The significance of Aleš Hrdlička's “Neanderthal phase of man”: A historical and current assessment

AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 56:435-459(1981)

The Significance of Ales Hrdlizka’s “Neanderthal Phase of Man”: A Historical and Current Assessment

FRANK SPENCER AND FRED H. SMITH Department of Anthropology, Queens College of the City University of New York, Flushing, New York 11367 (F.S.) and Department of Anthropology, University of Tennessee, Knotcville, Tennessee 37916 (F.H.S.)

KEY WORDS: Hrdlizka, Neandertal, Human evolution, History

ABSTRACT Alez Hrdlizka’s hypothesis on a Neandertal phase of human evolution is examined in light of current data and interpretations on Neandertals. Hrdlixka’s interpretations are related to his ideas regarding the peopling of the New World. A major early statement of Hrdlizka’s views on Neandertal was his Huxley Memorial Lecture of 1927. We assess this formulation and subsequent development of his hypothesis. Hrdlixka’s position is compared with the “pre- sapiens” and “pre-neandertal” hypotheses on the basis of current theory and data.

In 1980-1981, physical anthropologists celebrate two significant jubilees: the Fiftieth Annual Meeting of the American Association of Physical Anthropologists and the 125th an- niversary of the discovery of the Feldhofer Cave Neandertal. Certainly no single in- dividual can be more intimately associated with both the study of Neandertals and the birth of the AAPA than Alex Hrdlgka (1869-1943), the founder and first president of the association. Thus we felt that an appropriate way of marking the passage ofJoth jubilees would be an analysis of Hrdhcka’s ideas regarding the evolutionary significance of Neandertals and how those ideas fare in light of current evidence.

Such an analysis of Hrdlixka’s views is important and timely for several additional reasons. First, during his lifetime, HrdliEka stood virtually alone in his support of a unilineal approach to human evolution, which included a “Neanderthal phase of man” (Hrdlizka, 1914, 1927, 1930), as an alternative to the “pre-sapiens” hypothesis embraced by the vast majority of his contemporaries. In many respects he should be considered the founder of the modern unilineal approach to Late Pleistocene fossil hominid evolution. Sec- ond, many recently trained scholars fail to fully comprehend and appreciate the degree to which Hrdliyka’s diverse and often seemingly unrelated research interests and theoretical

perspectives were both interrelated and inter- dependent - a fact clearly demonstrated by Spencer (1979) -and therefore often misunder- stand his ideas. For example, his “Neanderthal phase of man” concept, the focus of this paper, certainly does not represent an isolated interest, but did, in fact, develop from his study of and interest in the antiquity of humans in the New World. Furthermore, his feelings on both of these issues were inextricably en- twined with his overall scheme for the peopling of the earth. Also, the existence of an archaic morphological stage between “Pithecan- thropus” and recent humans was a necessary outgrowth of his interpretations of how the evolutionary process operated. In the same vein, his belief in and use of what has come to be called morphological dating -for which he has been unjustly criticized (Stewart, 1949; see also Smith, 1977)-was clearly related to his conception of the evolutionary process and was not simply a convenient mechanism to disprove the antiquity of humans in the New World. Recognition of the “interrelatedness” in Hrdlizka’s work is particularly essential to understanding his perspectives on paleoanthropology. Finally, many of the points raised by Hrdlixka, particularly in his Huxley Memorial Lecture of 1927, both in support of a Neandertal stage of human evolution as well as in opposition to other existing hypotheses on the origin of modern humans, continue to

0002-948318115604-0435$07.00 0 1981 ALAN R. LISS, INC.

436 F. SPENCER AND F.H. SMITH

be valid and thus warrant discussion in the light of more recent data on Late Pleistocene hominids in the Old World.

