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necessary because the discovery of an individual with such a mosaic of features has profound implications. First, it rejects the extreme Out of Africa model of modern human emergence, which proposes that early moderns originating in Africa subsequently displaced all archaic humans in other regions. Instead the Lagar Velho child's anatomy supports a scenario that combines a dispersal of anatomically modern humans out of Africa with mixing between that population and the archaic populations it encountered. (For example, the African ancestry of early modern Europeans is reflected in their relatively long lower-leg bones, a tropical adaptation. Lagar Velho 1, however, has the short shins of the cold-adapted Neandertals.)

Lagar Velho 1 also provides insights into the behavioral similarities of Neandertals and early modern humans. Despite the paleontological evidence indicating anatomical differences between these two groups, their overall adaptive patterns, social behaviors and means of communication (including language) cannot have contrasted greatly. To their contemporaries, the Neandertals were just another group of Pleistocene hunter-gatherers, fully as human as themselves.

ERIK TRINKAUS is a paleoanthropologist at Washington University.

CIDALIA DUARTE is a researcher at the Portuguese Institute of Archaeology in Lisbon.

skilled hunters capable of killing even large animals such as rhinoceroses, according to University of Cambridge archaeologist Preston T. Miracle. And Shea's studies suggest that some Ne-andertals employed sophisticated stone-tipped spears to conquer their quarry—a finding supported in 1999, when researchers reported the discovery in Syria of a Neandertal-made stone point lodged in a neckbone of a prehistoric wild ass. Moreover, additional research conducted by Shea and investigations carried out by University of Arizona archaeologists

Mary C. Stiner and Steven L. Kuhn have shown that Neandertal subsistence strategies varied widely with the environment and the changing seasons.

Such demonstrations refute the notion that Ne-andertals perished because they could not adapt. But it may be that moderns were better at it. One popular theory posits that modern humans held some cognitive advantage over Neandertals, perhaps a capacity for the most human trait of all: symbolic thought, including language. Explanations such as this one arose from observations that after 40,000 years ago, whereas Neandertal culture remained relatively static, that of modern Europeans boasted a bevy of new features, many of them symbolic. It appeared that only moderns performed elaborate burials, expressed themselves through body ornaments, figurines and cave paintings, and crafted complex bone and antler tools—an industry broadly referred to as Upper Paleolithic. Neandertal assemblages, in contrast, contained only Middle Paleolithic stone tools made in the Mousterian style.

A CASE FOR NEANDERTAL CULTURE by joäo zilhäo and Francesco d'errico

EVER SINCE THE DISCOVERY nearly 150 years ago of the specimen that defined the Neandertals, researchers have tended to deny Neandertals the behavioral capabilities of modern humans, such as the use of symbols or of complex techniques for tool manufacture. Instead Neandertals were characterized as subhuman, stuck in primitive technical traditions impervious to innovation. And when sophisticated cultural remains were linked to late Neandertals at several sites in western Europe, the evidence was explained away. The most spectacular of these sites, a cave in north-central France named Grotte du Renne (one in a string of sites collectively known as the Arcy-sur-Cure caves), yielded a wealth of complex bone and stone tools, body ornaments and decorated objects, found in association with Neandertal remains. Other sites in France and along the Cantabrian and Pyrenean mountain ranges bore similar artifacts made in this tradition, called the Chatelperronian.

Because early modern Europeans had a comparable industry known as Aurignacian—which often appears at the same sites that contain Chatelperronian materials—some researchers have suggested that the archaeological layers were disrupted, mixing Aurignacian artifacts into the Neandertal-associated levels. Other scholars have interpreted this to mean that Neandertals picked up these ideas from moderns, either collecting or trading for items manufactured by moderns or imitating the newcomers' practices without really grasping the underlying symbolic nature of some of the objects.

