Athena Review, Vol. 3, No. 2  (2002)

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Background for the Peopling of the New World:

Old World Roots for New World Branches

    C. Loring Brace     University of Michigan

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The first Europeans to write about their impressions of the newly-encountered inhabitants of the Western Hemisphere were struck by the lack of marked visible differences between them. The phrase written by Antonio de Ulloa in 1772 declared that, “Visto un Indio de qualquier region, se puede decir que se hen visto todos in quanta al color y contextura” (quoted in Stewart and Newman 1951: 19). This got shortened in colloquial American English to read, “An Indian’s an Indian, if you’ve seen one, you’ve seen them all.” As more than one observer has noted, this may be something of an overstatement but one that has a germ of truth to it (Stewart 1960:259). Certainly there are differences between the Native Americans from Alaska and Ecuador, but that degree of difference is dwarfed by the distinctions of comparably distant peoples in the Old World. The contrast in appearance between the inhabitants of Oslo at sixty degrees north latitude and the natives of Nairobi on the equator is orders of magnitude greater than the contrast between the sixty degree north latitude Inuits of Nome Alaska and the equatorial residents of Quito in Ecuador. If we had no more evidence than this, we would have to conclude that the inhabitants of the Western Hemisphere had been in place for only the smallest fraction of time that the inhabitants of the Old World had been living at roughly the same latitudes where they are found today. In addition, we could guess that the New World denizens had all descended from people in a relatively restricted portion of the Old World. Sheer geographical propinquity makes Northeast Asia the logical can Fig.1: Ainu man and women at Piratori, on the Island of Yezo (photo: Bishop 1925, Smithsonian Ann. Rep.).didate as was realized by people such as Fray Jose de Acosta in the late sixteenth century and Thomas Jefferson in the 18th (Acosta 1589 [1880]; Jefferson 1787).

 When eastern Asia is mentioned, the people who immediately come to mind are the Chinese and those of similar appearance in Mongolia, Korea, and Japan as well as farther down in Southeast Asia. There is an immediate credibility problem in suggesting Chinese form as a model for the source of the appearance of Native Americans. Putting beads, a buffalo robe and a feather headdress on Mao Zedong would never make him look like a Plains Indian. Although the Chinese and their morphological relatives are the dominant presence in eastern Asia today, they are not the only people who live there, and there is reason to believe that they were not present at all at the northeast edge of the continent as the Pleistocene came to an end. The original human occupants of that territory are represented today by only a remnant, the Ainu of Hokkaido, the northeasternmost of the Japanese islands (fig.1). The long and high-bridged nose, the lack of epicanthic eye folds, the level upper margin of the eye sockets and the flat cheekbones of the Ainu are more compatible with stereotypic images of Plains Indians even if the effect is spoiled by the body hair and full beard that is found on many adult males. Still, the presence of an indigenous and very un-Chinese-looking people at the northeastern edge of the Asian continent warrants further contemplation when the question of the origins of the New World populations is being considered.  

Fig.1: Ainu man and women at Piratori, on the Island of Yezo (photo: Bishop 1925, Smithsonian Ann. Rep.).

Of course, assessing whether people “look” Chinese or “look” Native American is completely subjective no matter how much there may be to an underlying reality being so judged. There is also the question as to the affinities of the Ainu themselves. Their European appearance has been remarked upon by anthropologists over the last two centuries (Desmoulins 1826:289-290; Broca 1860:481; von Baelz 1901:245; Hooton 1946:586 among others). Genetic comparisons, however, have not confirmed the verdict of the eye (Omoto 1970, 1972; Omoto and Harada 1975). Still, the very un-Asian appearance of the Ainu suggests that some kind of quantified test of morphological similarities and differences should be worth pursuing. Previous attempts to do this have yielded various results (Yamaguchi 1982; Dodo 1986; Howells 1986; Hanihara 1992, 1993, 1994; Ishida and Dodo 1993). One of the reasons for the different pictures produced is the somewhat restricted distribution of the samples used for comparison. When a more comprehensive set of samples representing all of the major human population blocks in the world - as well as what can be gleaned from Late Pleistocene samples - were all treated at once, a broader perspective was gained (Brace et al. 2001).  

