Kostenki Man, reconstructed by Mikhail Gerasimov (1907-1970). An early European who was not yet phenotypically European.
Who were the first Europeans? We now have a better idea, thanks to a new paper about DNA from a man who lived some 38,700 to 36,200 years ago. His remains were found at Kostenki, a well-known Upper Paleolithic site in central European Russia (Seguin-Orlando et al., 2014).
Kostenki Man tells us several things about the first Europeans and, more broadly, the first non-African humans:
The Neanderthal encounter
Modern humans received their Neanderthal admixture when they were just spreading out of Africa some 54,000 years ago. At that time, they had not yet encountered the Neanderthals and were entering the territory of the Skhul/Qafzeh hominids, a semi-archaic people of the Middle East. So we may have got our Neanderthal admixture indirectly. The Skhul/Qafzeh hominids had probably interbred with their Neanderthal neighbors to the north, and our ancestors may have then picked up this admixture while in the Middle East.
When our ancestors spread farther north into Europe, some 45,000 to 42,000 years ago, they could have interbred directly with Neanderthals, but they didn't. Perhaps the two groups were just too different. They seem to have intermixed only via a third party that was neither fully modern nor fully archaic.
A strange detour ... and then another!
There was initially a large continuous population across northern Eurasia, perhaps composed of nomads who pursued wandering herds of reindeer across the European Plain and its eastward extension into central and northern Asia.
Not long before the time of Kostenki Man, these Northern Eurasians began to split into three regional groups: Western Eurasians, Eastern Eurasians, and the ancestors of Middle Eastern farmers. The degree of reproductive isolation is unclear, however, and gene flow may have continued between all three groups until the onset of the last ice age some 25,000 years ago. This may be why Kostenki Man does not fit perfectly into any of the three groups, although he is genetically closest to Western Eurasians.
Yes, Northern Eurasians were ancestral to the early farming peoples of the Middle East. It seems that early modern humans had to head north, learn to hunt reindeer, and then head south again before they could start farming. Sounds like a strange detour. Wouldn't it have been easier to stay put and do it locally? You know, Middle-Eastern hunter-gatherers becoming Middle Eastern farmers? Apparently not.
It gets even more convoluted. After some of those Northern Eurasians had gone south to the Middle East, some of their farming descendants "returned" to Europe and partially replaced its hunter-gatherers, particularly in southern and central Europe. This second detour has been greeted with disbelief. Dienekes (2014), for instance, has written: "I don't think many archaeologists would derive European farmers from Russia (Russia is actually one of the last places in Europe that became agricultural)."
True, but farming requires a mindset that may have come from those northern hunters (Frost, 2014). When Piffer (2013) looked at human variation in alleles at COMT, a gene linked to executive function, working memory, and intelligence, he found that northern hunting peoples had more in common with farming peoples than with other hunter-gatherers, "possibly due to the higher pressure on technological skills and planning abilities posed by the adverse climatic conditions."
That mindset made farming possible, but the first steps toward farming could not be taken in a cold climate. They had to be taken in a place with a long growing season and a wide variety of domesticable plants and animals, such as in the Middle East. Once farming had developed there, it could move back north, while taking along its technologies, its food crops, and its livestock species.
Farming can develop in the tropics with a "tropical" mindset, but it looks very different. The farming that arose in West Africa is overwhelmingly women's work and seems to have wholly developed out of female plant gathering. The guinea fowl is the only animal that has been domesticated for food consumption in sub-Saharan Africa.
The Ice Age was not so bad
The Upper Paleolithic humans of northern and eastern Europe did not die out during the last ice age, as was commonly thought. They survived the glacial maximum intact.
The European phenotype came later
Kostenki Man was dark-skinned, dark-eyed, and rather short. These details, curiously enough, appear not in the paper but in a review of the paper, published by the same journal, as well as in an interview with one of the authors (Associated Press, 2014; Gibbons, 2014).
