The Peopling of Europe

Where did the peoples of Europe come from? That thought has sparked curiosity for millennia. Tribes and nations developed origin myths for lack of better knowledge. Much that we would like to know was lost in the mists of prehistory.

The spread of Homo sapiens (The Wellcome Trust Sanger Institute)Scholars in anthropology, archaeology, linguistics and palaeontology have been labouring to shine a light into prehistory for the last 200 years. They have achieved much. Most scholars now accept that modern man emerged in Africa and spread around the world.1R. Klein, Out of Africa and the evolution of human behavior, Evolutionary Anthropology: Issues, News, and Reviews, vol. 17, no. 6 (2008), pp. 267-281. Yet trenchant disagreements remain over many of the particulars. Was agriculture spread through Europe by farming immigrants or by resident hunter-gatherers taking up agriculture? Why at the dawn of history were people from India to Iceland speaking languages of remarkable similarity? Did migrating Neolithic farmers bring with them the prototype of the Indo-European languages? Or did Copper and Bronze Age herders do so? Or can we explain this pattern without migration?

Simplified tree of Indo-European languages by Jack Lynch of Rutgers University. Click to go to his page.More recently the burgeoning field of population genetics has offered hope of finally resolving such wrangles. Within us all we carry evidence of our ancestors. Now that we can begin to read our own code, what stories of our past can it reveal? Ancient population movements can leave a trail in our DNA, pointing to distant relatives we didn't know we had. In fact it was clues from the genes of living people that provided the conclusive evidence not only that Homo sapiens spread out of Africa, but that the most likely route was across Arabia. 2A.Javed et al., Recombination networks as genetic markers in a human variation study of the Old World, Human Genetics, published online 18 October 2011 ahead of print; M. Melé et al., Recombination gives a new insightin 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 human genetic diversity: a survey of worldwide populations by high-density genotyping, Genomics, vol. 96, no. 4 (October 2010), pp. 199-210; G. Lavaletal., 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.Gutenkunst et al., Inferring the joint demographic history of multiple populations from multidimensional SNP frequency data, PLoSGenetics, vol. 5, no. 10 (1 October 2009), pp. 1-11; M. DeGiorgio, M.Jakobsson 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 foundereffect 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 ofthe 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 (May2008); Jakobsson, M. et al, Genotype, haplotype and copy-number variation inworldwide human populations, Nature, no. 451 (21 February 2008),pp. 998-1003; Q. Ayub et al., Reconstruction of human evolutionary tree using polymorphic autosomal microsatellites, American Journal of Physical Anthropology, vol. 122 (2003), pp. 259-268.

The eagerness with which some have rushed to popularise and commercialise their conclusions is understandable, but it is a prescription for confusion in this fast-moving field. Yesterday's ideas may reach television viewers just as they are being overturned. Commercial genetic testing is precariously balanced on the cutting edge of science. By the time a big genetic research project gets off the ground, the hypothesis upon which it was based may be out of date.3H.-J. Bandelt, Y.-G. Yao, M. B. Richards, and A. Salas, The brave new era of human genetic testing, BioEssays, vol. 30, no. 11-12 (2008), pp.1246-1251. Worst of all is a tendency to circular thinking: genetic results are interpreted in the light of the favoured archaeological model; then the conclusion is taken as proof of the archaeological model. Yet despite teething troubles, this new science is full of promise.

Over the last few years papers and books have poured out in a whirling stream. Some overturn long-held ideas. Others support them. For those trying to get a grip on the story of Europe's past, it has been the intellectual equivalent of white-water rafting - an exhilarating ride that leaves one breathless. Yet out of this seeming chaos a solid structure is emerging, piece by piece. Key publications have illuminated the great migrations in prehistory. Some are from archaeologists. Others are from population geneticists. Some of the most exciting work has yet to be published, but already different strands of evidence are being knitted into a complex answer to that simple question: where did Europeans come from?

