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NEandERthaL anatomy

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Neanderthal anatomy represents a mixture of primitive characters, derived characters that are shared with other hominins, and derived characters that are unique to Neanderthals (see figure 27.1). In general terms, Neanderthals

Figure 27.1 Neanderthal anatomy.

Primitive features

Shared, derived features

Unique, derived features

Primitive features a. Long, low cranial vault b. Well-developed supraorbital torus

c. Large face, broad nasal opening d. Large dentition e. Absence of chin f. Broad cranial base

Shared, derived features a. Lateral reduction of brow ridge b. Reduced occipital torus

c. Rounder occipital profile d. Large brain e. Reduced facial prognathism

Unique, derived features a. Spherical shape of cranial vault (seen in rear view)

b. Midfacial projection, large nose c. Teeth positioned forward d. Retromolar space

FICURE 27.2 Skull shape: The triangle in the Neanderthal skull (left) shows the spatial relationships between the forward edge of the first molar (C), the lower edge of the cheek bone (A), and the upper edge of the cheek bone (B). A similar relationship drawn in a modern human skull (right, with a Neanderthal outline shaded in) produces a much flatter triangle, illustrating the significant forward protrusion in the Neanderthal face.

FICURE 27.3 La Ferrassie: A 50,000-year-old Neanderthal from the site of La Ferrassie, France, discovered in 1908. (Courtesy of Margot Crabtree.)

may be described as being robustly built, heavily muscled, and short in stature. Evidence of the heavy musculature appears in the extremely large muscle attachments and the bowing of the long bones. This structure implies that, whatever the details of Neanderthal subsistence, this species' daily life involved routine, heavy work. The short, broad trunk is consistent with life in a cold environment (Bergmann's rule; see unit 11), as are the short forearm and lower leg relative to the humerus and femur (Allen's rule; see unit 11). For much of the time of the Neanderthals' existence (between 150,000 and just less than 30,000 years ago), Europe and the Middle East were indeed cold, reflecting the end of the Pleistocene Ice Age.

An aspect of the skull anatomy—the extreme protrusion of the upper face—has also been speculated to be related to cold adaptation. To see this relationship, imagine a normal human face, but made of rubber. Now hold the nose and pull it out several inches. This protrusion of the upper face and a broad nasal aperture combine to produce a large chamber in the nasal passage; according to University of New Mexico anthropologist Erik Trinkaus, this chamber would provide an effective way to warm frigid air before it enters the lungs. Two bony projections jut into the front of the nasal cavity from either side, an anatomical feature not seen in any other hominins. (See figures 27.2 and 27.3.)

Body weight for the Neanderthals is estimated at 63.5 kilograms for males and 50 kilograms for females; statures are estimated at 1.67 meters for males and 1.60 meters for females. Despite their short stature, Neanderthals had large brains, an average of 1450 cubic centimeters—some 100 cubic centimeters larger than the modern average. The significance of the larger brain remains a matter of speculation.

Inevitably, brain size impinges on the question of Neanderthals' capacity for spoken language. Does a human-size brain imply a human level of language capacity? Nothing in the neuroanatomy of Neanderthals would deny this capability. These specimens share the overall shape of brains of earlier humans, with the brain appearing low and broadest near the base, with small frontal lobes and large occipital lobes (at the back of the brain). The discovery of a hyoid bone at the cave site of Kebara, Israel, similarly offers no characters that would rule out language capacity. The hyoid attaches to the base of the tongue, and thus is important in the mechanics of verbalization. The Kebara hyoid is modern in all respects, implying no mechanical barrier to spoken language in this respect. Evidence of limited verbal skills does appear in the structure of the larynx, which is inferred from the shape of the cranial base (see unit 32).

In 1998, researchers from Duke University published their measurements on the size of the hypoglossal canal in Neanderthal crania, and compared the results with those of modern humans. The canal is the hole through which nerve fibers exit from the brain to the tongue. The sizes were comparable, the researchers said, indicating the control of fine tongue movements was similar in both hominins, which they took to indicate some degree of spoken language. (The canal is much smaller in other creatures.) However, David DeGusta and his colleagues at the University of California, Berkeley, published data a year later comparing canal size in a range of hominins, including australopithecines. They concluded that there was too much variation within and between species for a sound conclusion to be drawn. Most anthropologists assume that if Neanderthals did indeed have some form of vocalization that was more advanced than in nonhuman primates, it was probably rather basic.