Craniofacial Measurements Through Time: Before undertaking that review, however, there is another matter that should be dealt with. That is the question of the stability through time of a given morphological configuration. Ever since the demonstration by Franz Boas, early in the last century, showing how much the infamous “cephalic index” could change between one generation and the next following the relocation of the population being tested (Boas 1903), there has been skepticism concerning the utility of cranial metrics in documenting continuity through time (Lewis in press). What Boas actually demonstrated, however, was that the use of a proportion between two adaptively trivial dimensions (i.e. areas of the skull whose shapes are not controlled by selection) does not give a reliable indication of inherited relationships. Both professional scholars and other interested people have uncritically accepted the idea that substantial changes in configuration have taken place as a matter of course over time periods amounting to only a few thousand years.

The idea has been promoted that not only the bodily proportions but also the proportions of parts of the face and skull have undergone major changes because of very recent alterations in way of life. For example, it has been proposed that arm, body, and leg proportions and also the configuration of the craniofacial base and the elongation of the upper respiratory tract including the nasal skeleton in Polynesians have been the result of the impact of the selective forces that have affected them during the exploration and settling of Remote Oceania (Houghton 1996). The expansion of the upper respiratory tract was said to have impinged on the developmental processes which controlled the lower parts of the facial skeleton. That explanation was offered to account for the fact that the nasal and associated parts of the face in Polynesians are longer while the teeth and tooth-bearing parts are relatively reduced when they are compared to those of the peoples of mainland Asia, the presumed original source of the inhabitants of the Pacific. Since the Austronesian spread only goes back little more than 4,000 years and Remote Oceania has been settled for no more than 2,000 years (Pawley and Green 1975), this presumably demonstrated that the differentiation of a Polynesian from an essentially Chinese craniofacial configuration was accomplished in a matter of just a few thousand years.

 The case of the affinities of the contested prehistoric Kennewick skeleton from the State of Washington has raised similar matters. That particular individual (fig.2) is more than 9,000 years old according to the first reported radiocarbon dates (reported in Science May 22, 1998, p.3), and the initial impression of a number of observers (and I am one of them) was that its craniofacial form does not resemble that of the living Native Americans who wish to claim it as a direct ancestor, and who have declared their intent to rebury it without any effort to test it metrically to see whom it most resembles. To those of us who have said that it does not look like the current inhabitants of the area, the off-handed answer by representatives of the claimants is that craniofacial form has simply changed over the last 9,000 years. In their view this is just what should be expected and does not change what they assert to be the fact that it is their lineal ancestor (Preston 1997:74).

Fig.2: The skull of Kennewick Man (after drawing by Chatters; Smithsonian Arctic Studies website).

The question then needs to be addressed: how long do recognizable craniofacial configurations last through time, and how fast can such configurations change? The material available for testing such matters is relatively limited, often amounting to single specimens from different time periods and widely dispersed localities, and assertions of relationship and distinction are regularly made without any quantitative effort to test their probabilities. A few such tests have actually been undertaken. The famous Cro-Magnon skull (fig.3) of approximately 28,000 years ago (White 1989:93) has been compared using multiple discriminant functions (see glossary) to representatives of the major groups of the living populations of the world. The figures initially produced were posterior probabilities which can tell what populations the specimen being tested can be excluded from, but that are less certain as indicators of the populations to which the specimen actually will belong. 

What was demonstrated was that a configuration like that of Cro-Magnon could not be found in any of the populations of the world except those at the northwest edge of Europe (Brace 1991:189). Evidently a Cro-Magnon sort of craniofacial configuration has been identifiably European for nearly 30,000 years at the northwest edge of human habitation. When the same statistical procedure is used to test the affinities of the Cro-Magnon contemporary, Predmostí 3 from the Czech Republic, essentially the same conclusions were confirmed (Brace 1991:189). That is, the strongest affinities were with the living people of northwest Europe although there were weaker ties with the prehistoric inhabitants of Japan and with the living Polynesians and Native Americans along the US-Canada border. As will be shown later, the latter ties are notirrelevant hints.

Fig.3: Skull from Cro-Magnon, France (photo: Athena Review, from cast at the American Museum of Natural History).

