"LUCY" SKELETON represents the best-known species of early hominid, or human precursor, Australopithecus afarensis, often characterized as a "bipedal chimpanzee." The 3.18-million-year-old skeleton is from the Hadar region of Ethiopia.

By Ian Tattersall

humanity. But how many human did they emigrate?

bly a fairly accurate characterization, especially given the increasing evidence that early hominids favored quite heavily wooded habitats. Their preferred way of life was evidently a successful one, for although these primates were less adept arborealists than the living apes and less efficient bipeds than later hominids, their basic "eat your cake and have it" adaptation endured for well over two million years, even as species of this general kind came and went in the fossil record.

It is not even clear to what extent lifestyles changed with the invention of stone tools, which inaugurate our archaeological record at about 2.5 million years ago. No human fossils are associated with the first stone tools known, from sites in Kenya and Ethiopia. Instead there is a motley assortment of hominid fossils from the period following about two million years ago, mostly associated with the stone tools and butchered mammal bones found at Tanzania's Olduvai Gorge and in Kenya's East Turkana region. By one reckoning, at least some of the first stone toolmak-ers in these areas were hardly bigger or more advanced in their body skeletons than the tiny Lucy; by another, the first tools may have been made by taller, somewhat larger-brained hominids with more modern body structures. Exactly how many species of early hominids there were, which of them made the tools, and how they walked remain among the major conundrums of human evolution.

Physically, at least, the picture becomes clearer after about 1.9 million years ago, when the first good evidence occurs in northern Kenya of a species that is recognizably like ourselves. Best exemplified by the astonishingly complete 1.6-million-year-old skeleton known as the Turkana Boy, discovered in 1984, these humans possessed an essentially modern body structure, indicative of modern gait, combined with moderately large-faced skulls that contained brains double the size of those of apes (though not much above half the modern human average). The Boy himself had died as an adolescent, but it is estimated that had he lived to maturity he would have attained a height of six feet, and his limbs were long and slender, like those of people who live today in hot, arid African climates, although this common adaptation does not, of course, indicate any special relationship. Here at last we have early hominids who were clearly at home on the open savanna.

A long-standing paleoanthropologi-cal tradition seeks to minimize the num-

TURKANA BOY," an adolescent Homo ergaster dated to about 1.6 million years ago, is representative of the first hominids with an effectively modern body skeleton.

NEWLY DISCOVERED SPECIES: Australopithecus anamensis is the earliest well-documented hominid. This lower jaw from Kanapoi, Kenya, seen as it was found in the field, has been dated to around four million years ago. A. anamensis closely resembles A. afarensis in dental details, and a partial tibia (shinbone) indicates that it walked upright.

ber of species in the human fossil record and to trace a linear, progressive pattern of descent among those few that are recognized. In keeping with this practice, the Boy and his relatives were originally assigned to the species H. erectus. This species was first described from a skullcap and thighbone found in Java a century ago. Fossils later found in China— notably the now lost 500,000-year-old "Peking Man"—and elsewhere in Java were soon added to the species, and eventually H. erectus came to embrace a wide variety of hominid fossils, including a massive braincase from Olduvai Gorge known as OH9. The latter has been redated to about 1.4 million years, although it was originally thought to have been a lot younger. All these fossil forms possessed brains of moderate size (about 900 to 1,200 milliliters in volume, compared with an average of around 1,400 milliliters for modern humans and about 400 milliliters for apes), housed in long, low skull vaults with sharp angles at the back and heavy brow ridges in front. The few limb bones known were robust but essentially like our own.

Whether H. erectus had ever occupied Europe was vigorously debated, the alternative being to view all early human fossils from that region (the earliest of them being no more than about 500,000 years old) as representatives of archaic H. sapiens. Given that the Javan fossils were conventionally dated in the range of one million to 700,000 years and younger and that the earliest Chinese fossils were reckoned to be no more than

one million years old, the conclusion appeared clear: H. erectus (as exemplified by OH9 and also by the earlier Turkana Boy and associated fossils) had evolved in Africa and had exited that continent not much more than one million years ago, rapidly spreading to eastern Asia and spawning all subsequent developments in human evolution, including those in Europe.

Yet on closer examination the specimens from Kenya turned out to be distinctively different in braincase construction from those of classic eastern Asian H. erectus. In particular, certain anatomical features that appear specialized in the eastern Asian H. erectus look ancestral in the African fossils of comparable age. Many researchers began to realize that we are dealing with two kinds of early human here, and the earlier Kenyan form is now increasingly placed in its own species, H. ergaster. This species makes a plausible ancestor for all subsequent humans, whereas the cranial specializations of H. erectus suggest that this species, for so long regarded as the standard-issue hominid of the 1- to 0.5-million-year period, was in fact a local (and, as I shall explain below, ultimately terminal) eastern Asian development.