The Neanderthal pelvis is unique. In incomplete specimens, the pelvic canal appears to be unusually large, prompting speculation that the gestation was prolonged in this species and that the infant at birth was larger than in modern humans. When a more complete specimen came to light in 1987 (from Kebara), it revealed that the pelvic canal is not unusually large—just that the pubic bone is extraordinarily long.

NEANDERTHAL BEHAVIOR

Neanderthals lived by hunting and gathering, probably in small, nomadic groups, an existence that, judging from their extremely robust anatomy and large muscle attachments, evidently required extraordinary strength. Their tool technology employed the Levallois technique (see unit 23) to produce flakes that were then further worked to yield as many as 60 different implements, according to François Bordes, a French archeologist. For the Neanderthals, this Middle Paleolithic technology is termed the Mousterian technology, with the name being derived from a cave at Le Moustier, France. (See figure 27.4.) The Mousterian flakes could be used for many purposes, including cutting flesh, scraping hides, and working wood. Mousterian assemblages show little use of bone, antler, or ivory.

Toward the end of the Neanderthals' tenure, a second, more refined tool assemblage appears in western Europe. Known as the Chatelperronian, after a cave site near to Chatelperron in France, this technology was long a mystery to archeologists. (See figure 27.5.) While it is clearly Upper Paleolithic in character, having fine blades and artifacts made

FICURE 27.4 Mousterian tools: Neanderthals made stone tools using the Levallois technique, which involves striking flakes from a prepared core and then fashioning tools from the flakes. The French archeologist François Bordes has identified 20 different artifact types in the Mousterian.

from bone, antler, and ivory, this assemblage also includes at least 50 percent of flake tools, like those seen in the Mousterian. In 1979, Neanderthal remains were found in association with Chatelperronian tools at the cave site of Saint-Cesaire in France, which is dated at 36,000 years old. In 1996, a second such association was demonstrated at the site of Arcy-sur-Cure, also in France. In this case, part of a 33,000-year-old temporal bone of a child was identified as Neanderthal on the basis of the structure of the bony labyrinth (inner ear), in which Neanderthals are derived with respect to Homo erectus and modern humans.

These two associations support the conclusion that the Chatelperronian was a Neanderthal industry, produced when Neanderthals and modern humans coexisted in western Europe. Upper Paleolithic technology first appeared in western Europe 40,000 years ago; called the Aurignacian (see unit 30), it represented the work of modern humans. Whether the Chatelperronian industry is a home-grown invention of the late Neanderthals or a result of cultural contact between Neanderthals and modern humans remains unknown.

FICURE 27.5 Chatelperronian tools: Late in Neanderthal history, populations in western Europe manufactured tools that included many Upper Paleolithic elements, such as blades. This industry is called the Chatelperronian.

Another tool that the Neanderthals used routinely was their front teeth. This dentition is often worn with characteristic shelving, perhaps through repeated biting or pulling on hide or other soft but tough material.

Remains of Neanderthals have often been found in caves, sometimes in circumstances suggesting deliberate burial, as at the Kebara Cave in Israel, for example. (See figure 27.6.) A 40,000-year-old skeleton discovered in a cave site near La Chapelle-aux-Saints, France, was found together with a bison leg, other animal bones, and some flint tools. And a woman's skeleton was also found in an exaggerated fetal position in the cave of La Ferrassie. Many other examples are described in the literature, often with the assumption that burial was deliberate and associated with ritual practice.

Some of the "burials" can probably be explained by natural events, such as the collapse of cave roofs on occupants or abandoned bodies, and thus are devoid of ritual. But chance would have to be invoked in too many other cases to explain associations of bodies and stone tools, of alignments of bodies, and so on. The evidence is convincing that Neanderthals, and probably other archaic sapiens, occasionally buried their dead with a degree of ritual that we recognize as human. The act of burial is probably one reason why so many Neanderthal skeletons have been recovered.

FICURE 27.6 Neanderthal burial: This skeleton was recently recovered from the Kebara Cave, Israel, where it was evidently the subject of deliberate burial. (Courtesy of Ofar Bar-Yosef.)

