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date: 24 June 2021

Modern Human Behaviorfree

Modern Human Behaviorfree

  • Pamela R. WilloughbyPamela R. WilloughbyUniversity of Alberta


In evolutionary terms, a modern human is a member of our own species, Homo sapiens. Fossil skeletal remains assigned to Homo sapiens appear possibly as far back as 300,000 or 200,000 years ago in Africa. The first modern human skeletal remains outside of that continent are found at two sites in modern Israel, the Mugharet es Skhūl and Jebel Qafzeh; these date between 90,000 and 120,000 years ago. But this just represents a short, precocious excursion out of Africa in an unusually pleasant environmental phase. All humans who are not of direct sub-Saharan African ancestry are descended from one or more populations who left Africa around 50,000 years ago and went on to colonize the globe. Surprisingly, they successfully interbred with other kinds of humans outside of Africa, leaving traces of their archaic genomes still present in living people. Modern human behavior, however, implies people with innovative technologies, usually defined by those seen with the earliest Upper Paleolithic people in Eurasia. Some of these innovations also appear at various times in earlier African sites, but the entire Upper Paleolithic package, once known as the Human Revolution, does not. Researchers have had to split the origin of modern biology and anatomy from the beginnings of modern cultural behavior. The first clearly evolves much earlier than the latter. Or does it?


  • Archaeology
  • Biological Anthropology

The Context of Human Evolution

Archaeology can be defined as the study of past human cultural behavior. Archaeologists examine this past through the study of material remains; these are composed of individual artifacts, information about how they are produced and used, as well as the sites where past activities took place. The context in which artifacts are found is critical for discovering information about the evolution of technology and behavior over the course of human history.

The study of the earliest artifacts and the sites from which they have been recovered is also part of paleoanthropology, the biocultural study of human evolution. Paleoanthropology was developed by F. Clark Howell in the 1960s to describe the kind of research he and his colleagues were conducting in the Omo River valley in southern Ethiopia (Coppens et al. 1976). This work was both multi- and interdisciplinary, involving geologists, geochronologists, paleoenvironmental specialists, paleontologists, and archaeologists. It did not produce lots of fossil hominins, but it revolutionized how researchers study human evolution.

Archaeology and paleoanthropology overlap when one deals with the first stages of cultural evolution, the Paleolithic. The Paleolithic, or Old Stone Age, is the earliest stage of technology. It is subdivided into three phases—Lower (Oldowan and Acheulean), Middle, and Upper Paleolithic. In Africa south of the Sahara, researchers use Early, Middle, and Later Stone Age (ESA, MSA, and LSA) to refer to the similar Paleolithic stages. In the past two decades, from the early 2000s on, North African researchers have begun to adopt the same terms as have been used elsewhere in the continent.

The notion of Paleolithic stages was first centered on the study of the most abundant finds, flaked stone artifacts. Individual pieces of flaked stones are artifacts, a term used for anything made or used by humans. A group of artifacts from the same context makes up an assemblage, and groups of similar assemblages make up an industry or industrial complex. The kinds of flaked stone artifacts change over time and are used to define both assemblages and industries. In turn, this led to descriptions of the origin and evolution of key aspects of human cultural and social behavior.

What Does It Mean to be a Modern Human?

What it means to be modern in evolutionary terms depends on what one is looking at. Paleontologists and biological anthropologists describe skeletal remains that they label as modern. Archaeologists define stages of the evolution of modern technology. Primatologists study the evolution of both social behavior and tool use in non-human primates, notably chimpanzees and New World capuchin monkeys. As a result, there are many ways to describe modernity in evolutionary terms. In human evolutionary terms, there are both anatomical and behavioral definitions of modernity.

The biological classification of a modern human goes back to Carl von Linné or Carolus Linnaeus and the 10th edition of his Systema Naturae (Linnaeus 1758); in historical terms, this represents year one of biological classification. In his master classification of life, Linnaeus included living humans in a species he labeled Homo sapiens, or “wise man.” His basic definition was Nosce te ipsum, or “know thyself,” which is a Latin translation of the Greek phrase carved over the entrance to the Temple of Delphi. In other words, look around you, and when you see people, they’re representative of Homo sapiens. While he is rightly seen as the founder of biological systematics, Linnaeus was a creationist who thought that he was cataloguing God’s purpose in Creation (Chazan 1995; Ingold 1995). Later researchers included fossil examples within the species and defined Homo sapiens on anatomical features which distinguished them from earlier hominins. Interestingly, there is still no type specimen. The famous 19th-century American vertebrate paleontologist E. D. Cope left his body to science to be designated the type specimen, but it hasn’t happened. These features include (1) an average brain size of over 1,350 cc; (2) a vertical frontal (forehead); (3) a high cranial vault; (4) reduced brow ridges above the eyes; and (5) a face that is flat and tucked under. (6) The face is reduced in size and projection, possibly due to less use of teeth as tools. (7) The maxilla or upper jaw exhibits a canine fossa, and (8) mandibles vary in robusticity. (9) The postcrania (the rest of the skeleton minus the skull and mandible) are less robust than earlier, with long limbs and slender bodies. The latter reflects tropical body proportions (Holt and Formicola 2008; Lieberman 2011; Pearson 2000; Stringer 2016; Willoughby 1994, 2007).