A BRIEF HISTORY OF THE NEANDERTAL DEBATE By the time Hrdlizka entered the debate, a

considerable amount of controversy and numerous changes of opinion regarding the phylogenetic significance of the Neandertal fossil assemblage had already occurred. During the initial phase of the debate, scientific opinion ranged from the view that Neandertal fossils represented an earlier stage of human evolution (e.g., Huxley, 1863; King, 1864), to the view that they were simply examples of severe pathological conditions of one form or another (e.g., Mayer, 1864; Vir- chow, 1872,1882). By the turn of the century, however, enough Neandertal specimens were known and their probable antiquity established to the degree that most scholars followed the general appraisal of Gustav Schwalbe (1899,1901,1904,1906) that human evolution had essentially proceeded in a stepwise fashion through three distinct morphological stages of development, with the Neandertals representing the intermediary stage.

This appraisal was strongly opposed by Marcellin Boule, who emphasized certain “sim- ian” as well as other unique and primitive aspects of Neandertals in his analysis of the specimen from La Chapelle-aux-Saints (Boule, 1911-1913; Boule and Anthony, 1911) and concluded that Neandertals were not a reason- able antecedent of modern humans in Europe (see also Boule, 1921). Furthermore, Boule suggested, initially on the basis of some of the remains from Grotte des Enfants at Grimaldi (Boule, 1914), that more modern hominids had lived contemporaneously with Neandertals in Europe and that this “sapiens” lineage could be traced, independently of Neandertals, to an earlier period. The subsequent discovery of the infamous Piltdown remains provided Boule with a possible precursor of the “sapiens” lineage.

Following in the wake of Boule’s analysis of the La Chapelle skeleton and the discoveries at Piltdown (1911-1916) there was a general retreat from the Neandertal hypothesis in support of what later became known as the pre- sapiens theory (see below). This theory remained the most popular view of later hominid evolution until well after the Second World War.

There were those, however, who for various reasons continued to support some form of a basically unilineal concept and rejected the idea of a separate pre-sapiens lineage. Among this rather small group of workers - which included Rene Verneau (1924), Franz Wei- denreich (1928,1943,1947), and Hans Weinert (1932, 1953, 1955) -was, of course, Alex Hrdlizka (1914, 1927, 1930).

For reasons far too complex to discuss in this brief synopsis, a “compromise” interpretation of the origins of modern humans began to emerge in the thinking of many scientists in the years between the two world wars. This compromise, later known as the pre-neandertal hypothesis, postulated that certain “progressive’’ Neandertal groups were conceivably ancestral to modern humans but that other groups (especially the western European Wiirm Neandertals) were not. The fundamental difference between this theory and the pre-sapiens theory i s that the latter placed the divergence of lineages leading to Neandertals and modern Homo sapiens at an earlier date.’ While this approach is first encountered in the writings of Grafton Elliot Smith (1924), it did not become popular until the late 1940s and early 1950s and is perhaps best represented by the work of Sergio Sergi (1953a,b) and F. Clark Howell (1951, 1952, 1957).

The volume commemorating the 100th anni- versary of the 1856 Neandertal discovery (von Koenigswald, 1958) and other publications of this period (e.g., Boule and Vallois, 1957; Howells, 1959,1962; Patte, 1955) demonstrate both the continued lack of consensus among supporters of the pre-sapiens and various pre- neandertal hypotheses and the virtual absence of unilinealist views (barring those of Weinert) which postdate Weidenreich’s writings and the last opinion of Sir Arthur Keith (1949). In the early 1960s, however, C. Loring Brace (1962a, 1964) resurrected the unilineal approach, and from this point on through to the present, variations of all three of these major orienta- tions have had their supporters. But in spite of the continuing lack of consensus among paleoanthropologists regarding the phylogenetic significance of the European Neandertals, Hrdliaa’s views, though now necessarily somewhat dated, nevertheless continue to be

IThe views of Weinert (1953, 1955) and Weidenreich (1947. 1949) can in some ways be considered pre-neandertal rather than strictly unilined in perspective. And in some regards, the same can he said for HrdliKka. who clearly believed that only certain Neandertal groups evolved into modern Homo sapiens (1927:272).

