What have the revelations about Neanderthal DNA revealed about Homo sapiens?

Santiago Wolnei Ferreira Guimarães; Hilton P. Silva

doi:10.2478/anre-2020-0008

Authors

Santiago Wolnei Ferreira Guimarães Instituto de Filosofia e Ciências Humanas, Cidade Universitária José da Silveira Netto, Universidade Federal do Pará, Brasil
Hilton P. Silva Instituto de Filosofia e Ciências Humanas, Cidade Universitária José da Silveira Netto, Universidade Federal do Pará, Brasil

DOI:

https://doi.org/10.2478/anre-2020-0008

Keywords:

Human evolution, admixture, genetics, morphology

Abstract

Genetic studies have presented increasing indications about the complexity of the interactions between Homo sapiens, Neanderthals and Denisovans, during Pleistocene. The results indicate potential replacement or admixture of the groups of hominins that lived in the same region at different times. Recently, the time of separation among these hominins in relation to the Last Common Ancestor – LCA has been reasonably well established. Events of mixing with emphasis on the Neanderthal gene flow into H. sapiens outside Africa, Denisovans into H. sapiens ancestors in Oceania and continental Asia, Neanderthals into Denisovans, as well as the origin of some phenotypic features in specific populations such as the color of the skin, eyes, hair and predisposition to develop certain kinds of diseases have also been found. The current information supports the existence of both replacement and interbreeding events, and indicates the need to revise the two main explanatory models, the Multiregional and the Out-of-Africa hypotheses, about the origin and evolution of H. sapiens and its co-relatives. There is definitely no longer the possibility of justifying only one model over the other. This paper aims to provide a brief review and update on the debate around this issue, considering the advances brought about by the recent genetic as well as morphological traits analyses.

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References

Almeida F, Alves N, Ponce De León M, Pierson B, Duarte C, Bártolo P, et al. 2007. The Lapedo Child Reborn: Contributions of CT Scanning and Rapid Prototyping for an Upper Paleolithic Infant Burial and Face Reconstruction. The Case of Lagar Velho Interpretation Centre, Leiria, Portugal. In: D Arnold, F Iccolucci and A Chalmers, editors. The 8th International Symposium in Virtual Reality, Archaeology and Cultural Heritage Vast. Short and Project Papers from VAST. 69–73.
View in Google Scholar

Arnold M, Meyer A. 2006. Hybridization in primates: one evolutionary mechanism. Zool 109(4):261–76.
View in Google Scholar

Arntzen J, Wallis G. 1991. Restricted gene flow in a moving hybrid zone of the newts Triturus cristatus and T. marmoratus in western France. Evolution 45:805–26.
View in Google Scholar

Bons P, Bauer C, Bocherens H, De Riese T, Drucker D, Francken M, et al. 2019. Out of Africa by spontaneous migration waves. PLoS ONE 14(4):1–25.
View in Google Scholar

Bräuer G. 1982. Current problems and research on the origin of Homo sapiens in Africa. Humanbiol Budapest 9:69–78.
View in Google Scholar

Bräuer G. 1984a. A craniological approach to the origin of anatomically modern Homo sapiens in Africa and implications for the appearance of modern Europeans. In: FH Smith, F Spencer, editors. The Origins of Modern Humans. A World Survey of the Fossil Evidence. Liss, New York. 327–410.
View in Google Scholar

Bräuer G. 1984b. The “Afro-European sapiens-hypothesis” and hominid evolution in East Asia during the Late Middle and Upper Pleistocene. Cour Forsch Inst Senckenberg 69:145–65.
View in Google Scholar

Bräuer G. 1992. Africa’s place in the evolution of Homo sapiens. In: G Brauer, FH Smith, editors. Continuity or Replacement: Controversies in Homo sapiens Evolution. Rotterdam, Balkema. 83–98.
View in Google Scholar

Bräuer G. 2008. The origin of modern anatomy: by speciation or intraspecific evolution? Evol. Anthropol 17:22–37.
View in Google Scholar

