The Grisly Folk The Emergence of the Neanderthals

Most of the group had left the cave earlier that day. Only one elderly male was left behind. In his youth he had been attacked by a saber-tooth when part of a hunting group; with the help of his clan he had just managed to survive, his right arm torn off, his right leg crippled, his left eye put out as he fell heavily in the attack. He was now a cripple and depended on the generosity of his group to survive.

It was midafternoon when the valley and the surrounding region were rocked by a massive earthquake. The man in the cave could not escape before a large portion of the roof collapsed on him, killing him instantly. Members of the group had immediately rushed back to the cave from their hunting and foraging, only to find the cave entrance almost sealed off by rock debris. Their guttural yells into the cave were met with no reply. Within a few hours they had managed to open up the entrance so that at least they could enter. The old man could be partially seen, his body crushed by the rock debris that had until that afternoon been part of the cave ceiling. Members of the group mourned the loss of their kinsman. They removed the rock debris covering the old man and dug a shallow pit. They placed his body in the grave, turning him on his side, placing his arms across his chest, and fully extending his legs as though in a resting position. They then covered him with smaller pieces of the cave debris. They gathered whatever possessions they could find and moved out of the cave. While it was possible to clear away much of the debris and resettle the cave, all knew that the man's spirit would not enable them to stay. They needed to find another refuge. The leader of the group knew of another suitable cave only a few days distant. They set off down the valley.

In 1957, the skeleton of this elderly, crippled Neanderthal man was excavated from Shanidar, the cave in northern Iraq that had entombed him. This one cave was eventually to yield the remains of nine Neanderthals, some of them apparently buried — one, it is claimed, with a final gift of flowers placed in the grave. This area was just as unstable geologically in the recent past as it is today, and the roof fall was probably the result of an earthquake or major tremor. The significance of the Shanidar excavations and their later interpretation will be discussed presently.

In the previous chapter we examined the emergence of hominins within Europe and the likely role that H. heidelbergensis and the "Steinheim group" played in the later evolution of the Neanderthals. It is from around 200,000 to 25,000 years ago that we see the rise and fall of the "classic" Neanderthals that for a time dominated Europe, while slightly more gracile Neanderthal specimens have been recovered from eastern Europe (Croatia and elsewhere) and western Asia (Israel, Iraq, Uzbekistan). While Homo neanderthalensis flourished, populations of modern humans (H. sapiens) also appeared in western Asia and later in Europe; in numerous cases the two species coexisted, or interdigitated, for a short time at least.

The first fossil hominin to be recognized as not representing a modern human was the specimen from Feldhofer Cave in the Neander Valley, Germany (1856), found and recognized just three years before the publication of Charles Darwin's On the Origin of Species. This was not the first discovery of a Neanderthal specimen; unrecognized specimens had previously been discovered in Engis Cave in Belgium between 1829 and 1830 and in Forbes Cave in Gibraltar in 1848 (Stringer, 2000b). Not all accepted the significance of the Neander discovery. For example, some suggested that the robust nature of the specimen, with its thick supraorbital torus, merely represented an idiot who squinted a great deal (thus the developed brow ridges), while others argued that the "bowlegged" nature of the associated leg bone indicated that the individual was a Cossack who had chased Napoleon's army back into western Europe from his disastrous invasion of

Russia in 1812. The most staunch critique of the Feldhofer remains (or any other Neanderthal remains for that matter) as representing an early form of human was the man acknowledged as the father of paleopathology, Professor Virchow of Berlin University, who throughout the second half of the 19th century would characterize all of the unique morphological features of the Neanderthals as "pathological in nature," even when there was clearly no evidence for this (see Trinkaus & Shipman, 1993; Shreeve, 1995; Jordan, 1999). Some in England agreed with Virchow's argument. One commentator stated the following (quoted from Jordan 1999:16):

It may have been one of those wild men, half-crazed, half-idiotic, cruel and strong, who are always more or less to be found on the outskirts of barbarous tribes, and who now and then appear in civilized communities to be consigned perhaps to the penitentiary or gallows, when their murderous propensities manifest themselves.

The continuing discovery of similar fossils throughout Europe, including those from La Naulette and Spy in Belgium as well as from La Chapelle-aux-Saints (Figure 8.1), La Ferrassie, and La Quina in France showed that such interpretations could not stand up to the ever-increasing evidence, especially when these fossils were found with a distinct material culture as well as remains of extinct fauna. Many of the early researchers were now beginning to place these finds within a geological and archaeological context. In 1889 the first Neanderthals were being excavated from Krapina, in Croatia, and this site would soon yield the remains of two or three dozen Neanderthals. How could anyone now, including Virchow and his followers, still seriously argue that all of these remains represented a family or clan of pathological idiots! By the turn of the century, most accepted that the Neanderthals were a distinct nonpathological population, clearly different from ourselves. The argument now became whether they were our direct ancestors, or whether they represent a primitive human lineage that had become extinct.

