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The Peopling of Europe

June 14, 2012


The first Europeans

The earth’s climatic cycle of freeze to fry tugged humans hither and thither. Warm spells enticed early hominids out of Africa, while cold spells every 125,000 years or so drove them to extinction or withdrawal from northerly climes.1I.S. Castañeda et al., Wet phases in the Sahara/Sahel region and human migration patterns in North Africa, Proceedings of the National Academy of Sciences of the United States of America, (online before print November 12, 2009); B. Arredi, E.S. Poloni, C. Tyler-Smith, The peopling of Europe, in M. Crawford (ed.), Anthropological Genetics: Theory, method and applications (2007), p. 383.

Anatomically modern man (Homo sapiens sapiens) crossed into Europe from Asia some 45,000 years ago.2M. V. Anikovich et al, Early Upper Paleolithic in Eastern Europe and implications for the dispersal of Modern Humans, Science, vol. 315. no. 5809 (12 January 2007), pp. 223-226; P.Mellars, Archeology and the Dispersal of Modern Humans in Europe: Deconstructing the Aurignacian, Evolutionary Anthropology, vol. 15 (2006), pp. 167–182. Pictured here is a facial reconstruction by Richard Neave from the earliest skull of anatomically modern man found in Europe. He commented that the skull looked like a mixture of modern Western Eurasian, East Asian and Sub-Saharan African. The continental differences we see today had yet to evolve. (The skin colour can only be guesswork. For more on that see Pigmentation.) The 35,000-year-old skull was discovered in the Peştera cu Oase (The Cave with Bones) in Romania.3João Zilhão et al., The Pestera Cu Oasepeople, Europe’s earliest modern humans, in P. Mellars, K. Boyle, O. Bar-Yosef,and C. Stringer, (eds.), Rethinking the Human Revolution (2007); Hélène Rougier et al., Peştera cu Oase 2 and the cranial morphology of early modern Europeans, Proceedings of the National Academy of Sciences of the United States of America, vol. 104, no. 4 (23 January 2007), pp.1165–1170.

So how do we know that Homo sapiens arrived in Europe some 10,000 years before the man from Peştera cu Oase? Teeth from Grotta del Cavallo, southern Italy, have been recently reclassified as belonging to anatomically modern humans some 45,000–43,000 years old, while an anatomically modern human jaw from Kent’s Cavern, UK has been recently re-dated to between 44,200 and 41,500 years old.4S. Benazzi et al., Early dispersal of modern humans in Europe and implications for Neanderthal behaviour, and T. Higham et al., The earliest evidence for anatomically modern humans in northwestern Europe, both Nature, Published online 02 November 2011. In the main though the clues lie in the things our forefathers left behind. These early arrivals were hunter-gatherers using stone tools. (Our name for the period – Palaeolithic – comes from the Greek for old and stone.) Their ancestors had undergone a behavioural evolution long before in Africa, gradually accumulating the features we identify as human: tool use, self-decoration, clothing, burial of their dead, trading and other activities that require knowledge passed on within a community and so imply language.5O.Bar-Yosef, The dispersal of modern humans in Eurasia: a cultural interpretation, and Sally McBrearty, Down with the revolution, both in P.Mellars, K. Boyle, O. Bar-Yosef, and C. Stringer, (eds.), Rethinking the Human Revolution (2007). From the DNA of the clothing louse, scientists have deduced that clothing appeared possibly as early as 170,000 years ago, 70,000 years before modern humans started migrating to colder climates.6M.A. Toups et al., Origin of clothing lice indicates early clothing use by anatomically modern humans in Africa, Molecular Biology and Evolution, vol. 28, no. 1 (2011), pp. 29-32. 100,000 years ago in Blombos Cave in South Africa, our ancestors were grinding and mixing ochres to make red and yellow paint.7C.S. Henshilwood et al., A 100,000-Year-Old Ochre-Processing Workshop at Blombos Cave, South Africa, Science, vol. 334, no. 6053 (14 October 2011), pp. 219-222. Pea-sized Nassarius shells found at Blombos Cave and equally ancient sites at Oued Djebbana in Algeria and Skhul, Mount Carmel, Israel, were perforated as though they had been strung together like beads in necklaces or bracelets. There were signs of wear from a leather string. If the aim was personal adornment, then these are the earliest known pieces of jewellery.8M. Vanhaeren et al., Middle Paleolithic Shell Beads in Israel and Algeria, Science, vol. 312, no. 5781 (23 J une 2006), pp. 1785-1788. Art and craft are among the defining signs of modern human behaviour.

Modern Man had spread right across Asia and into Australia before a burst of warm weather made it possible to move north into the Levant and from there to Europe. Paul Mellars has tracked the tools they left along the way (see map above). Flint tools cannot be radiocarbon dated, but ancient people also used bone and antler, which can. A characteristic tool made by those spreading across Europe is the split-base antler point, first found at Aurignac in the Pyrenees, from which the technology was named Aurignacian. These split-based points appear earliest in the Levant. In fact they occur there as part of the Ahmarian tool-set, prior to the development of Aurignacian types. Crucially remains of a fully modern human were found in the Ahmarian layer at Ksar Akil. Split-based points occur next in South-East Europe, so we may guess that people crossed what was then a land bridge west of the Black Sea (then a lake). 9P.Mellars, Archeology and the Dispersal of Modern Humans in Europe: Deconstructing the Aurignacian, Evolutionary Anthropology, vol. 15 (2006), pp. 167–182. For a slightly different perspective see J.F. Hoffecker, The early upper Paleolithic of eastern Europe reconsidered, Evolutionary Anthropology: Issues, News, and Reviews, vol. 20, no. 1 (January/February 2011), pp. 24–39.


These first adventurers surely must have encountered Neanderthals – their distant genetic cousins who had been in Europe from about 400,000 years ago.10J. L. Bischoff et al., High-Resolution U-Series Dates from the Sima de los Huesos Hominids Yields 600+/–66 kyrs: Implications for the Evolution of the Early Neanderthal Lineage, Journal of Archaeological Science, vol.34, no. 5 (May 2007), pp. 763-770; J. J. Hublin, The origin of Neandertals, Proceedings of the National Academy of Sciences U.S.A., vol. 106 (2009), pp.16022-16027. The arrival of Modern Man in an area seems generally to signal the departure of Neanderthals – never a large population. It was thought that they survived longest in southwestern Iberia, where Modern Man arrived late. Neanderthals died out there around 37,000 years ago.11O. Joris and M. Street, At the end of the 14C time scale – the middle to upper paleolithic record of western Eurasia, Journal of Human Evolution, vol. 55 (2008), pp. 782–802; J. Zilhão et al., Pego do Diabo (Loures, Portugal): dating the emergence of anatomical modernity in westernmost Eurasia, PLoS ONE vol. 5, no. (January 2010): e8880; J. Martínez-Moreno, R. Mora and I. de la Torre, The Middle-to-Upper Palaeolithic transition in Cova Gran (Catalunya, Spain) and the extinction of Neanderthals in the Iberian Peninsula,Journal of Human Evolution, vol. 58, no. 3 (2010), pp. 211-226. So it was a surprise to discover a typical Neanderthal toolkit dated between 31 and 34 thousand years ago at Byzovaya, in subarctic Russia. This site in the Polar Urals may be one of the last refuges of the Neanderthals.12L. Slimak et al., Late Mousterian persistence near the Arctic Circle, Science, vol. 332, no. 6031 (13 May 2011), pp. 841-845.