Cann R, Mark S, Allan W. 1987. Mitochondrial DNA and human evolution. Nature 325(6099):31–6.
View in Google Scholar

Carbonell E, De Castro J, Parés J, Pérez-González A, Cuenca-Bescós G, Ollé A, et al. 2008. The first hominin of Europe. Nature 452(7186):465–69.
View in Google Scholar

Carneiro J, Rodrigues-Filho L, Schneider H, Sampaio I. 2016. Molecular data highlight hybridization in squirrel monkeys (Saimiri, Cebidae). Genet Mol Biol 39(4):539–546.
View in Google Scholar

Caspari R, Wolpoff M. 2013. The process of Modern Human Origins: the evolutionary and demographic changes giving rise to modern humans. FH. Smith, J Ahern, editors. The Origins of Modern – Biology Reconsidered – Chapter 11. New Jersey, John Wiley & Sons. 355–91.
View in Google Scholar

Coon C. 1962. The Origin of Races. New York, Knopf.
View in Google Scholar

Cortés-Ortiz L, Jr. T, Canales-Espinosa D, García-Orduña F, Rodríguez-Luna E, Bermingham E. 2007. Hybridization in large-bodied New World primates. Genetics 176: 2421–25.
View in Google Scholar

Crevecoeur I, Rougier H, Grine F, Froment A. 2009. Modern Human cranial diversity in the Late Pleistocene of Africa and Eurasia: evidence from Nazlet Khater, Peştera cu Oase, and Hofmeyr. Am J Phys Anthropol 140:347–58.
View in Google Scholar

D’errico F, Stringer C. 2011. Evolution, revolution or saltation scenario for the emergence of modern cultures. Philos Trans Royal Soc B 366(1567):1060–69.
View in Google Scholar

Dannemann M, Kelso J. 2017. The contribution of neanderthals to phenotypic variation in modern humans. Am J Hum Genet 101(4):578–89.
View in Google Scholar

Dannemann M, Prüfer K, Kelso J. 2017. Functional implications of Neandertal introgression in modern humans. Genome Biol 18:61.
View in Google Scholar

De Castro Jm, Arsuaga J, Carbonell E, Rosas A, Martínez I, Mosquera M. 1997. A hominid from the lower Pleistocene of Atapuerca, Spain: possible ancestor to Neandertals and modern humans. Science 276 (5317):1392–95.
View in Google Scholar

De Castro Jm, Martinón-Torres M, Gómez-Robles A, Prado-Simón L, Martín-Francés L, Lapresa M, and others. 2011. Early Pleistocene human mandible from Sima del Elefante (TE) cave site in Sierra de Atapuerca (Spain): a comparative morphological study. J Hum Evol 61:1–11.
View in Google Scholar

De Manuel M, Kuhlwilm M, Frandsen P, Sousa V, Desai T, Prado-Martinez J, et al. 2016. Chimpanzee genomic diversity reveals ancient admixture with bonobos. Science 354(6311):477–81.
View in Google Scholar

Dean D, Hublin JJ, Holloway R, Ziegler R. 1998. On the phylogenetic position of the pre-Neandertal specimen from Reilingen, Germany. J Hum Evol 34(5):485–508.
View in Google Scholar

Delson E. 1988. One source not many. Nature 332:206.
View in Google Scholar

Fernando A, Joshua G. 2019. Multiple episodes of interbreeding between Neanderthal and Modern humans. Nat ecol Evol 3:39–44.
View in Google Scholar

Frayer D, Wolpoff M, Pope G. 1993. Theories of modern human origins: the paleonto-logical test. Am Anthropol 95:14–50.
View in Google Scholar

Fu Q, Posth C, Hajdinjak M, Petr M, Mallick S, Fernandes D, et al. 2016. The genetic history of Ice Age Europe. Nature 534(7606):200–05.
View in Google Scholar

Gabounia L, De Lumley M, Vekua A, Lordkipanidze D, De Lumley H. 2002. Découverte d’un nouvel hominidé à Dmanisi (Transcaucasie, Geórgie). Comptes Rendus Palevol 1(4):243–53.
View in Google Scholar