The main story, originally all on this page, is now broken into separate pages, to make for faster loading and easier online machine translation.Linking text and conclusion remain on this page. The recommended sequence for new readers is:

The First Europeans

The first modern humans to arrive in Europe, the Neanderthals that they encountered there and their retreat from the Ice Age.

Optional extra: Early transport: people power

Mesolithic hunters and fishermen

The repopulation of most of Europe as the glaciers melted about 10,000 years ago.

Optional extra: Early transport: floating along

Optional extra: Pottery and climate

Interludes:

Genetic debate

Outlines for the general reader the debate over the findings from population genetics relating to the origins of the peoples of Europe.

Who do you look like?

Outlines for the general reader the latest findings from genetics on the genes that decide some of the most noticeable differences between human beings: pigmentation, height and head shape.

The first farmers

The arrival of farming in Europe from the Near East, stretching from the arrival on Cyprus c. 8,500 BC to the arrival in Scandinavia and the BritishIsles c. 4,000 BC.

Optional extra: The Near Eastern Neolithic

Optional extra: Who moved the megaliths?

The Copper Age

The beginnings of metallurgy in Europe c. 5,000 BC and how it spread; the new copper-using cultures and relationships between them.

Optional extra: the start of the use of metal in the Near East

The Indo-European family

An outline of the latest thinking on the European steppe homeland of the parent (spoken c. 4,000 BC) of the family of Indo-European languages. It marries the archaeological evidence of departures from the European steppe, carrying a package of new technologies, to linguistic evidence of the break-aways which ultimately resulted in the languages spoken today between Iceland and India.

Optional extra: Prehistoric transport: horse power

Optional extra: Prehistoric transport: rolling along

Optional extra: The linguistic arguments

Indo-European genetics

Meshes the evidence from modern populations and ancient DNA to deduce the genetic markers of the spread of speakers of Indo-Euopean languagues.

Optional extra: Ancient Western European DNA: a composite table of results from multiple studies. This is a large file and may be slow-loading.

Warning:

The Babel of ethnic names

Before we enter the realms of history, it is as well to brace yourself for the confusion caused by the ability to write things down. The modern name for apast people may not be the name that they used for themselves, or that turns up in the records of other people encountering them. One and the same people couldbe called by different names in different languages or at different times.

People could identify themselves in different ways for different purposes, just as we do today. For example someone today might see himself as a residentof Warsaw, a Pole, a Slav and a European, or even a resident of New York, a Pole, a Slav and an American. Today these multiple identities are no mystery. We are accustomed to the concept of nations, which may be viewed as the same as, or different from, ethnicity. We understand that language or religion may count more than birth-place in a person's self-identification. Those familiar with clans and castes find them easy to fathom.

But the past, as so famously said, is a foreign country. So it is necessary here and there in what follows to explain the path that is being picked through the fog. To go straight there: Identifying the Celts, Germani: who were they?, Slavs: the confusion of names.

Beaker Folk to Celts and Italics

Follows the trail (c. 3,000 BC) of an unusual type of memorial and a bell-shaped beaker, together with clues from population genetics and linguistics to present a new view of the origins of the Celts and the peoples of Italy.

Optional extra: The Basques: an outline of the debate on their origins, including new ideas.

Iron Age Cimmerians and Celts

Once again a people moving up the Danube from the European steppe brought a new technology into Central Europe, this time the Cimmerians bringingiron-working and chariot-horses. This influence filtered into existing cultures to create (from c. 750 BC) the Iron Age cultures that emerge into history as Celtic.

Optional extra: Celtic tribes of Britain and Ireland: lists by country and British region, with comments, maps and introductory essays.

Minoans and Mycenaens

Outlines the interaction of these two early Greek civilizations, one Indo-European, the other seemingly Near Eastern in origin, merging the archaeological, linguistic and genetic evidence.

Optional extra: The fruit of the vine: the earliest wine-making.