HRDLIEKAS NEANDERTHAL PHASE OF MAN 437

cited regularly, not only in historical but also in analytical contexts, by virtually all students of the later phases of human evolution.

THE DEVELOPMENT OF HRDLI~KA’S UNIFIED

In order to understand more clearly HrdliEka’s interpretation of the European fossil hominid record and its relevance to his thesis on the peopling of the Western Hemi- sphere, it is necessary to first consider the salient features of his particular conception of the evolutionary process.

The theoretical basis of Hrdlizka’s views on human evolution

Succinctly, Hrdlizka was convinced that modern humans were the product of what he called “an extraordinary progressive differentiation from some anthropogenic stock, which developed somewhere in the later Tertiary, among the primates” (Hrdlixka, 1912a:2; see also Hrdlixka, 1907~9). But unlike the vast majority of his contemporaries who saw this process terminating mysteriously at some unde- termined point in the late Tertiary or lower Quaternary (e.g., Kollmann, 1884; Wallace, 1887, 1889)2, Hrdlixka was of the opinion that it was an ongoing process (e.g., Hrdlizka, 1907:12-13, 1908, 1912a:3-5, 1918:21, 1921b), and for this reason he regarded human skeletal diversity (both contemporary and historic) as a reflection of the inherent “instability” of the human constitution. “Every organic feature,” he wrote:

HYPOTHESIS ON THE PEOPLING OF THE WORLD

of whatever consistency or importance, is the result of all the factors by which it is affected. With the skeletal parts, by far the strongest of these factors, in itself a very composite one, is the potentiality of heredity, next to which in importance comes habitual muscular action, particularly muscular use due to long established habits of whole groups of people. Heredity, however, especially in so far as it applies to the latest acquired characteristics of the skeleton, is subject to in- cidental irregularities as well as to gradual modifications. Habits of muscle action, on the other hand, change with environment and culture; such changes in activities may take place much more slowly in some localities than in others, yet they are

bound to manifest themselves everywhere in the course of ages and to be followed by corresponding and recur- ring structural alterations. The great skeletal diversity of mankind today can be accounted for in no other man- ner (Hrdliska, 1907: 12-1 3).

Taken at face value this statement can be interpreted either in a Mendelian or Lamarckian sense. However, it is evident that Hrdlixka belonged to neither camp. Essentially, he ad- vocated the reduction of environmental influences to the status of modifying rather than formative factors. That is to say, instead of employing Mendelian concepts to buttress essentially pregenetic racial generalizations, he was toying with the notion of how the genetic material of an organism is affected by the multitudinous forces of the environment. The causes of this “instability,” he later ex- plained:

are on the one hand, the nature of the human organism, which like every other organism is in its ultimate analysis a chemical complex, living by chemical change and subject to physical and chemical influences, and on the other the variability of these influences. So long as the chemical status of the organism, especially that of the developing organism and of the perpetuating or generative elements of the species, is not in absolute and lasting harmony with the environment, so long it is safe to say, will absolute fixedness of structure and form be impossible (Hrdlizka, 1912a:5).

Besides the general implications of his earlier work in the New York asylums (e.g., Hrdlicka, 1898) and the American Southwest (e.g., Hrdliska, 1908), the major evidence Hrdlizka used to support his argument against the prevailing notion of the “permanence of the modern human type” was from a study he had made at the Cairo Medical Museum, Egypt, in

2When Alfred Russel Wallace (1887) examined the evidence for human antiquity in the New World, he saw not only considerable ~ n - tiquity for the specimens but also a continuity of type. Attempting to equate this finding with evolutionary theory, Wallace (1889) suggested that once man had become morphologically differentiated from his “apish kin” (apparently in the mid-Tertiary), he had remained physically stable. This he justified by arguing that with the emergence of the human brain, Homo sapiens had become essentially impervious to the whims of natural selection, and thereby became a special creature in the biotic realm.