Finally, there have been many tantalizing hints over the years that Neanderthals practiced cannibalism. A detailed analysis of cutmarks and breakage patterns on 78 bones from the cave site of Moula-Guercy, in southern France, appears to confirm that suspicion. The bones of red deer and Neanderthals in the same cave appear to have been handled in identical fashion. "The circumstantial forensic evidence of cannibalism is excellent," commented Alban Defleur, head of the French/American team that carried out the study. "No mortuary practice has ever been shown to leave these patterns on the resulting osteological assemblages." In other words, the object of the ancient slicing and bone breaking was to get meat, not to observe ritual.

A BRIEF HISTORY OF DISCOVERY AND INTERPRETATION

In August 1856, quarry workers in the Neander Valley, Germany, unearthed humanlike bones in a cave, Feldhofer Grotto, above the Düssel River. The fossilized remains—the top of a cranium, some leg and arm bones—were taken to Carl Fuhlrott, a mathematics teacher and local historian known to be interested in natural curiosities. Clearly the remains of a bulky and powerfully muscled individual, they were unlike anything Fuhlrott had seen before, so he sought the more informed viewpoint of Hermann Schaaffhausen, a professor of anatomy at the University of Bonn.

The anthropological community's reaction to the Neanderthal bones was mixed: some believed they represented a primitive race of human; others thought they belonged to a diseased individual; few considered the Neanderthal to be part of human ancestry. William King, an Irish anatomist, was unusual among his colleagues for regarding the Feldhofer specimen as different from Homo sapiens, and in 1864 he gave it the species name Homo neanderthalensis.

By the end of the century, the discovery of more fossil individuals with the same suite of curious anatomical characteristics effectively undermined the notion that pathology explained the appearance of the Feldhofer individual. More important, Eugene Dubois's discovery of Pithecanthropus in the early 1890s forced serious consideration of what ancestral forms of human might have looked like (see unit 24).

Gustav Schwalbe, a professor at the University of Strasbourg, suggested at the turn of the century that both Pithecanthropus and Neanderthal were part of a steady progression from primitive to modern human beings—a pattern known as unilinear evolution. Under this view, Neanderthals are usually given a subspecies attribution, Homo sapiens neanderthalensis, while modern humans would be Homo sapiens sapiens.

Neanderthals' status as a direct human ancestor did not last long, for two reasons: (1) the 1908 discovery, and subsequent misinterpretation, of the famous Neanderthal skeleton from the site of La Chapelle-aux-Saints, and (2) the 1912 discovery, and subsequent misinterpretation, of parts of the Piltdown Man, which turned out to be one of the biggest scientific hoaxes of all time (see unit 3).

The La Chapelle-aux-Saints Neanderthal was virtually complete, offering anthropologists the first opportunity to compare in detail Neanderthal anatomy with that of modern humans. The skeleton was sent to the Museum of Natural History in Paris, where Marcellin Boule undertook a detailed study, beginning in 1908. The picture Boule sketched of the Old Man of La Chapelle—and by implication all Neanderthals —was less than flattering. Effectively, he described a slouching, bent-kneed, bent-hipped semi-idiot. Very quickly the anthropological establishment accepted Boule's characterization of Neanderthals, and pronounced the species to be an evolutionary specialization that went nowhere. (See figures 27.7 and 27.8.)

Boule's description of Neanderthals as an evolutionary dead end left modern man without an ancestor. In 1912, Piltdown Man appeared on the scene and was accepted by many (but not all) anthropologists as our direct ancestor, filling this void in our evolutionary past. The so-called pre-sapiens theory was developed at this point, which argued that there had been an ancient split in the human lineage which led to the early appearance of a relatively modern skeletal form alongside a more archaic hominid, represented in the fossil record by the Neanderthals. The pre-sapiens theory effectively dominated anthropological thinking for almost 50 years, despite various vigorous efforts to dislodge it. For instance, the American anthropologist Ales Hrdlicka tried, but failed, to resurrect the unilinear hypothesis in the 1920s, based on anatomical and archeological arguments. He called his hypothesis the "Neanderthal phase of Man."