Our reassessment of the evidence from the Grotte du Renne shows that the Neandertal-associated ornaments and tools found there did not result from a mixing of the strata, as demonstrated by the presence of finished objects and the by-products of their manufacture in the same stratigraphic level. Moreover, the Châtelperronian artifacts recovered at the Grotte du Renne and other sites, such as Quinçay, in the Poitou-Charentes region of France, were created using techniques different from those favored by Aurignacians. With regard, for example, to the pendants—modified bear, wolf and deer teeth, among others—Neandertals carved a furrow around the tooth root so that a string of some sort could be tied around it for suspension, whereas Aurignacians pierced their pendants. As archaeologist François Lévêque and a colleague have described, even when, as they did on occasion, Neandertals put a hole through a tooth, they took an unusual approach, puncturing the tooth. Moderns preferred to scrape the tooth thin and then pierce it.

Similarly, the new knapping techniques and tool types that appear among late Neandertals at other sites in France, Italy and Spain fail to show any influence from the Aurignacian. Instead they maintain affinities with the preceding local traditions, of which they seem to represent an autonomous development.

If the Neandertals' Chatelperronian culture was an outcome of contact with moderns, then the Aurignacian should predate the Chatelperronian. Yet our reanalysis of the radiometric dates for the archaeological sequences reveals that apart from a few debatable instances of mixture, wherever both cultures are

Yet hints that Neandertals thought symbolically had popped up. Neandertal burials, for example, are well known across Europe, and several, it has been argued, contain grave goods. (Other researchers maintain that for Neandertals, interment merely constituted a way of concealing the decomposing body, which might have attracted unwelcome predators. They view the purported grave goods as miscellaneous objects that happened to be swept into the grave.) Evidence for art, in the form of isolated pierced teeth and engraved bone fragments, and red and yellow ocher, has been reported from a few sites, too, but given their relative rarity, researchers tend to assign alternative explanations to these items.

The possibility that Neandertals might have engaged in modern practices was taken more seriously in 1980, when researchers reported a Neandertal from the Saint-Césaire rock-shelter in Charente-Maritime, France, found along with stone tools manufactured according to a cultural tradition known as the Châ-telperronian, which was assumed to have been the handiwork

of moderns. Then, in 1996, Hublin and his co-workers made a startling announcement. Excavations that began in the 1940s at the Grotte du Renne at Arcy-sur-Cure near Auxerre, France, had yielded numerous blades, body ornaments and bone tools and revealed evidence of huts and hearths—all hallmarks of the Upper Paleolithic. The scant human remains found amid the artifacts were impossible to identify initially, but using computed tomography to examine the hidden inner-ear region preserved inside an otherwise uninformative skull fragment, Hub-lin's team identified the specimen as Neandertal.

In response, a number of scientists suggested that Neandertals had acquired the modern-looking items by stealing them, collecting artifacts discarded by moderns or perhaps trading for them. But this view has come under fire, most recently from archaeologists Francesco d'Errico of the University of Bordeaux and Joao Zilhao of the University of Lisbon, who argue that the represented at the same site, the Chatelperronian always underlies the Aurignacian, suggesting its priority. Furthermore, consideration of the hundreds of datings available from this period in Europe and the Near East shows that wherever the context of the dated samples is well known, the earliest occurrences of the Aurignacian are apparently from no earlier than around 36,500 years ago. The same radiometric data, however, indicate that by then

Neandertals were already moving toward modernity on their own. In other words, the Chatelperronian and other late Neandertal cultures, such as the Uluzzian of Italy, emerged in Europe around 40,000 years ago, long before any moderns established themselves in those areas.

That this autonomous development included the manufacture and use of symbolic objects created for visual display on the body, as are often observed in traditional societies, reflects various

PENDANTS, BONE TOOLS AND KNIVES from the Grotte du Renne site seem to be the handiwork of Neandertals. That the advanced items underlie early modern human cultural remains from the same site and are manufactured according to methods different from those favored by the moderns suggests that some Neandertals independently developed a modern culture.