Interestingly enough, when the Italian Neanderthal from Monte Circeo of between 51,000 and 57,000 years ago (Schwarcz et al. 1991:316) is tested in the same fashion, it also shows that it is convincingly distinguished from all the living populations of the world except those at the northwest edge of Europe (Brace 1991:189). When European Neanderthals are treated as a group and compared with the living populations of the world using a neighbor-joining clustering procedure (see glossary), they always show a link to Europe before they tie in with any other population (Brace et al. 2001:10018). Evidently, despite the century-long insistence that Neanderthals are radically distinct from “anatomically modern” Homo sapiens, the European Neanderthals not only share something with living humans but the form shared is with the living humans who come from the same part of the world that the Neanderthals inhabited over 50,000 years ago.

For some time, now, the claim has been promoted that the more than 90,000-year-old fossils from Jebel Qafza in Israel can be called “Proto-Cro-Magnon” (Vandermeersch 1981:9; Schwarcz et al. 1988; Valladas et al. 1988). When Qafza 6 was tested by the multiple discriminant function procedure against the living populations of the world, the probability of its proportions occurring in a European population was 0.000. It was least likely to be distinguished from a sample of West Africans to which were added the Haya from Tanzania (0.662). When typicality probability levels were used to test the differences between Qafza 6 plus Qafza 9 and various African groups as well as with Cro-Magnon 1, the figures for the comparisons with Africans averaged around 0.5 while for Cro-Magnon they were 0.001 (Quintyn 1999:228-229). Evidently it is easier to get a “morphologically modern” European out of a Neanderthal predecessor than out of Qafza while it is easier to get a “morphologically modern” West African from Qafza than from a Neanderthal. The human remains at Qafza were associated with a fauna that represented an incursion into the Middle East from sub-Saharan Africa (Tchernov 1992:176). There is reason to suggest, then, that the Qafza people were immigrants from sub-Saharan Africa. If that can be considered the case, then Qafza represented contemporary sub-Saharan Africans. Continuity in craniofacial form in Africa, then, can be shown to have persisted for more than 90,000 years (i.e. supporting the idea that Paleoindian, or Kennewick-like skull traits could have persisted 9,000 years).

 Interestingly enough, the dental metrics tell the same story. Summary tooth size is 1503 mm2 for Qafza and 1281 mm2 for the living Ashanti (Vandermeersch 1981: 176; Brace et al. 1991). The figure for the “classic” Neanderthals is 1415 mm2 and for living Europeans is 1127 mm2 (Brace 1979). Although there is more than a 200 mm2 reduction in summary tooth size between Qafza and living Africans and between Neanderthals and living Europeans, the tooth size profile from I1 (the upper central incisor) to M3 (the upper third molar) is parallel for the Qafza/African comparison and also for the Neanderthal/ European comparison. The Qafza/European comparison and the Neanderthal/African comparison show that those profiles are quite different (Brace 1996). Evidently a reduction of all the teeth to proportionally the same extent would produce a modern European dental arch out of a Neanderthal one just as it would produce a modern African dental arch out of the condition represented by Qafza. A similar proportional reduction would not produce the European condition out of Qafza. In addition, the total quantity of reduction needed to produce a modern European dentition out of a Qafza-sized predecessor is greater than would be the case if one postulated a Neanderthal-sized predecessor. Qafza then is an unlikely candidate as a “Proto-Cro-Magnon” or the ancestor of modern Europeans. The realization that the indigenous Mousterian cultural traditions of Europe from the Atlantic coast all the way to Eastern Europe were the most likely sources of the succeeding Upper Paleolithic cultures makes it vanishingly unlikely that the bearers of those cultures were invaders from elsewhere (Svoboda 1986:240; Clark and Lindly 1989:640; Churchill and Smith 2000).

 The dental and craniofacial pattern comparison of Qafza with living Africans produces metric similarities and differences that are of exactly the same order of magnitude as the comparison of Neanderthals to living Europeans. Qafza, however, is accepted as “modern” in form while the Neanderthals are referred to as “archaic” and therefore not acceptable as possible European ancestors. This, however, is simply a subjective stance, and it contains within it a presumption that evolution could not have occurred. When one considers all the aspects of the Qafza skeletal remains vis-a-vis  modern Africans, it is clear that Qafza is every bit as archaic as are the European Neanderthals vis-a-vis living Europeans (Quintyn 1999). The final reason why Neanderthals were dismissed as possible ancestors of Europeans is the assumption that the change from Neanderthal to “modern” form was too abrupt to have happened by in situ evolution (Tattersall 2000:61). As one report put it, “Neanderthals were replaced by modern humans in Europe within too short a period for the former to have evolved into the latter” (McBrearty and Brooks 2000:454).

