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The fossil that gave the Neanderthals their name was found in a cave being quarried for limestone in Germany's Neander Valley in 1856. At least two Neanderthal fossils were discovered before the Neander Valley individual; however, neither was recognized as a member of an extinct human group until after the name "Neanderthal" was assigned. Many similar fossils have been found in scattered locations all over Europe and the Middle East since the Neander Valley discovery. Dates assigned to the various fossils indicate that the Neanderthals originated late in the Ice Age and became extinct a few thousand years before the last glacial retreat (from about 200,000 years ago to about 30,000 years ago). Thus the Neander Valley specimen lent its name to a fossil relative of modern humans that occupied Europe and the Middle East late in the Ice Age. Though the Neanderthals were very similar to modern humans, they had several distinctive characteristics. Neanderthals were short and exceptionally stout-bodied with broad supportive bones and joints. This body form suggests a life filled with intense physical effort. Perhaps the compact body also helped them cope with cold stress under Ice Age conditions. Their brains were somewhat larger than modern human brains. That size may have compensated for the more massive total body size of the Neanderthals, since large-bodied organisms generally have larger brains. Their foreheads sloped up from their exceptionally heavy eyebrow ridges, their jaws extended forward beyond the plane of the face, and their chins were weakly developed. These and several other characteristics are used to define a fossil find as a Neanderthal.

Structure and Behavior
Rudolph Virchow's initial interpretation of the Neander Valley fossil as a diseased human was popular for a time. Virchow held that the fossil was a modern human whose unique features were the result of disease. However, as more fossils with the same characteristics were discovered all around Europe and the Middle East, this explanation became untenable. Later, misinterpretation of the characteristics of Neanderthal fossils led Marcellin Boule and others to interpret Neanderthals as stooped, bent-kneed, apelike subhumans with an animal nature to match. Additional fossil discoveries, including evidence for toolmaking and burials, sometimes with flowers placed in the grave, caused anthropologists to rethink the presumed animal nature of the Neanderthals. Although the evidence for flowers has been challenged, the evidence for burials, presumably accompanied by mourning, is accepted by many anthropologists. In addition, fossils showed that some Neanderthals lived muchof their lives with deformed limbs and other disabilities, which would have made it difficult or impossible for them to fend for themselves. Yet they apparently lived many years in that condition, suggesting the support of other members of a social group. Such behavior was not in keeping with Boule's picture of the Neanderthals as nonhuman animals. Reinterpretation of the anatomic evidence also suggested that, instead of a bent-kneed, stooped posture, the Neanderthals walked on two rather straight legs and had hands capable of manipulating materials and making tools, much as modern humans do. All this indicated that the Neanderthals were more like modern humans than Boule's interpretation, and they came to be thought of in that light.

Taxonomic Relationship to Modern Humans
Neanderthals have always been recognized as close relatives of modern humans, but the specific taxonomy of the relationship is still a point of contention. They are placed in the same genus (Homo) as modern humans by almost all anthropologists, but researchers debate whether they were members of our species, Homo sapiens or belonged in their own species, Homo neanderthalensis. The discussion of the structure and behavior of Neanderthals bears directly on this question. If the Neanderthal characteristics were the result of disfigurement caused by disease, Neanderthals were simply aberrant humans and not especially interesting from the perspective of human evolution. However, if they were stooped, bent-kneed, and animal-like in behavior, they were probably a separate species, perhaps ancestral to modern humans, and therefore more interesting from the evolutionary perspective. On the other hand, if they were upright in stature, were skilled toolmakers, were supportive of their handicapped and elderly, and buried their dead with mementos such as flowers, they might earn the designation Homo sapiens and take on an even greater interest to the more modern members of that species. Such arguments are part of the practical taxonomy of the Neanderthals, but the real key to species identification and species separation is (at least theoretically) interbreeding. If the members of two groups can mate with each other and produce fertile offspring, and if these offspring can produce fertile offspring, the two groups are generally considered to be members of the same species. Therefore, the real taxonomic question becomes: Could Neanderthals and early modern humans interbreed? Because it is difficult to determine whether fossil groups interbred with one another, Neanderthal taxonomy has been primarily determined by anatomic and presumed behavioral characteristics, such as those already discussed. That taxonomy has vacillated with changing interpretations of those characteristics. Neanderthals have been placed in their own species (Homo neanderthalensis) for much of their history, but they have been identified as a human subspecies (Homo sapiens neanderthalensis) at other times. The latter designation implies that the Neanderthals and modern humans (Homo sapiens sapiens) were members of the same species and therefore could interbreed. Determination of the Neanderthals' taxonomic position is an integral part of arguments over the mechanism of the origin of modern humans. There are two main hypotheses for that origin: the replacement hypothesis of Christopher Stringer and the multiregional hypothesis vigorously supported by Milford Wolpoff. The replacement hypothesis is also designated "out-of-Africa" because it assumes that a population of African origin expanded throughout Africa, Europe, and Asia and rapidly replaced the more primitive humanlike species living there, including the Neanderthals. Whether this replacement was by competition or by more direct and violent means is undetermined. The multiregional hypothesis suggests that the widespread, more primitive humanlike populations evolved into modern humans rather than being replaced by new immigrants. Both hypotheses hold that the more primitive populations also originated in Africa and spread to Europe and Asia at a much earlier date. Because the Neanderthals are the best known and best understood early human group, an understanding of the Neanderthal relationship is critical to an understanding of the evolutionary history of humanity. A Neanderthal contribution to modern human ancestry would support the multiregional hypothesis, and the lack of such a contribution would be consistent with the replacement hypothesis.