The Paleolithic and the Evolution of Modern Behavior

When examining archaeological material, modernity meant archaeological material directly associated with Homo sapiens. The term “Paleolithic” was defined in the 19th century by Sir John Lubbock (later Lord Avebury) as the first stage of the Stone Age (Lubbock 1865). It initially referred to artifacts from European open air and cave sites which were made of flaked stone. (This contrasted them with the ground or polished stone tools more common in the subsequent Neolithic or New Stone Age.) The age of Paleolithic sites was initially inferred by association with extinct Ice Age animal remains. The earliest stages of the Paleolithic are clearly exclusively African. By 1.7 million years ago, hominins belonging to the species Homo erectus dispersed out of Africa and colonized large parts of Asia and Europe. They were followed by a second species, referred to as Homo heidelbergensis or “archaic Homo sapiens.” In Europe, Homo heidelbergensis evolved into Neanderthals (and into possibly other groups such as the Denisovans) while in Africa the same species evolved directly into what is referred to as anatomically modern Homo sapiens, ourselves (Stringer 2016).

The artifacts associated with some Homo erectus and Homo heidelbergensis fossils belong to the Acheulean Industrial Complex. Named for Saint Acheul in northwestern France, the Acheulean ranges in age from 1.8 Mya to around 200,000 years ago in Africa, and from toward the end of this range in Eurasia. Diagnostic tools of the Acheulean are bifaces manufactured on cores or flakes and classified as hand axes, cleavers, trihedral forms, and picks (De la Torre 2016). Simple modified cores or choppers persist from earlier Oldowan and pre-Oldowan industries, as do flakes detached from such cores (Toth and Schick 1986, 2018). Around 500,000 years ago, there is the first significant change in Acheulean technology. This is marked by the emergence of Levallois methods of flake tool production and is the first example of “prepared core” technology. In this case, the core is shaped in a predetermined way in order to produce standardized flakes, blades, or points. These are used without any further modification. Alternately, hominins struck flakes off of cores and then retouched them into finished tools. Both methods continue into the subsequent Middle Paleolithic (in Africa called the MSA or Middle Stone Age) (Wadley 2015). It ranges from at least 200,000 to 30,000 years ago. Middle Paleolithic artifacts are associated with Neanderthals and other groups, such as Homo heidelbergensis and the Siberian Denisovans, in Eurasia and the Middle East. But as became clear in the late 1980s, they are directly associated with anatomically modern Homo sapiens in Africa and at two sites in Israel, the Mugharet es Skhūl and Jebel Qafzeh; these date between 90,000 and 120,000 years ago (Valladas et al. 1988; Mercier et al. 1993). This created a new problem for archaeologists, as modern anatomy and modern behavior were supposed to evolve together.

Middle Paleolithic sites do show some cultural and technological innovations. The first intentional burials occurred at this time, and there are numerous examples of social welfare and group care, such as the survival of individuals with significant evidence of trauma at sites like Shanidar Cave in Iraqi Kurdistan (Trinkaus 1983). Like their late Acheulean predecessors, they hafted a variety of stone tools onto organic handles, creating the first composite tools, but their achievements have traditionally been compared to those from the Upper Paleolithic and found wanting. This is because Upper Paleolithic sites were long thought to be the earliest ones associated with anatomically modern humans like ourselves (Holt and Formicola 2008), and people like us should have been able to create the full range of material culture observed today. The Upper Paleolithic begins around 50,000 years ago and lasts until the end of the Pleistocene around 10,000 years ago. It was first identified in sites such as L’Aurignac and Cro-Magnon, in southern France. Comprehensive reviews of the Eurasian Upper Paleolithic have been written by a number of researchers such as Bar Yosef (2002) , Straus (1995) , and Zilhão (2007) . What all of them still emphasize are the innovations which are supposed to appear in the Upper Paleolithic for the first time. A list of these would include: (1) blade and bladelet technology; (2) microlithic technology; (3) the first portable art, comprising figurines of women (Venuses) and animals; (4) the first parietal (or wall) art, consisting of paintings and engravings in caves (Cook 2013); (5) the earliest bone, ivory, and antler tools; (6) ochre or other pigment processing; (7) specialization in hunting one or two species of animals; (8) the use of fish or shellfish; (9) more complex burials; (10) long-distance exchange of raw materials and finished items; (11) the first information networks and emergence of ethnic groups; and (12) the spread of people into new regions, such as Siberia, the Americas, and Australia. Based on worldwide hunter-gatherer ethnographic data, Binford (1980) also argued that only modern people could organize themselves using a collector strategy of mobility, where one brings resources to people. As a result, people could be permanently settled year-round in the same community, which is the opposite of what he called the forager strategy, where people moved seasonally to resources (Binford 1980, 1989). Thus only Upper Paleolithic “modern” hunter-gatherers could have permanent settlements.

The identification of a whole suite of innovations with the onset of the Upper Paleolithic led to the creation of a model that was unquestioned until the late 1980s. This model has been variously labeled the human revolution, the creative revolution, the dawn of belief, or the quantum leap (Dickson 1990; Mellars 1989; Mellars et al. 2007). For some researchers, the appearance of the Upper Paleolithic in Eurasia at the same time as modern humans was not a coincidence (Mellars 2005). But for others, most of the innovations said to be unique to this period had actually occurred earlier (D’Errico 2003; D’Errico et al. 2003; Nowell 2010). However, as it became clear that there were anatomically modern people in the African Middle Paleolithic as well as at Skhūl and Qafzeh, ideas started to change. While the Upper Paleolithic was still the period associated with modern innovations, it began to be recast as behavioral modernity.