1909.3 This study of a human skeletal collection excavated from cemeteries (representing a time span from predynastic times to the early Coptic period) at Lisht and Naga ed-Dbr, enabled him to write with conviction:

the susceptibility of the human organism to modification, even under . . . exceptionally uniform environmental conditions, has not been overcome, and numerous changes in the Egyptian skeleton between the predynastic and middle dynastic, and again between that and the Coptic period, excluding from consideration the influence of negrR infusion, are percep!iJble (Hrdhcka, 1912~5, see also Hrdhcka, 1909).

Thus, in contradistinction to workers such as Franz Boas (1911, 1912), who tended to employ the environm$. to the virtual exclu- sion of genetics, Hrdlicka was endeavoring to develop a concept (albeit a primitive one) of mutational theory to account for the apparent “instability” of the human organism (see Hrdlizka, 1935).

Applying these ideas to the questio3 of earlier hominid evolution, it was Hrdlicka’s developing conviction that there was a rough correlation between morphology and chronology. According to this view Hrdlixka saw the modern human form slowly dissolving in a mosaic of primitive features as one proceeded back through time. As such the Neandertal fossils appeared to represent a phase in this gradual transformation to a more pithecoid form represented in the fossil record by Dubois’s Pithecanthropus. This being the case, Hrdlixka believed that a closer examination of the European fossil hominid record would reveal the existence of a distinct morphological gradient between the hominid populations of the Middle-&ate Pleistocene and the early Holocene (Hrdlicka, 1907:12)-in which the morphological trend in the former would be seen to be in the general direction of “zoological inferiority, while in the latter there would be a tendency toward the modern human form. Hence, on the basis of these “au- thenticated and “geologically ancient crania” (which incidentally he did not examine first- hand until 1912), he felt justified in claiming that “the greater the separation of two skeletal populations in time, the more distinct would be the somatological differences between them” (Hrdli%ka, 1907:12).

This predicted morphological gradient was later confirmed, so Hrdlizka believed, by the skeletons found at Pyedmosti by Karel MaXka (see HrdliEka, 1912a, 1914; Matiegka, 1934, 1938). These fossils, he felt, partially bridged the morphological hiatus between Neander- tals and other Aurignacian-associated (but less Neandertal-like) specimens.

In the meantime, pursuing the above theme of functional adaptation in his work-particularly his$udies on the Eskimo of Smith Sound (Hrdlicka, 1910) and the natives of the Kharga Oasis (Hrdlizka, 1909:143-144, 1912b) -it occurred to Hrdlicka that perhaps one of the underlying mechanisms impinging on the evolution of the human skull had been a change in “masticatory function,” namely, a progressive change in human diet and dietary practices. In a nutshell, he believed that as a result of improved tool use and other technico- cultural techniques there had been a demonstrable reduction in the size of teeth in human evolution and an accompanying change in the size and conformation of the human jaw and cranium.

Following a preliminary study (Hrdliaa, 1911), in which he presented an outline of the above “dietary hypothesis,” Hrdlixka em- barked on a lengthy program of research designed to demonstrate the evolutionary significance of human dentition (see Spencer, 1979)4. Although all of this research is germane to Hrdizka’s Neandertal hypothesis, of particular interest are his initial observations on shovel-shaped incisors (e.g., Hrdlixka, 1907:55, 1908:562, 1910:251, 266) and his subsequent demonstration of their phylogenetic significance (HrdliEka, 1920). Through this study, along with his concurrent comparative studies of hominid and hominoid dentition (Hrdlicka, 1921a, 1922, 1923a,b, 1924), he was able to advance not only his case for a Neandertal phase of human evolution but also his hypothesis which attempted to recon- struct the events leading up to the emergence,

SHrdhxka’s reasons for choosing to do research in Egypt are not without historic interest. Since the early 1840s, when Samuel George Morton had used a collection of embalmed heads from Egyptian catacombs to substantiate his thesis on the ”immutability” and ‘“plurality” of the human races (Morton, 1844), the skeletal material from the Nile valley had been frequently cited as an example of the historic continuity of the human type (e.g., Schmidt, 1872; Thompson and Randall MacIver. 1905).

