• some level of gene flow between geographically separated populations prevented speciation, after the dispersal 
• all living humans derive from the species Homo erectus that left Africa nearly two million-years-ago 
• natural selection in regional populations, ever since their original dispersal, is responsible for the regional variants (sometimes called races) we see today 
• the emergence of Homo sapiens was not restricted to any one area, but was a phenomenon that occurred throughout the entire geographic range where humans lived 

• after Homo erectus migrated out of Africa the different populations became reproductively isolated, evolving independently, and in some cases like the Neanderthals, into separate species 
• Homo sapiens arose in one place, probably Africa (geographically this includes the Middle East) 
• Homo sapiens ultimately migrated out of Africa and replaced all other human populations, without interbreeding 
• modern human variation is a relatively recent phenomenon 

The multiregional view posits that genes from all human populations of the Old World flowed between different regions and by mixing together, contributed to what we see today as fully modern humans. The replacement hypothesis suggests that the genes in fully modern humans all came out of Africa. As these peoples migrated they replaced all other human populations with little or no interbreeding.

To understand this controversy, the anatomical, archaeological, and genetic evidence needs to be evaluated.

• In Asia these hominids evolved into Peking Man and Java Man, collectively referred to as Homo erectus. 
• In Europe and western Asia they evolved into the Neanderthals. 

Neanderthals lived in quasi isolation in Europe during a long, relatively cool period that even included glaciations. Neanderthals are distinguished by a unique set of anatomical features, including: 

• a large, long, low cranial vault with a well-developed double-arched browridge 
• a massive facial skeleton with a very projecting mid-face, backward sloping cheeks, and large nasal aperture, with large nasal sinuses 
• an oddly shaped occipital region of the skull with a bulge or bun 
• molars with enlarged pulp chambers, and large, often very heavily worn incisors 
• a mandible lacking a chin and possessing a large gap behind the last molar 
• a massive thorax, and relatively short forearms and lower legs
• although short in stature they possessed robustly built skeletons with thick walled limb bones 
• long clavicles and very wide scapulas 

In contrast, at roughly the same time, in Africa, a body plan essentially like our own had appeared. While these early Homo sapiens were anatomically modern they were not behaviorally modern. It is significant that modern anatomy evolved prior to modern behavior. These early sapiens were characterized by: 

• a cranial vault with a vertical forehead, rounded occipital and reduced brow ridge 
• a reduced facial skeleton lacking a projecting mid-face 
• a lower jaw sporting a chin 
• a more modern, less robustly built skeleton 

Hence, the anatomical and paleogeographic evidence suggests that Neanderthals and early modern humans had been isolated from one another and were evolving separately into two distinct species.

Homo sapiens exhibited technological skills around 50,000 years ago.

• little variation in stone tool types, with a preponderance of flake tools that are difficult to sort into discrete categories 
• over long periods of time and wide geographical distances there was general similarity in tool kits 
• a virtual lack of tools fashioned out of bone, antler or ivory 
• burials lacked grave goods and signs of ritual or ceremony 
• hunting was usually limited to less dangerous species and evidence for fishing is absent 
• population densities were apparently low 
• no evidence of living structures exist and fireplaces are rudimentary
• evidence for art or decoration is also lacking 

The archaeological picture changed dramatically around 40-50,000 years ago with the appearance of behaviorally modern humans. This was an abrupt and dramatic change in subsistence patterns, tools and symbolic expression. The stunning change in cultural adaptation was not merely a quantitative one, but one that represented a significant departure from all earlier human behavior, reflecting a major qualitative transformation. It was literally a "creative explosion" which exhibited the "technological ingenuity, social formations, and ideological complexity of historic hunter-gatherers."⁷ This human revolution is precisely what made us who we are today.