The Role of Genetic Research in Identifying Modern Humans

The separation of anatomical from behavioral modernity began with genetic data that first appeared in the 1960s and 1970s. Blood samples collected from people worldwide were analyzed for a variety of population-specific markers; however, the results did not fall into clear geographic categories, so-called human races. In fact, humans expressed very little between-group variation (Lewontin 1972; Cavalli-Sforza et al. 1994). W. W. Howells (1976) , a prominent biological anthropologist, used these data, along with a survey of fossil modern remains, to argue that there were only two models for the origins of Homo sapiens. Either there had been a single center of origin, or multiple centers. He labeled the first the Noah’s Ark hypothesis, the latter, the Candelabra model (Howells 1976; Willoughby 2007, 48-49). For him, the single center of origin made most sense in Ernst Mayr’s (1942) evolutionary context, which emphasized geographic isolation of a small subset of a species. This is a remarkable conclusion, one that was made more than a decade before new methods allowed for a resolution of the question.

About a decade after Howells wrote his seminal paper, it was genetic data that again reinforced the single-center model. A single study pointed to a solution for the origins of modern humans, both biological and cultural–behavioral. In 1987, Cann, Stoneking, and Wilson published a revolutionary paper (Cann et al. 1987). They examine mitochondrial DNA from a number of living people worldwide. Mitochondria are organelles in animal cells which convert sugar to energy. They contain DNA that is passed on from mother to child. Small bits of this DNA show variation that reflects ancestry. It is these bits that showed a common ancestry for all of the people sampled, some time between 143,000 and 288,000 years ago. Africans showed the most variation as a group, implying that they represented descendants of the original ancestral group. The first sign of a mitochondrial signature outside of Africa was much younger, eventually dated to around 50,000 years ago (Harpending et al. 1993; Relethford 2001; Reich 2018). In addition, when compared to the living great apes, the mitochondrial DNA of modern people exhibits very few differences in the “hypervariable” regions that reflect history and ancestry (Gagneux et al. 1999). Subsequent studies of both mitochondrial and nuclear DNA have confirmed the conclusions of the original authors.

Within a year of Cann et al.’s original mitochondrial paper, some paleoanthropologists were stating that the problem of modern human origins was solved (most notably, Stringer and Andrews, 1988). Chris Stringer (2015, 2016) then proposed the Out of Africa 2, or the replacement, theory, basically Howell’s single-center model reborn for an era with more testable data.

Studies of the DNA of ancient hominins began with extraction of mitochondrial DNA. The original Neanderthal specimen from the Feldhofer Cave in Germany yielded one of the first usable samples, but its mitochondrial DNA was no closer to that of living Europeans than of human populations elsewhere (Krings et al. 1997; see also Pääbo 2014). It was only when nuclear DNA could be extracted that a different picture of Neanderthals emerged. Now there was proof of interbreeding with early modern humans (Green et al. 2010). In addition, DNA from Denisova Cave in southern Siberia led to the identification of one or two new hominins: the Denisovans (whose entire fossil inventory was a few fragments) as well as some other “unknown” humans of East Asian ancestry. At Denisova Cave, it appeared that four kinds of humans coexisted and that they all interbred with each other (Prüfer et al. 2014; Reich et al. 2010). Geneticists still support the Out of Africa 2 model, but modify it to be replacement with hybridization rather than complete replacement of indigenous Eurasians by modern Africans.

Out of Africa 2 and the Origins of Modern Human Behavior

For archaeologists, this mitochondrial Eve or Out of Africa 2 model presented a novel issue. Suddenly there was a number of attempts to re-examine modern human biological and behavioral origins in the light of a recent African ancestry (e.g., Mellars 1990; Mellars and Stringer 1989; Mellars et al. 2007). At the time, Paleolithic archaeologists looked to Europe or the Levant for the origins of both modern anatomy and behavior. Modern anatomy clearly evolved in Africa. Could modern behavior have done so as well?

Various reviews of the African evidence have appeared in the past two decades, from the early 2000s. Some challenge the prevailing ideas about modern humans as simplistic (Shea 2011a, 2011b), while others have tried to identify Upper Paleolithic-like achievements in earlier African sites such as Klasies River and Blombos Cave (Henshilwood and Marean 2003; McBrearty and Brooks 2000; Marean 2015). An exclusively African origin was supported by re-dating of sites once thought to lie right at the Middle and Upper Paleolithic boundary. This boundary was also the effective limit of radiocarbon or 14C dating. Newer chronometric dating methods, introduced in the late 1980s (Reynolds 1991; Wintle 1996), showed that the earliest anatomically modern humans appeared quite early in Africa as well as at Skhūl and Qafzeh in the Levant (Valladas et al. 1987, 1988). The latter represented a short dispersal, dating to Marine Isotope Stage (MIS) 5e, the last time it was as warm as at present. They would not leave Africa again until the next interstadial, MIS stage 3 around 50,000 years ago (Hetherington and Reid 2011; Raymo and Hubers 2008). This time it would last, as Africans spread out to colonize the entire globe.