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FICURE 27.7 Boule's view of Neanderthals: The French prehistorian conducted a detailed study of the La Chapelle-aux-Saints skeleton, concluding that it was primitive in many ways. He drew this famous comparison between the Neanderthal (left) and a modern human.

New fossils discovered during the 1920s and 1930s in Europe and Asia initially failed to shake the pre-sapiens hypothesis, even though these specimens displayed Neanderthal-like features in these parts of the world at different times in prehistory. Later, the German anatomist Franz Weidenreich (see unit 24) invoked these fossils in a sophisticated elaboration of Hrdliccka's Neanderthal-phase hypothesis. According to Weidenreich, the pithecanthropines gave rise to the Neanderthals, which were directly ancestral to modern humans—in broad outline, a scheme reminiscent of the hypothesis put forth by Gustav Schwalbe 40 years earlier. Moreover, Weidenreich envisaged parallel evolutionary lineages

FICURE 27.8 Boule's influence: Marcellin Boule's conclusions about the primitiveness of Neanderthals influenced the profession's view, as seen here in a depiction of Neanderthal life drawn under the supervision of Henry Fairfield Osborn in 1915.

FICURE 27.9 Neanderthal rehabilitation: This depiction of Neanderthal, created by Carton Coon in 1939, was meant to show a "normal" Neanderthal, as evidenced by the street clothes.

FICURE 27.9 Neanderthal rehabilitation: This depiction of Neanderthal, created by Carton Coon in 1939, was meant to show a "normal" Neanderthal, as evidenced by the street clothes.

in various regions of the Old World, all leading through separate Neanderthal-like stages to the modern geographical variants of modern humans. Weidenreich's proposal was dubbed the candelabra model of modern human origins— drawn schematically, the long regional ancestries resemble an array of candles. This model, which was elaborated during the 1940s, is the precursor to a major position in the current debate (namely, the multiregional evolution model; see unit 28).

Despite Weidenreich's efforts, the unilinear point of view was slow to re-emerge. Eventually, a confluence of events through the 1940s, 1950s, and 1960s overturned the dominance of the pre-sapiens theory; instead, it became just one of several competing theories, which included in their number the unilinear model. The first of these events was the development of the synthetic theory of evolution (see unit 4), which allows for anatomical variation within species. The second was the exposure of the Piltdown fossils as a hoax, which removed this pillar of support in a single stroke. The third event was the re-evaluation of the La Chapelle-

aux-Saints skeleton, which showed that it was not brutish, as Boule had concluded.

The rehabilitation of Neanderthal was effectively completed by Loring Brace, of the University of Michigan, whose 1964 paper, "The Fate of the 'Classic' Neanderthals," was highly influential. Brace re-examined the La Chapelle-aux-Saints skeleton and, like Straus and Cave before him, concluded that Boule had described anatomical features that simply were not present. (See figure 27.9.) By the late 1960s, Neanderthals had been restored, in many people's eyes, to their rightful place: as direct ancestors of modern humans. The unilinear theory was at last successfully revived, now as one of a handful of competing theories, including the pre-Neanderthal hypothesis and a version of the pre-sapiens hypothesis.

Brace added an extra stage to Schwalbe's original, three-stage scheme to transform it: australopithecines to pithecan-thropines to Neanderthals to modern humans. According to Brace's so-called single-species hypothesis, only one species of hominin existed at any given period in human evolution—

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• Boundary of the Neanderthal world
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FIOU RE 17AO Geographical distribution: Neanderthal populations were confined to Europe, the Middle East, and western Asia. (Courtesy of Chris Stringer.)

the ultimate expression of the unilinear pattern (see unit 3). Milford Wolpoff, also at Michigan, joined Brace as a vigorous supporter of this hypothesis. In the mid-1970s, the discovery of the coexistence at Koobi Fora of a small-brained, highly robust individual (KNM-ER 406, Australopithecus boisei) and a large-brained, nonrobust individual (KNM-ER 3733, Homo ergaster) demonstrated that the single-species hypothesis was invalid, at least for that period of human prehistory (close to 2 million years ago).

