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by Michele A. Miller, PhD
Editor of Old World Archaeology, Athena Review
The fate of Neanderthals has always been hotly debated, beginning with the discovery of their first remains in the 19th century (see timeline, part 1). Discussions about Neanderthal existence and their relation to modern humans have at times been vehement, often leading to personal rivalry not only in the scholarly community, but also within the realm of popular culture. Yet what is it about these robust and stained old bones that has lead to so much discussion and dispute?
In part, the answer is that the way we view Neanderthals reflects how we view ourselves, as Neanderthals are the closest known relatives to anatomically modern humans. Are we truly the unique and especially blessed creations of God, or merely the latest branch in a tree which once blossomed with many other similar life-forms?
The last 150 years have seen continuous debate in which Neanderthals have been either dismissed as mere dead-end, distant cousins of modern Homo sapiens, or embraced as our direct ancestors. By the late 18th century, an age when many believed the Bible recounted the only true early history of humankind, recognition of extinct species, although only accepted as ante-diluvial, or destroyed in the Biblical flood, was a revolutionary step by Cuvier, Buffon, and other early paleontologists. The 19th century discovery of fossilized Neanderthal bones was, therefore, essential in the early debate over the antiquity of man and the ultimate acceptance of evolutionary theory.
Fig.1: Huxley's drawing of the original Neanderthal fossil found in the Neander Valley in 1856, showing the low but large skull, and accentuated brow ridges (T.H. Huxley, 1863).
The first Neanderthal remains, although not recognized as such for several decades, were found in 1830 in Engis Cave, Belgium and in 1848 at Gibraltar. A few years later, fossils found at Feldhofer Cave in the Neander Valley of Germany (fig.1) touched off considerable debate. While a very few championed the remains as the ancient species of Neanderthal Man, many others rejected them as merely the pathological skeleton of a deformed individual.
When Darwin published his theory of natural selection in 1859, he avoided applying the mechanism of evolution directly to humans, ending his manuscript only with the enigmatic words, and light will be thrown on the origin of man and his history. While Darwin did not yet press the issue, others were quick to see the implications of his work. Only a few years after the initial publication of On the Origin of Species, Thomas Henry Huxley, Darwins most prominent supporter, published the first scientific assessment of the Neanderthal and Engis fossils (fig.1).
The extinct human species, Homo neanderthalensis, was first proposed in a controversial paper read in 1864 by William King, a professor from Queens College in Ireland. King thought the Neanderthal skull more closely conforms to the braincase of a chimpanzee than a human.
Kings consignment of Neanderthals to a different species from humans was based on a comparison with hunting and gathering people from the Andaman Islands near India, whose physical traits, viewed in connection with the strictly human conformation of his cranium, ...specificially identify him with Homo sapiens. Psychical endowments of a lower grade than those of the Andamaner cannot be conceived to exist; they stand next to brute benightedness. (in Trinkaus and Shipman 1993, pp.88-89). T.H. Huxley, by contrast, on purely anatomical grounds compared two Neanderthal skulls with those of Scandanavians and Australian aborigines, and found Neanderthals to be within the human range.
Fig.2: Neanderthal skullcap (calotte) from Spy, Belgium showing characteristic brow ridges (Athena Review, from cast at AMNH).
As the theory of evolution gained acceptance, so did the recognition of Neanderthals as ancient, anatomically-different, humans. By 1866, the Neanderthal jaw discovered at Trou de la Naulette, Belgium, was declared, by the anatomist Paul Broca, to constitute the first fact that furnishes anatomical evidence to the Darwinists. It is the first link on a chain that extends from man to the apes (Trinkhaus and Shipman 1993, p.103).
