In terms of its population movements and descent, the facts appear to be that modern Europeans are a three-fold mix of three ancient populations as follows:
(1) Palaeolithic and Mesolithic hunter-gatherers who lived in Europe from c. 45,000 years ago;We can examine the history of these groups in greater detail as follows:
(2) Neolithic Anatolian and Aegean farmers who migrated into Europe from c. 6,500 BC–4,000 BC, and
(3) Indo-European-speaking Yamnaya-culture people who swept into Europe from the Russian steppe from 3,000 to 2,000 BC.
(1) Palaeolithic and Mesolithic hunter-gatherers from c. 45,000 years agoAll modern indigenous Europeans (e.g., those not descended from the later invaders from the Eurasian steppe like the Magyars or other later arrivals) have a mix of genes from these three types of ancient people (see here). The distinctive European traits of blue eyes (from hunter gatherers), lactose tolerance (from the Yamnaya people) and fairer skin spread by interbreeding and natural selection (see here and here).
These were the earliest members of Homo sapiens in Europe; they were a hunter-gatherer people who lived in Europe from about 45,000 years ago during the end of the last Ice Age (which lasted from about 108,000 to 10,000 BC). They came from the Middle East along a Mediterranean route. But Europe must have been sparsely populated by these people: in essence, the earliest European hunter-gatherers must have been a relatively small population. These Mesolithic hunter-gatherers have contributed to modern European genetics, though to a different extent in different regions.
It appears that some of them interbred with the Neanderthals (who had in turn evolved from Homo erectus populations) (see here). But, even if true, the Neanderthal genetic contribution to modern Europeans is low, maybe as low as 1.5–2.1% (Prüfer et al. 2014). (For a useful family tree, see here).
There also seems to be some evidence that the mysterious Homo sapiens denisova lived in Europe in the Stone age.
At any rate, the Palaeolithic and Mesolithic hunter-gatherers appear to have had dark skin, which lightened by Darwinian evolution over the centuries, but perhaps accelerated by the adoption of farming which involved a reduced intake of vitamin D. Blue eyes may have evolved amongst these early European hunter-gatherers as well (see here and here).
(2) Neolithic Anatolian farmers from c. 6,500 BC–4,000 BC
From c. 6,500 BC–4,000 BC, Neolithic Anatolian farmers from northern Greece and north-western Turkey started migrating into central Europe through the Balkan route and then by the Mediterranean route to the Iberian Peninsula. They brought sedentary agricultural communities and new domestic animals and plants. Modern southern Europeans still seem to have inherited much more of their genes from these people. The original Anatolian farmer phenotype was probably similar to that of the modern people of Sardinia (Hofmanová et al. 2016: 3), and, generally speaking, the swarthy phenotype of southern Europeans is the legacy of their greater descent from the Neolithic Anatolian farmers as opposed to northern Europeans. Genetic analysis of ancient farmers seems to show that after their arrival in Europe the Neolithic Anatolian farmers only mixed infrequently and at low levels with the hunter-gatherers, but increasingly from the later Neolithic period (Hofmanová et al. 2016: 4).
(3) Indo-European-speaking Yamnaya-culture people from 3,000 to 2,000 BC
From 3,000 to 2,000 BC, there was massive migration of people from the South Russian steppe into central Europe, and then into northern and western Europe, and these people were of the Yamnaya culture north of the Black Sea. These people were almost certainly proto-Indo-European speakers (Balter and Gibbons 2015), cattle herders, and probably had a phenotype with brown eyes, pale skin, and taller height. It is also interesting – and not surprising – that the Caucasian Yamnaya-culture people have bequeathed to modern Europeans the trait of persistent adult lactose tolerance (Allentoft et al. 2015: 171). The migration of the Yamnaya-culture people west and east spread the Indo-European languages (Allentoft et al. 2015: 171).
The genetic contribution of the Neolithic Anatolian farmers is important, but admittedly less so as you move northwards in Europe. However, even the Scandinavians have significant descent from the Neolithic Anatolian farmers, and even a marginal population like the Irish do as well (see here).
The further north you go in Europe, it appears the more is the genetic contribution of the Palaeolithic and Mesolithic hunter-gatherers.
And virtually everyone has some descent from the Indo-European-speaking Yamnaya-culture people. Linguistically, this third group is fundamentally important because virtually everyone in Europe now speaks an Indo-European language (apart from the Basques, Hungarians, Finns, Estonians, and other minor people).
The Indo-European Yamnaya-culture people of the steppe had themselves mixed with a population of hunter-gatherers isolated in the Caucasus region, so that the early Yamnaya pastoralists were a mix of Eastern European hunter gatherers and another group of hunter-gatherers from the Caucasus. These people then migrated back into Europe in a mass movement from c. 3,000 to 2,000 BC (Balter and Gibbons 2015: 815). For example, they flooded into eastern and central Europe and created the Corded Ware culture (c. 3100–1900 BC) (see the map here). Their descendants appear to have arrived in Greece from 2400–2000 BC bringing with them the Proto-Greek language that would evolve into Mycenaean Greek and then the later Greek dialects of Classical Greece.