Gittelman R, Schraiber J, Vernot B, Mikacenic C, Wurfel M, Akey J. 2016. Archaic hominin admixture facilitated adaptation to out-of-Africa environments. Curr Biol 26:3375–82.
View in Google Scholar

Green R, Krause J, Briggs A, Maricic T, Stenzel U, Kircher M, et al. 2010. A draft sequence of the Neandertal genome. Science 328:710–22.
View in Google Scholar

Groves C. 1989. A regional approach to the problem of the origin of modern humans in Australasia. In: P Mellars, C Stringer, editors. The Human Revolution. Edinburgh, Edinburgh University Press. 274–85.
View in Google Scholar

Groves C. 2007. Species concepts and speciation: facts and fantasies. In: W Henke, I Tattersall, editors. Handbook of Paleoanthropology. Springer. 1861–79.
View in Google Scholar

Habgood P. 1989. The origin of anatomically-modern-humans in Australia. In: P Mellars, C Stringer, editors. The human revolution: behavioural and biological perspectives on the origins of modern humans. Princeton, Princeton University Press. 245–74.
View in Google Scholar

Habgood P. 1992. The origin of anatomically modern humans in East Asia. In: G Bräuer, FH Smith, editors. Continuity or replacement: controversies in Homo sapiens evolution. Rotterdam, Balkema. 273–88.
View in Google Scholar

Harris K, Nielsen R. 2016. The genetic cost of Neanderthal introgression. Genetics 203:881–91.
View in Google Scholar

Harvati K. 2003. The Neanderthal taxonomic position models of intra- and inter-specific craniofacial variation. J Hum Evol 44:107–32.
View in Google Scholar

Harvati K. 2015. Neanderthals and their contemporaries. In: W Henke, I Tattersall, editors. Handbook of Paleoanthropology, 2nd Edition. Springer. 2243–79.
View in Google Scholar

Harvati K, Frost S, McNulty K. 2004. Neanderthal taxonomy reconsidered: implications of 3D primate models of intra and interspecific differences. Proc Natl Acad Sci U.S.A 101(5):1147–52.
View in Google Scholar

Howell F. 1952. Pleistocene glacial ecology and the evolution of “classic Neanderthal” man. Southwest J Anthropol 8:377–410.
View in Google Scholar

Howells W. 1975. Neanderthal man: facts and figures. In: RH Tuttle, editor. Paleoanthropology: Morphology and Paleoecology. The Hague: Mouton. 389–407.
View in Google Scholar

Howells W. 1976. Explaining modern man: evolutionists versus migrationists. J Hum Evol 5:577–96.
View in Google Scholar

Hublin J. 1985. Human fossils from the North African Middle Pleistocene and the origin of Homo sapiens. In: E Delson, editor. Ancestors: The Hard Evidence. New York, Liss. 282–88.
View in Google Scholar

Hublin J. 1992. Recent human evolution in northwestern Africa. Philos Trans Royal Soc B 337(1280):185–92.
View in Google Scholar

Hublin J. 2001. Northwestern Africa and Middle Pleistocene hominids and their bearing on the emergence of Homo sapiens. In: L Barham, K Robson-Brown, editors. Human Roots: Africa and Asia in the Middle Pleistocene. Bristol, CHERUB, Western Academic and Specialist Press. 99–121.
View in Google Scholar

Hublin J. 2009. The Origin of Neandertals. Proc Natl Acad Sci U.S.A 106(38):16022–27.
View in Google Scholar

Hublin J, Tillier A. 1981. The Mousterian juvenile mandible from Irhoud (Morocco): a phylogenetic interpretation. In: C Stringer, editor. Aspects of Human Evolution. London. Taylor and Francis. 167–85.
View in Google Scholar

Hublin J, Tillier A. 1992. L’Homo sapiens en Europe occidentale: gradualisme et rupture. JJ Hublin, AM Tillier, editors. Aux Origines d’Homo sapiens. Paris, Presses Universitaires de France. 291–327.
View in Google Scholar