PENDANTS, BONE TOOLS AND KNIVES from the Grotte du Renne site seem to be the handiwork of Neandertals. That the advanced items underlie early modern human cultural remains from the same site and are manufactured according to methods different from those favored by the moderns suggests that some Neandertals independently developed a modern culture.

social roles within Neandertal cultures. Thus, "modern" behavior seems to have emerged in different regions and among different groups of humans, as would happen later in history with the invention of agriculture, writing and state society.

An alternative explanation, taking into account the broadly simultaneous appearance of personal ornaments in many parts of the Old World, is that contacts between modern and archaic humans challenged each group's personal, social and biological identities, igniting an explosion of production of symbolic objects by all those involved. On the strength of the available data, however, we favor the hypothesis of independent invention.

Regardless of which is eventually proved correct, the behavioral barrier that seemed to separate moderns from Neandertals and gave us the impression of being a unique and particularly gifted human type—the ability to produce symbolic cultures—has definitively collapsed.

JOÁO ZILHÁO is professor of prehistoric archaeology at the University of Lisbon in Portugal.

FRANCESCO D'ERRICO is a CNRS researcher at the Institute of Prehistory and Quaternary Geology, University of Bordeaux, in France.

Chatelperronian artifacts at the Grotte du Renne and elsewhere, though superficially similar to those from the Aurignacian, reflect an older, different method of manufacture [see "A Case for Neandertal Culture," above].

Most researchers are now convinced that Neandertals manufactured the Chatelperronian tools and ornaments, but what prompted this change after hundreds of thousands of years is unclear. Cast in this light, "it's more economical to see that as a result of imitation or acculturation from modern humans than to assume that Neandertals invented it for themselves," reasons Cambridge archaeologist Paul A. Mellars. "It would be an extraordinary coincidence if they invented all these things shortly before the modern humans doing the same things arrived." Furthermore, Mellars disagrees with d'Errico and Zilhao's proposed order of events. "The dating evidence proves to me that

[Neandertals] only started to do these things after the modern humans had arrived in western Europe or at least in northern Spain," he asserts. Unfortunately, because scientists have been unable to date these sites with sufficient precision, researchers can interpret the data differently.

From his own work on the Grotte du Renne body ornaments, New York University archaeologist Randall White argues that these artifacts reflect manufacturing methods known—albeit at lower frequencies—from Aurignacian ornaments. Given the complicated stratigraphy of the Grotte du Renne site, the modern-looking items might have come from overlying Aurignacian levels. But more important, according to White, the Chatelperronian does not exist outside of France, Belgium, Italy and northern Spain. Once you look at the Upper Paleolithic from a pan-European perspective, he says, "the Chatelperronian becomes post-Aurignacian by a long shot."

Still, post-Aurignacian does not necessarily mean after contact with moderns. The earliest Aurignacian sites do not in-

THE FATE OF THE NEANDERTALS by fred h. smith

STRONG EVIDENCE has accumulated in recent years that the emergence of modern humans in Europe resulted largely from the immigration of peoples into the continent, probably from the Near East, starting sometime between 40,000 and 30,000 years ago. Most researchers envisioned these early modern populations as having moved into Anatolia and the Balkans, then up through the plains and valleys of central Europe, and finally into northern and western Europe. Meanwhile the indigenous Neandertals, it was thought, were systematically pushed into more peripheral and undesirable parts of the landscape by these expanding populations of moderns. The Neandertals' last bastion appeared to have been the Iberian Peninsula, where fossils from a Spanish site called Zafarraya have been dated to 32,000 years ago and tools attributed to Neandertals have been dated to around 28,000 years ago. A number of scholars argued that after this time no traces of Neandertals remained in Europe and that the Neandertals did not make any biological contributions to early modern humans.

It seemed that the Neandertals were sent into complete extinction by a superior human species—us.

Evidence from an important site in northwestern Croatia calls aspects of this conventional wisdom into question. By performing accelerator mass spectrometry dating directly on two Neandertal specimens from Vindija cave, my colleagues and I have demonstrated that Neandertals were living in some of the most desirable real estate in central Europe as late as 28,000 years ago. These dates, the most recent known for Neandertal fossils, show that these humans were not quickly relegated to the periphery; they competed quite well with intruding modern populations for a long time.

