Ancient History & Civilisation

CHAPTER 5

Getting Brainier

In contrast to the richness of the African fossil record before about 1.5 million years ago, the evidence for later hominid evolution in that continent thins out considerably. This is largely for reasons of geological accident, but it is also because the attentions of rather few paleoanthropologists have been spread out over a continent of vast size, so much of the record presumably remains unexplored. At the same time, for reasons of history as much as of their size and intrinsic importance, the Asian and European records have traditionally loomed larger in the hominid evolutionary story for the period following about 1.5 million years ago. Nonetheless, the logical place to begin our account of the phase of human evolution after the initial move out of Africa is still that continent, where a partial skull was found at the Ethiopian site of Bodo in 1976. This cranium boasts a brain volume of around 1250 cubic centimeters, substantially larger than anything attributed to Homo ergaster, and right at the maximum of the range for Homo erectus. And in its structure it resembles a species, Homo heidelbergensis, that was previously known best from Europe.

In 1908 Homo heidelbergensis, “Heidelberg man,’’ was described from a marvelously preserved lower jaw found in a gravel pit near the German village of Mauer, not far from the city after which the species was named. This jaw was unlike anything found before it (only Neanderthals, Homo erectus, and various ancient Homo sapiens were known at this point), and its discoverer had few qualms in attributing this curious specimen to a new species. The quite robust Mauer jaw possesses a ramus (the part that rises up to the jaw joint) that is notably wide from front to back but is also short from top to bottom. Its corpus (the tooth-bearing portion) is markedly tapered, decreasing in top-to-bottom thickness from front to back.

Later finds revealed that the Mauer specimen is quite unusual in these characteristics; but a whole suite of other features link it with a much better represented group of fossils from the southern French site of Arago. At about 400,000 years old, the latter group is in the same general time range as the best estimate for the Mauer jaw (500,000 years or so). The Arago site yielded not only several lower jaws and part of a pelvis but also an almost complete face with a matching parietal bone, the part forming the top and upper side of the cranium. The Arago cranium in turn recalls a number of other well-preserved skulls from sites around the world. These include not only the Bodo specimen but also crania from Petralona in Greece, Kabwe and Saldanha in southern Africa, and Dali and Jinniushan in China. Regrettably, none of these is well dated, but all do fit plausibly into the period between about 500,000 and 200,000 years ago.

There are certainly differences to be observed among these diverse fossils. For example, the Bodo skull has a large but low-set nasal opening, whereas the Kabwe skull, from what is now Zambia, has a much smaller and higher-set one. In the Bodo and Arago crania the front part of the brain lies much farther forward over the eyes than it does in the Kabwe and Petralona specimens. The shape of the rear of the skull varies somewhat. The eye sockets may be conformed a little differently. And within the whole group there is wide variation in the degree to which the craniofacial sinuses (cavities in the bone structure), and particularly the frontal sinus (the one over the eyes, where you experience ‘‘brain freeze’’ when you swallow a frozen drink too fast), are developed. Yet overall these fossils make a relatively homogeneous group; and for the present, at least, it makes a certain amount of sense to view Homo heidelbergensis as a highly successful species, probably of African origin, that became widespread throughout the Old World (Africa plus Europe and Asia).

This species had a brain that was relatively large, although not quite of a size comparable with the modern average. The face is big and projects forward, and it lies beneath distinctive and bulbous brow ridges that are thickest above the middle of each eye socket, and the front surface of which twists upward toward the sides. The lower jaw is long from front to back and bereft of anything resembling a chin. Interestingly, where the base of the skull of Homo heidelbergensis is preserved (best seen, perhaps, in the Bodo skull), it shows a distinct downward bend in front of the foramen magnum, the hole through which the spinal cord passes down to the vertebral column from the base of the brain. This is important, for the base of the skull is not only the bottom of the brain-case, but it is also the roof of the vocal tract, the space in which we form the sounds that emerge from our mouths as speech.

Language is perhaps the most striking possession of Homo sapiens among living creatures; and if we are fully to understand how various unique characteristics of modern human emerged, then it is important to discover when and how our ancestors became capable of speech. For even if the ability to produce the sounds of speech may exist independently of it, language as we know and use it could never have developed independently of the ability to produce speech. The basic vibrations that we manipulate to create the sounds that become speech are produced in our throats, at the vocal cords. But these vibrations are modified higher in the throat, by the muscles that surround the pharynx, a space that loops high above the larynx, or voice box, which contains the vocal cords.