Another fascinating part of forgotten history is how the Indo-European languages have displaced what almost certainly must have been non-Indo-European languages in Europe spoken by the Palaeolithic and Mesolithic hunter-gatherers and Neolithic Anatolian farmers and their descendants.
For example, as late as the early Roman Republic there appears to have been a large surviving group of non-Indo-European languages in Europe as follows:
(1) the proposed Vasconic group of languages, including the extinct Aquitanian language (from which the modern survivor Basque is derived), the Iberian languages, Tartessian, and possibly the Lusitanian language (see the map here);At any rate, it is now firmly accepted that the ancient non-Indo-European Aquitanian language was the direct ancestor of modern Basque, and Aquitanian was spoken in large areas of south-western France, northern Spain and in the Pyrenees (Trask 1995: 87; see the map here). In turn, it would appear plausible that the ancient non-Indo-European Iberian language in Spain was related to ancient Aquitanian, particularly on the basis of recent evidence relating to the numerals of both languages (and, more speculatively, to Tartessian as well).
(2) the Tyrsenian languages, including Etruscan, Raetic, and the Lemnian language;
(3) the Paleo-Sardinian language, the Sicanian language, the hypothetical non-Indo-European German substrate language, the pre-Greek substrate language, and the Eteocretan language, and perhaps the Pictish language.
These mysterious languages seem to have been the descendants of the ancient language of the Neolithic Anatolian farmers.
Of course, some scholars like Colin Renfrew have proposed the Anatolian hypothesis and argued that the Neolithic Anatolian farmers already spoke Proto-Indo-European and hence brought Indo-European languages to Europe (see Renfrew 2003).
If true, then the Indo-European Yamnaya-culture people must have brought a later offshoot, perhaps a proto-Balto-Slavic language (Balter and Gibbons 2015: 815), and the non-Indo-European language substrate in Europe must have been descended from the languages of the Palaeolithic and Mesolithic hunter-gatherers.
That would make the Basque language a truly ancient language descended from the ancient Mesolithic hunter-gatherer languages of Europe.
The question of the origins and classification of Basque is very interesting indeed. Anyone who has a decent knowledge of European languages can see that Basque is an alien and very weird language, unconnected to any other language in Europe, as, for instance, in a random selection of Basque words:
Arrigorriagakoa (a surname)These words look bizarre to the modern European eye because Basque is clearly derived from some ancient non-Indo-European language. (In fact, having myself done some ancient Near Eastern languages, these words remind me of Sumerian or Akkadian).
Goikoetxea (“high lying house,” a surname)
Etxandi (another surname)
But recent genetic study of both ancient and modern Basques suggests that they are mostly descended from the ancient Neolithic Anatolian farmers and so their mysterious language may well be derived from the ancient Neolithic Anatolian farmer language of the Middle East (Günther et al. 2015: 11920). If so, this suggests that the Anatolian hypothesis is wrong: the Yamnaya-culture people from the steppe were the proto-Indo-European speakers from whose language all other Indo-European languages in Europe have derived.
Allentoft, Morten E. et al. 2015. “Population Genomics of Bronze Age Eurasia,” Nature 522 (11 June): 167–172.
Balter, Michael and Ann Gibbons. 2015. “Indo-European Languages tied to Herders,” Science 347.6224: 814–815.
Gibbons, Ann. 2014. “Three-Part Ancestry for Europeans,” Science 345.6201 (5 September): 1106–1107.
Günther, Torsten et al. 2015. “Ancient Genomes link Early Farmers from Atapuerca in Spain to Modern-Day Basques,” Proceedings of the National Academy of Sciences 112.38: 11917–11922.
Haak, Wolfgang. 2015. “Massive Migration from the Steppe was a Source for Indo-European Languages in Europe,” Nature 522: 207–211.
Hofmanová, Zuzana et al. 2016. “Early Farmers from across Europe directly descended from Neolithic Aegeans,” Proceedings of the National Academy of Sciences of the United States of America, June 6
Jones, Eppie R. et al. 2015. “Upper Palaeolithic Genomes reveal Deep Roots of Modern Eurasians,” Nature Communications 6
Mathieson, Iain et al. 2015. “Genome-Wide Patterns of Selection in 230 Ancient Eurasians,” Nature 528.7583: 499–503.
Prüfer, K. et al. 2014. “The Complete Genome Sequence of a Neanderthal from the Altai Mountains,” Nature 505.7481: 43–49.
Renfrew, Colin. 2003. “Time Depth, Convergence Theory, and Innovation in Proto-Indo-European: ‘Old Europe’ as a PIE Linguistic Area,” in Alfred Bammesberger and Theo Vennemann (eds.), Languages in Prehistoric Europe. Universitätsverlag, Heidelberg. 17–48.
Trask, Robert Lawrence. 1995. “Origins and Relatives of the Basque Language: Review of the Evidence,” in José Ignacio Hualde, Joseba A. Lakarra, R. L. Trask (eds), Towards a History of the Basque Language. J. Benjamins, Amsterdam and Philadelphia. 65–99.