This overlap of Neandertal and early modern peoples for several millennia in the heart of Europe allowed considerable clude any human remains. Researchers have assumed that they belonged to moderns because moderns are known from younger Aurignacian sites. But "who the Aurigna-cians were biologically between 40,000 and 35,000 years ago remains very much an unanswered question," White notes.

He adds that if you look at the Near East around 90,000 years ago, anatomically modern humans and Neandertals were both making Mousterian stone tools, which, though arguably less elaborate than Auri-gnacian tools, actually require a considerable amount of know-how. "I cannot imagine that Neandertals were producing these kinds of technologically complex tools and passing that on from generation to generation without talking about it," White declares. "I've seen a lot of people do this stuff, and I can't stand over somebody's shoulder and learn how to do it without a lot of verbal hints." Thus, White and others do not buy the argument that moderns were somehow cognitively superior, especially if Neandertals' inferiority meant that they lacked language. Instead it seems that moderns invented a culture that relied more heavily on material symbols.

Researchers have also looked to brain morphology for clues to cognitive ability. According to Ralph L. Holloway of Co-

lumbia University, all the brain asymmetries that characterize modern humans are found in Neandertals. "To be able to discriminate between the two," he remarks, "is, at the moment, impossible." As to whether Neandertal anatomy permitted speech, studies of the base of the skull conducted by Jeffrey T. Laitman of the Mount Sinai School of Medicine suggest that if they talked, Neandertals had a limited vocal repertoire. The significance of such physical constraints, however, is unclear.

Fading Away

IF NEANDERTALS POSSESSED basically the same cognitive ability as moderns, it makes their disappearance additionally puzzling. But the recent redating of Neandertal remains from Vindija cave in Croatia emphasizes that this did not happen overnight. Loyola's Smith and his colleagues have demonstrated opportunity for various interactions, and Vindija may reflect some of them. Work by my Croatian colleagues Ivor KaravaniC of the University of Zagreb and Jakov RadovCiC of the Croatian Natural History Museum has revealed a combination of Mousterian and Aurignacian tools in the same stratigraphic level as the dated Neandertal fossils, indicating that Neandertals either made advanced implements or traded with moderns for

them. Morphologically, the Vindija Neandertals look more modern than do most other Neandertals, which suggests that their ancestors interbred with early moderns.

The likelihood of gene flow between the groups is also supported by evidence that Neandertals left their mark on early modern Europeans. Fossils representing early modern adults from central European sites such as Vogelherd in southwestern Germany and Mladec in Moravia (Czech Republic) have features that are difficult to explain unless they have some Neandertal contribution to their ancestry.

For example, Neandertals and early modern Europeans virtually all exhibit a projection of the back of the skull called an occipital bun (aspects of the shape and position of the buns differ between them because the overall skull shapes are not the

MOVEMENT OF MODERNS (purple) into Europe did not displace the Neandertals, who were still living in central and western Europe 28,000 years ago. A number of the early modern European specimens bear some Neandertal features, which suggests that during the long period of overlap the two populations mixed.

same). Yet fossils from the Near Eastern sites of Skhul and Qafzeh, which presumably represent the ancestors of early modern Europeans, do not have this morphology. It is hard to explain how the growth phenomenon responsible for this bunning could reappear independently and ubiquitously in early modern Europeans. Instead it is far more logical to recognize this morphology as a link to the Neandertals. The Portuguese child discovered late in 1998 in the Lapedo Valley offers more intriguing clues [see "The Hybrid Child from Portugal," on page 32].

I believe the evidence shows that the behavioral and biological interactions between Neandertal and early modern human populations were very complex—too complex for the origins of modern humans in Europe to have involved a simple, complete biological replacement of the Neandertals. Neandertals as organisms no longer exist, and Neandertal genes may not have persisted to the present day, but those genes were there in the beginnings of modern European biological history.