In apes (and in newborn humans), the larynx lies high in the throat, and the skull base is flat. In the resulting short pharynx, sounds cannot be modified much. As the infant human grows, however, the skull base bends and the larynx descends, producing a long pharynx in which a greater variety of sounds can be produced. At least in part, this is a key to the remarkable vocal gymnastics that we perform each time we utter a sentence. Neither apes nor newborn humans can produce the range of sounds necessary for this, and the contour of the base of the skull does seem to be a fairly reliable indicator of the vocal tract’s potential for producing the sounds necessary for speech, even though the shortness of the face also plays a role. On the evidence of the bending seen in the base of the Bodo skull, it seems that much of this potential may well have been present in Homo heidelbergensis, as long ago as 600,000 years. Still, with the face as yet unretracted to produce balanced proportions of the pharynx and oral cavity, it is doubtful that the full human vocal apparatus was in place in Homo heidelbergensis, and there is no other evidence to suggest that these hominids actually spoke.

As with earlier hominid species, the appearance of Homo heidelbergensis is not accompanied by any notable change in technological equipment. The sediments from which the Bodo cranium was derived contain mostly Mode 1 artifacts, although Mode 2 tools (handaxes) are also documented in them. Beyond this, though, there is not much to be said at this point about the lifestyle of the Bodo hominid, and we have to turn to Europe for a better behavioral record of H. heidelbergensis. And as it happens this record is quite impressive, even though it is mostly limited to a handful of sites in France and Germany.

One of these sites is the cave of Arago in southern France in which were found the various H. heidelbergensis fossils that allow us to link the Heidelberg lower jaw to a specimen with a face. At Arago the hominid fossils were mixed in with broken animal bones and crude artifacts of Mode 1 type, and it seems that this site was indeed a place where hominids at least periodically gathered and carried out daily activities, including the butchering of animals. What daily life was like for these hominids, however, is better indicated at the locality of Terra Amata, a little bit to the east of Arago on France’s Mediterranean coast. This site, a little bit younger than Arago at about 350,000 years old, is thought to represent a beach camp that was seasonally occupied by Ice Age hunters. Stone tools, animal bones, and ashy sediments attest to the activities of early hominids, and at the site there are also indications of what appear to have been shelters. These, along with similar features at the 350,000-year-old site of Bilzingsleben, in Germany, are the earliest artificial structures on record.

The archaeologists who excavated Terra Amata have reconstructed the best-preserved of the shelters as a hut consisting of an oval saplings implanted in the earth, reinforced around the perimeter with stones, and brought together in the middle to make a roof. Whether or not this structure was covered with animal hides to waterproof it is a matter for conjecture, though the excavators think this was not the case. Just inside the hut, where the ring of reinforcing stones is interrupted for the entrance, is a shallow, scooped-out depression containing ash, animal bones, and burned stones, evidence of a hearth where a fire once burned and meat was cooked. This is among the earliest evidence of the domestication of fire, something for which we find consistent evidence only after this time—although a couple of possible instances of hominid use of fire have been reported as early as about 1.6 million years ago, and at one site in Israel fire was consistently used over a long period of occupation some 790,000 years ago.

Hut-like structures like these were probably made by Homo heidelbergensis 400,000 years ago.

Hut-like structures like these were probably made by Homo heidelbergensis 400,000 years ago. These shelters, built on an ancient beach at Terra Amata, in southern France, were up to 25 feet long. The cutaway reveals an interior containing a circular hearth and stone tools. Artwork by Diana Salles, after a concept by Henry de Lumley. From Ian Tattersall, The Last Neanderthal, 1995.

The stone tools found at Terra Amata are rather more impressive than those from Arago and include crude handaxes and cleavers, as well as flakes of various kinds. Interestingly, Acheulean tools seem to have found their way to Europe rather late—and they barely made it to eastern Asia at all. Before a recent report from China, only the occasional handaxe had ever been found to the east of the Movius Line, a conceptual divide, first noted by the Harvard archaeologist Hallam Movius, that separates eastern and most of southern Asia from the rest of the continent. But once handaxes made it into Europe they became a consistent feature of tool kits, at least locally, until they were supplanted by a new method of toolmaking known as ‘‘prepared core.’’ Best known from the Levallois technique, named for the suburb of Paris where examples of it were first found, prepared-core toolmaking involved carefully preparing (shaping with numerous blows) a piece of stone (the ‘‘core’’) in such a way that a single final blow—probably effected by bashing the core on a stone anvil, rather than by striking it with a hammer-stone— could detach from it a thin, light, and effectively finished tool bearing a continuous cutting edge around its periphery.