An Eastern Asian Cul-de-Sac

THE PLOT THICKENED in early 1994, when Carl C. Swisher of the Berkeley Geochronology Center and his colleagues applied the newish argon/argon dating method to volcanic rock samples taken from two hominid sites in Java. The results were 1.81 and 1.66 million years: far older than anyone had really expected, although the earlier date did confirm one made many years before. Unfortunately, the fossils from these two sites are rather undiagnostic as to species: the first is a braincase of an infant (juveniles never show all the adult characteristics on which species are defined), and the second is a horrendously crushed and distorted cranium that has never been satisfactorily reconstructed. Both specimens have been regarded by most as H. erectus, more for reasons of convenience than anything else. Over the decades,

"PEKING MAN" is the name given to this skull of a male H. erectus from Zhoukoudian, near Beijing. The skull was reconstructed from fragments of various individuals, all probably around 500,000 years old.

TWO ACHEULEAN TOOLS, from St. Acheul, France, are probably around 300,000 years old, but implements of this kind began to be made in Africa as many as 1.5 million years ago. On the left is a pointed hand ax and on the right a blunt-ended cleaver.

REPLICA OF OLDOWAN BASALT CORE illustrates how sharp flakes were struck from the core to provide cutting implements. Tools of this kind were first made around 2.5 million years ago.

TWO ACHEULEAN TOOLS, from St. Acheul, France, are probably around 300,000 years old, but implements of this kind began to be made in Africa as many as 1.5 million years ago. On the left is a pointed hand ax and on the right a blunt-ended cleaver.

sporadic debate has continued regarding whether the Javan record contains one or more species of early hominid. Further, major doubt has been cast on whether the samples that yielded the older date were actually obtained from the same spot as the infant specimen. Still, these dates do fit with other evidence pointing to the probability that hominids of some kind were around in eastern Asia much earlier than anyone had thought.

Independent corroboration of this scenario comes, for instance, from the Dmanisi site in the former Soviet republic of Georgia, where in 1991 a hominid lower jaw that its describers allocated to H. erectus was found. Three different methods indicated that this jaw was as old as 1.8 million years, and with four crania from the site now in hand, there is ample evidence of an unexpectedly early hominid exodus from Africa. Even the most parsimonious reading of the admittedly imperfect record suggests that these pioneering emigrants must have been H. ergaster or something very much like it.

A very early hominid departure from Africa has the advantage of explaining an apparent anomaly in the archaeological record. The stone tools found in sediments coeval with the earliest H. ergaster (just under two million years ago) are essentially identical with those made by the first stone toolmakers many hundreds of thousands of years before. These crude tools consisted principally of sharp flakes struck with a stone "hammer" from small cobbles. Effective cutting tools though these may have been (experimental archaeologists have shown that even elephants can be quite efficiently butchered using them), they were not made to a standard form but were apparently produced simply to obtain a sharp cutting edge. Following about 1.5 million years ago, however, standardized stone tools began to be made in Africa, typified by the hand axes and cleavers of the Acheulean industry (first identified in the mid-19th century from St. Acheul in France). These were larger implements, carefully shaped on both sides to a teardrop form. Oddly, stone tool industries in eastern Asia lacked such utensils, which led many to wonder why the first human immigrants to the region had not brought this technology with them, if their ancestors had already wielded it for half a million years. The new dates suggest, however, that the first emigrants had left Africa before the invention of the Ach-eulean technology, in which case there is no reason why we should expect to find this technology in eastern Asia. Interestingly, in 1989 Robin W. Dennell of the University of Sheffield in England caused quite a stir by reporting very crude stone tools from Riwat in Pakistan that are older than 1.6 million years. Their great age is now looking decreasingly anomalous.

Of course, every discovery raises new questions, and in this case the problem is to explain what it was that enabled human populations to expand beyond Africa for the first time. Most scholars had felt that it was technological advances that allowed the penetration of the cooler continental areas to the north. If, however, the first emigrants left Africa equipped with only the crudest of stone-

SKULLCAP known as Olduvai Hominid 9 (OH9) was dated to 1.4 million years old; it was originally believed to have been much younger. Its affinities are still being debated.

SKULLCAP known as Olduvai Hominid 9 (OH9) was dated to 1.4 million years old; it was originally believed to have been much younger. Its affinities are still being debated.