Did our ancestors interbreed with archaic hominids like Neanderthals? One genetic model from modern DNA predicts two such events in human history which left a record in our code, one about 60,000 years ago in the eastern Mediterranean and one about 45,000 years ago in eastern Asia.13R. Dalton, Neanderthals may have interbred with humans, Nature News, (published online 20 April 2010). And see E.Y. Durand et al, Testing for ancient admixture betweenclosely related species, Molecular Biology and Evolution (online15 February 2011 before print). Yet an alternative model dispenses with any such events,14M.G.B. Blum and M. Jakobsson, Deep divergences of human gene trees and models of human origins, Molecular Biology and Evolution, vol. 28, no. 2 (February 2011), pp. 889-898. while an earlier study found strong evidence for ancient mixture in both a European and a West African population. Africa had no Neanderthals.15V. Plagnol and J. D. Wall, Possible Ancestral Structure in Human Populations, PLoS Genetics, vol. 2 (July 2006), pp. 972-979. And see M.F. Hammer et al., Genetic evidence for archaic admixture in Africa, Proceedings of the National Academy of Sciences of the United States of America, online September 6, 2011 before print.

Now that the Neanderthal genome has been sequenced from ancient DNA, it is possible to make direct comparisons. A preliminary comparison found that Neanderthals shared more genetic variants with the present-day people of both Europe and East Asia than with sub-Saharan Africans.16R.E. Green et al, A Draft Sequence of the Neandertal Genome, Science, vol. 328. no. 5979 (7 May 2010), pp. 710-722. That suggests that Neanderthals mixed with the ancestors of non-Africans before the future Europeans went one way and Asians another. Caution is needed though. The result might simply spring from insufficient sampling of the more diverse African population.17J.A. Hodson, C.M. Bergey and T. R. Diostell, Neandertal genome: the ins and outs of African genetic diversity, Current Biology, vol. 20, no. 12, R517-R519 (22 June 2010).

More crucially, Neanderthals are not the only candidate for archaic hominid mixture. A more plausible alternative is the North African archaic Homo sapiens lineage known as Aterian, which seems to have crossed into the Levant in an early wave of migration. As a much closer relative than Neanderthals to the next wave of Homo sapiens leaving Africa, the Aterians would be more likely to be able to successfully interbreed with them, with fertile offspring. Yet at the same time the lengthy separation of Aterians from their kin in Sub-Saharan Africa left them with more archaic traits similar to those of Neanderthals. A similar proposal has been put forward by Silvia Ghirotto and colleagues, who point out that the evidence from mtDNA is of no interbreeding at all between Neanderthals and early modern man. They feel that nuclear DNA and mitochondrial evidence might be reconciled if Neandertals shared a longer period of common ancestry with the ancestors of present-day non-Africans than with the ancestors of modern Africans.18S. Ghirotto et al., No evidence of Neandertal admixture in the mitochondrial genomes of early European modern humans and contemporary Europeans, American Journal of Physical Anthropology, vol. 146, no. 2 (October 2011), pp. 242–252.

Hardy hunters

The earliest DNA retrieved from a modern human comes from a 30,000-year-old man unearthed at Kostënki 14 (Markina Gora) in Russia. Scientists were able to study his mitochondrial DNA (mtDNA). This type of DNA is passed down unchanged from mother to child, unless a mutation arises. His haplogroup was U2.19Krause, J. et al., A complete mtDNA genome of an early modern human from Kostenki, Russia, Current Biology (online 31 December 2009). U2 today is scattered at low frequencies in populations from South and Western Asia, Europe and North Africa, with its oldest branches (U2a-c) in South Asia. That is a clue that people carrying U2 had spent a long time in the warm south before a group split off to travel north into Europe, where the mutation creating U2e probably occurred. U2e is mainly found in those of European descent.

Similarly the parent haplogroups M, N and R are all ancient in South Asia. This suggests that Modern Man crossed from East Africa to Arabia and then across the Persian Gulf into what is now Central Asia. There groups seem to have split off, some to populate Asia and move on to Australasia and the Americas, others to move westwards to the Levant and Europe. (See the first map on Peopling of Europe.)

The first Europeans did not simply live to hunt. They were creative. Aurignacian people carved simple flutes from mammoth and swan bone.20N. J. Conard, M. Malina and S.C. Münzel, New flutes document the earliest musical tradition in southwestern Germany, Nature, vol. 460, pp. 737-740 (6 August 2009). Their figurines of animals include the now extinct mammoth, carved in mammoth ivory.21C. Heckel, Physical characteristics of mammoth ivory and their implications for ivory work in the Upper Paleolithic, Mitteilungen der Gesellschaft für Urgeschichte, vol. 18 (2009), pp. 71-91.

The culture which followed the Aurignacian is known as the Gravettian, after La Gravette in France, where small, pointed blades used for big-game hunting were found; these became recognised as characteristic of the period from about 28,000–23,000 years ago in Western and Central Europe. The Gravettian tool-set appears earliest in Eastern Europe: the earliest radiocarbon dates so far come from Buran-Kaya, Crimea, Ukraine (31,900+240/−220 BP), Obłazowa cave, Poland (31,000±550 BP), Willendorf, Austria (30,500+900/−800 BP), and Molodova, Ukraine (29,650±1,320 BP).22S. Prat et al., The oldest Anatomically Modern Humans from far Southeast Europe: direct dating, culture and behavior, PLoS ONE, vol. 6, no. 6 (2011), e20834. A group of Gravettian people lived in Paglicci Cave in Italy, leaving cave paintings as well as tools behind them. David Caramelli and his colleagues tested three skeletons from the cave for mtDNA. One of the skeletons probably carried mtDNA haplogroup N*. N is the ancestor of all the common European maternally-inherited mtDNA haplogroups. It arose among the first group of modern humans to leave Africa. It is so old that it is seldom found in living people today without subsequent mutations.23D. Caramelli et al, Evidence for a genetic discontinuity between Neandertals and 24,000-year-old anatomically modern Europeans, Proceedings of the National Academy of Sciences of the United States of America, vol. 100, no. 11 (2003), pp. 6593-6597; Caramelli, D. et al., A 28,000 years old mtDNA sequence differs from all potentially contaminating modern sequences, PloS ONE, vol. 3, no. 3 (2008): e2700. doi:10.1371/journal.pone.0002700.

The relatively mild climate allowed people to roam far to the north. At Sunghir, on the East European Plain outside Vladimir, a group of reindeer hunters camped about 25,000 years ago. No doubt they had followed the herds as they moved north in the summer. These hunters were tall and massively broad-shouldered. We can picture them clad in skins and furs. Along with reindeer they hunted mammoth and arctic fox, whose pelt would make warm clothing and bedding. The astonishing thing about their dress was the degree of ornamentation. It must have taken many patient hours to create the thousands of ivory beads which were sewn on to every item of clothing, to judge by the finds in graves. Such tailored clothing was made possible by the invention of the needle about 40,000 years ago, made in these early days from bone. The burial at Sunghir that has excited most attention is that of two children. A boy about 13 years old and a girl about 10 years old were laid in the same grave. They were probably brother and sister, since they carried the same mtDNA. The richness of their dress and grave goods makes theirs the most spectacular of burials from this period. Each child had an outfit decorated with about 5,000 tiny ivory beads, as well as ivory pins, pendants and animal carvings, among them a simple image of a horse. The wild herds of horses on the plains added variety to the hunting.24J.F. Hoffecker, Desolate landscapes: Ice-Age settlement in Eastern Europe (2002), p.151, 183; V. Formicola, From the Sunghir children to the Romito dwarf: aspects of the Upper Paleolithic funerary landscape, Current Anthropology, vol.48, no. 3 (2007), pp. 446-452; T.I Alexeeva, et al., Homo Sungirensis, Upper Palaeolithic man: ecological and evolutionary aspects of the investigation (2008); I. Gilligan, The prehistoric development of clothing: archeological implications of a thermal model, Journal of Archaeological Method and Theory, vol. 17 (2010), pp.15–80.

