Neolithic radiations: H1r (5,200 years) and HV4a1a (6,500 years). They conclude "In short, we find no well-founded reasons to confirm that the H1 distribution in Europe reflects a human expansion centred on the Franco-Cantabrian area."22O. García et al., Using mitochondrial DNA to test the hypothesis of a European post-glacial human recolonization from the Franco-Cantabrian refuge, Heredity, vol. 106 (2011), pp. 37–45.
Ancient DNA gives mixed messages. In nine samples from Mesolithic sites in different parts of Portugal, one apparently carried mtDNA H1b, and another three H some type, while two were N and one each U4 and U5b1c2.23H. Chandler, B. Sykes and J. Zilhão, Using ancient DNA to examine genetic continuity at the Mesolithic-Neolithic transition in Portugal, in P. Arias, R. Ontanon and C.Garcia-Monco (eds.), Actas del III Congreso del Neolitico en la Peninsula Iberica (2005), pp. 781-86. The U4 was reported as U* and the U5b1c2 as U5. Yet so far there is no sign in ancient DNA that H of any type progressed outside Iberia in the Mesolithic. So prudence dictates that we should await more sophisticated testing of Mesolithic DNA from Iberia. Early studies, such as this one on remains from Portugal, often had problems of contamination with modern DNA.
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