A whole variety of flakes could be produced in this way, and these could in turn be retouched to various specifications. One of the resulting forms was the flake-based handaxe, generally a smaller tool than the Acheulean handaxe, sometimes itself made on a large flake, but with the same basic shape. Many of the tools manufactured in this way may have been attached to handles, creating compound tools that were much more complex in both concept and potential uses than simple handheld rock implements had been.

It is not easy to judge what this new kind of tool implied about the cognition and lifestyles of the hominids who made them. The underlying technological notion is a great deal more complex than anything involved in simply chipping a piece of stone into a particular shape. So here were hominids (plausibly Homo heidelbergensis or something like it) who were capable of quite intricate (though probably intuitive) reasoning, although nothing else in the archaeological record they left behind convincingly suggests that they had the symbolic mental processes and linguistic abilities that we have today. There is also some doubt among archaeologists whether these hominids were clever, guileful hunters along the lines of Homo sapiens. Indeed, by the middle 1990s it had become generally believed that hominids of this kind did not have any of the hunting sophistication of historically documented hunting-gathering peoples. However, an extraordinary discovery made in 1995 at the German site of Schoeningen may throw this assumption at least partially into doubt.

An archaeologist holds in his left hand a stone core that he has carefully prepared by shaping it with numerous blows to both sides.An archaeologist holds in his left hand a stone core that he has carefully prepared by shaping it with numerous blows to both sides. In his right hand he holds the sharp flake he has just struck from it with a single final blow. ‘‘Frepared-core’’ toolmaking represented a revolution in stoneworking technology when it was introduced about 300,000 years ago. Courtesy of Kathy Schick and Nicholas Toth, Stone Age Institute.

Wooden implements preserve very poorly, so they seldom make it into the archaeological record. Normally they rot away within a year or two, or at best within a few millennia, leaving no trace behind. But in a peat bog at Schoeningen, archaeologists discovered not only flint artifacts and cut-marked animal bones but also several miraculously preserved wooden spears dating from about 400,000 years ago, as well as some notched pieces of wood that had probably served as handles for stone-tipped implements. The spears, six to seven feet long, were made from individual spruce saplings, their carefully sharpened tips coming from the bottom of the tree, where the wood is hardest. Each spear was skillfully worked so that its weight, hence the center of balance, was concentrated two-thirds of the way forward. This is exactly the shape of a modern javelin, and it is claimed that these spears were made for throwing and not for thrusting—although their effectiveness as thrown weapons has been questioned. However, their very form does suggest that the hominids who made them may have been equipped for a hunting style that was considerably more sophisticated than many archaeologists had expected.

Before the Schoeningen find, the oldest reasonably complete wooden implement known was a 125,000-year-old spear point found at another German site, Lehringen, lying between the fossilized ribs of a straight-tusked elephant. This more recent spear has been interpreted as a handheld thrusting implement that would have to have been wielded close-up, a dangerous proposition at best. The Schoeningen hunters, on the other hand, might have hurled their spears at their prey from a safe distance— an enormous improvement in hunting technique.

Once again, we are reminded of—and frustrated by—how indirectly stone tools reflect actual behaviors. Had the Lehringen pachyderm been initially attacked with throwing spears, only finally being dispatched with the thrusted spear? If, as we must suppose, the Lehringen spear was wielded by a Neanderthal, a distinctive group of hominids that occupied Europe and the western part of Asia from some time before about 200,000 years ago until around 30,000 years ago, did the Neanderthals possess only thrusting spears? We can only guess at the answers to questions such as these; but somehow it doesn’t seem very likely that after the throwing spear had been invented, putatively well before Neanderthal times, it was promptly forgotten. Indeed, it has been suggested that the very presence of hominids in northern Europe could well have depended on the possession of efficient hunting techniques, because in this area of harsh living conditions large-bodied mammals might have been the only important source of sustenance available to hominids for most of the year.