Huxley J. 2010. [1942] Evolution: The Modern Synthesis. M Pigliucci, GB Müller. Cambridge, MIT Press.
View in Google Scholar

Douka K, Slon V, Jacobs Z, Ramsey C, Shunkov M, Derevianko A, et al. 2019. Age estimates for hominin fossils and the onset of the Upper Paleolithic at Denisova Cave. Nature 565(7741): 640–3.
View in Google Scholar

Klein R. 1995. Anatomy, behaviour and modern human origins. J World Prehist 9:167–98.
View in Google Scholar

Kuhlwilm M, Gronau I, Hubisz M, De Filippo C, Prado-Martinez J, Kircher M, et al. 2016. Ancient gene flow from early modern humans into Eastern Neanderthals. Nature 530(7591): 429–33.
View in Google Scholar

Lahr M. 1994. The Multiregional Model of modern human origins: a reassessment of its morphological basis. J Hum Evol 26:23–56.
View in Google Scholar

Lahr M. 1996. The Evolution of Modern Human Diversity. Cambridge, Cambridge University Press.
View in Google Scholar

Lahr M, Foley R. 1994. Multiple dispersals and modern human origins. Evol Anthropol 3:48–60.
View in Google Scholar

Lahr M, Foley R. 1998. Towards a theory of modern human origins: geography, demography, and diversity in recent human evolution. Yearb Phys Anthropol 41:137–76.
View in Google Scholar

Lieberman D. 1995. Testing hypotheses about recent human evolution from skulls. Curr Anthropol 36: 159–97.
View in Google Scholar

Lieberman D. 2011. The evolution of the human head. Belknap Press of Harvard University Press.
View in Google Scholar

Li ZY, Wu XJ, Zhou LP, Liu W, Gao X, Nian XM, and others. 2017. Late Pleistocene archaic human crania from Xuchang, China. Science 355:969–72.
View in Google Scholar

Manderscheid E, Rogers A. 1996. Genetic admixture in the Later Pleistocene. Am J Phys Anthropol 100:1–5.
View in Google Scholar

Manzi G. 2011. Before the emergence of Homo sapiens: overview on the Early-to-Middle Pleistocene fossil record (with a proposal about Homo heidelbergensis at the subspecific level). Int J Evol Biol, Review Article:1–12.
View in Google Scholar

Meyer M, Kircher M, Gansauge M, Li H, Racimo F, Mallick S, et al. 2012. A high-coverage genome sequence from an archaic Denisovan individual. Science 338:222–6.
View in Google Scholar

Meyer M, Fu Q, Aximu-Petri A, Glocke I, Nickel B, Arsuaga JL, et al. 2014. A mitochondrial genome sequence of a hominin from Sima de los Huesos. Nature 505(7483):403–18.
View in Google Scholar

Mondal M, Bertranpetit J, Lao O. 2019. Approximate Bayesian computation with deep learning supports a third archaic introgression in Asia and Oceania. Nature Communications 10:246.
View in Google Scholar

Mourthe I, Trindade R, Aguiar L, Trigo T, Bicca-Marques J, Bonatto S. 2019. Hybridization between neotropical primates with contrasting sexual dichromatism. Int J Primatol 40(1):99–113.
View in Google Scholar

Nei M, Roychoudhury A. 1993. Evolutionary relationships of human populations on a global scale. Mol Biol Evol 10: 927–43.
View in Google Scholar

Pearson O, Grine F, Barham L, Stringer C. 2000. Human postcranial remains from the Middle and Later Stone Age of Mumbwa Caves. L Barham, editor. The Middle Stone Age of Zambia, South Central Africa. Bristol, Western Academic & Specialist Press. 149–64.
View in Google Scholar

Peyrégne S, Slon V, Mafessoni F, de Filippo C, Hajdinjak M, Nageli S and others. 2019. Nuclear DNA from two early Neandertals reveals 80,000 years of genetic continuity in Europe. Sci. Adv. 5: eaaw5873.
View in Google Scholar