While it came to be accepted that Neanderthals were an ancient form of human, what remained to be determined, and what is still hotly debated, is the exact relationship of these robust and (as many consider) primitive-looking beings with modern humans. By the early 1900s, scholars like Gustav Schwalbe and Ale Hrdlicka believed Neanderthals were direct ancestors of modern humans, while others like Arthur Keith and Marcellin Boule suggested Neanderthals were an evolutionary side-branch, an extinct species that had nothing to do with human ancestry. These same evolutionary ideologies still exist today, in the form of the multi-regional continuity vs. replacement theories.
In part, where Neanderthals were placed on the human evolutionary tree was largely influenced by how their anatomy and behavior were viewed. Boule, for instance, who studied the Neanderthal remains from La Chapelle-aux-Saints (fig.3) and La Ferassie, described Neanderthals (based on several incorrect anatomical analyses) as stooping in posture, with a slouching, bent-kneed gait-altogether too apish to be ancestoral to modern humans. Boule sought to distance modern humans from what he saw as brutish beings, arguing forcefully that the Neanderthals had been annihilated by the more elegant inventive Homo sapiens, a scene retold in various 20th century novels such as William Goldings The Inheritors.
Fig.3: Neanderthal skull from La Chapelle-aux-Saints in France (Athena Review, from cast at AMNH).
In a similar way, Karl Gorjanovic-Kramberger, the Croatian paleontologist who excavated and analyzed the large sample of Neanderthals from Krapina, distanced modern humans from Neanderthals by proposing fragmented skeletons from the site provided evidence of cannibalism. These men ate their fellow tribesmen, and whats more they cracked open the hollow bones and sucked out the marrow, (Pracovjek iz Krapine, 1918). The specter of the Neanderthal as a cannibal survives today in the recent research at Moula-Guercy Cave in France by Defleur et al. (1999).
These interpretations of Neanderthal behavior fed neatly into a prevalent pre-sapien theory of human origins proposed by Boule and Keith, who believed the human lineage had split at some remote time in the past (ie., during the Pliocene, 5-2 mya) into two separate branches. One of these branches led to modern humans, while the other ended in extinct species, including Neanderthals. A general belief in this theory may have helped many otherwise astute scientists accept the authenticity of the infamous Piltdown skull in Kent, England (timeline, phase 3). It was far more comfortable to consider our ancestors to have large modern-like brains, if a rather ape-like jaw, then to look for human origins in the primitive-looking (if large-brained) skulls of Neanderthals or the much older, smaller-brained fossil hominids uncovered between 1924-1927 in Africa (Australopithecus; fig.4) and China (Homo erectus).
Surprisingly, it was another claim of Neanderthal cannibalism that actually lead to the view that Neanderthals were more human-like than previously believed. Alberto Blanc interpretated a skull from Monte Circeo, Italy in 1939 as indicating cannibalistic but also mortuary practices. Along with an enlarged opening at the base of the braincase, which Blanc claimed occurred as the brain was extracted for a feast, he also proposed that the skull was found buried in a circle of stones. While Blanc's evidence for Neanderthal ritual was later questioned, his theories lead to renewed belief in the humanity of Neanderthals.
Even before Blanc's discovery, various researchers had suggested that Neanderthals exhibited ritualistic behaviors. For instance, excavations from 1917 to 1921 at Drachenloch (Dragon's Lake) in the Churfirsten Mountains of Switzerland uncovered Mousterian tools alongside numerous bones of Ursus spelaeus, the huge cave bears that became extinct 50,000-40,000 years ago. The amateur archaeologist Dr. Emil Bächer described the bear bones as found within several stone cists capped with stone slabs or deliberately piled up behind heaps of stones. Although later analyses revealed the accumulations resulted from the natural deaths of the bears during hibernation, Bächer saw them as evidence for Neanderthal cave bear cults, in which the bears were the subject of worship and perhaps sacrifice, an idea employed in 20th century popular literature, including Clan of the Cave Bear by Jean Aeul.