Further south, flax fibres have been found in a cave used by man in Georgia dating back 30,000 years. People probably used them to make linen and thread for clothes and cords.25E. Kvavadze et al, 30,000-Year-Old Wild Flax Fibers, Science, vol. 325, no. 5946 (11 September 2009), p. 1359.

The Ice Age

Climate change almost evicted the first Europeans. As the last glacial gripped Europe, glaciers advanced, while plants and animals retreated. Eventually ice sheets miles thick covered much of northern Europe. Even before this, the population of Europe was tiny by comparison with today. It has been estimated from archaeological data at 4400–5900 inhabitants. 26J.-P. Bocquet-Appel, P.-Y. Demars, L. Noiret and D. Dobrowsky, Estimates of Upper Palaeolithic meta-population size in Europe from archaeological data, Journal of Archaeological Science, vol. 32, no. 11 (2005), pp. 1656-1668. The climatic clampdown reduced Europeans to the status of an endangered species.

It was not just areas of the globe close to the poles that suffered. Levels of rainfall dropped, expanding deserts and reducing forests everywhere. On every continent mankind was squeezed into shrunken zones that could support human life. During the iciest period, the expanded Sahara cut off any escape route from Asia to the tropical refuge in western Africa. Meanwhile the Himalayan Mountains and swathes of desert and semi-desert surrounded a reduced rainforest in the north-east of the Indian subcontinent.

Europeans could take refuge in southern Europe and Asia Minor. Archaeologists find that as settlements disappeared in Northern Europe, they increased three-fold in Cantabrian Spain. This area was clearly a major refuge. Italy and the Balkans also remained partly forested. In a belt to the north of the forested areas, the steppe offered rich grazing in summer for animal herds. Some hunter-gatherer bands developed a pattern of wintering in the sheltered valleys of Lower Austria and Moravia, but moving 170 km or more into the steppe in summer, to follow the herds. A similar pattern of summer hunting on the steppe and tundra is seen right across Siberia. Forested areas around the Black and Caspian Seas may have provided winter refuges for some of these hunters.27H.T. Wright, Humanity at the Last Glacial Maximum: A cultural crisis, chap. 6 in P. N. Peregrine, I. Peiros and M. Feldman, Ancient Human Migrations: A multidisciplinary approach (2009); P.E. Tarasov et al., Last glacial maximum biomes reconstructed from pollen and plant macrofossil data from northern Eurasia, Journal of Biogeography, vol. 27 (2000), pp. 609-620.

The Younger Dryas

Though the climate gradually improved after the ice sheets reached their maximum extent around 20,000 years ago, the ice warrior made one more attack. The big freeze came with devastating speed. The first warning was a period in which the climate oscillated from warm to cold. Then in a single year, 12,679 years ago, northern Europe went from a temperate climate to glacial conditions.28A.Brauer, G.H. Haug, P. Dulski, D. M. Sigman and J.F.W. Negendank, An abrupt windshift in western Europe at the onset of the Younger Dryas cold period,Nature Geoscience vol. 1 (2008), pp. 520-523; Mini ice age took hold of Europe in months, New Scientist, no 2734 (11 November 2009). Once more Europeans were threatened with extinction, but managed to survive, though in some cases by ceasing to be Europeans. It appears that some took advantage of the lower sea level to flee across the Straits of Gibraltar to North-West Africa, contributing mtDNA haplogroup U5b1b to the present-day Berbers.29M. Alcaraz Castano, El Ateriense del Norte de África y el Solutrense peninsular: ¿contactos transgibraltareños en el Pleistoceno Superior?, Munibe (Antropologia-Arkeologia), no. 58 (2007), pp.101-126; 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

Did people fleeing Europe feel that they were changing their identity? Who knows? We don’t know when the concept of Europe arose. Europe is not a separate landmass. Nor is Africa. Yet the crossing-point from Africa to Asia being narrow, it makes sense to think of them as two different continents. Quite why Europe and Asia, which form one landmass, are perceived as separate is less clear. Geographically Europe might be better classed as a subcontinent of Asia. The boundary was the Don River in antiquity, but is now the mountain range of the Urals.30K. Wilson and J. van der Dussen (eds.), The History of the Idea of Europe (1995), p. 2. People have moved across that boundary, and across the Mediterranean, from time immemorial, so Europeans are closely related to their nearest neighbours.

Mesolithic hunters and fishermen

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).

With so much water locked into glaciers, the sea level was still low enough at the start of this period for people to be able to walk to Britain across a land-bridge from Continental Europe. From there they entered Ireland – the first humans to do so. However the British Isles and Scandinavia were initially less appealing than more southerly regions of Europe; signs of human activity are sparse so far north until after 7,000-6,000 years ago. It is sheer chance that the earliest evidence of carpentry in Europe comes from a lake-side site in England. The boggy conditions at Star Carr in Yorkshire preserved remnants of a brushwood platform over reed-beds at the lake edge.2B. Weninger et al., A radiocarbon database for the Mesolithic and early Neolithic in Northwest Europe, chapter 9 in P. Crombé et al. (eds.), Chronology and evolution within the Mesolithic of North-West Europe: Proceedings of an international meeting, Brussels May 30th–June 1st 2007 (2009), pp. 143-176; C. Conneller et al., The temporality of the Mesolithic landscape: new work at Star Carr, ibid, pp. 77-94.

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.3M.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.4F. 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.5K. 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.6M. 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.7T. 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.8J.R. Huyghe, A genome-wide analysis of population structure in the Finnish Saami with implications for genetic association studies, European Journal of Human Genetics, vol. 19, no. 3 (March 2011), pp. 347-52..

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.9M. 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.10R. 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.11O. 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 corralled by man in the more distant past.12K.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.

European Mesolithic Y-DNA

Since it is far more difficult to extract Y-DNA from ancient remains than mtDNA, we still have no Y-DNA from this period. 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. 13Dating 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.14D. 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.15B. 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 Low Countries about 5000 BC.16P. 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.17P.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.

Another Y-DNA haplogroup which may have spread up the Volga in the Mesolithic period is R1a1a (M17). This seems to have been carried by hunter-gatherers on the eastern border-lands of Europe, around the Urals. Its big expansion occurred from there in the Copper Age, but some earlier R1a1a adventurers may have left northern traces in R1a1a*, R1a1a1* and a subclade of the latter defined by the L664 marker.

European Mesolithic mtDNA

With mtDNA we are on firmer ground. U5 is one of the oldest European mtDNA haplogroups.18P. 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.19B. 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. 20B. 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. 21L 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.22H. 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.”23O. 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 U5b1c2.24H. 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.

The genetic debate

In the days before we had much ancient DNA, it was easy to assume that the distribution of mitochondrial (mtDNA) haplogroups we see today is largely due to the Mesolithic recolonisation of Europe. Bryan Sykes argued this case in popular and scholarly works at the start of the present millennium. Initially the only mtDNA haplogroup that he linked to the later spread of Neolithic farmers into Europe from the Near East was J. He calculated that 80% of native Europeans could trace their ancestry to European hunter-gatherers. 1B. Sykes, The Seven Daughters of Eve (2001); M. Richards, B. Sykes et al, Tracing European founder lineages in the Near Eastern mtDNA pool, American Journal of Human Genetics, vol. 67, no. 5 (2000), pp. 1251-76. At around the same time a similar conclusion was drawn from Y-DNA.2O. Semino et al, The genetic legacy of paleolithic Homo sapiens sapiens in extant Europeans: a Y chromosome perspective, Science, vol. 290 (2000), pp. 1155-59; and see J.T. Lell and D.C. Wallace, The peopling of Europe from the maternal and paternal perspectives, American Journal of Human Genetics, vol. 67, no. 6 (2000), pp.1376–1381.