As with the argument that Neanderthal form was just too different to serve as a source for modern humans, the claim that there just was not enough time for the change to take place has just as little basis in demonstrable fact. The common assumption that the Neanderthals were replaced in relatively categorical fashion is based on the visual image conjured up when a fully developed Neanderthal of over 50,000 years ago is juxtaposed with the skull of the “Old Man” of Cro-Magnon of half that age or even with the skull of a living human being. A nice quantitative index of the change from the Neanderthal to the “modern” condition can be seen in the record of tooth size starting with the early Neanderthals from Krapina in Croatia 130,000 years ago (Rink et al. 1995) and proceeding via the “classic” and Late Neanderthals, the Early and Late Upper Paleolithic, the Neolithic, and the living in Europe (Brace et al. 1987). 

Fig.4: Summary tooth size figures in mm2 for Early Neanderthals represented by Krapina in Croatia, “Classic” Neanderthals, Late Neanderthals represented by Hortus in France, Early Upper Paleolithic represented by Predmostí in the Czech Republic, Late Upper Paleolithic (Prayer 1976), and the European Neolithic (Brace 1979).

This is depicted in Figure 4 where the vertical axis is Summary Tooth Size or TS, the sum total of the cross-sectional areas of each tooth category both upper and lower. As can be seen in Figure 4, the big jump is not between Neanderthal and modern but between the early Neanderthals at Krapina in Croatia and the “Classic” Neanderthals of Western Europe dating to around 50,000 years ago. The time gap between Krapina and the “Classic” Neanderthals is also the largest interval. The difference between the “Classic” Neanderthals and the Late Neanderthal/Early Upper Paleolithic is not so great as the difference between the Early and the Late Upper Paleolithic. When the regression of tooth size through time is plotted, dental reduction from Krapina to the end of the Pleistocene was proceeding at the rate of 1% every 2,000 years. Starting with the Neolithic, however, the rate doubled to become 1% every 1,000 years (Brace et al. 1987).

In any case, the business of converting a Neanderthal head into one of modern form involved a significant reduction in the size of the teeth and of course the jaws that held them (fig.5). The overlap at the end of what is called Neanderthal and the beginning of  “modern” is so complete that they are essentially the same. The earliest “moderns,” then, are really half way in between Neanderthals and the living in both form and time. This can be visually represented by equating nasion-opisthocranion length (see box 1) and comparing the rest of the craniofacial outline as shown in Figure 5. The specimens compared are La Ferrassie I, Predmostí 3, and a recent male from the Faeroe Islands. The cranial outline is essentially similar, but what changes is the face, and it is largely accomplished by the reduction of the jaws and the tooth-bearing part of the facial skeleton. The brow ridge at the top of the face also undergoes a reduction that keeps in step with the reduction of the teeth and jaws. 

Fig.5: Superimposed cranial outlines of a Classic Neanderthal (La Ferrassie A), Early Upper Paleolithic “modern” (Predmostí 3), and recently living male from the Faeroe Islands (Drawn by Kay Clahassey from photographs taken by the author; with permission of Dr. Jean-Louis Heim of the Musée de l’Homme in Paris and Dr. Pia Bennike at the Panum Institute, Copenhagen).

Figures 4 and 5 provide graphic support for the interpretation of the Predmostí remains offered by Hrdlicka early in the last century: “The writer has seen this collection on two occasions and he regards it as by far the most important assemblage of material from the transitional period between earlier and the latest Paleolithic forms. It represents in a measure the much searched-for bridge between the Neanderthal and recent man” (Hrdlicka 1914:551).

The persistent refusal to consider Neanderthals as representatives of a previous stage in human evolution is mainly a survival of anti-evolutionary thinking in paleoanthropology. In many respects, the mind-set of the field has never quite caught up with the establishment of evolutionary biology in the 20th century, and it has a way to go before it catches up with square one in the 21st.  