Fig.4: Type specimen of Australopithecus africanus with incomplete skull and limestone endocast from Taung, South Africa, found by Raymond Dart in 1924 (Athena Review, from cast at AMNH)
As Neanderthal behavior began to appear more human, some scholars accepted the idea that they were our own ancestors. For instance, at this time Ale Hrdlicka suggested a unilinear theory of human evolution, in which Neanderthals were merely one stage leading to Homo sapiens. The idea (perhaps originating with T.H. Huxley) that Neanderthals were not very physically different from modern humans eventually began to gain wider acceptance.
After World War II, some of the greatest barriers to the acceptance of Neanderthals humanity collapsed. Thanks to fluorine dating tests conducted by Oakley in the early 1950s there was conclusive proof that the troublesome Piltdown remains were a forgery consisting of an ancient cranium of an anatomically modern human with the purposely stained and filed jaw of an orangutan. Only a few years later, two major re-analyses of the skeletal material from La Chapelle by C. Arambourg, and Straus and Cave, finally put to rest the notion of the bent-kneed, bent-over, apish Neanderthals. Moreover, Etienne Patte demonstrated the anatomical characteristics observed in the Neanderthal fossils were statistically not much different than those seen in the full range of modern humans. Following Coon (1939), Straus and Cave observed that, if bathed, shaved and dressed, Neanderthals would hardly attract attention on a New York subway. This was reinforced by Garrods discovery in the 1930s at Mount Carmel of so-called progressive Neanderthals.
It was during this phase of synthesis among possible species that Franz Weindenreich developed the multi-regional theory of human origins that still is accepted by several prominent researchers today. According to this theory, the regional variations in modern human form that we call racial differences originated in geographically separated populations of Homo erectus. An ultra-lumper, Weindenreich believed that the various ancient hominid fossils which had heretofore been split into many different species (and even genera) should all be considered as a single species, Homo sapiens.
In his 1947 publication, Weidenreich proposed that a single, ancient form of this species had long ago spread across the Old World, developing separately to form modern humans. Although this allowed for localized features to develop in humans in each area, Weidenreich also postulated that there was considerable interbreeding and therefore genetic exchange between the various regional groups, which can be better visualized as the intersecting diagonal lines between regional groups in an evolutionary diagram (fig. 4). One corollary of his theory is that Neanderthals would be considered merely one of many regional representatives of a particular stage of human development, with similar ancient human stages apparent in geographic areas outside Europe.
Fig.5: "Candelabra" model of human evolution (arrows show genetic intermixing).
In ensuing debate by 1959, William Howell challenged Weidenreich's ideas with a Noahs Ark model in which humans developed from a single, recent origin and replaced the earlier hominids without interbreeding or genetic exchange. Three years later, Carleton Coon postulated a highly controversial theory for the origin of modern humans similar to Weidenreichs but adding that Homo erectus not only developed into Homo sapiens separately in each region, but did so at different rates. In the politically charged sixties, Coons suggestion that Caucasoids (Europeans) and Mongoloids (Asians) had developed into Homo sapiens before native African and Australian peoples was rejected as racist.
Many of Coons other ideas were better accepted, especially his proposal that the particularities of Neanderthal anatomy reflected adaptation to the cold, glacial climate of Pleistocene Europe. For instance, he noted that the short-limbed stocky build of Neanderthals mirrored that of modern Eskimo tribes, while human populations closer to the equator tended to be more slender and longer-limbed, otherwise known as Allens rule. A more squat, stocky build enables a mammal to conserve body heat since it minimizes the surface area of the body relative to volume. Similarly, with his radiator theory of Neanderthal noses, Coon noted that the Neanderthals large nasal apertures and prominent nose served to warm the air as it was inhaled, thus protecting the delicate brain stem from the cold air of their surroundings (for recent, contrary findings, see Mowbray and Gannon this issue).
In 1946 Earnest Hooton brought attention to the heterogeneity between classic European Neanderthals and progressive central European and Near Eastern Neanderthals, making interspecific comparisons more complex.