“This work, which was carried out between 1915 and 1925, constitutes a major contribution to the development of modem dental anthropology. Besides developing a rigorous method of measuring teeth with calipers of his own design, which are still in use today, Hrdlizka’s work from this period also demonstrated the utility of ranked scales for describing trait expressivity, as well as encouraging the study of crown minutia and various other dental features.

HRDLI~KA’S NEANDERTHAL PHASE OF MAN 439

dispersion, and differentiation of modern Homo sapiens in the Old World.

HrdliEka’s rejection of glacial man in the New World

When Hrdlizka began his career in anthropology at the close of the nineteenth century there was already a considerable accumulation of evidence that seemingly supported the arrival of man in the New World during either glacial or preglacial times (e.g., Abbott, 1872, 1888; Ameghino, 1878, 1879, 1908,1909; Lyell, 1863; Mercer, 1892; Putnam, 1897, 1906, 1909; Salisbury, 1897; Wright, 1892, 1897; Whitney, 1879). Although the favorable reception received by these early discoveries in the Western Hemisphere was undoubtedly encouraged by the fact that the battle for “Glacial Man” in Europe had been essentially fought and won, it appears that the willingness with which workers from this period engaged in enthusiastic and often reckless speculation about the age of human bones and implements found in apparent association with extinct faunal remains was due largely to a lack of understanding of either continental or regional geology (Hrdlicka, 1907-1912a; see also Clewlow, n.d.; Willey, 1968; Wilmsen, 1965).

Between 1899 and 1912 Hrdlizka made a thorough study of all the available evidence attributed to early man in the New World. The results of this protracted investigation were summarized in two major works published in 1?07 and 1912. From this investigation Hrd- licka realized quite correctly that archeologists had developed little in the way of methodology during the past century and as such were largely responsible for the plethora of unsub- stantiated claims and absurd opinions that abounded in the literature on the subject of early man in the New World. In an effort to correct this state of affairs, he mounted an energetic campaign during the first quarter of this century, calling for the institution of more rigorous scientific methods and recommended, among other things, the enlisted aid of other disciplines zuch as geology and paleontology (e.g., Hrdlicka, 1907:ll-12, 56-57, 1912a:55-

The prescription Hrdlizka laid down for the identification of geologically ancient human remains demanded “indisputable stratigraphi- cal evidence, some degree of fossilization of the bones, and marked serial somatological distinctions in the most important osseous parts” (Hrdlizka, 1907:13, see also Hrdlixka,

97, 99-125, 1917).

1912a:3-5). Using these criteria, Hrdlizka reported that without exception he did not find a single human bone or artifact that could be unequivocally assigned to a glacial context. Furthermore, all of the osteological material fell within the established range of normal variation of modern American Indians. Tak- ing these findings into account, and guided by the knowledge that the “first traces of man’s appeapnce” were confined to the Old World, Hrdlicka arrived at the conclusion: “that America was peopled by immigration . . . [from Asia], which could not have taken place until after great multiplication and wide distribution of the human species and the development of some degree of culture” (HrdliEka, 1907:9). And, according to the available evidence, Hrdlie’ka said this migration from the Old World had propably occurred toward the close of the Pleistocene or the beginning of the Holocene (Hrdlizka,

During the next decade Hrdlizka’s ideas on the peopling of the New World matured. Criti- cal to this development had been his study of shovel-shaped incisors, which provided not only a genetic basis for the Asiatic origin of the American Indians but also evidence to support the thesis that the European Neandertals were the progenitors of modern human populations. In essence, Hrdlizka (1920) found the highest frequencies of shovel- shaped incisors among the aboriginal populations of the New World and Asia, while among contemporary European and African populations the frequencies were of a much lower order. Upon extending this investigation, he found this dental character in Neandertals as well as in the nonhuman primabees and a variety of other mammals (Hrdlicka, 1920: 460, 46$-464). Commenting on these findings, Hrdlicka said, “The whole study affords one of the clearest and most remarkable illustrations of the common origin of m y with the rest of the mammals” (Hrdlicka, 1920:464).