Optional extra: The chariot: its invention on the Western Asian steppe and spread into Egypt and Europe.

Optional extra: the Phoenicians, the great trading rivals of the Greeks in the Mediterranean.

Etruscans and Romans

Again a great European civilization emerged from the interaction of two cultures, one Indo-European, the other Near Eastern in origin. This outline of the origin of the Romans and mobility within the Empire combines the latest findings from population genetics, archaeology and linguistics.

The Great Wandering: The Migration Period

An overview of the upheaval in Europe in Late Antiquity, outlining the problems of understanding the evidence, key elements of the debate, and theengines of change. Specific migrations are covered on their own pages:

The Germani

Today the speakers of Germanic languages are spread far and wide. These languages include English, Dutch and the Scandinavian languages, as well as theobvious German. Where did they come from? The answer may be surprising.

Optional extra: Goths and Vandals

Enter the Slavs

The explosive spread of the Slavs (from c. 500 AD) over Eastern Europe and the Balkans from their homeland on the Middle Dnieper.

The Vikings

The wide wanderings of Scandinavians from c. 800-1100 AD form a coda to the Migration period. From hit-and-run raiders, they amalgamated to form armies and establish new kingdoms.

Conclusion

These pages have been constantly revised and expanded since the first went online. Even so it is far from comprehensive. I have not mentioned the wanderings of Jews,4A. Gusev et al., The architecture of long-range haplotypes shared within and across populations, Molecular Biology and Evolution, (online 6 October 2011 before print); F. L. Mendez et al., Increased Resolution of Y Chromosome Haplogroup T DefinesRelationships among Populations of the Near East, Europe, and Africa, Human Biology, vol. 83, no. 1 (Feb 2011), pp. 39-53; S.M.Bray et al, Signatures of founder effects, admixture, and selection in the AshkenaziJewish population, Proceedings of the National Academy of Sciences of the United States of America(Published online before print August 26, 2010); J.B. Listman et al., Identification of population substructure among Jews usingSTR markers and dependence on reference populations included, BMC Genetics, vol. 11 (2010), 48; L. Zelinger et al., An ancient autosomal haplotype bearing a rare achromatopsia-causing founder mutation is shared amongArab Muslims and Oriental Jews, Human Genetics, (online first June 14, 2010); D. Behar et al., The genome-wide structure of the Jewish people,Nature, advance online publication 9 June 2010; G. Atzmon et al., Abraham's Children in the Genome Era: Major Jewish Diaspora PopulationsComprise Distinct Genetic Clusters with Shared Middle Eastern Ancestry, The American Journal of Human Genetics, (03 June 2010); M.F. Hammer et al., Jewish and Middle Eastern non-Jewish populations share a commonpool of Y-chromosome biallelic haplotypes, Proceedings of the National Academy of Sciences of the United States of America, vol. 97 (2000), no. 12, pp. 6769-6774; A. Nebel et al, The Y Chromosome Pool of Jews as Part ofthe Genetic Landscape of the Middle East, The American Journal of Human Genetics, 69 (5), (2001), pp. 1095-1112 ; D. Behar et al, Contrasting patterns of Y chromosome variation in Ashkenazi Jewish and host non-JewishEuropean populations, Human Genetics, vol. 114 (2004), pp. 354–365; D. M. Behar et al., Counting the Founders: The MatrilinealGenetic Ancestry of the Jewish Diaspora, PLoS ONE, vol. 3, no.4 (2008). and Roma,5D. Gresham et al, Origins and divergence of the Roma (Gypsies), American Journal of Human Genetics, vol. 69 (2001), no.6, pp. 1314–1331; I. M. Klari et al., Dissecting the molecular architecture and origin of Bayash Romanipatrilineages: Genetic influences from South-Asia and the Balkans, American Journal of Physical Anthropology, vol. 138, no. 3 (March 2009), pp. 333–342; A. Gusmão et al., A Genetic Historical Sketch ofEuropean Gypsies: The Perspective From Autosomal Markers, American Journal of Physical Anthropology, vol. 141 (2010), pp. 507–514; M. Regueiro et al., Divergent patrilineal signals in three Roma populations,American Journal of Physical Anthropology (online 27 September 2010 before print); A. Zalán et al., Paternal genetic history of the Vlax Roma,Forensic Science International: Genetics (online 24 September 2010); I. Mendizabal et al., Reconstructing the Indian origin and dispersal ofthe European Roma: a maternal genetic perspective, PLoS ONE 6, no.1 ( 2011), e15988.. and only lightly touched on some of the folk-movements and trade routes of historic times, which need to be taken into account when looking at the genetic and linguistic maps of Europe. The restless peoples of Europe have stirred the gene pool many a time, overlaying thesignatures of more ancient population movements. The resulting palimpsest cannot be read in an instant.