As controversies with racial comparisons continued, exciting new discoveries made by Ralph and Rose Solecki at Shanidar Cave in Kurdistan, Iraq added fuel to the idea that Neanderthals were culturally more similar to ourselves than previously acknowledged (fig. 6). Between 1953 and 1960 nine Neanderthal skeletons were found at Shanidar, all more similar in anatomy to the classic Neanderthals of Europe than the more progressive type found at Tabun cave in Israel. While four of these individuals seem to have been buried in the cave due to natural causes (probably caught under rock fallen from the cave ceiling), the other five appeared to have been intentionally buried. Further, the discovery of large amounts of pollen in the soil by the skeleton of one elderly male suggested to Solecki that the individual had been buried with offerings of flowers. Adding to such signs of humanity in the Shanidar remains, analysis of the bones of another male Neanderthal showed he had lived well past maturity despite sustaining a number of injuries earlier in life , which may have left him both blind and considerably crippled, without the use of his right arm. In his 1971 book about the cave discoveries, tellingly titled Shanidar: The First Flower People, Solecki pointed out that such a significantly handicapped individual would not have survived without the care and protection of his fellow Neanderthal community.
With Neanderthals looking more and more human in their behavioral characteristics, scholars once again argued to place them on the human lineage. At the same time that Coon's theories were being debated among American anthropologists, C. Loring Brace refocused attention back to the Neanderthal question, arguing that Neanderthal anatomy was less highly specialized than had been formerly proposed. Moreover, he asserted that a gradual change could be seen in tool-making traditions from the Mousterian into the Upper Paleolithic, thus contradicting the sudden replacement of Neanderthals by modern humans. Today the replacement versus continuity debate remains prominent (see Shea, this issue).
Fig.6: Neanderthal cranium from Shanidar, Iraq (Athena Review, from cast at AMNH).
In a later paper Brace claimed that most of the differences between Neanderthals and modern humans were in the dimensions of their face and teeth, and that this reduction in size of the face and teeth that occurred over time could be explained as the by-product of cultural evolution. As specialized tools replaced the use of teeth for various tasks, for instance, the size of teeth and facial dimensions would eventually be reduced. In essence, Brace suggested that for humans, culture was more important than biology in determining the direction of evolutionary change. Later, Brace argued that not only was culture the human ecological niche but, as according to ecological theory, only one species can occupy a certain niche at a time, so only one human species could occur at one time. This version of the unilineal theory of hominid evolution soon became widely accepted as the single species theory during the late 1960s and early 1970s. With the fossil discoveries at the end of that decade leading to proposals of several early hominid species (i.e., Homo ergaster, H. habilis and H. rudolfensis) living contemporaneously in Africa, however, new formulations were required.
By the early 1970's Milford Wolpoff had reinstated Weidenreichs theory of multiregional continuity as a mechanism for the evolution of modern humans. Seeing evidence within the fossil record for continuity of certain physical characteristics within geographic regions, Wolpoff reasoned that these specific variations originated with Homo erectus. Unlike Coon, however, Wolpoff recognized the key role of gene flow between otherwise isolated human populations. Without this genetic exchange it would be impossible to propose that each population would have evolved into the same species, Homo sapiens. This continuity of anatomical characteristics is perhaps least obvious in the comparison of classic European Neanderthal skeletal material to modern humans. However, in an effort to prove his position, Wolpoff compared teeth from the more transitional Krapina Neanderthals with fossil teeth from early Homo sapiens, finding a long-term evolutionary trend for increasing size in molars and decreasing size in incisors from former to later.