SUCCESSIVE WAVES of early humans exited from Africa to all parts of the Old World. The record of these emigrations is incomplete, but it is evident that this history is much longer and more complex than has traditionally been believed.

working technologies, we have to look to something other than technological prowess for the magic ingredient. And because the first human diaspora apparently followed hard on the heels of the acquisition of more or less modern body form, it seems reasonable to conclude that the typically human wanderlust emerged in concert with the emancipation of hominids from the forest edges that had been their preferred habitat. Of course, the fact that the Turkana Boy and his kin were adapted in their body proportions to hot, dry environments does nothing to explain why H. ergaster was able to spread rapidly into the cooler temperate zones beyond the Mediterranean; evidently the new body form that made possible remarkable endurance in open habitats was in itself enough to make the difference.

The failure of the Acheulean ever to diffuse as far as eastern Asia reinforces the notion, consistent with the cranial specializations of H. erectus, that this part of the world was a kind of paleo-anthropological cul-de-sac. In this region, ancient human populations largely followed their own course, independent of what was going on elsewhere in

the world. Further datings tend to confirm this view. Swisher and his colleagues reported in 1996 dates for the Ngandong H. erectus site in Java that center on only about 40,000 years ago. These dates, though very carefully obtained, have aroused considerable skep ticism; but, if accurate, they have considerable implications for the overall pattern of human evolution. For they are so recent as to suggest that the long-lived H. erectus might even have suffered a fate similar to that experienced by the Nean-dertals in Europe: extinction at the hands

FOSSILS FROM LONGGUPO, such as the lower jaw fragment (side and top views at left), may indicate the presence of hominids in China as many as 1.9 million years ago.

Millions of Years Ago

Less than 0.1 0.1 to 0.5 H 0.5 to 1.0 1.0 to 1.5 H 1.5 to 2.0 More than 2.0

Millions of Years Ago

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of late-arriving H. sapiens. Here we find reinforcement of the gradually emerging picture of human evolution as one of repeated experimentation, with regionally differentiated species, in this case on opposite sides of the Eurasian continent, being ultimately replaced by other hom-

inid lineages that had evolved elsewhere.

At the other end of the scale, in 1996 an international group led by Huang Wanpo of Academia Sinica in Beijing reported a remarkably ancient date for Longgupo Cave in China's Sichuan Province. This site had previously yielded an incisor tooth and a tiny lower jaw fragment with two teeth that were initially attributed to H. erectus, plus a few very crude stone artifacts. Huang and his colleagues concluded that the fossils and tools might be as many as 1.9 million years old, and their reanalysis of the fossils suggested to them a closer resemblance to the earliest African Homo species than to H. erectus.

This latter claim has not gone unex-amined. As my colleague Jeffrey H. Schwartz of the University of Pittsburgh and I pointed out, for instance, the teeth in the jaw fragment resemble African Homo in primitive features rather than in the specialized ones that indicate a special relationship. What is more, they bear a striking resemblance to the teeth of an orangutan-related hominoid known from a much later site in Vietnam. And although the incisor appears hominid, it is fairly generic, and there is nothing about it that aligns it with any particular human species. Future fossil finds from Long-gupo will, with luck, clarify the situation; meanwhile the incisor and stone tools are clear evidence of the presence of humans in China at what may be a very early date indeed. These ancient eastern Asians were the descendants of the first emigrants from Africa, and, whatever the hominids of Longgupo eventually turn out to have been, it is a good bet that Huang and his colleagues are right in guessing that they represent a precursor form to H. erectus rather than that species itself.

All this makes sense, but one anomaly remains. If H. erectus was an indigenous eastern Asian development, then we have to consider whether we have correctly identified the Olduvai OH9 braincase as belonging to this species. If we have, then H. erectus evolved in eastern Asia at quite an early date (remember, OH9 is now thought to be almost 1.4 million years old), and one branch of the species migrated back to Olduvai in Africa. But if these new Asian dates are accurate, it seems more probable that as we come to know more about OH9 and its kind we will find that they belonged to a different species of hominid altogether.

The opposite end of the Eurasian continent was, as I have hinted, also isolated from the human evolutionary mainstream. As we saw, humans seem to have arrived in Europe fairly late. In this region, the first convincing archaeological sites, with rather crude tools, show up at about 800,000 years ago or thereabouts (although in the Levant, within hailing distance of Africa, the site of 'Ubeidiya has yielded Acheulean tools dated to

IAN TATTERSALL was born in England and raised in East Africa. He is chair of the department of anthropology at the American Museum of Natural History in New York City. His latest books include The Monkey in the Mirror: Essays on the Science of What Makes Us Human (Harvard Books, 2003), Becoming Human: Evolution and Human Uniqueness (Har-court, 1998) and The Fossil Trail: How We Know What We Think We Know about Human Evolution (Oxford University Press, 1995).

around 1.4 million years ago, just about as early as any found to the south). The problem has been the lack of a sign of the toolmakers themselves.

This gap began to be filled by finds made by Eudald Carbonell of the University of Tarragona in Spain and his coworkers at the Gran Dolina Cave site in the Atapuerca Hills of northern Spain.