Wolpoff, now a major protagonist in the current debate on the origin of modern humans, nevertheless insists that the unilinear hypothesis holds for the later stages of human prehistory. A scientific tradition carrying the names of Schaaffhausen, Schwalbe, Hrdliccka, Weidenreich, and Brace is therefore continued by Wolpoff. In unit 28, we will see how the modern version of this tradition measures up against the modern version of denying direct ancestry between Neanderthals and modern humans. (See figure 27.10.)

implications of neanderthal dna

One of the more dramatic developments in the study of Neanderthal prehistory came in mid-1997, with the report of the extraction of mitochondrial DNA from the fossilized bones of the type specimen, discovered in the Neander Valley in 1856. Comparison of a short (328 base pair) sequence of Neanderthal mitochondrial DNA with that from a large selection of modern individuals showed it to be very different. For instance, the average number of nucleotide differences in this sequence among modern humans is eight; by contrast, the Neanderthal sequence differed in 28 nucleotide positions, implying that it was genetically very distant from modern humans, and could not have been ancestral to them. Based on the number of differences in nucleotide positions between modern humans and Neanderthals, the joint German/ American team that did the work calculate that the last common ancestor of modern humans and Neanderthals lived about 600,000 years ago, and that modern humans originated in Africa, as the "out of Africa" hypothesis argues (see units 28 and 29). Further reports of comparisons of Neanderthal and modern human DNA have strengthened the original conclusion. For instance, DNA extracted from a 29,000-year-old Neanderthal fossil from Mezmaiskaya cave, in the northern Caucasus, shows similarities with the Feldhofer samples, but no modern human sequences were detected.

KEY QUESTIONS

• What aspects of Neanderthal anatomy imply an adaptation to cold environments?
• What is the most likely origin of the Chatelperronian tool industry?
• Why did so much resistance arise against accepting Neanderthals as a form of ancient human when they were first discovered?
• How is the current taxonomic status of Neanderthals best described?

KEY REFERENCES

Beerli P, Edwards SV. When did Neanderthals and modern humans diverge? Evol Anthropol 2002;11(suppl. 1):60-68.

DeGusta D, et al. Hypoglossal canal size and hominid speech. Proc Natl Acad Sci USA 1999;96:1800-1804.

Gargett R. Grave shortcomings: the evidence for Neanderthal burial. Curr Anthropol 1989;30:157-177.

Hublin J-J, et al. A late Neanderthal associated with Upper Paleolithic artifacts. Nature 1996;381:224-226.

Kay R. The hypoglossal canal and the origin of human vocal behavior. Proc Natl Acad Sci USA 1998;95:5417-5419.

Klein RG. Neanderthals and modern humans in West Asia. Evol Anthropol 1995/96;4:187-193.

Krings M, et al. Neanderthal DNA sequences and the origin of modern humans. Cell 1997;90:19-30.

--. A view of Neanderthal genetic diversity. Nature Genetics

2000;26:144-146.

Orchinnikov IV. Molecular analysis of Neanderthal DNA from north Caucasus. Nature 2000;404:490-493.

Schwartz JH, Tattersall I. Significance of some previously unrecognized apomorphies in the nasal region of Homo neanderthalensis. Proc Natl Acad Sci USA 1996;93:10852-10854.

Shea JJ. Neanderthals, competition, and the origin of modern human behavior in the Levant. Evol Anthropol 2003;12:173-197.

Smith FH. The Neanderthals: evolutionary dead ends or ancestors of modern people? J Anthropol Res 1991;47:219-238.

Spencer F. The Neanderthals and their evolutionary significance: a brief historical survey. In: Smith F, Spencer F, eds. The origins of modern humans. New York: Alan R. Liss, 1984:1-49.

Stringer C. New perspectives on the Neanderthals. Evol Anthropol 2002;11(suppl. 1):58-63.

Stringer CB, Gamble C. In search of the Neanderthals. London: Thames and Hudson, 1993.

Tattersall I. The last Neanderthal. New York: Macmillan, 1995.

Tattersall I, Schwartz JH. Hominids and hybrids: the place of Neanderthals in human evolution. Proc Natl Acad Sci USA 1999;96:7117-7119.

Trinkaus E, Shipman P. The Neanderthals. New York: Alfred A. Knopf, 1993.

White T. Once were cannibals. Sci Am 2001;265:58-65.

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