The African Origins of Modern Humans

Is it only anatomical modernity, or behavioral modernity too, that evolved in Africa? The first modern Africans are associated with Middle Paleolithic (Modern Stone Age, or MSA) artifacts, not Upper Paleolithic (or Later Stone Age, LSA) ones (Willoughby 2009). African MSA humans were originally seen as the people without culture, modern anatomically but Middle Paleolithic culturally, more like Neanderthals (Willoughby 2000). Before new dating methods confirmed that they belonged to the MSA, it was felt that these sites were of Upper Paleolithic age. As a result, it was argued that modern Africans had failed to develop the same abilities as their Eurasian counterparts, but the new focus on Africa led to a number of publications that reviewed the evidence for Upper Paleolithic-like innovations in the MSA (Henshilwood and Marean 2003; McBrearty and Brooks 2000; Willoughby 1993, 2007). These include features such as the following: (1) the use of the coast and coastal resources at sites like the Abdur Limestone locality in Eritrea (Walter et al. 2000) as well as in South Africa (Henshilwood and Marean 2003); (2) the presence of ostrich eggshell beads at sites like Enkapune ya Muto, Kenya (Ambrose 1998b) and Magubike, Tanzania (Miller and Willoughby 2014), as well as decorated ostrich eggshell containers at Diepkloof Cave in South Africa (Texier et al. 2010); (3) Nassarius marine shell beads in a number of sites in both North Africa and South Africa (Steele et al. 2019); (4) stylistic variation in stone tool technology in industries such as the Stillbay and Howieson’s Poort in South Africa (Henshilwood and Dubreuil 2011) and the Aterian in North Africa (Scerri 2013; Scerri et al. 2018); (5) shellfish and fish use at a variety of South African coastal sites; (6) organic tools including bone points at sites like Sibudu, Klasies River, Blombos, and Peers Cave, all in South Africa (Henshilwood 2007), as well as in North Africa (Bouzouggar et al., 2018); (7) worked red ochre common in MSA sites, and some engraved pieces have been recovered, such as the distinctive hatch-marked one from Blombos; (8) the critical association of modern human skeletal remains at a number of sites, including one of the author’s—Magubike rock shelter in southern Tanzania (Willoughby et al. 2018; Grine 2016).

Was There a Human Revolution?

There are elements of what can be recognized as analogous to Eurasian Upper Paleolithic innovations in a variety of African MSA sites but these came and went over time (Stringer 2015), and do not comprise a single package. The archaeological sign in the Levant and Europe of the arrival of modern humans (the Aurignacian) is not found anywhere in Africa. Thus it is possible that what appears as the human revolution outside of Africa could have evolved in response to the successful dispersal of modern people out of the continent. Perhaps their encounters with new peoples, Neanderthals, Denisovans, and possibly others, stimulated technological, social, and behavioral changes.

One question remains. If modern humans evolved in Africa by at least 200,000 years ago, why is there no sign of them anywhere else until around 50,000 years ago? Some researchers feel that they had to undergo some neurological or biological change before they truly became modern (Klein 1998, 2008). Others feel that environmental stress in the Late Pleistocene affected population size and distribution, effectively blocking any significant dispersals (Mellars 2006; Willoughby 2012). This could have been the result of the Mount Toba eruption on the island of Sumatra in modern Indonesia around 74,000 years ago (Ambrose 1998a), which is described as the most significant volcanic event since the meteorite hit the earth and wiped out the dinosaurs and other organisms 65 million years ago. Or it could be a product of the regular environmental stress which occurred time and again in the sub-Saharan African Upper Pleistocene (Cohen et al. 2007; Scholz et al. 2007). It was Harpending et al. (1993) who first proposed that there was a significant bottleneck or almost-extinction event possibly associated with the Out of Africa 2 migration.

Whatever the cause, it is clear that modern anatomy evolved well before modern behavior, if one takes the Upper Paleolithic model as the benchmark. Whatever else, the process of becoming behaviorally modern in evolutionary terms was a complex one which will take a lot of additional research to elucidate. Africa is the continent in which the earliest hominins and the oldest material culture are found. It is also the place where the origins of our own species occurred. This discovery has totally changed the focus of much of African Stone Age research. Meanwhile, it is worthy of note that if European scholars had not been the first to study the Paleolithic, our notion of what it means to be modern may have been quite different.

Further Reading

  • Bar-Yosef, Ofer. 2002. “The Upper Palaeolithic Revolution.” Annual Review of Anthropology 31: 363–393.
  • Calvin, William. 2002. A Brain for All Seasons: Human Evolution and Abrupt Climate Change. Chicago: University of Chicago Press.
  • Cook, Jill. 2013. Ice Age Art: Arrival of the Modern Mind. London: The British Museum Press.
  • Drell, Julia R. R. 2000. “Neanderthals: A History of Interpretation.” Oxford Journal of Archaeology 19 (1): 1–24.
  • Henshilwood, Christopher S., and Francesco D’Errico. 2005. “Being Modern in the Middle Stone Age: Individuals and Innovation.” In The Hominid Individual in Context, edited by Clive Gamble and Martin Porr, 244–264. London: Routledge.
  • Henshilwood, Christopher S., and Francesco D’Errico, eds. 2011. Homo Symbolicus: The Dawn of Language, Imagination and Spirituality. Amsterdam: John Benjamins.
  • Lieberman, Daniel E. 2011. The Evolution of the Human Head. Cambridge, MA: Harvard University Press.
  • Mellars, Paul. ed. 1990. The Emergence of Modern Humans: An Archaeological Perspective. Ithaca, NY: Cornell University Press.
  • Mellars, Paul, Katie Boyle, Ofer Bar-Yosef, and Chris Stringer, eds. 2007. Rethinking the Human Revolution: New Behavioural and Biological Perspectives on the Origin and Dispersal of Modern Humans. Cambridge, UK: McDonald Institute for Archaeological Research.
  • Mellars, Paul, and Chris Stringer, eds. 1989. The Human Revolution: Behavioural and Biological Perspectives on the Origins of Modern Humans. Princeton, NJ: Princeton University Press.
  • Noble, William, and Iain Davidson. 1996. Human Evolution, Language and Mind. Cambridge, UK: Cambridge University Press.
  • Pääbo, Svante. 2014. Neanderthal Man: In Search of Lost Genomes. New York: Basic Books.
  • Reich, David. 2018. Who We Are and How We Got Here: Ancient DNA and the New Science of the Human Past. New York: Pantheon.
  • Smith, Fred H., and James C. M. Ahern, eds. 2013. The Origins of Modern Humans: Biology Reconsidered. Hoboken, NJ: Wiley.
  • Smith, Fred H., and Frank Spencer, eds. 1984. The Origins of Modern Humans: A World Survey of the Fossil Evidence. New York: Alan R. Liss.
  • Stringer, Chris B. 2011. The Origin of Our Species. London: Allen Lane.
  • Stringer, Chris, and Robin McKie. 1996. African Exodus: The Origins of Modern Humanity. London: Jonathan Cape.
  • Trinkaus, Erik, and Pat Shipman. 1993. The Neandertals: Changing the Image of Mankind. New York: Knopf.
  • Willoughby, Pamela R. 2007. The Evolution of Modern Humans in Africa: A Comprehensive Guide. Lanham, MD: AltaMira Press.
  • Wintle, Ann G. 1996. “Archaeologically Relevant Dating Techniques for the Next Century: Small, Hot and Identified by Acronyms.” Journal of Archaeological Science 23 (1): 123–138.