Could this be true? It seemed a triumphant vindication of those who had long held that agriculture was spread culturally, rather than by mass migration. Television programmes took up the tale. The Channel 4 series The Face of Britain (2007) and RTE’s The Blood of the Irish (2009) presented the Celts of Britain and Ireland as descended from Mesolithic colonisers from the Continent. But by the time they were shown, population genetics had moved on.

Clines and waves

Geneticists realised that where they find the greatest genetic diversity of a haplogroup is most likely to be its point of origin, since the longer a lineage has been in a place, the longer it has had to accumulate mutations. That changed the picture. The Mesolithic re-colonisers moved from south to north, but the dominant Y-DNA genetic cline in Europe is from east to west. That suggests that later migrations bringing farming and/or metallurgy were more important in creating the present paternal lineages of Europe.3B. Arredi, E.S. Poloni, C. Tyler-Smith, The peopling of Europe, in M. Crawford (ed.), Anthropological Genetics: Theory, method and applications (2007).

A study by Patricia Balaresque and colleagues relied heavily upon upon one type of calculation of genetic diversity – that based on STRs within the Y-Chromosome. They concentrated upon one particular subclade of R1b, concluding that it could be linked to the spread of farming into Europe.4P. Balaresque et al., A predominantly neolithic origin for European paternal lineages, PLoS Biology, vol. 8, no. 1 (2010). Their approach was dismantled the following year by a reassessment which found no particular differences in diversity.5G.B.J. Busby et al., The peopling of Europe and the cautionary tale of Y chromosome lineage R-M269, Proceedings of the Royal Society B: Biological Sciences, published online before print, August 24, 2011. Yet Kristian Herrera and colleagues show the variance of the whole R1b haplogroup to be clearly highest in Western Asia.6K. J. Herrera et al., Neolithic patrilineal signals indicate that the Armenian plateau was repopulated by agriculturalists, European Journal of Human Genetics, advance online 16 November 2011. Only by ignoring the Asian data can one see little variance.

Not that diversity is always a reliable guide. A present-day population could have acquired Y-DNA diversity from diverse waves of immigrants, or by a mass movement from place A to place B, taking with it the full diversity that it had at place A. So diversity is most convincing when supported by other evidence. In this case the chain of SNP mutations within R1b runs from east to west, with those which occurred earliest being most prevalent towards the east. On the global level, it is the chain of mutations within mtDNA and Y-DNA that enabled geneticists to work back to a genetic Adam and Eve in Africa at the root of the human family tree. The people closest to this Adam and Eve exhibit the highest genetic diversity in the world across the genome.7M. Melé et al., Recombination gives a new insight in the effective population size and the history of the Old World human populations, Molecular Biology and Evolution (online 1 September 2011 ahead of print); J. Xing et al., Toward a more uniform sampling of humangenetic diversity: a survey of worldwide populations by high-density genotyping, Genomics, vol. 96, no. 4 (October 2010), pp. 199-210; G. Laval et al., Formulating a historical and demographic model of recent human evolution based on resequencing data from noncoding regions, PLoS ONE,vol.5, no. 4 (2010): e10284; 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; R.N. Gutenkunstet al., Inferring the joint demographic history of multiple populations from multidimensional SNP frequency data, PLoS Genetics, vol. 5, no. 10 (1 October 2009), pp. 1-11; M. DeGiorgio, M. Jakobssonet and N.A. Rosenberg,Explaining worldwide patterns of human genetic variation using a coalescent-based serial founder model of migration outward from Africa, Proceedings of the National Academy of Sciences of the United States of America, vol. 106, no. 38 (Sep 2009), pp. 16057-16062; O. Deshpande, S.Batzoglou, M.W. Feldman and L.L. Cavalli-Sforza, A serial founder effect model for human settlement out of Africa, Proceeding of the Royal Society B: Biological Sciences, vol. 276 (2009), pp. 291-300; I. Ionita-Laza, C.Lange and N.M. Laird, Estimating the number of unseen variants in the human genome, Proceedings of the National Academy of Sciences of the United States of America, vol. 106, no. 13 (March 2009), pp. 5008-5013; J. Z.Li et al, Worldwide human relationships inferred from genome-wide patterns of variation, Science, vol. 319, (2008), pp. 1100-04; G. Hellenthal, A. Auton, D. Falush, Inferring human colonization history using a copying model, PLoS Genetics, vol. 4, no. 5 (May 2008); Jakobsson, M. etal, Genotype, haplotype and copy-number variation in worldwide human populations, Nature, no. 451 (21 February 2008), pp. 998-1003; R.Klein, Out of Africa and the evolution of human behavior, Evolutionary Anthropology: Issues, News, and Reviews, vol. 17, no. 6 (2008), pp. 267- 281; Q. Ayub et al., Reconstruction of human evolutionary tree using polymorphic autosomal microsatellites, American Journal of Physical Anthropology, vol. 122 (2003), pp. 259-268.

Jacques Chiaroni and his colleagues described the way that different kinds of genetic spread leave characteristic patterns. They have christened one the Surfing Effect. A mutation that occurs in the wave front of an expanding population will have an advantage. It will have a better chance of becoming predominant within the breeding group, because that is where the migrating population is smallest. A successful mutation will surf the wave and end up at saturation level where the expanding population meets a geographical barrier. A good example is the commonest Y-DNA haplogroup of western Europe – R1b1b2 (M269). It flooded over Europe from the east, spawning subclades as it went, until it was stopped by the Atlantic Ocean. On the Atlantic seaboard it reaches saturation level.8J. 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; N.M Myres et al., A major Y-chromosome haplogroup R1b Holocene era founder effect in Central and Western Europe, European Journal of Human Genetics, vol. 19, no. 1 (January 2011), pp. 95–101. How this might work in practice is demonstrated by a study of a population expansion in historic times. Genealogical analysis showed that majority of the present population of Saguenay Lac Saint-Jean in Quebec can be traced back to ancestors having lived directly on or close to the wave front of 17th-century expansion.9C. Moreau et al., Deep human genealogies reveal a selective advantage to be on an expanding wave front, Science, published online November 3 2011.