The Shaping and Spread of “Modern” Human Form: The mechanism producing the reductions both in the facial and the postcranial skeleton have been discussed elsewhere and will only be mentioned briefly here (Brace 1995, 2000). In both instances, the reductions were the consequences of the relaxation of selective force pressures which followed innovations in the cultural realm. The development of effective projectiles represented by the invention of Levallois points in the African Acheulean over 240,000 years ago (McBrearty et al. 1996) improved the hunting capabilities of Middle Pleistocene hominids and reduced some of the necessity for dealing with prey animals through sheer brute strength (Shea 1995; Boeda et al. 1999). Charles Darwin was the first to articulate the idea that the relaxation of selective forces maintaining a particular structure was followed in time by the reduction of the structure itself (Darwin 1859:134-139, 454).

Darwin, of course, did not know what a gene was and how it worked. The full realization of that had to await the birth of molecular genetics nearly a century after his seminal work was published. It is now evident that random changes in the basic genetic material have non-random consequences, most of which are detrimental to the development of a trait if that trait is necessary for survival. If circumstances change, however, and the trait is no longer necessary, i.e. if it becomes neutral, then chance mutations affecting its form will not be selected against. Most mutations interfere with the development of the trait to which they contribute, and, if they are allowed to accumulate, will lead to the failure of that trait to develop to the full extent that it had done during the period when it had significant survival value. The phenomenon has been called the Probable Mutation Effect, which states that the most likely effect of the most likely mutation is structural reduction (Brace 1963, 2000). This is the mechanism that accounts for those reductions observed by Darwin in instances where selective force relaxation had occurred.

In humans, the development of effective hafted projectile points took away some of the selective force favoring the maintenance of postcranial robustness and muscularity. Even more effective, however, was the significance of the discovery and application of string (Brace 1995:272-273). With that initially “unobvious” innovation, it was possible to devise nooses for snares that allowed the capture of prey with no major effort on the part of the hunter. A hook attached to the end of a string gave access to denizens of the aquatic world again with the expenditure of relatively little effort. Finally the use of string to construct netting gave the hunter access to fish and birds in great quantities and made available an enormous biomass that could not be tapped by earlier hunters. Again this could be gained with the expenditure of relatively little effort. The reduction in skeletal and muscular robustness that becomes obvious in the Late Neanderthals just continues in the early “moderns” of the initial Upper Paleolithic and goes on to the present day.  

The primary use of the teeth is in the processing of food. Non-dental food processing practices reduced the amount of previously necessary chewing and relaxed the selective pressures that had kept tooth size at approximately the same level since the beginning of the Pleistocene nearly two million years ago. Habitation in the North Temperate Zone was made possible after the Middle Pleistocene by the use of fire, not only for warmth but also for thawing meat that had frozen before it could be consumed. Fire used for food preparation - cooking - reduced the amount of mandatory mastication, and led to the reduction in tooth and jaw size that converted Neanderthal into “modern” face form (Brace 1979, 1995, 2000; Brace et al. 1987, 1991). It is no accident that the greatest average manifestation of dental reduction from Middle Pleistocene levels is found in just those populations who have longest resided in the North Temperate Zone.

Fig.6: Course of the eastward spread of Levallois tool-making people about 200,000 years ago (after Brace).

Starting late in the Middle Pleistocene, the Levallois technique diffused rapidly to the occupied parts of the western and northern sections of the Old World (Schick 1998), but it was not adopted in much of the Indian subcontinent or farther east in China, Korea, Japan or Southeast Asia (Rajendran et al. 1977-1978; Matsufuji 2000). Initially it was evidently adopted by the people who were already living in the Temperate Zone, but its spread along the northern edge of human habitation and eastwards at the level of the 55th parallel towards Lake Baikal in Kazakhstan well over 200,000 years ago (fig.6) (Derev’anko 1998:342) had to be the result of the movement of people who were entering territory that had not previously been occupied. Even though the evidence is restricted to one juvenile lower second molar and a permanent upper central incisor (Shapkova and Derev’anko 2000:129), it is not inconsistent with the idea that the people who moved into that area bringing the Levallois industry with them were Neanderthals of the same kind as those making Levallois tools further to the west.