To explain the distribution of this ubiqui- tous “dental character in the Hominidae, Hrdlicka argued that its development had probably been an adaptive response -a “call

1907:9-10)5.

SAs shown by Fig. 20 Hrdlirka allowed Born 20,000 to 70,000 years for the onset of the postglacial period (e.g., Hrdldka, 1907:9-11). Although present estimates, based on radiocarbon dating techniques, have reduced these dates to something like 11.000 to 10,000 years, it is interesting to note that in spite of this human remains in the New World heve nct been shown to predate Hrdlirka’s estimates for the commencement of the Holocene.


for strengthening the dentition”- but with the gradual improvement in food preparation techniques and tool technology, the need for stronger incisors steadily decreased, leading to their subsequent replacement with the “weaker” flat-surfaced tooth (Hrdlixka, 1920: 464-465). Hence, since the expectation was to find lower frequencies of shoveling among the descendants of those people who had solved, by cultural means, the problems that shovel- shaped incisors had solved biologically, and who had done so for the longest period of time, to Hrdlizka the only satisfactory explanation for the present high frequency of the trait among the peoples of Asia and the New World was that these groups must have remained committed to an Upper Paleolithic way of life long after those groups in the western sector of the Old World (Hrdlizka, 1920:465).

Unfortunately, because Hrdlixka failed to elaborate on this latter point, the reader unfa- miliar with his work will miss the implication of this statement. However, given his commit- ment to the notion that Europe had been the “cradle” and site of dispersion of the human genus, the inference of the above statement becomes that much more apparent. As will be discussed in the next section, i t was Hrdlizka’s contention that Asia and the New World had been peopled exclusively from the western sector of the Old World, principally Europe. Ac- cording to Hrdlixka’s scheme, some of these early itinerant populations from Europe had moved slowly northeastward across the Siberian tundra into East Asia, and from there into the New World, while others moved eastward to the south of the Himalayas. Thus, in marked contrast to those populations south of the Himalayas, this northern branch of humanity had, because of prevailing environmental conditions, remained committed to a way of life not far removed from that of their western Paleolithic ancestors (Neandertals).

After 1920, Hrdlizka’s researches were di- rected primarily to the task of vindicating the “Neanderthal phase of man” and thereby dem- onstrating that the emergence and dispersion of modern Homo sapiens had occurred during the terminal stages of the Pleistocene in Europe, and that it was not until after this event that man had entered Asia and the New World. The results of this work were summarized in his 1927 Huxley Memorial Lecture. Following this summation, the focus of Hrdlizka’s work shifted to Beringia, where he assigned his efforts to the pursuit of evidence to document the thesis that it had been in this

region that the first Americans had entered the New World.

Hrdlizka’s scheme for thepeopzing of the world During the first three decades of this century

the idea that Asia had been the offcina gentium, the “cradle and nursery” of the genus Homo, had been popular among both American and Europe- an paleoanthropologists, particularly after Boule’s censure of the European Neandertals and insistence that modern Homo sapiens had evolved outside of Europe, probably somewhere in Central Asia. Indeed, it had been largely for this reason that Davidson Black (e.g., 1925) had been so eager to work in China, and why Henry Fairfield Osborn had vigorously promoted paleontological research in Mongolia (see Andrews, 1926).

