Homo sapiens began to disperse out of Africa via a large-scale wave, serving as the migration from which modern non-Africans descend. However, there were smaller, earlier dispersals previous to this large-scale wave that did not impact Eurasia genetically.¹

The Ranis/Zlaty Kun Population

The oldest individual from an earlier migration that has been studied as of now is the Zlaty Kun woman from Zlaty Kun, Czechia. Found in the Koneprusy cave system with evidence that her skull (the only remains found) was gnawed on by hyenas, this individual was discerned to be 45,000 years old or older. The artifacts found with her were associated with the Upper Palaeolithic, but could not be assigned to any specific technological complex. Zlaty Kun carried approximately 3% Neanderthal ancestry. The stretches of her Neanderthal segments were longer than those of other European hunter-gatherers, and the date of her Neanderthal admixture was estimated to be between 60 and 80 generations before Zlaty Kun's birth.² Though it is unknown if Neanderthal admixture with modern humans occurred in a single short burst, other studies have indicated that gene flow between humans and Neanderthals occurred around 60-50 KYA, indicating a more prolonged period of admixture potentially consistent with the estimated date of Zlaty Kun's Neanderthal admixture.³ Zlaty Kun, or the gene pool that she was a part of, did not contribute to later Eurasian populations. This discontinuity could be attributed to a volcanic eruption that affected the climate of West Eurasia.⁴ Phylogenetically, Zlaty Kun is basal to the split of East and West Eurasians,⁵ despite sharing more alleles with East Eurasians (though this discrepancy can be reconciled due to the admixture incurred by West Eurasians from a deeply-diverged Eurasian population referred to as Basal Eurasians.⁶

Multiple individuals from Ilsenhohle in Ranis, Germany, associated with the Lincombian-Ranisian-Jerzmanowician technocomplex and having lived between 42 and 49 KYA, were relatives of Zlaty Kun, and were likely derived from the same population as her. The Ranis population consisted of six individuals who were (at least partly) familially related. Additionally, two of the Ranis individuals (Ranis12 and Ranis4) were found to be fifth or sixth-degree relatives of Zlaty Kun, though Ranis12 and Ranis4 were less closely related to the other Ranis individuals. Additionally, the Ranis individuals and Zlaty Kun are derived from the same population, with the genetic evidence from the Ranis individuals indicating that the Ranis/Zlaty Kun population was small. This population lived after a reduction of population size associated with the Out-Of-Africa event, but the Ranis/Zlaty Kun population apparently experienced an additional reduction in population size. Additionally, evidence of consanguinity was apparent in the Ranis13 individual and Zlaty Kun, who were both dated to be ~45KY old, and potentially older.¹⁷ Phylogenetically, the Ranis/Zlaty Kun population (as had been previously established via analyses of Zlaty Kun alone) form a basal Out-Of-Africa branch, representing an early split from the population ancestral to other non-African populations (see Hub Eurasians). However, as had been previously shown, the Ranis/Zlaty Kun population did not contribute to later Out-of-African groups.¹⁸

The Neanderthal admixture identified originates from a single admixture event that occurred ~45-49KYA and is reflected in all non-Africans, indicating that all non-Africans resided within a common population at the time. This also indicates that human remains found outside Africa dating before 50KYA are derived from a different non-African population. ¹⁹

Ust'-Ishim

Similarly to Zlaty Kun, the ~45,000-year-old Ust'-Ishim man, found in Omsk Oblast, Russia, lived prior to (or simultaneous with) the split of Eastern and Western Eurasians, likely during a warm period,⁷ and did not impact subsequent populations.⁸ Phylogenetically, Ust'-Ishim is close to the population from which East and West Eurasians split (see Hub Eurasians), nearly forming a trifurcation between East and West Eurasians.⁹ Like Zlaty Kun, he shared more derived alleles with East Eurasians than West Eurasians. He also does not share more derived alleles with East Eurasians than with other prehistoric samples, meaning he is equally related to prehistoric Europeans and modern East Eurasians. As indicated by Zlaty Kun and Ust-Ishim's genetic affinities with East Eurasians, present-day West Eurasians derive some of their ancestry from a population that was not involved in the first widespread dispersal into Eurasia. Though he carried a similar amount of Neanderthal ancestry as modern-day Eurasians, his Neanderthal segments are longer than modern-day humans and indicate that he had a Neanderthal ancestor about 7-13 thousand years before he was born. This fact indicates that admixture between Neanderthals and humans had already occurred before 45KYA.¹⁰ Ust'-Ishim also possesses additional Neanderthal admixture from 30-50 generations before he lived, unlike the Ranis/Zlaty Kun population, whose Neanderthal ancestry was derived from a single admixture event. ²⁰