By the middle of the 1970's a new crop of scholars were adding to the debate. The first of these was Christopher Stringer, whose 1974 dissertation focused on the quantitative analysis of Neanderthal crania. Stringer observed too many differences between Neanderthal and modern human crania to believe that Neanderthals could have been our direct ancestors, and thus became a strong proponent of the replacement hypothesis. After noting that several skulls found in Ethiopia (Omo 1 and 2), dated to between 130 and 40 thousand years ago, appeared anatomically modern, Stringer proposed that Homo sapiens originated in Africa. Only two years later, however, Fred Smith presented his dissertation with remarkably different conclusions that, based on a new quantitative study of the Krapina Neanderthal material, supported the multiregional hypothesis that Neanderthals had evolved into modern humans within Europe.
By the middle of the decade, the opposing camps in the Neanderthal debate were clear. Smith and Wolpoff followed in the footsteps of Brace in asserting that Homo sapiens evolved indigenously in Europe from Neanderthals. Stringer, on the other hand, just as adamantly declared that modern humans evolved outside Europe, migrating there and replacing Neanderthals.
In 1976 Stringer gained an ally. In that year Gunter Brauer presented the results of his meticulous study of various archaic human remains in Africa, such as those of Homo heidelbergensis found at Bodo, Ethiopia (fig.7) and the more modern-looking finds at Broken Hill in Zambia. Only a few years earlier, on the basis of associated tool-types, the Broken Hill skeletons were still assumed to be 30,000-20,000 years old; that is, contemporary with the Upper Paleolithic in Europe. New radiocarbon dates for the remains, however, revealed that they were much older some as old as 200,000 years or more. Armed with his knowledge of these remains, Brauer developed the out of Africa theory, in which he argued that modern Homo sapiens evolved first in Africa and then migrated from there to populate the remainder of the world. Notably, however, Brauer also allowed for some interaction of these new humans with the native populations. For instance, he argued that these humans were likely to have interbred with the European Neanderthals, thus allowing for the possibility that Neanderthals had contributed somewhat to modern populations.
Two years later, Jean-Jacques Hublin became another supporter of the replacement model. Based on his study of the anatomy of the rear and base of Pleistocene hominid crania, he noted that there were great morphological differences between Neanderthal and Homo sapiens crania, and that these occured abruptly with the introduction of the latter in Europe. He thus essentially concurred with Stringer that modern humans could not have evolved from Neanderthals but must have evolved outside of Europe.
In 1983, Erik Trinkaus suggested a possible mechanism for Homo sapiens replacement of Neanderthals. Analyzing several pelvic fragments from Shanidar Cave, he suggested that Neanderthals had a larger birth canal than humans, possibly indicating that Neanderthal babies were larger with larger brains upon birth. Larger brains, however, would have taken longer to grow in utero, suggesting that Neanderthals had a longer gestation period, perhaps up to a year. If this were true, then modern humans may have had the advantage of shorter birth spacing, thus outbreeding their Neanderthal neighbors. Although hotly debated, the theory was discarded rather quickly when results of the discovery of new Neanderthal remains from Kebara cave in Israel, came to light. This first nearly complete Neanderthal pelvis, although of a male, suggested that Neanderthal birth canals were no larger than those of modern humans.
Fig.7: Homo heidelbergensis skull from Bodo, Ethiopia (Athena Review, from cast at AMNH).
As Trinkaus was revealing new insights into Neanderthal anatomy and behavior from the Shanidar remains, new dating techniques were radically altering the picture of other Near Eastern early human remains. The first hint that Near Eastern hominid dating assumptions were incorrect came from Eitan Tchernovs 1981 study of small mammals from different levels at various local Middle Palaeolithic sites. He found that rodents associated with the Qafzeh remains were of archaic types similar to those from Lower Palaeolithic levels at Tabun (i.e., below the Neanderthal remains). Therefore, Tchernov suggested that the archaic humans from Qafzeh were older than the Neanderthals from the other Israeli sites.
New dating techniques, such as thermoluminescence and electron spin resonance (ESR), later applied to the same remains confirmed Tchernov's conclusions. While Neanderthal remains from Kebara and Amud were dated to ca. 60,000 and ca. 40,000 years ago respectively, modern humans from Qafzeh and Skhul were dated to ca. 90,000 and ca. 80,000 years ago (see Shea this issue). Other dating methods indicated that the anatomically modern humans could even be as old as 130,000 years, with the two groups overlapping in the region for 60,000 years or more as Neanderthals moved south from Europe in accordance with colder climactic shifts.