By 1920 the case for an Asiatic origin of Homo rested on the evidence of Dubois’s Pithecanthropus and a scant collection of anthropoid fossils from the Siwalik Hills in northwest India. According to Guy E. Pilgrim (1915), the Punjab fossils represented two distinct types of Miocene apes. One, he claimed, closely resembled the European dryopithecine material and morphologically stood much closer to the modern hominoids, whereas the other possessed a number of anatomical features which placed it directly in the hominid line of primate evolution. In fact, Pilgrim (1915:2) even went so fa r as to suggest that it had been from this latter stock that Dubois’s Pithecanthropus had been derived.

While not denying the evolutionary significance of either Dubois’s or Pilgrim’s specimens, Hrdlixka was nonetheless strongly opposed to the idea that Asia had been the “cradle-land of the genus Homo. In his opinion all of the available evidence pointed to a western rather than an eastern origin. This opinion was based on two simple facts: (1) that after “Pithecanthropus the next beings in the line of man’s ascent” were confined to Europe, and (2) that there had not been a single piece of evidence recovered thus far on the Asiatic mainland suggesting the presence of man prior to the Neolithic period (Hrdlixka, 1921b: 536). In the light of these apparent facts, Hrdlixka conceded two possible arguments to account for the presence of Dubois’s Pithecan- thropus in Southeast Asia. First, it could be argued that for reasons “doubtless environmental” the “Pithecanthropines” had been prompted to migrate westward into Europe, where they underwent subsequent development. Although willing to admit that such a

441 V

HRDLICKA’S NEANDERTHAL PHASE OF MAN

migration was not beyond the realm of possibility, noting that there was sufficient time separating “Pithecanthropus” from the “next being in the line of man’s ascent”-that is, “Homo heidelbergensis”- for the $tter to have evolved from the former, Hrdlicka seriously doubted that this had been the case. In his opinion, a more plausible explanation was that the “Pithecanthropines” had not been confined to Southeast Asia but had been widely distrib- uted throughout the Old World. “Unless man’s origin should be regarded as a pure accident, which seems unjustifiable,” he said, “it may well be assumed that conditions such as favored the differentiation from anthropoids towards man in one locality, existed also in other regions” (Hrdlizka, 192Jb3541). Assum- ing this to be the case, Hrdlicka was thereby able to propose that the eastern “Pithe- canthropines” never acquired a lasting foothold in Asia, and that in all probability it had been a “Pithecanthropine” population located somewhere in Africa from which later forms of humanity, represented in the European fossil hominid record. had zprung.

Essentially, Hrdlicka’s reason for deman- ding the demise of Dubois’s Pithecanthropus was because its geographical location opened the door to the possible arrival of man at the Bering landbridge at a time much earlier than he was willing to entertain. Although the geological evidence at this time evidently preclud- ed the possibility of a northern extension of hominid populations during the Quaternary period (Hrdlizka, 1921b:536), Hrdlizka had no evidence from which to argue that the same conditions had prevailed during the Tertiary, and as he well realized, there was every indication that Dubois’s Pithecanthropus had emerged in Java during the late Pliocene. In fact it was precisely for this reason that Hrdlixka (1930) had rejected Davidson Black’s assessment of the early Pleistocene hominid found at Choukoutien. Black was convinced this specimen, which he had dubbed “Sinan- thropus pekinensis,” was a preneandertal, representing an intermediary form between the anthropoid apes and Homo (see Hrdlixka, 1930:366-367). Viewing this diagnosis a3 a direct threat to his earlier position, Hrdhcka endeavored to downgrade Black’s “Sinan- thropus” to that of a “Neanderthaler.” Accord- ing to Hrdlixka (1930:365-368; see also Spencer, 1979:592-602), the specimen fell within the known range of variation of the Neandertal phenotypic pattern, and as such indicated that Neandertals had spread much

farther eastward than hitherto suspec$d. This view was later substantiated, so Hrdlicka believed, by the discovery in 1938 of a Neandertal skeleton at Teshik-Tash, near Baisun, Uzbekistan. Seizing on this discovery, he wrote:

Thus, unexpectedly, Central Asia fur- nishes a first-rate piece of evidence of early man, which is bound to have a great bearing on the concepts of human prehistory in the Old World, and will necessitate a material revi- sion of notions relating to the Nean- derthal phase of human antiquity. The eastern most localities of Neanderthaloid remains hitherto known were Palestine, Crimea and the Caucasus - the Uzbekistan find extends the territory far beyond this, and halves the distance from the western Neanderthals to Peking Man (Hrdlizka, 1939:297).