Now since their subsistence and food-processing practices were the same as those of their relatives farther west, and if it were those practices that led to the transformation of Neanderthal into “modern” form in Europe, the same processes should have led to the transformation of those eastern Neanderthals into “modern” form as well. Furthermore, that “modern” form should bear a stronger relationship to European morphology than to the characteristic morphology that we associate with the indigenous peoples of other parts of the world such as Africa, Australia and the core of mainland East Asia. This also has to be taken into account when we consider the nature of the people who served as the source for the first migrants into the New World.

 Fig.7: A plot of the first two canonical discriminant function scores for samples representing the main regions of the Old World plus European Neanderthals and Upper Paleolithic representatives. Just over 90% of common variance is accounted for in the two discriminant functions plotted (after Brace et al. 2001, Figure 1 B).

Representatives of the major divergent population blocks of the Old World can be compared with each other and with European Neanderthal and Upper Paleolithic samples using canonical discriminant function scores. These were generated from a battery of 21 craniofacial measurements made on samples of each of the groups tested. The measurements used and the sample sizes are specified in more detail elsewhere (Brace et al. 2001; see box 1). A comparison based on the first two discriminant functions is depicted in Figure 7. Africa and Australia are quite close together while China and Europe are about as far away from them and from each other as it is possible to be. The Neanderthal sample is located not far from the Upper Paleolithic and modern Europeans. When a third discriminant function is added (representing less than 10% of the total variance), the Neanderthals are separated from the rest of the other samples by a very considerable gap.

Removing the Neanderthals and adding more representatives from East Asia including Mongols, Southeast Asians, the Ainu from northern Japan, plus a sample made up of the 29,000-year-old Upper Cave of Zhoukoudian in China and the 18,000 year-old Minatogawa specimens from Okinawa,  a neighbor-joining dendrogram produces the relationships in Figure 8. 

Fig.8: A neighbor-joining tree based on 1,000 bootstrap samplings showing similarities and differences between various modern and European Upper Paleolithic groups compared in fig.12, to which are added Late Pleistocene Asian (Upper Cave/Minatogawa) samples plus three more Asian samples. Because the Asian prehistoric specimens were all males, the representatives of the other samples were also restricted to males only (after Brace et al. 2001; Fig. 2).

Previous tests had shown that the Upper Cave and Minatogawa specimens behaved in similar fashion when compared with the other populations of the world which is why they were lumped for comparison here (Brace 1991:453). The Upper Cave and Minatogawa sample ties to the Ainu and then to the European Upper Paleolithic before these all link to modern Europeans. This is what would be expected if the Jomon and their relatives were indeed descendants of a stratum of related peoples who extended across the northern edge of the Old World from Europe to Siberia in the latter part of the Middle Pleistocene.

 Fig.9: A Chinese Neanderthal from Jinniushan. (Drawn by Kay Clahassey from Lü 1990; reproduced as Fig. 13-12 in Brace 1995).

If that stratum had been characterized by a Neanderthal degree of robustness, then it should be legitimate to postulate an ancestor of the aboriginal populations of the northeast edge of Asia that looked like European Neanderthals. The 1984 discovery of a more than 200,000-year-old skeleton at Jinniushan in Liaoning Province some 400 km northeast of Beijing fits that description adequately (fig.9; Lü 1990; Chen et al. 1994). Following up on that cue, Figure 10 shows a view of the French Neanderthal from La Chapelle-aux-Saints juxtaposed with a recent Ainu from Hokkaido in the north of Japan. There is no reason why a reduction in the size of the jaws and teeth of the same order of magnitude as that depicted in Figure 4 could not have produced the Ainu configuration out of a Neanderthal of that appearance over the course of 50,000 years.

Now if the other more distant populations such as Africa, South Asia and Australia are removed, and Polynesians and the prehistoric Jomon from Japan are added, we get the picture shown in Figure 12. The Polynesians are a step closer to mainland Asia than are the Ainu, and the latter are a step closer than are the Jomon - the obvious ancestors of the Ainu of Japan. One can read Figure 12 as an indicator of the increasing contribution of an indigenous Asian population to the genetic makeup of its neighbors as a result of growth and expansion made possible by agriculture of Chinese origin within the last 10,000 years.