The earliest fossil evidence for the emergence of full H. neanderthalensis is the French mandible from Montmaurin, which has the characteristic retro-molar space typical of Neanderthals and is dated to between 130,000 and 190,000 years ago. The recently redated skulls from Saccopastore in Italy come in next, at between 120,000 and 130,000 years old. The terminal point for the Neanderthals, at around 27 Ka (thousand years ago), is the leg bone fragment and lower jaw bone from Zafarraya in Spain. Most European

Figure 8.1 ► The classic La Chapelle-aux-Saints Neanderthal specimen from France.

and western Asian Neanderthal specimens fall between 40,000 and 65,000 years ago; these include the specimens from France, La Chappelle-aux-Saints (47-56 Ka), La Quina (35-65 Ka), La Ferrassie (<60 Ka), Regourdou (<60 Ka), Le Moustier (40-42 Ka); and Peche de L'Aze (45-55 Ka); from Italy, Grotta Guattari (50-60 Ka); Archi (<60 Ka); and Saccopastore (about 120 ka) (see Figure 8.2); from Belgium, Spy (<60 Ka) and Engis (<60 Ka); from Germany, the Feldhofer Cave in the Neander Valley itself (40 Ka); from Gibraltar, Forbes Cave (50 Ka); from Croatia, Vindija Cave (42 Ka, but some specimens may

Forbes Cave
Figure 8.2 ► One of the Italian Neanderthal specimens from Saccopastore.

actually be as young as 28,000 years); from Hungary, Subalyuk (60 Ka?); from its farthest eastern record, Uzbekistan, Teshik-Tash (70 Ka); from Iraq, Shanidar Cave (<50 Ka); from Syria, Dederiyeh (perhaps as much as 75 Ka); and from Israel, Kebara and Amud (<60 Ka). The dating of the Tabun Neanderthals, from the Mount Carmel foothills in Israel, remains problematic, but most consider them to be from between 70,000 and perhaps 120,000 years ago; if so, they currently represent the earliest penetration of western Asia by Neanderthal populations (see Trinkaus, 1983; Grün & Stringer, 1991; Grün et al., 1991; Stringer, 1998; Tchernov, 1998; Bar-Yosef, 1998; Tattersall & Schwartz, 2000; Schwartz & Tattersall, 2002, 2003; F.H. Smith, 2002).

The anatomical features of H. neanderthalensis are quite distinct from those of both the pre-Neanderthal populations of the middle Pleistocene (taking into account primitive retentions) and the later early specimens of our own lineage, H. sapiens (Figure 8.3). While they share primitive features with H. heidelbergensis, including the elongated cranium, low frontal, and developed supraorbital tori, the shape of the braincase is distinct in the two species: The cranial vault in H. neanderthalensis is higher and more rounded, with laterally projecting and rounded parietals, with a rounded and posteriorly projecting occipital bone, that is, an occipital "bun," and with undeveloped mastoids, though a marked mastoid crest. Seen from the back, the Neanderthal braincase has a characteristic cylindrical shape, described as "en bombe." The Neanderthals also had an

Braincase wider at middle or bottom

Low forehead (flatter frontal bone)

High rounded_

orbits

High, wide, and voluminous nose

Inflated cheeks -— (no canine fossa)

Large front teeth

Shorter, flatter parietal bones

Voluminous, long, wide, and low braincase

Low forehead (flatter frontal bone)

Shorter, flatter parietal bones

High rounded_

orbits

High, wide, and voluminous nose

Inflated cheeks -— (no canine fossa)

Large front teeth

Bones The Eye Fossa

Voluminous, long, wide, and low braincase

P^Retromolar gap behind 3rd molar Mental foramen (hole) usually under 1st molar

Shorter bulging occipital bone with suprainiac fossa

Larger juxtamastoid eminence Smaller mastoid process

P^Retromolar gap behind 3rd molar Mental foramen (hole) usually under 1st molar

Voluminous, long, narrower, and higher braincase

Longer curved occipital

Longer curved parietal bones

Voluminous, long, narrower, and higher braincase

Longer curved parietal bones

Longer curved occipital

Prominent Frontal Ridge

Brow ridge smaller or absent (especially at sides)

More prominent mastoid process (especially in males)

No retromolar gap behind 3rd molar Mental foramen (hole) usually

Figure 8.3 ► (a) The Chapelle-aux-Saints skull, indicating Neanderthal features. (b) The skull of anatomically modern Cro-Magnon 1, with the features characteristic of Homo sapiens labeled. Taken from Stringer and Gamble (1993), pp. 76-77.