  • Ambrose, Stanley H. 1998a. “Late Pleistocene Human Population Bottlenecks, Volcanic Winter, and Differentiation of Modern Humans.” Journal of Human Evolution 34 (6): 623–651.
  • Ambrose, Stanley H. 1998b. “Chronology of the Later Stone Age and Food Production in East Africa.” Journal of Archaeological Science 25 (4): 377–392.
  • Bar-Yosef, Ofer. 2002. “The Upper Palaeolithic Revolution.” Annual Review of Anthropology 31: 363–393.
  • Binford, Lewis R. 1980. “Willow Smoke and Dog's Tails: Hunter-Gatherer Settlement Systems and Archaeological Site Formation.” American Antiquity 45 (1): 4–20.
  • Binford, Lewis R. 1989. “Isolating the Transition to Cultural Adaptations: An Organizational Approach.” In The Emergence of Modern Humans, edited by Erik Trinkaus, 18–41. Cambridge, UK: Cambridge University Press.
  • Bouzouggar, Abdeljalil, Louise T. Humphrey, Nick Barton, Simon A. Parfitt, Laine C. Balzan, Jean-Luc Schwenninger, Mohammed Abdeljalil El Hajraoui, Roland Nespoulet, and Silvia M. Bello. 2018. “90,000-Year-Old Specialised Bone Technology in the Aterian Middle Stone Age of North Africa.” PLoS One 13 (10): 0202021.
  • Cann, Rebecca L., Mark Stoneking, and Allan C. Wilson. 1987. “Mitochondrial DNA and Human Evolution.” Nature 325 (6099): 31–36.
  • Cavalli-Sforza, Luigi Luca, Paolo Menozzi, and Alberto Piazza. 1994. The History and Geography of Human Genes. Princeton, NJ: Princeton University Press.
  • Chazan, Michael. 1995. “The Meaning of Homo sapiens.” In Ape, Man, Ape Man: Changing Views Since 1600, edited by Raymond Corbey and Bert Theunissen, 229–240. Leiden, The Netherlands: Leiden University Press.
  • Cohen, Andrew S., Jeffery R. Stone, Kristina R. M. Beuning, Lisa E. Park, Peter N. Reinthal, David Dettman, Christopher A. Scholz, Thomas C. Johnson, John W. King, Michael R. Talbot, Erik T. Brown, and S. J. Ivory. 2007. “Ecological Consequences of Early Late Pleistocene Megadroughts in Tropical Africa.” Proceedings of the National Academy of Sciences USA 104 (42): 16422–16427.
  • Cook, Jill. 2013. Ice Age Art: Arrival of the Modern Mind. London: The British Museum Press.
  • Coppens, Yves, F. Clark Howell, Glynn Ll. Isaac, and Richard E. F. Leakey, eds. 1976. Earliest Man and Environments in the Lake Rudolf Basin. Chicago: University of Chicago Press.
  • De la Torre, Ignacio. 2016. “The Origins of the Acheulean: Past and Present Perspectives on a Major Transition in Human Evolution.” Philosophical Transactions of the Royal Society B 371: 20150245.
  • D’Errico, Francesco. 2003. “The Invisible Frontier: A Multiple Species Model for the Origin of Behavioral Modernity.” Evolutionary Anthropology 12 (4): 188–202.
  • D’Errico, Francesco, Christopher Henshilwood, Graeme Lawson, Marian Vanhaeren, Anne-Marie Tillier, Marie Soressi, Frédérique Bresson, Bruno Maureille, April Nowell, Joseba Lakarra, Lucinda Backwell, and Michèle Julien. 2003. “Archaeological Evidence for the Emergence of Language, Symbolism, and Music–An Alternative Multidisciplinary Perspective.” Journal of World Prehistory 17 (1): 1–70.
  • Dickson, Bruce. 1990. The Dawn of Belief. Tucson: University of Arizona Press.
  • Gagneux, Pascal, Christopher Wills, Ulrike Gerlof, Diethard Tautz, Phillip A. Morin, Christophe Boesch, Barbara Fruth, Gottfried Hohmann, Oliver A. Ryder, and David S. Woodruff. 1999. “Mitochondrial Sequences Show Diverse Evolutionary Histories of African Hominoids.” Proceedings of the National Academy of Sciences USA 96 (9): 5077–5082.
  • Green, Richard E., Adrian W. Briggs, Tomislav Maricic, Udo Stenzel, Martin Kircher, Nick Patterson, Heng Li, Weiwei Zhai, Markus Hsi-Yang Fritz, Nancy F. Hansen, Eric Y. Durand, Anna-Sapfo Malaspinas, Jeffrey D. Jensen, Tomas Marques-Bonet, Can Alkan, Kay Prüfer, Matthias Meyer, Hernán A. Burbano, Jeffrey M. Good, Rigo Schultz, Ayinuer Aximu-Petri, Anne Butthof, Barbara Höber, Barbara Höffner, Madlen Siegemund, Antje Weihmann, Chad Nusbaum, Eric S. Lander, Carsten Russ, Nathaniel Novod, Jason Affourtit, Michael Egholm, Christine Verna, Pavao Rudan, Dejana Brajkovic, Željko Kucan, Ivan Gušic, Vladimir B. Doronichev, Liubov V. Golovanova, Carles Lalueza-Fox, Marco de la Rasilla, Javier Fortea, Antonio Rosas, Ralf W. Schmitz, Philip L. F. Johnson, Evan E. Eichler, Daniel Falush, Ewan Birney, James C. Mullikin, Montgomery Slatkin, Rasmus Nielsen, Janet Kelso, Michael Lachmann, David Reich, and Svante Pääbo. 2010. “A Draft Sequence of the Neandertal Genome.” Science 328 (5979): 710–722.