Refining by subclade

Early studies painted with a broad brush. Only a few mtDNA haplogroups had been discovered at that time; each was given a letter to identify it. Then researchers attempted to make sense of their distribution. The gradual process of breaking these parent groups down into subclades has created a more subtle picture. For example mtDNA T had been seen as Palaeolithic, yet sub-haplogroup Tl is one of the clearest examples of a lineage cluster with a much earlier expansion in Western Asia than in Europe.10K. 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), pp. 454-5. Both T and K have been found in the DNA of early farmers in the Levant and Europe.11E. Fernández, et al., Mitochondrial DNA genetic relationships at the ancient Neolithic site of Tell Halula, Forensic Science International: Genetics Supplement Series, vol.1, no. 1 (2008), pp. 271–273; W. Haak et al, Ancient DNA from the First European Farmers in 7500-Year-Old Neolithic Sites, Science, vol. 310, no. 5750 (2005), pp. 1016-1018. Similarly, a closer look at mtDNA H reveals its complexity. H itself was born in the Near East and spread into Europe. H3 is largely limited to Europe and North Africa. Both H1 and H3 have their densest distribution in Iberia. From this it was argued that these two subclades spread from the Franco-Cantabrian glacial refuge as the climate warmed in the Mesolithic period.12A. Achilli et al, The molecular dissection of mtDNA Haplogroup H confirms that the Franco-Cantabrian glacial refuge was a major source for the European gene pool, American Journal of Human Genetics, vol. 75 (2004), pp.910–918; L 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. This would be highly significant for the peopling of Europe. Not only is H itself the predominant haplogroup in Europe, carried by almost half of the population, but its commonest subclade is H1. It represents about 30% of H and 13% of the total European mtDNA pool. Yet breaking down H1 itself into subclades revealed that some are barely present in Iberia. H1a and H1b are densest in Eastern Europe (see Slavic genetic markers). Even more significantly, H1 and H3 have a low diversity in Cantabrian Spain and in particular among the Basques. Instead the highest diversity and allelic richness of H1 and H3 in Europe is found in north-eastern and north-central regions, while the Near East has the greatest overall diversity of H1, and North Africa that for H3. The once-popular idea of the Basques as the source population for most of modern-day Europe is not supported by this closer examination. 13O. 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; H. Ennafaa et al., Mitochondrial DNA haplogroup H structure in North Africa, BMC Genetics, vol.10 (2009), no. 8.

Instead it suggests that H1 and possibly H3 arrived in Europe with the first farmers. Against this is one sample of H1b reported in Mesolithic DNA in Portugal.14H. 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. However this comes from a study carried out some years ago, when ancient DNA studies were less reliable. The parent and grandparent of H (HV and R0) also have their origins in Western Asia. HVa, R0a, U7 and U3 have frequency peaks there, so their appearance in Europe may be another clue to the spread of farming.15A. Achilli et al, Mitochondrial DNA variation of modern Tuscans supports the Near Eastern origin of Etruscans, American Journal of Human Genetics, vol. 80, no. 4 (2007), pp. 759–768.

H5* is most frequent and diverse in the western Caucasus, while H5a has a stronger European distribution. H5a is thought to be only 7000-8000 years old, so its wide, though low, spread over Europe suggests that significant migration took place even after the initial spread of farming.16U. Roostalu et al, Origin and expansion of haplogroup H, the dominant human mitochondrial DNA lineage in West Eurasia: the Near Eastern and Caucasian perspective, Molecular Biology and Evolution, vol. 24, no. 2 (2007), pp. 436-448.


However debate has swung to and fro over the dating of haplogroups. Geneticists in recent years have often used Zhivotovsky’s evolutionary effective dating method for Y-DNA, which adjusts the calculated pedigree (genealogical) mutation rate, since in some populations the latter produced unexpectedly late dates.17L.A. Zhivotovsky et al., The effective mutation rate at Y chromosome short tandem repeats, with application to human population-divergence time, American Journal of Human Genetics, vol. 74 (2004), pp. 50–61; L. A. Zhivotovsky, Difference between evolutionarily effective and germ line mutation rate due to stochastically varying haplogroup size, Molecular Biology and Evolution, vol. 23, no. 12 (2006), pp. 2268-2270. Unfortunately this ad-hoc adjustment seems generally misapplied, producing dating estimates two or three times too old. For example Marcin Woźniak and colleagues point out that the pedigree mutation rate for R1a1a1g [M458] is more consistent with the archaeological record for the Slavs.18M. Woźniak et al., Similarities and distinctions in Y Chromosome gene pool of Western Slavs, American Journal of Physical Anthropology, vol. 142, no. 4 (2010), pp. 540-548. A study of the Caucasus region found that genealogical estimates gave a good fit with the linguistic and archaeological dates, while the evolutionary effective rates fell far outside them.19O. Balanovsky et al., Parallel Evolution of Genes and Languages in the Caucasus Region, Molecular Biology and Evolution, published online ahead of print 13 May 2011. Another approach is directly genealogical. Both surnames and Y-DNA haplogroups are passed down in the male line. A group of men with a surname of the same origin should have a common ancestor at the time of surname development. One study found that they mainly did, using a mutation rate similar to the genealogical rather than the evolutionary.20T.E. King and M.A. Jobling, Founders, drift, and infidelity: the relationship between Y chromosome diversity and patrilineal surnames, Molecular Biology and Evolution, vol. 26, no. 5 (May 2009), pp.1093-1102.

However problems with the pedigree rate remain. Dating estimates will vary according to which microsatellite loci are used, since some mutate faster than others. New approaches take this factor into account, but are only very recently reaching publication.21G. B. J. Busby and C. Capelli, Microsatellite choice and Y chromosome variation: attempting to select the best STRs to date human Y chromosome lineages, paper read at the European Human Genetics Conference at Amsterdam May 28 – 31 2011; G. B. J. Busby et. al., The peopling of Europe and the cautionary tale of Y chromosome lineage R-M269, Proceedings of the Royal Society B: Biological Sciences, published online before print, August 24, 2011; C. Burgarella and M. Navascués, Mutation rate estimates for 110 Y-chromosome STRs combining population and father–son pair data, European Journal of Human Genetics, vol. 19 (2011), pp. 70–75; W. Shi et al., A worldwide survey of human male demographic history based on Y-SNP and Y-STR data from the HGDP-CEPH populations, Molecular Biology and Evolution, vol. 27, no. 2 (2010), pp. 385-393. Attempts to calibrate the human mtDNA clock are no less controversial.22B. M. Henn et al., Characterizing the time dependency of human mitochondrial DNA mutation rate estimates, Molecular Biology and Evolution, vol. 26 (2009), no. 1, pp. 217-230; Endicott et al., Evaluating the mitochondrial timescale of human evolution, Trends in Ecology and Evolution vol. 24, no. 9 (2009), pp. 515-521; S. Rosset et al., Maximum-likelihood estimation of site-specific mutation rates in human mitochondrial DNA from partial phylogenetic classification, Genetics, vol. 180 (November 2008), pp. 1511–1524; M.P. Cox, Accuracy of molecular dating with the rho statistic: deviations from coalescent expectations under a range of demographic models, Human Biology, vol. 80, no 4 (2008), pp.335-357; C.D. Millar et al., Mutation and evolutionary rates in Adélie Penguins from the Antarctic, PLoS Genetics vol. 4, no. 10 (October 2008). To cap the confusion, a recent study found substantial variance in sex-specific mutation rates between families,23D.F. Conrad et al., Variation in genome-wide mutation rates within and between human families, Nature Genetics, Published online 12 June 2011 ahead of print. throwing a cloud of doubt over all dating estimates.

Modern versus ancient DNA

Another problem arises from the common assumption that a modern population reflects an ancient local gene pool. That would be convenient. It is a lot easier to obtain blood or saliva samples from the living, than to retrieve DNA from skeletons. Yet several studies of ancient DNA (aDNA) have discovered no relationship between ancient people and those who now occupy the same area.24L. Melchior, Evidence of Authentic DNA from Danish Viking Age Skeletons Untouched by Humans for 1,000 Years, PLoS ONE 3(5): e2214; Töpf et al., Ancient human mtDNA genotypes from England reveal lost variation over the last millennium, Biology Letters, vol. 3, no. 5 (2007), pp. 550–553; Price et al., The impact of divergence time on the nature of population structure: an example from Iceland, PLoS Genetics vol 5, no. 6 (2009): e1000505; S. Guimaraes et al., Genealogical discontinuities among Etruscan, Medieval and contemporary Tuscans, Molecular Biology and Evolution, published online on July 1, 2009: doi:10.1093/molbev/msp126; S.E. Smith et al., Inferring population continuity versus replacement with aDNA: a cautionary tale from the Aleutian Islands, Human Biology, vol. 81, no. 4 (August 2009); B. Bramanti et al, Genetic Discontinuity Between Local Hunter-Gatherers and Central Europe’s First Farmers, Science, (published online September 3, 2009): DOI: 10.1126/science.1176869; 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. For example the greatest modern density and diversity of H5* appears in the Western Caucasus, suggesting that it spread from there. Yet the Caucasus attracted Neolithic farmers from the Near East and numerous later waves of incomers, which might confuse the picture. H5 was present in Neolithic Syria, 25E. Fernández et al., Mitochondrial DNA genetic relationships at the ancient Neolithic site of Tell Halula, Forensic Science International: Genetics Supplement Series, vol.1, no. 1 (2008), pp. 271–273. so the Near East is the more likely point of origin.