Braincase widest higher up

Brow ridge smaller or absent (especially at sides)

Lower, squarer orbits Lower, narrower nose

— Canine fossa (hollowed cheeks)

Smaller front teeth

More prominent mastoid process (especially in males)

No retromolar gap behind 3rd molar Mental foramen (hole) usually

Figure 8.3 ► (a) The Chapelle-aux-Saints skull, indicating Neanderthal features. (b) The skull of anatomically modern Cro-Magnon 1, with the features characteristic of Homo sapiens labeled. Taken from Stringer and Gamble (1993), pp. 76-77.

enormous cranial capacity, with males averaging around 1600 cc (a greater average than that of modern humans), though there is some evidence that the cranial capacity of females was much less than that of males. The orbits are also round, as opposed to being more rhomboid. And while the supraorbitals in H. neanderthalensis are strongly developed, like those of H. heidelbergensis, they are different in shape, with distinct tori above each orbit as opposed to a single torus "shelflike" structure, and the bony thickening curves without a break down the lateral margins of the orbits. The midface is more prognathic and has often been described as though one had got hold of the nose and somehow stretched the midface forward, and this results in the zygomatics (cheekbones) appearing to be swept backward away from the midface. They have a well-developed nasal aperture, (broad, high, and prominent with marked internal nasal crest), a capacious maxillary sinus system, and a large anterior dental complex (mainly the incisor teeth). The mandible is also distinctive, with its retromolar space (a wide space between the last lower third molar and the ramus), a result of the forward-standing position of the dentition, so that the mental foramen appears to sit farther back, underneath the first molar, and the ascending ramus slopes back.

In body build, the Neanderthals were characteristically different both from their predecessors and from modern humans (Figure 8.4). The postcranial bones are robust and stout, with broad rib cage, long clavicle, wide pelvis, and relatively short and robust limbs that have well-developed muscle markings (see Stringer & Trinkaus, 1981; Trinkaus, 1983; Stringer & Gamble, 1993;

Large and wide rib cage

Large clavicle

Wide scapula with more muscle attachments along rear edge

Large shoulder joint

Large and thick patella

Large ankle joint

Wide and strong toe bones

Large and wide rib cage

Large clavicle

Wide scapula with more muscle attachments along rear edge

Large shoulder joint

Large elbow joint Bowed and short forearm Wide hips Large hip joint, rotated outward Hand with strong grip and wide fingertips

Long, thin superior pubic ramus Rounded, curved, and thick-walled femur shaft

Large and thick patella

Short, flattened, and thick-walled tibia

Large ankle joint

Wide and strong toe bones

Large elbow joint Bowed and short forearm Wide hips Large hip joint, rotated outward Hand with strong grip and wide fingertips

Long, thin superior pubic ramus Rounded, curved, and thick-walled femur shaft

Short, flattened, and thick-walled tibia

Tibia Fibula

Tarsals

Metatarsals

Phalanges

Facial bones

Scapula Sternum Rib

Humerus

Vertebra

Pelvis

Sacrum

Superior pubic ramus

Radius

Ulna

Carpals

Metacarpals

Phalanges

Femur

Tibia Fibula

Tarsals

Metatarsals

Phalanges

Figure 8.4 ► The skeleton of a "classic" Neanderthal (a) indicating typical features compared against the skeleton of an anatomically modern Homo sapiens (b).

Taken from Stringer and Gamble (1993), p. 79.

Facial bones

Howell, 1998; B.A. Wood & Richmond, 2000; Tattersall & Schwartz, 2000; Agusti & Antón, 2002; Schwartz & Tattersall, 2002). Indeed, Ruff et al. (1997) and Arsuaga (2002), using an association between the breadth of the hip bones (biiliac breadth) and overall estimated height, suggest that the average Neanderthal would have weighed around 168 pounds (75 kg), while many males would have exceeded 175 pounds (80 kg). Applying brain weight to body weight suggests that the Neanderthal brain was reduced as compared to H. sapiens because they have a projected brain that is 4.8 times the size expected for a mammal of its bulk, while H. sapiens has a brain that is around 5.3 times its expected weight (see Ruff et al., 1997; Arsuaga, 2002). The hand anatomy was strikingly modified: H. neanderthalensis had a powerful grip, with a broad, long palm, short fingers, and deep grooves for the interosseous muscles.