The stage is thus set for a prolonged debate about the hunting prowess of early Europeans and about how good an indication stone tools are of other aspects of technology, let alone of broader lifestyles. Nonetheless, within the stoneworking domain there is little doubt that the prepared-core technique was explored to its utmost by the Neanderthals.

Homo neanderthalensis was the very first kind of extinct human to be found and thus occupies a very special place in the history of paleoanthropology. In 1856 lime miners emptied a cavity (the Little Feldhofer Cave) in Germany’s Neander Valley near Dusseldorf, unearthing part of a hominid skeleton in the process. Its limb bones were humanlike although robust, but it was the skullcap that was really peculiar. For although the brain evidently had been very large (of modern human size), it had been contained within a very distinctive skull vault: long, low, bulging at the rear, and adorned in front with large brow ridges that arched separately over each eye socket.

This discovery was made three years before Charles Darwin published On the Origin of Species. In the absence of the idea that these bones might represent an extinct relative of mankind, there was almost no option but to consider that this odd, big-brained skull was that of a strange version of Homo sapiens, the only hominid then known on the planet. The apparent possibilities boiled down to two: either these remains were pathological, those of a diseased and deformed individual; or they were the bones of a member of one of the ‘‘barbarous’’ tribes that had formerly occupied Europe (and about whom Roman chroniclers had complained at great length). Almost everyone who entered the initial debate about the Feldhofer specimen took one or the other of these positions. Even the comparative anatomist and evolutionist Thomas Henry Huxley, later known as ‘‘Darwin’s Bulldog’’ for his tenacious defense of Darwin’s ideas, opted for one of these choices, interpreting the specimen as that of a rather brutish form of modern human. For although in his 1864 book Evidences as to Man’s Place in Nature he referred to the Neanderthal cranium as the ‘‘most pithecoid [ape-like] of known human skulls,’’ he also saw it as being linked to ‘‘the highest and best developed of human crania.’’ The sole exception to this chorus was the Irish anatomist William King, who assigned this strange material to its own new species, Homo neanderthalensis.

Since 1856, hundreds of fossils comparable with the Feldhofer specimen have been found and given the Neanderthal name. They come from dozens of sites, from Uzbekistan in the east all the way to the Atlantic coast of Europe in the west, and from Wales and Germany in the north down to Gibraltar and Israel on the Mediterranean. All Neanderthals share a distinctive anatomy that is very different from our own; but still the old notion lingers that these hominids somehow represent a bizarre (and by implication inferior) version of Homo sapiens. Perhaps, at least today, this is because warm-hearted paleoanthropologists find it somehow discriminatory to exclude a large-brained hominid such as H. neanderthalensis from the ‘‘privilege’’ of belonging to H. sapiens. But it bears repeating that if morphology means anything at all in our assessment of fossils (and if it doesn’t, what are we left with?), the Neanderthals were an evolutionary entity entirely separate from us. And they thus need to be understood on their own terms, not ours, and to be accorded their own separate identity.

Since the middle of the twentieth century it has frequently been claimed that Neanderthals were ancestors of Homo sapiens because various late Neanderthal fossils show ‘‘advanced’’ features and various early modern specimens show ‘‘primitive’’ ones. Such claims do not, however, stand up to close scrutiny. The fallback position from this is that Neanderthals and modern humans, belonging to the same species, interbred when they came into contact with each other; but again, the evidence in favor of this is scant at best. In 1999 it was claimed that the skeleton of a young child found at the Portuguese site of Lagar Velho represents a descendant of an intermixed Neanderthal/modern population. However, the anatomical evidence for this imaginative interpretation has been fairly described as ‘‘at best ambiguous’’; in addition, this child died just 24,500 years ago, long after the Neanderthals were extinct. William King may have based his claim for Homo neanderthalensis as a distinct species principally on the rather dubious grounds of his intuition that ‘‘the thoughts and desires that once dwelled within [the Feldhofer skull] never soared beyond those of a brute,’’ but there can be little doubt that his resulting classification was entirely correct.