To create firm links between haplogroups and long-gone cultures there is no substitute for aDNA. Yet we should be cautious. The problem of contamination with modern DNA bedevilled ancient DNA study in its early years. Little weight can be placed on the results from early studies unless they have been replicated. More recent studies usually report their methodology in reassuring detail. However samples tend to be too small for statistical significance. Conclusions about the entire population of Europe cannot be drawn from a handful of individuals from the same grave, very probably related. Far greater statistical weight can be placed on those studies which sample more widely and achieve a higher number of results, and on the collective results of multiple studies. A body of knowledge is gradually building up, which has started to suggest the dates that particular haplogroups first arrived in Europe.

However, even if we can prove by ancient DNA that a particular haplogroup had arrived in a particular place by a given date, that doesn’t rule out its arrival again in later waves of incomers. Indeed it could be arriving again right now on the latest plane. People will move about! This takes us back to a previous point. We need to discriminate between various subclades of the parent haplogroups. Ancient DNA tested recently to a high degree of resolution is the most valuable.

It might seem a statement of the obvious that present populations descend from ancient ones. However at least one author has put the cart before the horse and attempted to guess the origin of ancient DNA from that of present people. This is singularly foolish. The present spread of haplogroups tells a story of the migrations of the descendants of those ancient people. It is equally interesting if we find that they had no descendants. Many lineages will have died out simply by chance. Yet we may find some cultures, classes or individuals more prolific in their offspring.

Genome-wide vs sex-specific

Mapping the first human genome was a huge project, which took over a decade and cost three billion US dollars. Today both the time and cost of sequencing have dropped so sharply that a subsequent project to sequence 1000 genomes is well under way, while the International HapMap Project has reached a similar stage with over 1000 genomes. Both make it possible to compare full genomes from different ethnic groups, and additional genomes are available from regional projects. So genome-wide population comparisons are increasingly popular. They give a broad picture of the genetic make-up of a population and its affinities.

However these can only tell us if present-day population X has a similarity with present-day population Y. They cannot tell us how this arose. Sex-specific markers remain the clearest guide to migration, since mtDNA and Y-DNA are passed down from parent to child without recombination. The accumulation of spontaneous mutations along these lineages provide clear evidence of direction of flow. They also make it possible to detect sex-biased migration, for example male-dominated bands of traders taking local wives.


It will take time to tease out the many strands of migration and mobility that have created the present genetic kaleidoscope in Europe. Over the last decade there has been rapid progress. Where once there were simple trees for mtDNA and Y-DNA with a few bare branches, now there are bushy structures with a multitude of twigs, kept up to date online at ISOGG YDNA tree and PhyloTree(mtDNA). The breakdown into smaller and smaller subclades is crucial in distinguishing between different origins. Improved techniques of extracting ancient DNA give hope of driving down its cost and making the results more reliable. The quest for ancient DNA is becoming ever more ambitious, with entire genomes extracted fo

Who do you look like?

When a new baby arrives, relatives love to spot resemblances. She has her father’s eyes, her mother’s mouth, grandmother’s nose and so on. So it seems logical, when trying to guess more distant ancestry, to home in on appearances. If you read that the Vikings were tall and blond, and you are short and dark, you feel that you can rule out any Vikings in your ancestry. But is it that simple? Traits such as height, facial features and colouring are inherited autosomally. That means that the code for them is found on the 22 pairs of chromosomes that males and females share. (The other pair is two X-chromosomes for a woman, and an X and a Y for a man.) Genetic coding from both parents recombines into a new set of chromosomes for a new baby. Suppose you have one tall, fair-haired great-grand-parent and seven who were short and dark. Would you expect to be tall and blond?

The University of California: Berkeley provides a quick, animated guide to Sex and Genetic Shuffling: The Details. Because of the re-mixing in every generation of autosomal inheritance, it is too complicated to use in tracking human migration. That is why population geneticists use mitochondrial DNA, which passes down unchanged from mother to child, and Y-DNA, which passes unchanged from father to son, except for the occasional spontaneous mutation in both cases. These haplogroups have nothing to do with the inheritance of height, or colouring or other traits.1M.Jobling, M.E. Hules and C. Tyler-Smith, Human Evolutionary Genetics (2004), chapter 2.


Colouring leaps to mind when Europeans think about appearances. Whereas most of the peoples of the planet have uniformly black hair and dark eyes, people with origins in Europe, Western Asia and North Africa have a wider range of colouring. Why is that? Dark skin protects people from ultra-violet light, but for that very reason makes it more difficult for their skin to synthesise vitamin D, essential for bone growth and activation of the immune system. Pale people in sunny places are at higher risk of skin cancer and folate deficiency, while dark people in cooler climes are prone to problems of low vitamin D. So it has long been supposed that the range of skin colours we see today arose through natural selection.2M.Jobling, M.E. Hules and C. Tyler-Smith, Human Evolutionary Genetics (2004), section 13.3 and box 13.4 provide an introduction to the subject, updated by A. Juzeniene et al., Development of different human skin colors: A review highlighting photobiological and photobiophysical aspects, Journal of Photochemistry and Photobiology B: Biology, vol. 96, no. 2 (3 August 2009), pp. 93-100; M. Rode von Essen et al, Vitamin D controls T cell antigen receptor signaling and activation of human T cells, Nature Immunology (online ahead of print 7 March 2010).

How does that work? Since our genes code for functions, a mutation generally results in loss of function. The code has become faulty. So it is with the mutations that cause paler colouring. Mutations in specific genes prevent the body from producing melanin – the most important pigment influencing skin and hair colour. In places drenched in ultra-violet light, such mutations would be a disadvantage. People carrying them could die before they had a chance to reproduce at all. Or they could have fewer surviving offspring. But in cloudier climes such mutations give their bearers an advantage, so gradually they would gain ground in a population.

Interestingly it seems that the Neanderthals could have been ahead of us in developing pale skin and red hair (though by a different genetic route than Homo sapiens), as part of a range of pigmentation.3C. Lalueza-Fox et al., A melanocortin 1 receptor allele suggests varying pigmentation among Neanderthals,Science (October 25, 2007); C. C. S. Cerqueira et al., Predicting homo pigmentation phenotype through genomic data: From neanderthal to James Watson, American Journal of Human Biology, published online 12 March 2012 ahead of print. Neanderthals spent millennia under the often cloudy skies of Europe. But why the red hair? Pigmentation is a puzzle, and we may not have all the pieces yet. There are at least 11 genes involved. So far it looks as though one biological path to paler skin does not interfere with the ancestral dark hair and eyes, while another throws up red hair as a side effect, with another you get blond hair, and mixtures give in between shades. 4F. Liu et al., Digital quantification of human eye color highlights genetic association of three new loci, PLoS Genetics, vol. 6, no. 5 (2010), e1000934; J. Mengel-From et al., Genetic determinants of hair and eye colour in the Scottish and Danish populations, BMC Genetics vol. 10 (December 2009) 88; R.A. Sturm, Molecular genetics of human pigmentation diversity, Human Molecular Genetics, vol. 15, no. 18(R1) (April 2009), R9-17; W. Branicki, Interactions between HERC2, OCA2 and MC1R may influence human pigmentation phenotype, Annals Human Genetics, vol. 73, no. 2 (Mar 2009), pp.160-70. By giving a statistical weighting to 13 single or compound genetic markers from those 11 genes, it is possible to predict hair colour with a high degree of accuracy.5W.Branicki et al., Model-based prediction of human hair color using DNA variants, Human Genetics, online 3 January 2011 before print. There is a completely separate gene for blonde hair which crops up among some Melanesians.6E.E. Kenny et al., Melanesian blond hair is caused by an amino acid change in TYRP1, Science, vol. 336, no. 608 (4 May 2012), p. 554.