Whether H. neanderthalensis was cable of complex speech remains an endless, ongoing topic of debate. We currently have not the slightest idea whether Neanderthals processed the necessary neural adaptations for complex speech, for these simply do not fossilize (B.A. Wood & Richmond, 2000). Based on a study of Neanderthal mandibular form and basicranial morphology, Philip Lieberman (1989, 1991) argued that the Neanderthals were unlikely to have had a fully developed vocal tract and thus had limited abilities for speech. Arensburg (1989), in his study of the Kebara Neanderthal hyoid bone, however, has refuted much of Lieberman's argument. Some additional inferences have been made from the fossil record in relation to the size of the hypoglossal and vertebral canals, though whether they are developed enough for the necessary innervation of the tongue and breathing to control movements related to speech remains problematic (Aiello & Dean, 1990; Kay et al., 1998; DeGusta et al., 1999; MacLarnon & Hewitt, 1999; B.A. Wood & Richmond, 2000).

Much has been said about the unique morphology of H. neanderthalensis, and the functional adaptation of the body plan has been discussed ad nauseam. Their robust body structure suggests that the upper limbs were likely adapted to heavy foraging, such as spear thrusting, while the lower limbs are those of long-range bipeds (B.A. Wood & Richmond, 2000). The large size of the anterior dentition and their observed wear patterns (much more heavily worn on the labial surfaces) suggest that they were used as "dental tools" in food preparation and perhaps as a vice, helping to grip mammal hides in their teeth while skinning them with stone tools (Trinkaus, 1983; F.H. Smith, 1983).

Many have considered the rise of classic H. neanderthalensis from western Europe to be associated with an adaptation to cold climates, while the slightly less robust Neanderthals from eastern Europe and western Asia might suggest tolerance of the slightly more temperate conditions in these areas. We think that the suggested cold adaptation of the "classic" neanderthals was an exaptation, prefigured in some of the morphologies of H. heidelbergensis and enabling the earliest representatives of the lineage to become increasingly adapted to the cold while not restricting them to the periglacial habitats of Ice Age Europe. We must remember, too, that not all morphological features are the result of functional adaptation, but can be related to genetic drift (random change), which must have impacted to some degree the small and isolated populations. Neanderthal children from a very early age display typically robust Neanderthal features, and they are already quite different from modern humans of the same age; the typical Neanderthal features continued to develop in slow progression from gracile juvenile to robust adult (see Trinkaus, 1983, 1986; Stringer et al., 1990). Tattersall and Schwartz (2000:207) insisted that the most remarkable thing about the Neanderthals was their extraordinary adaptability. With this we can only agree. The Neanderthal populations within Europe and western Asia do not bracket a consistently cold time period, and in Syria, Israel, Iraq, and southeastern Europe they lived in a very tolerably warm, productive climate — temperate at worst! — and only those from the younger European sites lived under the harsh Ice Age regimes to which our minds try to adapt them. And yet we must remember that they did not penetrate the tropics, because someone else was, however slightly, better adapted to the tropics; and to this crucial point we will return.

So how would we likely view the Neanderthals today if we saw a family of them walking down the main street of a busy city? Carleton Coon in the 1960s suggested that if we dressed up a Neanderthal man in a suit and gave him a shave and a shower, he would look a little different, but overall would not stand out in the crowd. We take the opposite stance and note, with a sad acknowledgment of the persistence of human racial prejudice and xenophobia, the stance of an unknown commenter who suggested that if a Cro-Magnon got on the same train and sat down next to you, you would probably change seats, but if a Neanderthal sat down next to you, you would change trains. The Neanderthals were very different from ourselves. Not one whit less intelligent, imaginative, dextrous, or adaptable, probably— merely different. As emphasized by Arsuaga (2002: 67):

The Neanderthals were not simply primitive versions of ourselves ... [They] continued to evolve and ... developed their own distinctive characteristics, just as we did elsewhere. The Neanderthals were not living fossils. They did not belong to the past, and they were not anachronistic. In their particular epoch, they were just as "modern" as our ancestors, the Cro-Magnons, were. The two species were simply different.

With the coming of the last interglacial (oxygen isotope Stage 5) in northern Europe around 130,000 years ago, we see the recolonization of high latitudes by the warmth-loving plant and animal communities that had previously been driven south. Pollen sequences show brief episodes of birch and pine, on to elm and oak, then alder, hazel, yew, and hornbeam, and finally returning to the beginning of the colder conditions with pine, spruce, and silver fir (see Mellars, 1996). It is also at the start of the previous interglacial that Europe was dominated by warm fauna, including forest rhino, hippo, straight-tusked elephant, lion, and hyena, even in northern Europe (Turner & Antón, 1997; Jordan, 1999; Agusti & Antón, 2002).