Ponce De León M, Zollikofer C.P. 2001. Neanderthal cranial ontogeny and its implications for late hominid diversity. Nature 412(6846):534–8.
View in Google Scholar

Prüfer K, Racimo F, Patterson N, Jay F, Sankararaman S, Sawyer S, et al. 2014. The complete genome sequence of a Neanderthal from the Altai Mountains. Nature 505(7481):43–54.
View in Google Scholar

Racimo F, Sankararaman S, Nielsen R, Huerta-Sánchez E. 2015. Evidence for archaic adaptive introgression in humans. Nat Rev Genet 16:359–71.
View in Google Scholar

Relethford J. 2001. Genetics and the search for modern human origins. New York, Wiley-Liss.
View in Google Scholar

Rightmire GP. 1996. The human cranium from Bodo, Ethiopia: evidence for speciation in the Middle Pleistocene? J. Hum Evol 31(1):21–39.
View in Google Scholar

Rightmire GP. 1998. Human evolution in the Middle Pleistocene: the role of Homo heidelbergensis. Evol Anthropol 6(6):218–27.
View in Google Scholar

Rightmire GP. 2008. Homo in the middle pleistocene: hypodigms, variation, and species recognition. Evol Anthropol 17(1):8–21.
View in Google Scholar

Rogers A, Bohlender R, Huff C. 2017a. Early history of Neanderthals and Denisovans. Proc Natl Acad Sci U.S.A 114(37):9859–63.
View in Google Scholar

Rogers A, Bohlender RJ, Huff C. 2017b. Reply to Mafessoni and Prüfer: Inferences with and without singleton site patterns. Proc Natl Acad Sci U.S.A 114(48):E10258–60.
View in Google Scholar

Rougier H, Milota S, Rodrigo R, Gherase M, Sarcina L, Moldovan O. And others. 2007. Peştera cu Oase 2 and the cranial morphology of early modern Europeans. Proc Natl Acad Sci U.S.A 104(04):1165–70.
View in Google Scholar

Sankararaman S, Patterson N, Li H, Päabo S, Reich D. 2012. The data of interbreeding between Neanderthals and Modern Humans. PLoS Genetics 8(10):1–9.
View in Google Scholar

Sankararaman S, Mallick S, Dannemann M, Prüfer K, Kelso J, Pääbo S et al. 2014: The genomic landscape of Neanderthal ancestry in present-day humans. Nature 507(7492):354–7.
View in Google Scholar

Sankararaman S, Mallick S, Patterson N, Reich D. 2016. The Combined Landscape of Denisovan and Neanderthal Ancestry in Present-Day Humans. Curr Biol 26:1241–7.
View in Google Scholar

Schwartz J, Tattersall I. 2002 Bodo and the concept of Homo heidelbergensis. H Seidler, K Begashaw, editors. 25 years of Bodo, Proceedings of the 4th Phillip V Tobias Lecture on Human Evolution. Addis Ababa: National Museum 107–27.
View in Google Scholar

Schwartz J, Tattersall I. 2005. The Human Fossil Record, Craniodental Morphology of Early Hominids (Genera Australopithecus, Paranthropus, Orrorin) and Overview (Volume 4). New York, Wiley Liss.
View in Google Scholar

Schwartz J, Tattersall I. 2010 Fossil evidence for the origin of Homo sapiens. Yearb Phys Anthropol 53:94–121.
View in Google Scholar

Serre D, Pääbo S. 2004. Evidence for gradients of human genetic diversity within and among continents. Genome Res 14:1679–85.
View in Google Scholar

Simonti C, Vernot B, Bastarache L, Bottinger E, Carrell D, Chisholm R, and others. 2016. The phenotypic legacy of admixture between modern humans and Neandertals. Science 351:737–741.
View in Google Scholar

Sjödin P, Sjöstrand A, Jakobsson M, Blum M. 2012. Resequencing data provide no evidence for a human bottleneck in Africa during the penultimate glacial period. Mol Biol Evol 29(7):1851–60.
View in Google Scholar