Oase 1

Like the aforementioned individuals, Oase 1, a 42-37 thousand-year-old human found in Pestera cu Oase, Romania,¹¹ was basal to the split of East and West Eurasians.¹² This individual also had a high amount of Neanderthal admixture compared to modern humans. Like the other individuals mentioned, the length of the Neanderthal segments indicates that his Neanderthal ancestry was quite recent.¹³ However, unlike the other individuals mentioned, he had a significantly more recent Neanderthal ancestor (4-6 generations before his own).¹⁴ Oase 1 carried more Neanderthal ancestry (~6-9%) than any other modern human currently sequenced.¹⁵

Legacy

As previously stated, all three of these individuals did not contribute to later Eurasian populations. The transition between the Upper and Middle Palaeolithic, beginning 47KYA, coincided with the spread of Homo sapiens into Eurasia and the disappearance of Neanderthals,¹⁶ significantly impacting the genetic landscape of Eurasia.

Sources

NOTE: The footnotes in this article do not follow numerical order to reflect the fact that this article was reposted in light of new genetic evidence that is extremely significant to this topic. Also note that names in this article may be written slightly differently, as this site currently does not support diacritics.

^{1, 5, 9} Vallini, L., Zampieri, C., Shoaee, M.J. et al. The Persian plateau served as hub for Homo sapiens after the main out of Africa dispersal. Nat Commun 15, 1882 (2024). https://doi.org/10.1038/s41467-024-46161-7

^{2, 4, 6, 8, 12, 15} Prufer, K., Posth, C., Yu, H. et al. A genome sequence from a modern human skull over 45,000 years old from Zlaty kun in Czechia. Nat Ecol Evol 5, 820-825 (2021). https://doi.org/10.1038/s41559-021-01443-x

^{3, 16}Hajdinjak, M., Mafessoni, F., Skov, L. et al. Initial Upper Palaeolithic humans in Europe had recent Neanderthal ancestry. Nature 592, 253-257 (2021). https://doi.org/10.1038/s41586-021-03335-3

^{7, 10}Fu, Q., Li, H., Moorjani, P. et al. Genome sequence of a 45,000-year-old modern human from western Siberia. Nature 514, 445-449 (2014). https://doi.org/10.1038/nature13810

^{11, 14}Jean-Jacques Hublin, Nikolay Sirakov, Vera Aldeias, Shara Bailey, Edouard Bard, et al.. Initial Upper Palaeolithic Homo sapiens from Bacho Kiro Cave, Bulgaria. Nature, 2020, 581 (7808), pp.299-302. ff10.1038/s41586-020-2259-zff. Ffhal-03321929f

¹³Fu, Q., Hajdinjak, M., Moldovan, O. et al. An early modern human from Romania with a recent Neanderthal ancestor. Nature 524, 216-219 (2015). https://doi.org/10.1038/nature14558

Poznik, G., Xue, Y., Mendez, F. et al. Punctuated bursts in human male demography inferred from 1,244 worldwide Y-chromosome sequences. Nat Genet 48, 593-599 (2016). https://doi.org/10.1038/ng.3559

^{17, 18, 19, 20}Sumer, A.P., Rougier, H., Villalba-Mouco, V. et al. Earliest modern human genomes constrain timing of Neanderthal admixture. Nature (2024). https://doi.org/10.1038/s41586-024-08420-x