The rainbow look of Europeans suggests strong selection for a cold climate. It may seem a logical deduction that the process began as man left Africa. Yet scientists calculate that a new allele causing paler skin cropped up on gene SLC24A5 around 5300 to 6000 years ago. It is nicknamed the golden gene, as it also causes golden stripes in zebrafish.7R.L. Lamason et al, SLC24A5 affects pigmentation in zebrafish and man, Science vol. 310 (2005), pp.1782-1786; A. Gibbons, European skin turned pale only recently, gene suggests, Science, vol. 316, no. 5823 (20 April 2007), p.364. While the golden gene and two others (SLC45A2 and KITLG) cause most of the paleness of Europeans and their relatives, East Asians have their own colour-drainer (His615Arg in OCA2), as well as KITLG, showing independent evolution after their ancestors moved deep into Asia. However the similar distribution of alleles in the KITLG gene within Western Eurasians and East Asians suggests that some selection for paler skin is older. 8M. Edwards et al., Association of the OCA2 polymorphism His615Arg with melanin content in East Asian populations: further evidence of convergent evolution of skin pigmentation, PLoS Genetics, vol. 6, no.3 (March 2010); H.L. Norton, Genetic evidence for the convergent evolution of light skin in Europeans and East Asians, Molecular Biology and Evolution, vol. 24 (2007), pp.710-722.

So the evolution of colour variation may have begun in Africa, but took a leap in the era of the earliest farmers, probably because their diet was lower in vitamin D. Early Europeans boosted their vitamin D intake by eating fatty fish. Salmon bones, fish hooks, and paintings of salmon, trout, and pike have been found in caves they occupied.9G.E. Adán et al., Fish as diet resource in North Spain during the Upper Paleolithic, Journal of Archaeological Science, vol. 36, no. 3 (March 2009), pp. 895-899. The start of farming in the Near East created a higher reliance on cereals. The range of the most recent mutations for lighter colouring suggests that they were first spread by the early farmers. They are found everywhere that farmers migrated from the Near East: Europe, Western Asia and North Africa. For example SLC24A5 is found in 60-70% of the population in Tunisia and Morocco.10G. Lucotte et al., A Decreasing Gradient of 374F Allele Frequencies in the Skin Pigmentation Gene SLC45A2, from the North of West Europe to North Africa, Biochemical Genetics, vol. 48, nos. 1-2 (2010), pp. 26-33. The present range of hair colouring appears to be shown in the rock painting (above) of around 3000 BC from the Tassili n’Ajjer plateau, Algeria.11S. di Lernia and M. Gallinaro, The date and context of Neolithic rock art in the Sahara: engravings and ceremonial monuments from Messak Settafet (south-west Libya), Antiquity, vol. 84, no. 326 (December 2010), pp. 954–975.

Worshippers at Sumerian temples could be depicted with either blue or brown eyes. The same is true of ancient Egyptians of the same period. There is no reason to think that blue eyes predominated in these populations, but every reason to accept that they existed. The predominant gene for blue eyes (rs12913832 GG) appeared between 6,000 and 10,000 years ago. Professor Hans Eiberg and his team analysed the DNA of people with blue eyes in Denmark, Turkey and Jordan. All of them had exactly the same DNA sequence covering half the HERC2 gene. That suggests a common ancestor. Eiberg and his colleages surmised that The mutations responsible for the blue eye color most likely originate from the Near East area or northwest part of the Black Sea region, where the great agriculture migration to the northern part of Europe took place in the Neolithic periods about 6–10,000 years ago.12H. Eiberg et al, Blue eye color in humans may be caused by a perfectly associated founder mutation in a regulatory element located within the HERC2 gene inhibiting OCA2 expression, Human Genetics, vol.123, no 2 (Mar 2008), pp. 177-87. Interestingly there are two other mutations which produce blue eyes and are found world-wide (apart from central/southern Africa), though also concentrated in northern Europe. These presumably are older. The rs12913832 GG allele shows strong signals of positive selection in Northern Europe.13M.P. Donnelly et al., A global view of the OCA2-HERC2 region and pigmentation, Human Genetics, online 7 November 2011 ahead of print. What could be the advantage of blue eyes in northern climes? The lower melanin in blue eyes makes it easier to absorb light in winter and so protect against Seasonal Affective Disorder (SAD).14S. Higuchi et al., Influence of eye colors of Caucasians and Asians on suppression of melatonin secretion by light, American Journal of Physiology – Regulatory, Integrative and Comparative Physiology, vol. 292, no. 6 (June 2007), pp. R2352-R2356.

Red hair

There are two types of melanin. Black and brown pigments are formed from eumelanin. Red and yellow result from pheomelanin. Mutations on the MC1R gene, causing loss of only eumelanin, result in yellow or red coat colours in many mammals. Man is no different. Some of these mutations appear in redheads.15Helgi B. Schiöth et al., Loss of function mutations of the human melanocortin 1 receptor are common and are associated with red hair, Biochemical and Biophysical Research Communications, vol. 260, no. 2, (5 July 1999), pp. 488-491. To complicate matters the various red hair alleles on the MC1R gene can have different effects. Some are classed as highly penetrant. Most red-haired individuals (84%) have two highly penetrant alleles (one from each parent), but various other combinations can also result in shades of red.16Niamh Flanagan et al., Pleiotropic effects of the melanocortin 1 receptor (MC1R) gene on human pigmentation, Human Molecular Genetics, vol. 9, no. 17 (2000), pp. 2531-2537; Jonas Mengel-From et al., Genetic determinants of hair and eye colours in the Scottish and Danish populations, BMC Genetics, vol. 10 (2009), 88. Red hair is comparatively rare everywhere today. Partly that is because a person needs to inherit an allele for it from both parents in order to have red hair. A dark-haired person could be quite unaware that he or she is carrying a red hair allele, if he or she has a functioning gene producing eumelanin from the other parent. Having a red-headed child could come as a surprise. Professor Jonathan Rees, Professor of Dermatology at the University of Edinburgh gives a fuller account online for the layman: non-expert guide to the genetics of red hair (and freckles).

Since there are so many different alleles for red hair, it is highly unlikely that they all cropped up within just one population. Red hair is often considered a Celtic characteristic. Think of Queen Boudica’s striking mane of red hair.17Cassius Dio, Roman History, 62.1-12. Certainly Tacitus reported red- heads in Caledonia, long before the inrush of Anglo-Saxons and Vikings. Yet he also saw red hair as a characteristic of the Germani.18Tacitus, Agricola, 11. A recent study proves him right. Yorkshire, Denmark and South-East Wales seem to be harbouring more such alleles than Ireland.19E. Røyrvik, Western Celts? A genetic impression of Britain in Atlantic Europe, in B. Cunliffe and J.T. Koch (eds.), Celtic from the West (2010) pp. 83-106.