  • Grine, Frederick E. 2016. “The Later Quaternary Hominins of Africa: The Skeletal Evidence from MIS 6–2.” In Africa from MIS 6–2: Population Dynamics and Paleoenvironments, edited by Sacha C. Jones and Brian A. Stewart, 323–381. New York: Springer.
  • Harpending, Henry, Stephen T. Sherry, Alan R. Rogers, and Mark Stoneking. 1993. “The Genetic Structure of Ancient Human Populations.” Current Anthropology 34 (4): 483–496.
  • Henshilwood, Christopher S. 2007. “Fully Symbolic Sapiens Behaviour: Innovations in the Middle Stone Age at Blombos Cave, South Africa.” In Rethinking the Human Revolution: New Behavioural and Biological Perspectives on the Origin and Dispersal of Modern Humans, edited by Paul Mellars, Katie Boyle, Ofer Bar-Yosef, and Christopher Stringer, 123–132. Cambridge, UK: McDonald Institute for Archaeological Research.
  • Henshilwood, Christopher Stuart, and Benoît Dubreuil. 2011. “The Still Bay and Howiesons Poort, 77–59 ka: Symbolic Material Culture and the Evolution of the Mind during the African Middle Stone Age.” Current Anthropology 52 (3): 361–400.
  • Henshilwood, Christopher Stuart, and Curtis W. Marean. 2003. “The Origin of Modern Human Behavior.” Current Anthropology 44 (5): 627–651.
  • Hetherington, Renée, and Robert G. B. Reid. 2011. The Climate Connection: Climate Change and Modern Human Evolution. Cambridge, UK: Cambridge University Press.
  • Holt, Brigitte M., and Vincenzo Formicola. 2008. “Hunters of the Ice Age: The Biology of Upper Paleolithic People.” Yearbook of Physical Anthropology 51: 70–99.
  • Howells, William W. 1976. “Explaining Modern Man: Evolutionists versus Migrationists.” Journal of Human Evolution 5 (5): 477–495.
  • Ingold, Tim. 1995. “People Like Us—The Concept of the Anatomically Modern Human.” In Ape, Man, Ape Man: Changing Views Since 1600, edited by Raymond Corbey and Bert Theunissen, 241–262. Leiden, The Netherlands: Leiden University Press.
  • Klein, Richard G. 1998. “Why Anatomically Modern People Did Not Disperse from Africa 100,000 Years Ago.” In Neanderthals and Modern Humans in Western Asia, edited by Takeru Akazawa, et al., 509–521. New York: Plenum Press.
  • Klein, Richard G. 2008. “Out of Africa and the Evolution of Human Behavior.” Evolutionary Anthropology 17 (6): 267–281.
  • Krings, Matthias, Anne Stone, Ralf W. Schmitz, Heike Krainitzki, Mark Stoneking, and Svante Pääbo. 1997. “Neandertal DNA Sequences and the Origin of Modern Humans.” Cell 90 (1): 19–30.
  • Lewontin, Richard. 1972. “The Apportionment of Human Genetic diversity.” Evolutionary Biology 6: 381–398.
  • Lieberman, Daniel E. 2011. The Evolution of the Human Head. Cambridge, UK: Harvard University Press.
  • Linnaeus, Carl. 1758. Systema Naturae per Naturae Regna Tria, Secundum Classes, Ordines, Genera, Species cum Characteribus, Synonmymis, Locis, 10th ed. Stockholm: Laurentii Sylvii.
  • Lubbock, Sir John [Lord Avebury]. 1865. Pre-Historic Times, as Illustrated by Ancient Remains, and the Manners and Customs of Modern Savages. London: Williams and Norgate.
  • Marean, Curtis W. 2015. “An Evolutionary Anthropological Perspective on Modern Human Origins.” Annual Review of Anthropology 44: 433–556.
  • Mayr, Ernst. 1942. Systematics and the Origin of Species from the Viewpoint of a Zoologist. New York: Columbia University Press.
  • McBrearty, Sally, and Alison S. Brooks. 2000. “The Revolution That Wasn’t: A New Interpretation of the Origin of Modern Human Behavior.” Journal of Human Evolution 39 (5): 453–563.
  • Mellars, Paul ed. 1990. The Emergence of Modern Humans: An Archaeological Perspective. Ithaca, NY: Cornell University Press.
  • Mellars, Paul. 2005. “The Impossible Coincidence: A Single Species Model for the Origins of Modern Human Behavior in Europe.” Evolutionary Anthropology 14 (1): 12–27.
  • Mellars, Paul. 2006. “Why Did Modern Human Populations Disperse from Africa ca. 60,000 Years Ago? A New Model.” Proceedings of the National Academy of Sciences USA 103 (25): 9381–9386.
  • Mellars, Paul, Katie Boyle, Ofer Bar-Yosef, and Christopher Stringer, eds. 2007. Rethinking the Human Revolution: New Behavioural and Biological Perspectives on the Origin and Dispersal of Modern Humans. Cambridge, UK: McDonald Institute for Archaeological Research.