In the Upper Paleolithic of Eurasia, or the Late Stone Age as it is called in Africa, the archaeological signature stands in strong contrast to that of the Middle Paleolithic/Middle Stone Age. It was characterized by significant innovation: 

• a remarkable diversity in stone tool types 
• tool types showed significant change over time and space 
• artifacts were regularly fashioned out of bone, antler and ivory, in addition to stone 
• stone artifacts were made primarily on blades and were easily classified into discrete categories, presumably reflecting specialized use 
• burials were accompanied by ritual or ceremony and contained a rich diversity of grave goods 
• living structures and well-designed fireplaces were constructed
• hunting of dangerous animal species and fishing occurred regularly 
  higher population densities 
• abundant and elaborate art as well as items of personal adornment were widespread 
• raw materials such as flint and shells were traded over some distances 

Homo sapiens of the Upper Paleolithic/Late Stone Age was quintessentially modern in appearance and behavior. Precisely how this transformation occurred is not well understood, but it apparently was restricted to Homo sapiens and did not occur in Neanderthals. Some archaeologists invoke a behavioral explanation for the change. For example, Soffer¹¹ suggests that changes in social relations, such as development of the nuclear family, played a key role in bringing about the transformation. 

Shortly after fully modern humans entered Europe, roughly 40,000 years ago, the Neanderthals began a fairly rapid decline, culminating in their disappearance roughly 30,000 years ago. Neanderthals were apparently no match for the technologically advanced fully modern humans who invaded Europe and evidence for interbreeding of these two types of hominids is equivocal. 

Africans display higher genetic variation than other populations, supporting the idea that they were the first modern humans.

• Studies of contemporary DNA, especially mitochondrial DNA (mtDNA) which occurs only in the cellular organelles called mitochondria, reveal that humans are astonishingly homogeneous, with relatively little genetic variation.^1,5 
• The high degree of similarity between human populations stands in strong contrast to the condition seen in our closest living relatives, the chimpanzees.² In fact, there is significantly more genetic variation between two individual chimpanzees drawn from the same population than there is between two humans drawn randomly from a single population. Furthermore, genetic variation between populations of chimpanzees is enormously greater than differences between European, Asian and African human populations. 
• In support of an African origin for Homo sapiens the work of Cann and Wilson¹ has demonstrated that the highest level of genetic variation in mtDNA occurs in African populations. This implies that Homo sapiens arose first in Africa and has therefore had a longer period of time to accumulate genetic diversity. Using the genetic distance between African populations and others as a measure of time, they furthermore suggested that Homo sapiens arose between 100,000 and 400,000 years ago in Africa. 
• The low amount of genetic variation in modern human populations suggests that our origins may reflect a relatively small founding population for Homo sapiens. Analysis of mtDNA by Rogers and Harpending¹² supports the view that a small population of Homo sapiens, numbering perhaps only 10,000 to 50,000 people, left Africa somewhere between 50,000 and 100,000 years ago. 
• Scientists recently succeeded in extracting DNA from several Neanderthal skeletons.⁸ After careful analysis of particularly the mtDNA, but now also some nuclear DNA, it is apparent that Neanderthal DNA is very distinct from our own. In assessing the degree of difference between DNA in Neanderthals and modern humans, the authors suggest that these two lineages have been separated for more than 400,000 years. 

Although in its infancy, such genetic studies support the view that Neanderthals did not interbreed with Homo sapiens who migrated into Europe. It is, therefore, highly likely that modern humans do not carry Neanderthal genes in their DNA.

Neanderthals probably did not breed with modern humans but they borrowed some of their tools and skills.