As these dates were creating new challenges for palaeoanthropologists, new evidence from an unexpected field further stirred the great debate. Three molecular biologists at the University of California at Berkeley published a 1987 paper in Nature based on their study of mitochondrial DNA in modern human populations, claiming a relatively recent and common origin for all modern humans (Cann, Stoneking, and Wilson 1987). The study was based on the presumed fact that the DNA within an intracellular structure called mitochondria is inherited from the mother only (for other, contrary evidence see AR 2,2 p.5), unlike the DNA from the cell's nucleus, which comes from both parents. Therefore, mtDNA is passed down from mother to daughter, generation to generation, without being broken up or combined, except for the natural accumulation of mutations. By studying the mtDNA from a large group of individuals from populations around the world they were able to show that the number of mutations that differentiated the mtDNA of different racial groups was very small, thus indicating that the different races must have separated relatively recently. In addition, the biologists used a statistical method to create an evolutionary tree showing modern humans first evolved in Africa and then migrated from there to populate the rest of the world. Finally, using a rate of mutation based on the date of the first migration of modern humans to Australia and New Guinea, the researchers were actually able to calculate the approximate date of when Homo sapiens or specifically a group of Homo sapiens including our ancestral Mitrochondrial Eve first evolved, sometime between 280,000 and 140,000 years ago.
Obviously this study was welcomed by proponents of the out of Africa theory, but it did not go unchallenged. Critics pointed out flaws not only in the original sample of individuals used for the study, but also in the theoretical and statistical methods used to create the phylogenies, or evolutionary trees. Antagonists to the genetic studies raised several key issues, such as the effect of bottlenecking, or the possibility of an extended period of time without change in DNA makeup. They also noted that the evolutionary clocks, which are the basis for chronological interpretation of DNA, are built upon assumptions of the DNAs rate of change, potentially rendering any subsequent timeline inaccurate.
Ten years after the original mtDNA study, new research published by Svante Pääbo and his team appeared to confirm the replacement scenario. The DNA for this study was retrieved directly from the original Neanderthal from Feldhofer Cave in Germany. Comparisons of this DNA with modern human populations suggested Neanderthals were an entirely different species from Homo sapiens, with little or no interbreeding between the two. The study also pointed to a date of 690,000-550,000 years ago for the split between Neanderthals and modern human lineages. These results were essentially duplicated in the study of DNA from a Neanderthal infant from Mezmaiskaya cave Russia. (Ovchinnikov et al. 2000, and this issue).
More recently, however, a comprehensive analysis by Allen Templeton (2002) of ten genetic haplotypes indicates that there have been at least three major human migrations out of Africa (one at 1.7 mya with Homo erectus spreading to Eurasia and Southern Asia; a second between 840,000 and 420,000 years ago (a period that includes the evolution of archaic Homo sapiens including Neanderthals), and the third between 150,000 and 80,000 years ago, the time of the proposed migration by "Mitochondrial Eve's" descendants. Yet Templeton also found there has been considerable intermixing between the various populations used in the haplotype studies, which seem to retain traces even of early genomes. From a genetic standpoint, this appears to refute any complete "replacement" scenarios, including that which has been applied to Neanderthals, and may effectively reinstate a kind of "candelabra" model of ongoing genetic intermixing after the original migration of Homo erectus from Africa (fig.5).
These more recent molecular studies, for all their quantitative appearance of certainty, have not put a damper on the debates surrounding the fate of the Neanderthals or their relationship to modern humans. While many palaeoanthropologists hold that Homo sapiens did in some way replace their closest cousins, how this occurred is much disputed, with some claiming that out-and-out violence between the two groups was a factor, others arguing that competition for resources was more important, and still others proposing that interbreeding and genetic swamping was to blame for their demise.