FRED H. SMITH is a paleoanthropologist at Loyola University of Chicago.

that Neandertals still lived in central Europe 28,000 years ago, thousands of years after moderns had moved in [see "The Fate of the Neandertals," above]. Taking this into consideration, Stringer imagines that moderns, whom he views as a new species, replaced Neandertals in a long, slow process. "Gradually the Neandertals lost out because moderns were a bit more innovative, a bit better able to cope with rapid environmental change quickly, and they probably had bigger social networks," he supposes.

On the other hand, if Neandertals were an equally capable variant of our own species, as Smith and Wolpoff believe, long-term overlap of Neandertals and the new population moving into Europe would have left plenty of time for mingling, hence the mixed morphology that these scholars see in late Neandertals and early moderns in Europe. And if these groups were exchanging genes, they were probably exchanging cultural ideas, which might account for some of the similarity between, say, the Chatelperronian and the Aurignacian. Neandertals as entities disappeared, Wolpoff says, because they were outnumbered by the newcomers. Thousands of years of interbreeding between the small Neandertal population and the larger modern human population, he surmises, diluted the distinctive Neandertal features, which ultimately faded away.

"If we look at Australians a thousand years from now, we will see that the European features have predominated [over those of native Australians] by virtue of many more Europeans," Wolpoff asserts. "Not by virtue of better adaptation, not by virtue of different culture, not by virtue of anything except many more Europeans. And I really think that's what describes what we see in Europe—we see the predominance of more people."

From the morass of opinions in this contentious field, one consensus emerges: researchers have retired the vision of the shuffling, cultureless Neandertal. But whether these ancient hominids were among the ancestors of living people or a closely related species that competed with our own for the Eurasian territory and lost remains to be seen. In either case, the details will be extraordinarily complicated. "The more we learn, the more questions arise, the knottier it gets," muses archaeologist Lawrence G. Straus of the University of New Mexico. "That's why simple explanations just don't cut it." S3

Kate Wong is editorial director of ScientificAmerican.com out of

Africa is the birthplace of species evolved there? And when

It all used to seem so simple. The human lineage evolved in Africa. Only at a relatively late date did early humans finally migrate from the continent of their birth, in the guise of the long-known species Homo erectus, whose first representatives had arrived in eastern Asia by around one million years ago. All later kinds of humans were the descendants of this species, and almost everyone agreed that all should be classified in our own species, H. sapiens. To acknowledge that some of these descendants were strikingly different from ourselves, they were referred to as "archaic H. sapiens," but members of our own species they were nonetheless considered to be.

Such beguiling simplicity was, alas, too good to last, and over the past few years it has become evident that the later stages of human evolution have been a great deal more eventful than conventional wisdom for so long had it. This is true for the earlier stages, too, although there is still no reason to believe that humankind's birthplace was elsewhere than in Africa. Indeed, for well over the first half of the documented existence of the hominid family (which includes all upright-walking primates), there is no record at all outside that continent. But recent evidence does seem to indicate that it was not necessarily H. erectus who migrated from Africa—and that these peregrinations began earlier than we had thought.

A Confused Early History recent discoveries in Kenya of fossils attributed to the new species Australopithecus anamensis have pushed back the undoubted record of upright-walking hominids to about 4.2 to 3.9 million years ago. The most recent finds in Kenya and Chad may push this back to six million years ago or more. The A. anamensis fossils bear a strong resemblance to the later and far better known species Australopithecus afarensis, found at sites in Ethiopia and Tanzania in the 3.9- to 3.0-million-year range and most famously represented by the "Lucy" skeleton from Hadar, Ethiopia.

Lucy and her kind were upright walkers, as the structures of their pelvises and knee joints particularly attest, but they retained many ancestral features, notably in their limb proportions and in their hands and feet, that would have made them fairly adept tree climbers. Together with ape-size brains and large, protruding faces, these characteristics have led many to call such creatures "bipedal chimpanzees." This is proba-

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