Homo neanderthalensis is by far the best-documented of all extinct hominid species. And it is clear that, just as modern humans do today, individual Neanderthals (and populations from different times and places) differed from each other in their bony structure. Some had lower skull vaults than others, for example, or the bridges of their noses projected more or less horizontally. Some had retreating jawlines; others had more vertical ones. But equally clearly, these variations were on a different theme than ours. They do not simply represent various extremes of the Homo sapiens spectrum. Aside from such variations, all Neanderthals had large brains (about 1,200 to 1,740 cubic centimeters in volume; ours range between 1,000 and 2,000 cubic centimeters). And these brains were enclosed in relatively long and flattish skull vaults bearing low foreheads behind distinctive double-arched brow ridges. In profile, you can see that these ridges are smoothly curved as they rise from the roofs of the eye sockets and flow to the frontal bone behind.

In all of these respects the Neanderthal skull contrasts with that of Homo sapiens, in which the cranial profile is high and rounded, the dome rising directly above generally small brow ridges that are divided above each eye into distinct central and lateral portions separated by an oblique groove—which you can easily feel above your own orbits. Whereas the rear of the H. sapiens skull, as seen from the side, is commonly smoothly curved, that of H. neanderthalensis tends to protrude, sometimes in a distinct ‘‘bun.’’ High up at the back of the skull, in the midline, there is a curious area of pitted bone, called a suprainiac fossa, which modern humans lack. Seen from the back, the braincase of Neanderthals is smoothly rounded on the sides, rather than having rather vertical side walls as ours does. Unlike the face of modern humans, which is small and tucked beneath the front of the braincase, the Neanderthal face protrudes forward in the midline and is swept back at the sides, with sharply receding cheekbones. The area between the eyes and the mouth is puffed out by large sinuses. The Neanderthal nose is huge, and just within its opening lie some sideways-pointing structures unknown in humans (or in any other primates, for that matter) that are called medial projections and may reflect an unusual configuration of the respiratory system.

The lower jaw may or may not have minor swellings in the midline, but no Neanderthal known has anything like the specialized structure of the human chin. There are even substantial differences in the morphology of the teeth. This list of distinctions between H. neanderthalensis and H. sapiens skulls could go on and on, but the point should already be clear: Neanderthals and modern humans are exceedingly different in the way they are structured from the neck up.

How about from the neck down? Same story. Although H. neanderthalensis was built along the same basic plan as H. sapiens, it nonetheless showed numerous differences from us. Some of those differences are, indeed, striking. The Neanderthal pelvis, for example, is broad, flaring widely to the sides. The collarbones are very long, and the rib cage, narrow at the top, broadens greatly to the bottom. There is hardly any waist where the wide lower part of the rib cage meets the flaring pelvis. The limb bones are very thick-walled. They tend to be bowed out somewhat and have very large joint surfaces at their ends that swell out noticeably from the shafts. Again, there is much more detail that could be mentioned, but the bottom line is that the Neanderthals would have presented a very different figure than modern humans do. This would have included the way they moved, for the peculiarities of the Neanderthals' pelvis and torso would have affected their gait, too. Early modern humans arriving for the first time on the Neanderthals' territory were probably impressed by the ‘‘so near yet so far’’ appearance of these obviously related but equally obviously different hominids.

A reconstructed Neanderthal skeleton (left), as compared with a modern human, reveals the Neanderthal’s tapering rib cage and wide, flaring pelvis, among many other differences.

A reconstructed Neanderthal skeleton (left), as compared with a modern human, reveals the Neanderthal’s tapering rib cage and wide, flaring pelvis, among many other differences. Although the brains of both individuals were around the same size, they were enclosed in cranial vaults of very different shape, and the faces of both were strikingly different in size and structure. Photo by K. Mowbray, AMNH.

Just how closely related are the Neanderthals to Homo sapiens? Paleoanthropologists have tended not to look too closely at this question, often preferring either to look upon these hominids as a form simply en route to ourselves or as an extreme variant of Homo sapiens as we know it. But if we recognize a separate species Homo neanderthalensis, then we have to ask ourselves where this distinctive hominid type came from. And we can seek the answer in the fact that the Neanderthals did not exist in isolation. The earliest Neanderthal fossils we know are dated to perhaps as much as 200,000 to 250,000 years ago; but they are rare, and the Neanderthal record becomes relatively good only when we approach more recent times. Still, a scattering of hominid fossils is known from Europe from the period between the appearance of Homo heidelbergensis, about 500,000 years ago, and the time at which the first Neanderthals appear.