  • Mellars, Paul, and Christopher Stringer, eds. 1989. The Human Revolution: Behavioural and Biological Perspectives on the Origins of Modern Humans. Princeton, NJ: Princeton University Press.
  • Mercier, Norbert, Hélène Valladas, Ofer Bar-Yosef, Bernard Vandermeersch, Christopher Stringer, and J. L. Joron. 1993. “Thermoluminescence Date for the Mousterian Burial site of es Skūhl, Mt. Carmel.” Journal of Archaeological Science 20 (2): 169–174.
  • Miller, Jennifer Midori, and Pamela R. Willoughby. 2014. “Radiometrically Dated Ostrich Eggshell Beads from the Middle and Later Stone Age of Magubike Rockshelter, Southern Tanzania.” Journal of Human Evolution 74 (1): 118–122.
  • Nowell, April. 2010. “Defining Behavioral Modernity in the Context of Neandertal and Anatomically Modern Human Populations.” Annual Review of Anthropology 39: 437–452.
  • Pääbo, Svante. 2014. Neanderthal Man: In Search of Lost Genomes. New York: Basic Books.
  • Pearson, Osbjorn M. 2000. “Postcranial Remains and the Origin of Modern Humans.” Evolutionary Anthropology 9 (6): 229–247.
  • Prūfer, Kay, Fernando Racimo, Nick Patterson, Flora Jay, Sriram Sankararaman, Susanna Sawyer, Anja Heinze, Gabriel Renaud, Peter H. Sudmant, Cesare de Filippo, Heng Li, Swapan Mallick, Michael Dannemann, Qiaomei Fu, Martin Kircher, Martin Kuhlwilm, Michael Lachmann, Matthias Meyer, Matthias Ongyerth, Michael Siebauer, Christoph Theunert, Arti Tandon, Priya Moorjani, Joseph Pickrell, James C. Mullikin, Samuel H. Vohr, Richard E. Green, Ines Hellmann, Philip L. F. Johnson, Hélène Blanche, Howard Cann, Jacob O. Kitzman, Jay Shendure, Evan E. Eichler, Ed S. Lein, Trygve E. Bakken, Liubov V. Golovanova, Vladimir B. Doronichev, Michael V. Shunkov, Anatoli P. Derevianko, Bence Viola, Montgomery Slatkin, David Reich, Janet Kelso, and Svante Pääbo. 2014. “The Complete Genome Sequence of a Neanderthal from the Altai Mountains.” Nature 505 (7481): 43–49.
  • Raymo, Maureen E., and Peter Huybers. 2008. “Unlocking the Mysteries of the Ice Ages.” Nature 451 (7176): 284–285.
  • Reich, David. 2018. Who We Are and How We Got Here: Ancient DNA and the New Science of the Human Past. New York: Pantheon.
  • Reich, David, Richard E. Green, Martin Kircher, Johannes Krause, Nick Patterson, Eric Y. Durand, Bence Viola, Adrian W. Briggs, Udo Stenzel, Philip L. F. Johnson, Tomislav Maricic, Jeffrey M. Good, Tomas Marques-Bonet, Can Alkan, Qiaomei Fu, Swapan Mallick, Heng Li, Matthias Meyer, Evan E. Eichler, Mark Stoneking, Michael Richards, Sahra Talamo, Michael V. Shunkov, Anatoli P. Derevianko, Jean-Jacques Hublin, Janet Kelso, Montgomery Slatkin, and Svante Pääbo. 2010. “Genetic History of an Archaic Hominin Group from Denisova Cave in Siberia.” Nature 468 (7327): 1053–1060.
  • Relethford, John H. 2001. Genetics and the Search for Modern Human Origins. New York: Wiley.
  • Reynolds, T. E. G. 1991. “Revolution or Resolution? The Archaeology of Modern Human Origins.” World Archaeology 23 (2): 155–166.
  • Scerri, Eleanor M. L. 2013. “The Aterian and Its Place in the North African Middle Stone Age.” Quaternary International 300: 111–130.
  • Scerri, Eleanor M.L., Mark G. Thomas, Andrea Manica, Philipp Gunz, Jay T. Stock, Christopher Stringer, Matt Grove, Huw S. Groucutt, Axel Timmermann, G. Philip Rightmire, Francesco d’Errico, Christian A. Tryon, Nick A. Drake, Alison S. Brooks, Robin W. Dennell, Richard Durbin, Brenna M. Henn, Julia Lee-Thorp, Peter deMenocal, Micheal D. Petraglia, Jessica C. Thompson, Aylwyn Scally, and Lounès Chikhi. 2018. “Did Our Species Evolve in Subdivided Populations Across Africa, and Why Does It Matter?” Trends in Ecology and Evolution 33 (8): 582–594.
  • Toth, Nicholas, and Kathy Schick. 1986. “The First Million Years: The Archaeology of Protohuman Culture.” Advances in Archaeological Method and Theory 9: 1–96.
  • Toth, Nicholas, and Kathy Schick. 2018. “An Overview of the Cognitive Implications of the Oldowan Industrial Complex.” Azania 53 (1): 3–39.
  • Scholz, C. A., Thomas C. Johnson, Andrew S. Cohen, John W. King, John A. Peck, Jonathan T. Overpeck, Michael R. Talbot, Erik T. Brown, Leonard Kalindekafe, Philip Y. O. Amoako, Robert P. Lyons, Timothy M. Shanahan, Isla S. Castañeda, Clifford W. Heil, Steven L. Forman, Lanny R. McHargue, Kristina R. Beuning, Jeanette Gomez, and James Pierson. 2007. “East African Megadroughts between 135 and 75 Thousand Years Ago and Bearing on Early-Modern Human Origins.” Proceedings of the National Academy of Sciences USA 104 (42): 16416–16421.