So we now have an upper bound for the emergence of the European phenotype, i.e., light skin and a diverse palette of hair and eye colors. The lower bound has been set by the remains of a Swedish hunter-gatherer, dated to 8,000 years ago, who had the "European" allele for light skin at the gene SLC24A5 (Skoglund et al., 2014).
My main criticism centers on the dating to 38,700 - 36,200 years ago. At the Kostenki site, the radiocarbon dating used to be some 10,000 years younger. It was then recalibrated to an older range of dates when a layer of volcanic ash at the site was attributed to a volcano that had erupted in southern Italy some 39,000 years ago. This recalibration was initially controversial, but the controversy has since subsided (Sinitsyn and Hoffecker, 2006). I would not rule out a subsequent re-recalibration.
By retrieving ancient DNA from an early modern human, we have made a key advance in human paleogenetics, perhaps more so than by sequencing the Neanderthal genome. We again see that evolution did not slow down with the emergence of anatomically and behaviorally modern humans some 60,000 years ago. It actually began to speed up, as humans began to enter not only new natural environments but also new cultural environments of their own making.
Associated Press (2014). DNA study dates Eurasian split from East Asians, The Columbus Dispatch, November 6http://hosted2.ap.org/OHCOL/07e34bb59e064cedb7e2776e8db4b4f7/Article_2014-11-06-EU--Eurasian%20Split/id-ae36fa368c634c7383d807942bd5fe67
Dienekes (2014). Genome of Kostenki-14, an Upper Paleolithic European (Seguin-Orlando, Korneliussen, Sikora, et al. 2014), Dienekes' Anthropology Blog, November 7http://dienekes.blogspot.ca/2014/11/genome-of-kostenki-14-upper-paleolithic.html
Frost, P. (2014). The first industrial revolution, Evo and Proud, January 18http://evoandproud.blogspot.ca/2014/01/the-first-industrial-revolution.html
Gibbons, A. (2014). European genetic identity may stretch back 36,000 years, Science, News, November 6http://news.sciencemag.org/archaeology/2014/11/european-genetic-identity-may-stretch-back-36000-years
Piffer, D. (2013). Correlation of the COMT Val158Met polymorphism with latitude and a hunter-gather lifestyle suggests culture-gene coevolution and selective pressure on cognition genes due to climate, Anthropological Science, 121, 161-171.https://lesacreduprintemps19.files.wordpress.com/2014/01/correlation-of-the-comt-val158met-polymorphism-with-latitude-and-a-hunter-gather-lifestyle-suggests-culturee28093gene-coevolution-and-selective-pressure-on-cognition-genes-due-to-climate.pdf
Seguin-Orlando, A., T.S. Korneliussen, M. Sikora, A.-S. Malaspinas, A. Manica, I. Moltke, A. Albrechtsen, A. Ko, A. Margaryan, V. Moiseyev, T. Goebel, M. Westaway, D. Lambert, V. Khartanovich, J.D. Wall, P.R. Nigst, R.A. Foley, M.M. Lahr, R. Nielsen, L. Orlando, and E. Willerslev. (2014). Genomic structure in Europeans dating back at least 36,200 years, Science, Published online 6 November 2014http://www.sciencemag.org/content/early/2014/11/05/science.aaa0114
Sinitsyn, A.A., and J.F. Hoffecker. (2006). Radiocarbon dating and chronology of the Early Upper Paleolithic at Kostenki, Quaternary International, 152-153, 164-174.http://www.sciencedirect.com/science/article/pii/S1040618206000206
Skoglund, P., H. Malmstrom, A. Omrak, M. Raghavan, C. Valdiosera, T. Gunther, P. Hall, K. Tambets, J. Parik, K-G. Sjogren, J. Apel, E. Willersley, J. Stora, A. Gotherstrom, and M. Jakobsson. (2014). Genomic diversity and admixture differs for stone-age Scandinavian foragers and farmers, Science, 344 (6185), 747-750.http://www.sciencemag.org/content/344/6185/747.short