Smith F. 2010. Species, populations, and assimilation in later human evolution. In: CS Larsen editor. A Companion to Biological Anthropology. Oxford: Wiley-Blackwell. 357–78.
View in Google Scholar

Smith F, Ahern J. 2013. Introduction: thoughts on Modern Humans origins: from 1984 to 2012. In: FH Smith, C Ahern, editors. The Origins of Modern Humans, Biology Reconsidered. New Jersey, John Wiley & Sons. XI–XXV.
View in Google Scholar

Smith T, Tafforeau P, Reid D, Grün R, Eggisn S, Boutakiout M. et al. 2007. Earliest evidence of modern human life history in North African early Homo sapiens. Proc Natl Acad Sci U.S.A 104(15):6128–33.
View in Google Scholar

Stewart J, Stringer C. 2012. Human evolution out of Africa: the role of refugia and climate change. Science 335:1317–21.
View in Google Scholar

Stock J, Lahr M, Kulatilake S. 2007. Cranial diversity in South Asia relative to modern human dispersals and global patterns of human variation. In: MD Petraglia, B Allchin, editors. The Evolution and History of Human Populations in South Asia. Springer 245–68.
View in Google Scholar

Stoneking M, Sherry ST, Redd AJ, Vigilant L. 1992. New approaches to dating suggest a recent age for the human mtDNA ancestor. Philos Trans Royal Soc B 337:167–76.
View in Google Scholar

Stringer C, Andrews P. 1988. Genetic and fossil evidence for the origin of modern humans. Science 239:1263–8.
View in Google Scholar

Stringer C. 2002. Modern human origins: progress and prospects. Philos Trans Royal Soc B 357(1420):563–79.
View in Google Scholar

Stringer C. 2006. The origins of modern humans 1984–2004. In: H Soodyall, editor. The prehistory of Africa. Johannesburg, Jonathan Ball 10–20.
View in Google Scholar

Stringer C. 2012. The status of Homo heidelbergensis (Schoetensack 1908). Evol Anthropol 21:101–7.
View in Google Scholar

Stringer C. 2014. Why we are not all multiregionalists now. Trends Ecol Evolut 29:248–51.
View in Google Scholar

Stringer C. 2016. The origin and evolution of Homo sapiens. Philos Trans Royal Soc B 371(1698):1–12.
View in Google Scholar

Tattersall I. 1986. Species recognition in human paleontology. J. Hum Evol 15:165–75.
View in Google Scholar

Tattersall I. 2007. Neanderthals, Homo sapiens, and the question of species in paleo-anthropology. Journal of Anthropological Species 85:139–46.
View in Google Scholar

Tattersall I, Schwartz J. 2000. Extict Humans. Boulder, Westview Press CO.
View in Google Scholar

Tattersall I, Schwartz J. 2006. The distinctiveness and systematic context of Homo neanderthalensis. In: K Harvati, T Harrison, editors. Neanderthals Revisited. New York, Springer. 9–22.
View in Google Scholar

Thorne A, Wolpoff M. 1981. Regional continuity in Australasian Pleistocene hominid evolution. Am J Phys Anthropol 55:337–49.
View in Google Scholar

Thorne A, Wolpoff M. 1992. The multiregional evolution of humans. Sci Am 266:76–83.
View in Google Scholar

Trinkaus E. 1981. Neanderthal limb proportions and cold adaptation. C Stringer, editor. Aspects of Human Evolution. London, Taylor & Francis 187–224.
View in Google Scholar

Trinkaus E. 1983. The Shanidar Neandertals. New York, Academic.
View in Google Scholar

Trinkaus E. 1986. The Neandertals and modern human origins. Annu Rev Anthropol 15:193–218.
View in Google Scholar

Trinkaus E. 1992. Morphological contrasts between the Near Eastern Qafzeh-Skhūl and late archaic human samples: grounds for a behavioral difference? In: T Akazawa, K Aoki, T Kimura, editors. The Evolution and Dispersal of Modern Humans in Asia. Tokyo, Hokusen-Sha Publishers 277–94.
View in Google Scholar