  • Shea, John J. 2011a. “Refuting a Myth about Human Origins.” American Scientist 99: 128–135.
  • Shea, John J. 2011b. “Homo sapiens Is as Homo sapiens Was.” Current Anthropology 52 (1): 1–35.
  • Straus, Lawrence Guy. 1995. “The Upper Palaeolithic of Europe: An Overview.” Evolutionary Anthropology 4 (1): 4–16.
  • Steele, Teresa E., Esteban Álvarez-Fernández, and Emily Hallett-Desguez. 2019. “A Review of Shells as Personal Ornamentation during the African Middle Stone Age.” Paleoanthropology 2019: 24–51.
  • Stringer, Chris B. 2015. “Why We Are Not All Multi-regionalists Now.” Trends in Ecology and Evolution 29 (5): 248–251.
  • Stringer, Chris B. 2016. “The Origin and Evolution of Homo sapiens.” Philosophical Transactions of the Royal Society B 371: 20150237.
  • Stringer, Chris, and Peter Andrews. 1988. “Genetic and Fossil Evidence for the Origin of Modern Humans.” Science 239 (4845): 1263–1268.
  • Texier, Pierre-Jean, Guillaume Porraz, John Parkington, Jean-Philippe Rigaud, Cedric Poggenpoel, Christopher Miller, Chantal Tribolo, Caroline Cartwright, Aude Coudenneau, Richard Klein, Teresa Steele, and Christine Verna. 2010. “A Howiesons Poort Tradition of Engraving Ostrich Eggshell Containers Dated to 60,000 Years Ago at Diepkloof Rockshelter, South Africa.” Proceedings of the National Academy of Sciences USA 107 (14): 6180–6185.
  • Trinkaus, Erik. 1983. The Shanidar Neandertals. New York: Academic Press.
  • Valladas, Heléne, Jean-Louis Joron, Georges Valladas, Baruch Arensburg, Ofer Bar-Yosef, Anna Belfer-Cohen, Paul Goldberg, H. Laville, Lilliane Meignen, Yoel Rak, E. Tchernov, Anne-marie M. Tillier, and B. Vandermeersch. 1987. “Thermoluminescence Dates for the Neandertal Burial Site at Kebara in Israel.” Nature 330 (6144): 159–160.
  • Valladas, Heléne, Jean-Louis Reyss, Jean-Louis Joron, Georges Valladas, Ofer Bar-Yosef, and Bernard Vandermeersch. 1988. “Thermoluminescence Dating of Mousterian ‘Proto-Cro-Magnon’ Remains from Israel and the Origin of Modern Man.” Nature 331 (6157): 614–616.
  • Wadley, Lyn. 2015. “Those Marvellous Millennia: The Middle Stone Age of southern Africa.” Azania 50 (2): 155–226.
  • Walter, Robert C., Richard T. Buffler, J. Henrich Bruggemann, Mireille M. M. Guillaume, Seife M. Berhe, Berhane Negassi, Yoseph Libsekel, Hai Cheng, R. Lawrence Edwards, Rudo von Cosel, Didier Neraudea, and Mario Gagnon. 2000. “Early Human Occupation of the Red Sea Coast of Eritrea during the Last Interglacial.” Nature 405 (6782): 65–69.
  • Willoughby, Pamela R. 1993. “The Middle Stone Age in East Africa and Modern Human Origins.” African Archaeological Review 11: 3–20.
  • Willoughby, Pamela R. 1994. “The Origin and Dispersal of Modern Humans.” In Strength in Diversity: A Reader in Physical Anthropology, edited by Ann Herring and Leslie Chan, 235–258. Toronto: Canadian Scholars’ Press.
  • Willoughby, Pamela R. 2000. “Archaeologists, Palaeoanthropologists and the People without Culture.” In The Entangled Past: Integrating History and Archaeology, edited by Matthew. Boyd, John C. Erwin, and Mitch Hendrikson, 281–291. Calgary, Canada: Chacmool Archaeological Association.
  • Willoughby, Pamela R. 2007. The Evolution of Modern Humans in Africa: A Comprehensive Guide. Lanham, MD: AltaMira Press.
  • Willoughby, Pamela R. 2009. “From the Middle to the Later Stone Age in Eastern Africa.” In Sourcebook of Paleolithic Transitions: Methods, Theories and Interpretations, edited by Marta Camps and Parth R. Chauhan, 301–314. New York: Springer.
  • Willloughby, Pamela R. 2012. “The Middle and Later Stone Age in the Iringa Region of Southern Tanzania.” Quaternary International 270: 103–118.
  • Willoughby, Pamela R., Tim Compton, Silvia M. Bello, Pastory M. Bushozi, Anne R. Skinner, and Chris B. Stringer. 2018. “Middle Stone Age Human Teeth from Magubike Rockshelter, Iringa Region, Tanzania.” PLOS One 13 (7): e0200530.
  • Wintle, Ann G. 1996. “Archaeologically Relevant Dating Techniques for the Next Century: Small, Hot and Identified by Acronyms.” Journal of Archaeological Science 23 (1): 123–138.
  • Zilhão, João. 2007. “The Emergence of Ornaments and Art: An Archaeological Perspective on the Origins of ‘Behavioral Modernity.’” Journal of Archaeological Research 15 (1): 1–54.