• Cave sites in Israel, most notably Qafzeh and Skhul date to nearly 100,000 years and contain skeletons of anatomically modern humans. Furthermore, Neanderthal remains are known from sites such as the 110,000-year-old Tabun cave, which predates the earliest Homo sapiens by about 10,000 years in the region. 
• The presence of Neanderthals at two other caves in Israel, Amud and Kebara, dated to roughly 55,000 years means that Neanderthals and Homo sapiens overlapped in this region for at least 55,000 years. Therefore, if Homo sapiens were in this region for some 55,000 years prior to the disappearance of the Neanderthals, there is no reason to assume that Neanderthals evolved into modern humans.
• Archaeological evidence from Europe suggests that Neanderthals may have survived in the Iberian Peninsula until perhaps as recently as 30,000 to 35,000 years ago. Fully modern humans first appear in Europe at around 35,000-40,000 years ago, bringing with them an Upper Paleolithic tool tradition referred to as the Aurignacian. Hence, Neanderthals and fully modern humans may have overlapped for as much as 10,000 years in Europe. Again, with fully modern humans on the scene, it is not necessary to have Neanderthals evolve into modern humans, further bolstering the view that humans replaced Neanderthals. 
• The situation in southern France is, however, not quite as clear. Here, at several sites, dating to roughly 40,000 years there is evidence of an archaeological industry called the Châtelperronian that contains elements of Middle and Upper Paleolithic artifacts. Hominids from these sites are clearly Neanderthals, sparking speculation that the Châtelperronian is an example of Neanderthals mimicking the culture of modern humans. The lack of anatomical intermediates at these sites, suggests that if Neanderthals did encounter and borrow some technology from Homo sapiens, they did not interbreed. 
• A potential 24,500-year-old Neanderthal/sapiens hybrid was announced from the site of Lagar Velho, Portugal.⁴ This 4-year-old has a short, squat body like a Neanderthal, but possesses an anatomically modern skull. There are a number of problems with interpreting this find as a Neanderthal/sapiens hybrid.¹⁴ First of all, as a hybrid it should have a mixture of traits throughout its body and not possess the body of a Neanderthal and skull of a modern human. For example, if we look at hybrids of lions and tigers they do not possess the head of one species and the body of the other, but exhibit a morphological mixture of the two species. Secondly, and more importantly, acceptance of this specimen as a hybrid would suggest that Neanderthal traits had been retained for some 6,000 to 10,000 years after Neanderthals went extinct, which is highly unlikely. This is theoretically unlikely since Neanderthal traits would have been genetically swamped by the Homo sapiens genes over such a protracted period of time. 
• Proponents of the Multiregional Model, such as Milford Wolpoff, cite evidence in Asia of regional continuity. They see an evolutionary link between ancient Homo erectus in Java right through to Australian aborigines. A possible problem with this view is that recent dating of late surviving Homo erectus in Indonesia suggests that they survived here until 50,000 years ago, which is potentially when fully modern humans may have arrived in the region from Africa. 
• China may contain the best evidence for supporting the Multiregional Model. Here there are discoveries of a couple of skulls dated to roughly 100,000 years ago that seem to possess a mixture of classic Homo erectus and Homo sapiens traits. Better geological dating and more complete specimens are needed to more fully assess this possibility. 

About the author: Paleoanthropologist Donald C. Johanson, is professor of anthropology and Director of the Institute of Human Origins at Arizona State University. He is best known for his discovery of "Lucy", a 3.2 million-year old Australopithecus afarensis skeleton he found in 1974 in Ethiopia. His books include Lucy: The Beginnings of Humankind and, most recently, From Lucy to Language. Dr. Johanson hosted the Emmy-nominated NOVA television series In Search of Human Origins.
http://www.anthro.mankato.msus.edu/information/
biography/fghij/johanson_donald.html

                 

Institute of Human Origins
The Institute of Human Origins' web site is a comprehensive site on human origins and has a special segment on the emergence of modern humans.
http://www.becominghuman.org

University of Oslo
Investigate human evolution at the Palaeontological Museum of the University of Oslo in Norway.
http://www.toyen.uio.no/palmus/galleri/montre/english/early_man_liste_e.htm

Human evolution chart
Click on this interactive chart of human evolution, stretching from 5 million years ago to the present, to learn about different hominid species.
http://www.geocities.com/palaeoanthropology/timeline.html

Read a book  
» From Lucy to Language by D.C. Johanson and B. Edgar (Simon and Schuster, 1996).
» The Fossil Trail: How We Know What We Think We Know about Human Evolution by I. Tattersall (Oxford University Press, 1995).
» Extinct Humans by I. Tattersall and J. H. Schwartz (Westview Press, 2000).

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