Underhill P, Passarino G, Lin A, Shen P, Foley R, Lahr M. 2001. The phylogeography of Y chromosome binary haplotypes and the origins of modern human populations. Ann Hum Genet 65:43–62.
View in Google Scholar

Vallois H. 1954. Neanderthals and presapiens. J Royal Anthropol Inst 84:11–30.
View in Google Scholar

Vernot B, Akey J. 2014. Resurrecting surviving Neandertal lineages from modern human genomes. Science 343:1017–21.
View in Google Scholar

Vernot B, Tucci S, Kelso J, Schraiber J, Wolf A, Gittelman R, et al. 2016. Excavating Neandertal and Denisovan DNA from the genomes of Melanesian individuals. Science 352:235–9.
View in Google Scholar

Vervaecke H, Elsacker V. 1992. Hybrids between common chimpanzees (Pan troglodytes) and pygmy chimpanzees (Pan paniscus) in captivity. Mammalia 56(4):667–9.
View in Google Scholar

Vigilant L, Stoneking M, Harpending H, Hawkes K, Wilson A. 1991. African populations and the evolution of human mitochondrial DNA. Science 253:1503–8.
View in Google Scholar

Weaver T. 2012: Did a discrete event 200.000–100.000 years ago produce modern humans? J Hum Evol 63:121–6.
View in Google Scholar

Weaver T, Hublin JJ. 2009. Neandertal birth canal shape and the evolution of human childbirth. Proc Natl Acad Sci U.S.A 106 (20): 8151–8156.
View in Google Scholar

Wei-Haas M. 2019. Ancient DNA reveals new twists in Neanderthal migration: genetc surprises pulled from 120.000-year-old bones showcase the nuanced history of this close human relative. Nature-Science & Innovation. Retrieved from: https://www.nationalgeographic.com/science/2019/06/neanderthals-spread-across-europe-asia-gets-new-twist-from-ancient-dna/
View in Google Scholar

Weidenreich F. 1947. Fact and speculations concerning the origin of Homo sapiens. Am Anthropol 49:39–48.
View in Google Scholar

Wolpoff M. 1980a. Paleoanthropology. New York, Knopf.
View in Google Scholar

Wolpoff M. 1980b. Cranial remains of Middle Pleistocene European Hominids. J Hum Evol 9(5):339–58.
View in Google Scholar

Wolpoff M. 1986. Describing anatomically modern Homo sapiens: a distinction with-out a definable difference. V.V. Novotný, A. Mizerová (Eds.): Fossil Man: New Facts, New Ideas, Papers in Honor of Jan Jelínek’s Life Anniversary. Anthropos (Brno) 23:41–53.
View in Google Scholar

Wolpoff M, Thorne A, Smith F, Frayer D, Pope G. 1994. Multiregional evolution: a worldwide source for modern human populations. M Nitecki, D Nitecki, editors. Origins of anatomically modern humans. New York, Plenum. 175–99.
View in Google Scholar

Wolpoff M, Hawks J, Frayer D, Hunley K. 2001. Modern human ancestry at the peripheries: a test of the replacement theory. Science 291:293–7.
View in Google Scholar

Wood B. 1991. Koobi Fora Research Project, Volume 4: Hominid Cranial Remains. Oxford, Clarendon Press.
View in Google Scholar

Zilhão J, Trinkaus E. 2002. A brief history. In: J Zilhão, E Trinkaus, editors. Portrait of the artist as a child: the gravettian human skeleton from the Abrigo do Lagar Velho and its archaeological context. Ministério da Cultura: Instituto Português de Arqueologia. 13–27.
View in Google Scholar

Zilhão J, Constantin S, Danciu A, Rodrigo R, Trinkaus E, Gherase M, et al. 2007. The Peştera cu Oase people, Europe’s earliest modern humans. In. P Mellars, K Boyle, O Bar-Yosef, C Stringer, editors. Rethinking the Human Revolution. Cambridge Mc-Donald Institute for Archaeological Research. 249–62.
View in Google Scholar