So red hair is not restricted to Celts. It is not even restricted to Indo-Euopeans. In the 4th century BC Herodotus described a nomadic, foraging tribe called the Budini with piercing grey eyes and bright red hair, who lived in the forest east of the River Don and 15 days journey north of the Sea of Azov.20Herodotus, The Histories, 4.21-2, 108-9. They sound like the Udmurts, who claim as many red-heads as the Irish. The Udmurt Republic lies in Russia, in the forest zone between two tributaries of the Volga. The Udmurts speak a Finno-Ugric language. They now celebrate their rufosity each September with the Red Festival. The Dutch followed suit with Red Head Day. So far there has been no world-wide scientific sampling to settle the vexed question of which nation really has the highest percentage of red-heads.

Equally tricky is the mummy debate. Some mummies from Egypt and the Tarim Basin have red hair. Is this just the result of fading after death? Or was henna used? Or are some of these mummies the genuine copper-haired article? The answer could be a combination of all three. The earliest mummies are the result of natural preservation in desert sands. The British Museum houses a chap once affectionately nicknamed Ginger, buried about 3400 BC in predynastic Egypt. Could the red locks that gave him his nickname be faded from brown? The mummies now emerging from another predynastic cemetery may provide answers. At Hierakonpolis 43 (c.3600-3400BC), most of the hair found on mummies is very dark brown, showing that this colour can be preserved for millennia, and that it was the most common. Yet male burial no. 79 had natural wavy, red hair.21J. Fletcher, The Secrets of the Locks Unraveled. Nekhen News, Vol 10, (Milwaukee Public Museum 1998), pp. 7-8. Ramsess II (d.1213 BC) used henna to cover his grey hair in old age, but fragments of pigmentation in the roots indicated that it was originally a natural red. If red hair ran in his family, that might explain the occurance of the name Seti among them, red being associated with the dangerous god Set. 22L. Balout and C. Roubet (eds.), La Momie de Ramsès II: Contribution Scientifique a l’Egyptologie 1976-1977, (Paris: Éditions Recherche sur les Civilisations/Muséum National d’Histoire Naturelle/Musée de l’Homme 1985).

Red hair was evidently rare in Ancient Egypt, as it is in North Africa today. But it crops up occasionally among the Berbers of Algeria and Morocco, who seem to be descended from the first farmers to arrive in North Africa, depicted with a range of hair colour in the rock painting at Tassili. There is no reason to imagine that the pharaohs of the 19th dynasty were foreign to Egypt.

The issue of fading pigments does arise with ancient depictions too. The rock painting of the Tassili ladies above is convincing, as it shows a variety of hair colours. The brown has not faded too much. Rock paintings are hard to date precisely, but this may be the earliest image of a red-head. Similarly this painting of Buddhist monks from the Tarim Basin makes a clear distinction between the red-brown hair and beard of the presumably Indo-European (Tocharian) man on the left and the grey or faded black hair of the monk on the right.


It has long been observed that tall people tend to have tall children, but is the explanation genes or lifestyle? Scientists have managed to disentangle the two by twin studies. Identical twins reared in different households tend to be similar in height, but not absolutely identical. About 80 percent of the difference in height between individuals within a population is determined by genetic factors, and scientists are close to pinning down exactly which genes are involved. The rest of the variation can be explained mainly by nutrition. 23M. B. Lanktree et al., Meta-analysis of dense genecentric association studies reveals common and uncommon variants associated with height, The American Journal of Human Genetics, (online 30 December 2010 ahead of print); H.L. Allen et al., Hundreds of variants clustered in genomic loci and biological pathways affect human height, Nature, (advance online publication 29 September 2010); J. Yang et al., Common SNPs explain a large proportion of the heritability for human height, Nature Genetics, (published online ahead of print 20 June 2010); Å. Johansson et al., Common variants in the JAZF1 gene associated with height identified by linkage and genome-wide association analysis, Human Molecular Genetics, vol. 18, no. 2 (2009), pp. 373–380; P.M. Visscher, Sizing up human height variation, Nature Genetics, vol. 40 (2008), pp. 489-490. Yet these figures apply to twins brought up in the same era and generally within the same country, so their diet is unlikely to be dramatically different. What happened when people shifted from hunting to farming? It meant a huge change in diet from one heavy on meat to one heavy on cereals. Archaeologists in Europe can see the result in the human skeletons they find. The early farmers were shorter and slighter than their hunting forebears. Contributory factors may have been higher fertility and early weaning among the settled farmers. Later dairy farming created a cheap and regular source of protein in milk, raising average heights among pastoralists.24J. Piontek, B. Jerszyńska, S. Segeda, Long bones growth variation among prehistoric agricultural and pastoral populations from Ukraine (Bronze era to Iron age), Variability and Evolution, vol. 9, (2001), pp. 61-73; A. Mummert et al., Stature and robusticity during the agricultural transition: Evidence from the bioarchaeological record, Economics and Human Biology, vol. 9, no. 3 (July 2011), pp. 284-301. In North America, European colonists encountered nomadic buffalo-hunters on the Great Plains. Systematic measurements by Franz Boas in 1892 showed that these Native Americans were the tallest people in the world at that time for whom we have reliable statistics. With an average height of 172.2 cm, they were 3 to 11 cm taller than contemporary Europeans, and slightly taller than European-Australians. His work came just in time. The buffalo herds were in decline and a way of life was almost over.25R.H. Steckel and J.M. Prince, Tallest in the World: Native Americans of the Great Plains in the Nineteenth Century, American Economic Review, vol. 91, no.1 (March 2001), pp. 287-294.

Another way in which humans vary is the shape of the skull. Normally we only have to think about this if we are selecting a helmet, or a custom-made hat. Crania can be dolichocephalic (long from back to front), mesocephalic (moderate) or brachycephalic (broad). Skull variation caught the attention of pioneers in anthropology. By the pre-war period elaborate classifications of skull types were in use, but gradually unease developed about seeing these as inherited. Could infant head-binding, diet or other environmental factors be more important in determining head-shape? Head-binding has appeared in a number of cultures. The Mangbetu people of the Congo were still elongating the skulls of their infants to the 1950s, so the technique could be observed. The same type of head-shaping was common in the Near East in the 6th and 5th millennia BC. It produced skulls so long that they appear alien.26K.O. Lorentz, Ubaid headshaping, in R.A. Carter and G. Philip, Beyond the Ubaid (2010), pp. 125-148.

However in present populations recent studies suggest that the heritability of craniofacial traits is actually quite high.27A. Jelenkovic, Contribution of genetics and environment to craniofacial anthropometric phenotypes in Belgian nuclear families, Human Biology vol. 80, no.6 (2008), pp.637-654; N. Martínez-Abadías et al, Heritability of human cranial dimensions: comparing the evolvability of different cranial regions,Journal of Anatomy, vol. 214, no. 1, (January 2009), pp. 19-35. So how did these variations arise? There is enough of a correlation between an extremely cold climate and brachycephaly to suggest that natural selection favoured this type of skull in cold conditions, as it minimised heat loss, thanks to the reduced surface/mass ratio. However the distribution of brachycephaly today can mainly be explained by genetic drift. 28L. Betti et al, The relative role of drift and selection in shaping the human skull, The American Journal of Physical Anthropology (2009). So it may be of use in tracing migration. But since these are traits inherited autosomally, they could change over the generations as people mix. This is another Gordian knot that genetics has the potential to slice through. Ancient DNA can tell us with certainty who is descended from whom.

r a few individuals.

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