Additional evidence has been drawn into the debate by Zilhão (Duarte et al. 1999; see also this issue), with the discovery in November, 1998 of the complete skeleton of a four-year-old child at the Lagar Velho rock shelter in the Lapedo Valley of Portugal. While large parts of the site had been bulldozed, a remnant of the original deposit contained charcoal, animal remains, and stone tools typical of the Solutrean Period of the Upper Palaeolithic (ca. 22,000-20,000 BP). The child burial was found in the intact deposits lying 2.5 m below this hanging remnant. Many features of the burial characterized it as a typical human burial of the Gravettian Period of the Upper Paleolithic (ca. 27,000-22,000 BP) and it was indeed dated by radiocarbon to between 25,000-24,500 BP. Certain characteristics of the child's anatomy, however, have led Erik Trinkaus, who studied the remains, to propose it presented a mosaic of Neanderthal and modern human traits. These features include Neanderthal-like limb proportions in addition to a characteristically modern protruding chin. Based on these findings, the excavator João Zilhão and Trinkaus have suggested the child represents the genetic mixing of Neanderthal and modern human populations.
Neither the Lapedo Valley child nor the new mtDNA evidence has gone unchallenged. Debate on the relationship between modern humans and Neanderthals continues with many basic questions unanswered. We have more sophisticated means of evaluating Neanderthal anatomy, and no longer picture our closest human relatives as shuffling, bent-kneed, slack-jawed brutes. Yet scholars still do not agree on how similar Neanderthals were to anatomically modern humans in terms of cognitive ability and behavior. Were Neanderthals capable of symbolic thought? Did they have language, rituals and culture? Were they innovators, or merely imitators of their more quick-brained cousins?
Closely related to these questions is how we place Neanderthals in consideration of human origins. Could they have been merely another race in the hugely variable and polytypic species known as Homo sapiens? Or were they an entirely separate species, not capable of interbreeding with our own? And what was the nature of interaction between humans and Neanderthals? We know that in areas such as the Levant they must have lived in relatively close proximity for thousands of years, and it is hard to accept that they did not interact on some level. But was it through love (ie., interbreeding), a mixing of genetic material that is evidenced in the fossil record as hybrids,or was it through war, the replacement of one species by another, by violence or competition for resources? One thing certain is that, ultimately, Neanderthals disappeared from the earth, leaving modern humans alone to contemplate their fate.
Bibliography and references:
AMNH: American Museum of Natural History, New York, NY.
Cann, R. , M. Stoneking, and A. Wilson. 1987. "Mitochondrial DNA and Human Evolution." Nature 325: 31-36
Duarte, C., J. Mauricio, P.B. Pettit, P.Souto, E. Trinkaus, H. Van der Plict, and J. Zilhão. 1999. "The Early Upper Paleolithic Human skeleton from the Abrigo do Lagar Velho (Portugal) and Modern Human Emergence in Iberia." Proceedings of the National Academy of Sciences, 96: 7604-7609.
Huxley, T.H. 1863. On Some Fossil Remains of Man.
Krings, M. Geisert, H. Schmitz, R.W., Krainitzki, H. and Paabo, S. 1999. "DNA Sequence of the mitochondrial hypervariable region II from the Neandertal type specimen." Proceedings of the National Academy of Sciences, 96: 5581-5585.
Ovchinnikov, I.V, Gotherstrom, A., Romanova, G.P., Kharitonov, V.M., Liden, K., and Goodwin, W. 2000. "Molecular analysis of Neanderthal DNA from the northern Caucasus." Nature 404: 490-493.
Templeton, A. 2002. "Out of Africa again and again." Nature 416: 45-51.
Trinkaus, E., and P. Shipman. 1993. The Neandertals: Changing the Image of Mankind. New York, Knopf.
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