The aim here is to give a taste of the exciting convergence of ideas and evidence that may ultimately give us a clearer answer to the question of where we Europeans came from. It may not be a simpler answer. What emerges from this survey is that visions of stability over millennia must give way to a more dynamic picture of Europe's prehistory. The continent was not barred to incomers after the arrival of the earliest human beings. On the contrary, the tracks of Neolithic arrivals from the Near East can be seen in DNA. Nor was the Neolithic wave of migration the last one of importance. Movements in the ages of metal had a massive impact, as did those after the fall of Rome. Travellers have criss-crossed Europe in complex patterns over the centuries. From that mixing and mingling we have gained much.

Geographical clusters found in autosomal DNA. Click to enlarge in new window.Yet each European country has had its own history. Despite the high degree of genetic similarity among Europeans, it is still possible to findnational clusters by testing a huge array of variable loci in the human genome. The largest variations correspond to geographical relationships. This is consistent with migration ... and geographic physical boundaries being the most critical or common factor in determining genotypic patterns.6J.Novembre et al., Genes mirror geography within Europe, Nature, vol. 456, no. 7218 (6 November 2008), pp. 98-101. Migrations left a different imprint on each country. Then settled, inter-marrying populations have had time to develop their own genetic quirks. So to some extent genes mirror geography today.7C. Tian et al, European population genetic substructure: further definition of ancestry informative markers for distinguishing among diverse European ethnic groups, Molecular Medicine, vol. 15, nos. 11-12 (November 2009), pp. 371–383; C. Tian et al., Analysis and application of European genetic substructure using 300 K SNP information, PLoS Genetics, vol. 4, no. 1 (2008): e4; P. Drineas, J. Lewis and P. Paschou, Inferring geographic Ccordinates of origin for Europeans using small panels of ancestry informative markers, PLoS ONE, vol. 5, no. 8: e11892 (August 2010).