Fig.10: The French Neanderthal, La Chapelle-aux-Saints, upper right, and a recent Ainu, lower left. (Drawn by Mary L. Brace with the permission of Jean-Louis Heim at the Musee de l’Homme in Paris, and of Professor Takeru Akazawa of the University Museum, University of Tokyo).

The Jomon used here are made up of both Middle (5,000 years old) and Late (3,000 years old) Jomon individuals (Tsukada 1986). Gene flow from mainland Asia can be expected to have had an increasing impact as time went on. The Ainu would have been more affected than the Jomon which would account for their slightly closer link to the mainland Asian groups in Figure 11. If the Polynesians had their roots in the Japanese archipelago as has been suggested (Brace et al. 1989), their movement via Taiwan and the Philippines approximately 4,000 years ago would have brought them into contact with agricultural populations of mainland origin, and this would account for the slightly closer link shown in Figure 11.

Fig.11: Neighbor-joining dendrogram of prehistoric and recent groups representing the northern edge of the Old World from Europe to Japan, including SE Asia and Polynesia. The pattern displayed was produced after 1,000 bootstrap samplings (after Fig.3 of Brace et al. 2001).

The Western Hemisphere: Adding representative New World samples to the groups used in Figure 11 and arranging them with the aid of the neighbor-joining procedure gives us the configuration visible in Figure 12. A Bronze Age sample from western Mongolia and a modern Japanese sample were also added. The sample labeled United States/Canadian Border is made up of Blackfoot plus the Juntunen ossuary from northern Michigan and the Ossossane ossuary from Ontario northwest of Toronto. When those samples were run as separate twigs, they were very close to each other which is why they were combined here to make a single case. As can be seen in Figure 12, this grouping is closer to Europe, the European Upper Paleolithic 

and the Mongolian Bronze Age than to anything else. The Pueblo sample from the American Southwest ties closely with Mexico and highland South America when run separately, so they were combined into a single sample to run (fig.13). The Pueblo/ Mexico/Peru sample is about equidistant between the Europe-US/Canadian Border and the Jomo n/Ainu groups but is linked least with the Mainland Asian samples.

Fig.12: Neighbor-joining dendrogram using Fig.12 groups and a series of New World samples plus separate Mongolian Bronze Age and Japanese samples.

The Athabascan and the combined Inuit/Aleut samples are the only New World groups who tie more closely to the core mainland Asians than to Europeans and their relatives. Representatives of the most recent entrants from the northeastern edge of the Old World are part of a movement that goes back no more than 4,000 or 5,000 years, so their ancestors had to have been affected by the burgeoning of those who had developed an agricultural mode of subsistence in mainland Asia. The people who did that, of course, were the Chinese. It should be no surprise, then, that there are evident links between Chinese morphology and that shown in the Inuit/Aleut and the Athabascans.  

The earlier entrants into the New World, dating back to 15,000 years ago and perhaps more, had spread eastwards toward Alaska at a time when there was no Bering Strait. Because of the lower sea level in the last glaciation, the continents simply coalesced (Hopkins ed. 1967). Immigrants could have spread generation by generation into unoccupied territory until they had become the occupants of a new continent without even knowing it. Since this was long before agriculture had allowed the expansion of Chinese and Chinese-related populations, there was much less genetic impact by the Chinese on those who constituted the first New World entrants. The residual hint of that northern tie running all the way from Europe eastwards and showing up in the Great Lakes region of the United States and Canada can be seen in the distribution of the X haplotype of mitochondrial DNA (Macaulay et al. 1999; Schurr 2000 and pp.62-75, this issue).

The suggested route of movement of people into the New World approximately 15,000 years ago is depicted by the solid arrows in Figure 13. Subsequent population movements about 6,000 years ago were set in motion by the population expansion made possible by the development of agriculture as well as by the technological and resource utilization that derived from that even in areas where agriculture was not possible. The dotted arrows show the movement of wheat cultivating people westwards from the Middle East both north and south of the Mediterranean and towards the Indian subcontinent. They also show  population movement from China down into Southeast Asia by rice cultivating people with a later incursion into Korea and Japan.