From 115,000 to around 75,000 years ago there was a succession of cold and slightly more temperate pulses, and then at 75,000 years ago there was a plunge into another long glacial period (Stage 4). It was during this period that glaciers covered most of present-day Scandinavia, and the British Isles and tundra/prairies, or "polar deserts," covered much of northern and central Europe. There was a brief return to the glacial and temporal pulses (Stage 3), but around 32,000 years ago the cold set in once more (Stage 2); at 21,000-17,000 years ago there was a return to maximum glacial conditions (the last glacial maximum, LGM), with glaciers covering much of present-day Poland, northern Germany, Great Britain, and Ireland (see Mellars, 1996; Agusti & Antón, 2002). The shorter interstadial periods during this time were in most cases either too brief or not sufficiently marked to show up in paleontological and geological climatic records, and probably also too brief for trees and associated faunal groups to migrate into northern Europe from their southern refuge areas (Mellars, 1996). That is not to say that these were necessarily hard times. These tundra/prairies would provide an abundance of herd game, which could probably be more easily hunted without the requirements of stalking and chasing individual deer and boars in the thick forests of the south. Some of these more temperate pulses around 30,000-40,000 years ago encouraged the migration of H. sapiens into

Europe. Then the Neanderthals lost out to the newcomers while populations of H. sapiens maintained and expanded their grip on these more favorable habitats. But that is another story, to which we shall return in the last section.

Molecular evidence has now offered strong support for a long separation of the Neanderthals from H. sapiens. In 1997 mtDNA was successfully extracted from the Neanderthal specimen from Feldhofer Cave in the Neander Valley, Germany (see Kahn & Gibbons, 1997; Krings et al., 1997; R. Ward & Stringer, 1997). Only small fragments of Neanderthal mtDNA were recovered. After copying these strands using the polymerase chain reaction technique, researchers were able to identify 378 base pairs. Comparison of these with samples of recent humans showed that they differ in around 27 base pairs (around 7%), while the variability within the modern human sample averages only 8 base pairs (2%). Subsequently, mtDNA has been extracted from two other Neanderthals: specimens from Vindija, in Croatia (Krings et al., 2000), and from Mezmaiskaya, in the Russian Caucasus (Ovchinnikov et al., 2000), proved as similar to the Feldofer samples as three random H. sapiens samples are to each other. It must be acknowledged that Gutiérrez et al. (2002) have raised questions about these results, both as to the analytical methodologies and the possibility of postmortem changes; yet the amplified sequence of a 12,000-year-old skull from Cheddar Gorge in southwest England showed only two deviations from people living in the area today. As such, modern humans and those living over 12,000 years ago are almost identical in their genetic structure, perhaps allaying some of the problems raised by Gutiérrez et al. (2002) and, in the process, further underlining the distinctiveness of the Neanderthals (Sykes, 2001). Likewise, the mtDNA of 8000- to 12,000-year-old Australian skeletons from Kow Swamp, in northern Victoria, as analyzed by Adcock et al. (2001), fits easily into the modern aboriginal Australian range of variation (we will comment in a later chapter about the Mungo DNA, which has been claimed to be different). But it is as well to remember that, as yet, no DNA has been extracted from any Cro-Magnon specimen — the H. sapiens that succeeded Neanderthals in Europe — so we cannot exclude the possibility that they, too, might have been "different."

Be that as it may, Neanderthals and modern humans are perfectly distinct. Modern humans arose recently in Africa as a distinct species and replaced the Neanderthals, who contributed few if any genes to modern human populations. In other words, there was little or no gene flow between them (see Kahn & Gibbons, 1997; Krings et al., 1997, 2000; R. Ward & Stringer, 1997; Klein, 1999), although occasional hybridization may indeed have occurred — see later! Thus there is nothing within the molecular data to support the idea that modern humans evolved from an earlier Neanderthal population. Indeed, using a "molecular clock," the last common ancestor between the modern human and Neanderthal lineages lived around 500,000-700,000 years ago. This is at around the same time that H. heidelbergensis appears in the fossil record.

Homo neanderthalensis made stone tools that were considerably more sophisticated than those observed in either H. heidelbergensis or their presumed ancestors from the "Steinheim group": the Mousterian industry (Figure 8.5). This toolkit is usually defined as part of a Mode 3 Technology, which makes its first appearance in the archeological record around 250,000 years ago (Clark, 1977; Foley & Lahr, 1997; Agusti & Anton, 2002).

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