Interestingly, the fossils from this temporally intermediate era all have a certain number of the features that we associate with Neanderthals—but not all of them. It seems, in fact, that the Neanderthals were part of a larger group of hominid species that diversified in Europe subsequent to the appearance there of Homo heidelbergensis or perhaps even of Homo antecessor. This is a classic example of the ‘‘adaptive radiation’’ that typically happens when a new kind of animal successfully invades a new territory—as western Europe was to hominids. Once again we see that local diversification has been a major element in the evolutionary history of hominids, as in those of so many other kinds of animal.

The prime example here, perhaps, is a 225,000-year-old cranium found in Steinheim, Germany, in 1933. Despite having been somewhat distorted after burial, this specimen resembles the Neanderthals in, among other things, its brow shape, its large nasal opening, the shape of its eye sockets, its (small) suprainiac fossa, and a hint of medial projections in the nose. But it differs, again among other things, in being relatively small-brained and in lacking the rounded braincase walls and facial puffiness seen in Neanderthals. Many have thus perceived ‘‘preNeanderthal'' features in this specimen, though it is rare to argue that it is actually a Neanderthal. The best interpretation seems to be that the Steinheim cranium represents a species that had recently shared an ancestor with the Neanderthals yet also belonged to its own separate species.

Another example of diversification is provided by the amazing series of hominid fossils, around 500,000 years old, that have been found in the extraordinary ‘‘Pit of the Bones’’ at Atapuerca in Spain. Here, at the bottom of a deep shaft opening inside a large cave, the broken remains of at least twenty-eight individuals have been found. These hominids are quite distinctive in their own  right but have brow ridges, large noses, and other features that are Neanderthal-like. At the same time they lack medial projections, puffed-out faces, and the special features of the rear of the skull that characterize Neanderthals. Yet other European fossils, contemporaneous with those from the Pit of the Bones, share none of their Neanderthal-like features. In this larger context the Neanderthals, rather than seeming to be part of an isolated hominid lineage in Europe, start to fit quite comfortably into a routine scenario of evolutionary experimentation following the first successful hominid incursion into Europe. For whatever reasons, it was the Neanderthals who emerged triumphant from this process of diversification. And this suggests that their related rivals might have suffered, at the Neanderthals’ hands, the same kind of fate that awaited the Neanderthals themselves when Homo sapiens finally arrived on the European scene.

The best-preserved hominid skull from the Sima de los Huesos (“Pit of the Bones’’) at Atapuerca, Spain, is some 500,000 years old.The best-preserved hominid skull from the Sima de los Huesos (“Pit of the Bones’’) at Atapuerca, Spain, is some 500,000 years old. Although it looks similar in certain ways to a Neanderthal skull, the differences in the shape of the braincase and in the lower areas of the face show that it belonged to a different species. Photo by K. Mowbray, AMNH.

The notion that the Neanderthals were part of a radiation of hominids in Europe that was separate from the radiation in Africa that gave rise to Homo sapiens fits nicely with exciting new information that is becoming available from the molecular record. In the last few years, molecular biologists have succeeded in extracting fragments of mitochondrial DNA (mtDNA) from a few late Neanderthal bones, with instructive results. Mitochondrial DNA is not the same as the nuclear DNA that resides in the nuclei of the body’s cells and makes up the chromosomes. Rather, while remaining within the cell’s outer membrane, mtDNA resides outside the nucleus in a cellular organelle called the mitochondrion, which is often described as the ‘‘powerhouse’’ of the cell because it is central to the extraction of the energy contained in nutrients.

Mitochondria have their own DNA because the complex cells of which our bodies are built were originally formed a couple of billion years ago by the ‘‘capture’’ of one kind of simple single-celled organism—the ancestor of mitochondria—by another, the progenitor of the rest of the cell. This symbiosis of components from two different lineages of organisms turned out to be highly advantageous, for these complex cells burn fuel up to twenty times more efficiently than other types of cells. Scientists who study the biological histories of modern human populations find mtDNA especially interesting for two reasons. First, it accumulates changes (mutations) much faster than nuclear DNA does, which means that very recent evolutionary events can be detected. Second, it is transmitted exclusively through the mother, because eggs contain mitochondria but sperm do not. This means that the mtDNA gets passed along intact from one generation to the next rather than being reshuffled, as nuclear DNA is, when the genomes of the two parents are combined. As a result, all changes in this type of DNA must be the result of mutations, and mtDNA can thus be used quite simply to trace ancestries through the female lineage within species, as well as to make comparisons among species.