 Fig.13: The solid arrows indicate the likely route of initial entry into the New World approximately 15,000 years ago. The dotted arrows indicate population movements initially driven by the effects of the development of agriculture and principally occurring 5,000 years ago and less (after Brace et al. 2001).

While rice could not be cultivated in Siberia and over into Beringia, the techniques of extracting nourishment from all sorts of unlikely plant resources that were ultimately derived from the knowledge gained by those who had originally developed Asian agriculture allowed people to spread into parts of the north that were not being exploited by those concentrating on hunting. The mind-boggling ingenuity shown in the use of those unlikely resources is abundantly demonstrated in one of the more remarkable anthropological treatises, namely Franz Boas’ chapter, “Recipes,” in his monograph on the Kwakiutl of the American Northwest Coast (Boas 1921). The course of the movement within the last 5,000 years of people sustaining themselves by various kinds of specialized resource-utilization ingenuity is depicted by the dotted arrows in Figure 13 with larger spaces between the dots. Obviously the coastal people at the northern edge of that distribution had focused their ingenuity on marine rather than plant resources since nothing much grows on land there.

The last movement depicted in Figure 13 is the spread of marine resource-using specialists latterly coupled with root-crop agriculture out into the Pacific. This is indicated by the dashed-line arrows. The beginnings from the Japanese archipelago some 4,000 years ago led to the designation of these people as members of what was called the “Jomon-Pacific” cluster (Brace et al. 1989:105). The final movement out into “Remote Oceania,” as with the Na-Denè movement down into the American Southwest, only happened within the last 2,000 years.  

Conclusion: From the morphological analysis presented here as well as from the molecular genetic evidence, it seems only appropriate that the first entrants into the New World should properly be referred to as Eurasians. Although the second evident wave of immigrants, the Aleut/Inuit and the Athabascans, show a much stronger connection to the living populations of mainland Asia, the presence of an obvious residuum of European form fully qualifies them also to be called Eurasian. If it was the legacy of gracilized northern Neanderthals that contributed to the significant difference between the original human populations of the New World and the post-Pleistocene entrants, the question still remains concerning the form of the antecedents of the modern inhabitants of the Asian mainland and their relatives. At present, however, the almost complete absence of hominid fossils from the Pleistocene in central and southern China means that we have no way of dealing with such a question. The archaeological record shows that people were there (Clark and Schick 1988), but the one specimen that is well enough preserved to give a hint as to morphology, the Dali skull found not far from Xian and over 200,000 years old (Chen and Yuan 1988), has a shape that recalls Homo erectus more than Homo sapiens. As a consequence, it is not of much help in suggesting the course of the emergence of modern Asian craniofacial form. The solution to the problem will only come with discoveries yet to be made.

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Acknowledgements: Support for the work on which this report is based was provided by the Committee on Scholarly Communication with the People’s Republic of China in 1980 and 1985; by the L. S. B. Leakey Foundation in 1986; by the National Science Foundation BNA- 9616298 in 1986-1988, and INT-9107991 in 1992-1993; and by the University of Michigan Museum of Anthropology Research Fund in 1986, 1990-1992. Efforts to incorporate added material from European and Russian Paleolithic and Mesolithic specimens and from the Brazilian Lagoa Santa material in Copenhagen were denied support by the National Science Foundation in 1992, 2000-2001, and by the L. S. B. Leakey Foundation and by the Wenner-Gren Foundation for Anthropological Research in 2001-2002. Crucial craniofacial measurements were contributed for Mongolian and Native American samples by A. Russell Nelson of the Department of Anthropology at the University of Wyoming and by Pan Qifeng of the Institute of Archaeology in the Chinese Academy of Social Sciences in Beijing. Essential computer assistance was provided by Noriko Seguchi of the Museum of Anthropology at the University of Michigan.



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C. Loring Brace is Professor of Anthropology at the University of Michigan, Ann Arbor, and Curator of Biological Anthropology at the University of Michigan Museum of Anthropology. He has documented the impact of changing selective forces and how they have shaped the course of human evolution. This has also been extended into documenting the reasons for the biological differences visible in the living human populations of the world. The study of the latter has led to the realization that one cannot understand the nature of human biological variation if one uses “race” as a starting point. He is the author of The Stages of Human Evolution (5th ed. 1995) and Evolution in an Anthropological View (2000).




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