Advances of the Late Twentieth Century
By the 1990's, the Neanderthals were well established as a group related to modern humans, but the questions remained: How close was the relationship? Did the two groups interbreed? Were Neanderthals a part of the evolutionary heritage of modern humans? During the 1990's, improved techniques and additional fossil discoveries led to greater understanding of the Neanderthals but little consensus on these questions. Afew examples will illustrate the situation. In a 1996 study, Jean-Jacques Hublin and several coworkers determined that Neanderthals found at an archeological site in France made bone tools and wore decorative emblems on their bodies, behaviors not uncovered with older Neanderthal fossils. They concluded that the Neanderthals were influenced by early modern humans who lived in the same area at the same time and that a reasonably elaborate cultural exchange must have occurred between the two groups. However, based on the strikingly different anatomy of the two groups' inner ears, they also concluded that the Neanderthals and modern humans did not interbreed. The investigators reasoned that if interbreeding had occurred, the two groups would have shared a common ear structure. In 1997, Matthias Krings, Svante Paabo, and their colleagues isolated and engineered deoxyribonucleic acid (DNA) from the mitochondria of Neanderthal bones and compared it toDNAfrom modern human mitochondria. They found the Neanderthal DNA to be quite different from that of modern humans and concluded not only that the two groups were different species but also that Neanderthals were not ancestral to modern humans. In 1998, Daniel Lieberman proposed that a reduction in the length of the sphenoid bone during embryology can explain most differences between the two groups' skulls. The sphenoid is a bone in the skull of both Neanderthals and modern humans, and Lieberman showed it to be shortened in modern humans but not in Neanderthals. Hehypothesized that the impact of shortening the sphenoid resulted in the modern human skull characteristics, while the longer sphenoid resulted in the Neanderthal skull. Based on the fundamental nature of the change, he concluded that Neanderthals do not belong to the same species as modern humans and were probably not ancestral to modern humans. In 1999, CidГ lia Duarte, Erik Trinkaus, and several colleagues discovered the buried remains of a four-year-old child in southern Spain. The skeleton was estimated to be about 24,500 years old, and they interpreted its anatomy to be a mixture of modern human and Neanderthal characteristics. Most anthropologists agree that southern Spain supported Neanderthal populations longer than other parts of the world, perhaps as late as 27,000 years ago, and that modern humans and Neanderthals coexisted in the region. Duarte, Trinkaus, and their group suggested that the skeleton they found demonstrated that the two groups did interbreed and that Neanderthals were part of the ancestry of Homo sapiens.

Consideration of this short list of studies in the 1990's demonstrates the state of knowledge about the Neanderthals' place in human evolution. Viewed alone, each study seems to clinch the position of its authors. In fact, the first three reinforce one another so well that Neanderthals would seem to be eliminated from direct participation in the evolution of modern humans. However, Duarte and Trinkaus's study would seem to clinch the opposite position, that Neanderthals were direct participants in the evolution of modern humans. This situation symbolizes the absence of consensus in the field. There are also established scientists with alternative viewpoints for each of these studies. Lieberman himself is a coauthor of a letter that criticizes his own conclusions about the sphenoid and points to the need for a better understanding of the development of primate skulls to help clarify the situation. Anumber of anthropologists have pointed out that Krings and Paabo's conclusions are extrapolated from a single, short segment of the mitochondrial DNA and that more extensive studies, including studies of DNA from the nucleus, are necessary before definitive conclusions can be drawn. In fact, nuclear DNA studies of modern humans have suggested that modern human DNAcomes from a number of sources rather than a single African source as in the out-of-Africa hypothesis. Clearly, extensive DNA comparisons would be helpful; however, DNA from fossils is difficult to find and difficult to work with, so an extensive collection of such studies is not likely to accumulate. Ian Tattersall, who rejects the Neanderthals as direct contributors to modern human evolution, has criticized Duarte and Trinkaus's data and their interpretation of the data. The verbal exchange has been bitter, not an unusual circumstance for disagreements in this field. Although anthropologists have learned an enormous amount about the Neanderthals, their relationship to modern humans continues to escape consensus. This is, without question, a result of the difficulty of the problem and the tentative nature of the evidence. Most agree that the Neanderthals were a successful group closely related to modern humans. Everyone's hope is that more fossils, improved technology, and fresh insight will clarify the question because understanding the Neanderthals is likely to contribute to an understanding of humanity.

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Principal Terms

deoxyribonucleic acid (DNA): the chemical that carries the instructions for all living things; closely related organisms have very similar DNA
genus: the first part of the scientific name of an organism; members of the same genus but different species are closely related, but cannot mate and produce fertile offspring
mitochondria: subcellular structures containing DNA used to estimate the relationships between groups of organisms; the more similar the DNA, the more closely related the groups
species: the second part of the scientific name of an organism; members of the same species can mate and produce fertile offspring
subspecies: the third part of a scientific trinomial, assigned to one of two groups that can mate and produce fertile offspring, but that have some strikingly different characteristics
taxonomy: the science of classifying and naming living and fossil organisms, or the classification and scientific name of a living or fossil group

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