By comparing a short stretch of mtDNA extracted from the original (Feldhofer Cave) Neanderthal specimen with samples obtained from apes and from a variety of modern humans from around the world, investigators arrived at a number of conclusions. First, the DNA sequence that was obtained from the Neanderthal specimen was quite distinct from those of all the modern humans sampled, although it was closer to them than to apes. It showed 27 differences from us, compared with the average of 8 differences that separate members of modern human populations from different areas of the world—and to the 55 that distinguish the average Homo sapiens from a chimpanzee. What’s more, although the Neanderthal sampled had lived in Europe, its mtDNA was no more similar to that of modern Europeans than to that of any other modern population. Clearly, the Neanderthal specimen was strongly distinguished from all lineages of modern humans and showed no closer resemblances to Europeans of the kind that would have been expected if ancestral European humans and Neanderthals had interbred.

None of this proves conclusively that Neanderthals belonged to the separate species Homo neanderthalensis, but it all does point very strongly in that direction. From the mtDNA differences they observed, the investigators calculated a tree of relatedness among the various modern human populations that had been sampled. This analysis indicated an African origin for the modern human gene pool. The scientists also used their data to derive a date for the last common ancestor of Neanderthals and modern humans, which they calculated had lived between 690,000 and 550,000 years ago. This fits pretty well with what we know from the fossil record, since although recognizable Neanderthals only began to show up about 200,000 years ago, the larger grouping to which they belonged had much deeper roots than that. Subsequent to the analysis of the Feldhofer DNA, mtDNA has been extracted from several other Neanderthal specimens with generally similar results (though they do show expected variation among individuals), demonstrating that the initial Feldhofer findings were not just a flash in the pan.

Nobody knows what kinds of groups the Neanderthals lived in, although from the sizes of the sites at which their physical and archaeological remains have been found it seems that social units were typically fairly small, possibly consisting of no more than 15 to 30 individuals of both sexes and all ages. Small bands like this roamed over vast tracts of the sparsely populated landscape, camping in one place for short periods until the local resources were exhausted, then moving on. What those resources were would have varied from time to time with changing climates, and from place to place as groups moved from valley to upland and back again. It has been pointed out that, in contrast to the tropics where plant resources would have been relatively consistent year-round, in Ice Age Europe plant foods that could sustain hominids would have been relatively scarce and more affected by seasonal change.

For this reason, many archaeologists are coming around to the view that meat formed a very important component of the Neanderthals’ diet. This again implies that their hunting techniques may have been quite advanced—something that apparently also may be implicit in those spears from Schoeningen. The picture of Neanderthals as predominantly meat-eaters is also supported by the few studies that have been done of the way Neanderthal teeth wore and of the chemistry of their bones (indeed, one study suggested that the Neanderthal examined had been a specialized hunter of extremely large-bodied mammals such as woolly rhino and mammoth). Additionally, frequent close encounters with unfriendly animals may account for the claim that the pattern of fractured and healed bones in Neanderthal skeletons resembles that among rodeo riders today. On the other hand, the only modern humans who historically depended primarily on animal proteins and fats possessed technologies that were highly specialized for obtaining these foods, something not evident in the Neanderthal toolkit.

What is possibly more important about the Neanderthals than their specializations, however, was their considerable adaptability. They survived numerous climatic changes over a huge span of time in a vast and topographically varied area. They could not have been so successful if their behavior patterns had not been highly flexible; and indeed, the evidence strongly suggests that this was the case. In one Italian locale archaeologists excavated some cave deposits with evidence of Neanderthal occupation that dated from 120,000 years ago, when the climate was relatively warm, and others from 50,000 to 40,000 years ago, when conditions were much colder. At the earlier time, occupations seem to have been quite brief, and animal remains were mostly skulls of old individuals. These observations were interpreted to suggest that the Neanderthals had scavenged what remained of the carcasses of aged animals who had died natural deaths. During the more recent period, remains consisted of many different body parts from animals in their prime, and the interpretation is that the Neanderthals had employed ambushhunting techniques to obtain entire carcasses, during longer stays in the neighborhood. These conclusions are entirely reasonable, but it is impossible to say whether the differences are due to an improvement in hunting techniques over time or whether they simply reflect responses to changing conditions.