Notes

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  1. 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.
  2. A. Javed et al., Recombination networks as genetic markers in a human variation study of the Old World, Human Genetics, published online 18 October 2011 ahead of print; M. 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 human genetic 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. Gutenkunst et 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. Jakobsson 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. 16057-1606; 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. et al, Genotype, haplotype and copy-number variation in worldwide human populations, Nature, no. 451 (21 February 2008), pp. 998-1003; Q. Ayub et al., Reconstruction of human evolutionary tree using polymorphic autosomal microsatellites, American Journal of Physical Anthropology, vol. 122 (2003), pp. 259-268.
  3. H.-J. Bandelt, Y.-G. Yao, M. B. Richards, and A. Salas, The brave new era of human genetic testing, BioEssays, vol. 30, no. 11-12 (2008), pp. 1246-1251.
  4. A. Gusev et al., The architecture of long-range haplotypes shared within and across populations, Molecular Biology and Evolution, (online 6 October 2011 before print); F. L. Mendez et al., Increased Resolution of Y Chromosome Haplogroup T Defines Relationships among Populations of the Near East, Europe, and Africa, Human Biology, vol. 83, no. 1 (Feb 2011), pp. 39-53; S.M.Bray et al, Signatures of founder effects, admixture, and selection in the Ashkenazi Jewish population, Proceedings of the National Academy of Sciences of the United States of America (Published online before print August 26, 2010); J.B. Listman et al., Identification of population substructure among Jews using STR markers and dependence on reference populations included, BMC Genetics, vol. 11 (2010), 48; L. Zelinger et al., An ancient autosomal haplotype bearing a rare achromatopsia-causing founder mutation is shared among Arab Muslims and Oriental Jews, Human Genetics, (online first June 14, 2010); D. Behar et al., The genome-wide structure of the Jewish people, Nature, advance online publication 9 June 2010; G. Atzmon et al., Abraham's Children in the Genome Era: Major Jewish Diaspora Populations Comprise Distinct Genetic Clusters with Shared Middle Eastern Ancestry, The American Journal of Human Genetics, (03 June 2010); M.F. Hammer et al., Jewish and Middle Eastern non-Jewish populations share a common pool of Y-chromosome biallelic haplotypes, Proceedings of the National Academy of Sciences of the United States of America, vol. 97 (2000) , no. 12, pp.  6769-6774; A. Nebel et al, The Y Chromosome Pool of Jews as Part of the Genetic Landscape of the Middle East, The American Journal of Human Genetics, 69 (5), (2001), pp. 1095-1112 ; D. Behar et al, Contrasting patterns of Y chromosome variation in Ashkenazi Jewish and host non-Jewish European populations, Human Genetics, vol. 114 (2004), pp. 354–365; E. Levy-Coffman, A mosaic of people: the Jewish story and a reassessment of the DNA evidence, Journal of Genetic Genealogy, vol. 1 (2005), no. 1, p. 12-33; D.M. Behar et al., Counting the Founders: The Matrilineal Genetic Ancestry of the Jewish Diaspora, PLoS ONE, vol. 3, no.4 (2008).
  5. D. Gresham et al, Origins and divergence of the Roma (Gypsies), American Journal of Human Genetics, vol. 69 (2001), no.6, pp. 1314–1331; I. M. Klari et al., Dissecting the molecular architecture and origin of Bayash Romani patrilineages: Genetic influences from South-Asia and the Balkans, American Journal of Physical Anthropology, vol. 138, no. 3 (March 2009), pp. 333–342; A. Gusmão et al., A Genetic Historical Sketch of European Gypsies: The Perspective From Autosomal Markers, American Journal of Physical Anthropology, vol. 141 (2010), pp. 507–514; M. Regueiro et al., Divergent patrilineal signals in three Roma populations, American Journal of Physical Anthropology (online 27 September 2010 before print); A. Zalán et al., Paternal genetic history of the Vlax Roma, Forensic Science International: Genetics (online 24 September 2010); H. Pamjav et al., Genetic structure of the paternal lineage of the Roma People, American Journal of Physical Anthropology (online 4 January 2011 ahead of print); I. Mendizabal et al., Reconstructing the Indian origin and dispersal of the European Roma: a maternal genetic perspective, PLoS ONE 6, no.1 ( 2011), e15988.
  6. J. Novembre et al., Genes mirror geography within Europe, Nature vol. 456 (6 November 2008), pp. 98-101.
  7. C. Tian et al, European population genetic substructure: further definition of ancestry informative markers for distinguishing among diverse European ethnic groups, Molecular Medicine, vol. 15, nos. 11-12 (November 2009), pp. 371–383; C. Tian et al., Analysis and application of European genetic substructure using 300 K SNP information, PLoS Genetics, vol. 4, no. 1 (2008): e4; P. Drineas, J. Lewis and P. Paschou, Inferring geographic coordinates of origin for Europeans using small panels of ancestry informative markers, PLoS ONE, vol. 5, no. 8: e11892 (August 2010).