In 1994 excavations at that site produced numerous simple stone tools, plus quite a few human fossil fragments, the most complete of which is a partial upper face of an immature individual. All came from a level that was dated to more than 780,000 years ago. No traces of Acheulean technology were found among the tools, and the investigators noted various primitive traits in the fossils, which they provisionally attributed to H. heidelbergensis. This is the species into which specimens formerly classified as archaic H. sapiens are increasingly being placed. Carbonell and his colleagues see their fossils as the starting point of an indigenous European lineage that gradually evolved into the Neandertals. These latter, large-brained hominids are known only from Europe and western Asia, where they flourished in the period between about 200,000 years and 30,000 years ago, when they were extinguished in some way by invading H. sapiens.

This is not the only possibility, however. With only a preliminary description of the very fragmentary Gran Dolina fossils available, it is hard to be sure, but it seems at least equally possible that they are the remains of hominids who made an initial foray out of Africa into Europe but failed to establish themselves there over the long term. Representatives of H. heidelbergensis are known in Africa as well, as long ago as 600,000 years ago, and this species quite likely re-colonized Europe later on. There it would have given rise to the Neander-tals, whereas a less specialized African population founded the lineage that ultimately produced H. sapiens.

At another site, just a kilometer from Gran Dolina, Juan-Luis Arsuaga of Complutense University in Madrid and his colleagues have discovered a huge cache of exquisitely preserved human fossils, about 400,000 years old. These are said to anticipate the Neandertals in certain respects, but they are not fully Neandertal by any means. And although they emphasize that the Neandertals (and possibly other related species) were an indigenous European development, these fossils from Sima de los Huesos ("Pit of the Bones") do not establish an unequivocal backward connection to their Gran Dolina neighbors.

Born in Africa

EVERY LONGTIME READER of Scientific American will be familiar with the competing models of "regional continuity" and "single African origin" for the

GRAN DOLINA CAVE in the Atapuerca Hills of northern Spain has produced the earliest human fossils yet found in Europe. These fossils, dated to about 780,000 years ago and initially attributed to H. heidelbergensis, may in fact represent a distinct form. The mature cranium (below) is from Sima de los Huesos, about one kilometer from Gran Dolina, where a huge trove of mostly fragmentary but exquisitely preserved human fossils is dated to about 300,000 years ago.

GRAN DOLINA CAVE in the Atapuerca Hills of northern Spain has produced the earliest human fossils yet found in Europe. These fossils, dated to about 780,000 years ago and initially attributed to H. heidelbergensis, may in fact represent a distinct form. The mature cranium (below) is from Sima de los Huesos, about one kilometer from Gran Dolina, where a huge trove of mostly fragmentary but exquisitely preserved human fossils is dated to about 300,000 years ago.

emergence of our own species, H. sapiens [see "The Multiregional Evolution of Humans," on page 46; and "The Recent African Genesis of Humans," on page 54]. The first of these models holds that the highly archaic H. erectus (including H. ergaster) is nothing more than an ancient variant of H. sapiens and that for the past two million years the history of our lineage has been one of a braided stream of evolving populations of this species in all areas of the Old World, each adapting to local conditions, yet all consistently linked by gene exchange. The variation we see today among the major geographical populations of humans is, by this reckoning, simply the latest permutation of this lengthy process.

The other notion, which happens to coincide much better with what we know of evolutionary processes in general, proposes that all modern human populations are descended from a single ancestral population that emerged in one place at some time between about 150,000 and 100,000 years ago. The fossil evidence, thin as it is, suggests that this place of origin was somewhere in Africa (although the neighboring Levant is an alternative possibility); proponents of this scenario point to the support afforded by comparative molecular studies for the notion that all living humans are descended from an African population.

In view of what I have already said about the peripheral roles played in human evolution by early populations both in eastern Asia and Europe, it should come as no surprise that between these two possibilities my strong preference is for a single and comparatively recent origin for H. sapiens, very likely in Af-rica—the continent that, from the beginning, has been the engine of mainstream innovation in human evolution. The rise of modern humans is a recent drama that played out against a long and complex backdrop of evolutionary diversification among hominids, but the

LEADING THEORIES of the origins of modern humans are contrasted in these diagrams. According to the notion of "regional continuity," all modern human populations trace their beginnings to H. erectus, but each regional population evolved along its own distinctive lines, exchanging enough genes with its neighbors [arrows represent gene exchange) to remain part of the same species; all eventually became H. sapiens. The "single origin" theory holds that H. sapiens descended from a single ancestral population that emerged in one place, probably Africa.

H. neanderthalensis

H. neanderthalensis

H. heidelbergensis

H. ergaster

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