The social organization of Neanderthals remains a mystery, although a study at one French site did lead to the suggestion that males and females may have led largely separate lives. But the truth is, we simply do not know. The Neanderthals controlled fire, as their predecessors had for some time, but most evidence for this comes not from deliberately constructed hearths lined with stones but from simple ash deposits. And even where hearths were made, we can be pretty sure that Neanderthals did not sing songs and tell each other stories around them, because it's a good bet that they didn't have language. Language is a symbolic activity, and the Neanderthals left behind no symbolic artifacts (engravings, notations, figurines, and so forth) of the kind that were so typical of their successors, the Cro-Magnons. Cro-Magnon is the name we give to the first Homo sapiens who occupied Europe; they are named after the site in southwestern France, ‘‘Magnon's Shelter,'' at which their remains were first found. Nonetheless, there can be little doubt that Neanderthals possessed some form of quite sophisticated vocal communication, presumably supplemented with an extensive repertoire of gestures. And, significantly, at some time before 50,000 years ago the Neanderthals invented the tradition of burying their dead. Neanderthal burial, however, was both occasional and very simple, without the grave goods and other paraphernalia so characteristic of later Cro-Magnon burials (though these apparently did not begin until well after the first Cro-Magnon incursions into Europe).

Of course, deliberate interment of the dead almost certainly did not mean to the Neanderthals what it means to most modern humans, with its overtones of spirituality and future life; but it does suggest some kind of empathy with the deceased. And at the Iraqi site of Shanidar the remains of an individual who survived to an advanced age (maybe 40 years) despite being severely handicapped by a useless arm, perhaps since birth, suggests that such individuals received the long-term support of their groups. Recent studies at other sites have reached similar conclusions. There are many different ways of being hominid, and almost certainly the Neanderthals’ way was not ours. But it is nonetheless evident that the Neanderthals were complex beings, who perceived and interacted with the world around them in their own characteristic and sophisticated ways.

The record of hominid evolution in the later Ice Age is better in Europe and the eastern shores of the Mediterranean (the Levant, specifically Israel) than it is in the rest of the world. Homo heidelbergensis fossils are known from various sites in southern Africa and eastern Asia following Bodo times some 600,000 years ago, but most of them have not been definitively dated, and none is accompanied by anything like the evidence of lifestyles we have available from France and Germany. In eastern Asia H. erectus, or something like it, seems to have survived on the island of Java until as late as around 40,000 years ago, the time at which we can surmise that H. sapiens arrived there. In mainland China, on the other hand, H. erectus seems to have been replaced by H. heidelbergensis or an equivalent well before H. sapiens showed up. In Africa, apart from the Bodo specimen, H. heidelbergensis is not well dated. However, H. heidelbergensis seems to have been broadly succeeded by a rather heterogeneous assortment of hominids represented by crania found in locations as far-flung as Florisbad in South Africa, Ngaloba in Tanzania, and Guomde in Kenya.

To minimize the number of species names while also acknowledging the many ways in which they differ from our living species, many paleoanthropologists have been in the habit of lumping together specimens such as these under the designation ‘‘archaic Homo sapiens.’ However, this is a category of convenience more than anything else, and it has had the unfortunate effect of disguising a much more complex underlying pattern of descent than the linear one the ‘‘archaic'' designation implies. As a result, it is still unclear what the actual pattern was, which is a pity because it was almost certainly among African hominids in this general time frame that true Homo sapiens eventually emerged.

On the technological front, it was almost certainly also in Africa that prepared-core tool technology was originally invented; and it is in that continent, too, that long, slender blade tools such as those made by Cro-Magnons appear to have first been made, well over a quarter-million years ago. Of course it is important to bear in mind, when thinking about technologies, that the story of technological development and innovation has been no more linear than that of the hominids themselves. New inventions have appeared, faded, and been replaced by apparently more archaic forms, only to reappear at later times. Indeed, our cultural evolution has very likely been even more complex and tortuous than hominid physical evolution—which is something that we should probably expect, given that cultural traditions can be transferred sideways among contemporaries as well as being transmitted down from one generation to the next.

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