In the meantime, however, having established a case for hominid origins in the western sector of the Old World, Hrdlizka was able to proceed with the orchestration of his scheme for the emergence of modern Homo sapiens and the peopling of the earth. Succinctly, Hrdlizka’s scheme can be divided conveniently into four stages. The first stage is characterized by the dispersal of Neandertals throughout southern Europe and North Africa during the Middle- Late Pleistocene. This stage terminates essentially with the emergence of what he called ‘‘westzrn Paleolithic man.” From his work on the Predmosti assemblage and other Aurigna- cian skeletal material, Hrdlixka believed that there was every reason to suggest that the emergent modern human form had been a “generalized type,” analagous somewhat to the Australian aborigine (Hrdlizka, 1921b:541). In the second and subsequent stages, which cov- ered a time period from the “early Aurigna- cian” to “the early Neolithic,” Hrdlizka contended that “western Paleolithic man” became separated into a number of discrete geographical breeding units, a process which resulted in their subsequent differentiation into the various racial types“ characterizing modern Homo sapiens (Hrdlicka, 1921b: 542-544). I t was during the second stage that “western Paleolithic man” established a foothold in Asia. According to Hrdlixka, this had probably been achieved by two routes: (1) a southern route along the Mediterranean Basin to


Asia Minor, and (2) a northern route into the Caucasus and the Caspian-Aral-Turkestan region. In the third stage, those populations which had entered Asia via the northern route moved slowly eastward where they eventually populated northeast Asia and the New World, while those populations in Asia Minor gradually spread south into the Arabian peninsula and Northwest India. In India some of these early itinerant populations either moved further south or continued their trek eastward that was to terminate with their arrival in the geographic cul-de-sac of Australia. The final stage in Hrdlixka’s scheme involved the peopling of Micronesia and Polynesia, and the continuing differentiation of the peoples of the Old and New World into their present ethnic units.

The Piltdown obstacle At the beginning of the twentieth century

there was considerable support for Neander- tals representing a phylogenetic intermediary between Pithecanthropus and modern Homo sapiens (e.g., GorjanoviC-Kramberger, 1906; Keith, 1911; Schwalbe, 1899, 1901, 1904, 1906; Sollas, 1908). However, by the beginning of the second decade this trend was dramatically reversed. To a large extent this reversal in scientific opinion can be traced to the influence of Marcellin Boule, though it is also evident that the discovery of the Pilt- down “fossils” was also an important contributing factor in that it essentially consolidated Boule’s “pre-sapiens” position.

As mentioned earlier, Boule’s contribution to the Neandertal debate dates essentially from 1908 when he was charged with the analysis of the recently discovered Neandertal skeletons of La Chapelle-aux-Saints (1908) and La Ferrassie (1909-1910). In a series of com- munications extending through the sixth, seventh, and eighth volumes of the Annales de Paleontologie, and culminating in their repub- lication as a large independent monograph in 1913, Boule established the classic description of a Neandertal (Boule, 1908, 1911-1913; see also Boule and Anthony, 1911).6 Summarizing this work in a paper presented at the XIVth International Congress of Prehistoric Anthro- pology and Archaeology in Geneva in 1912, Boule (1914) contended that the Neandertals were an archaic and extinct species, and therefore urged their immediate removal from the human phylogenetic tree. At the same time he axed Dubois’s Pithecanthropus, declaring the specimen to be nothing more than a giant gibbon!

At this juncture Boule left the question of the precursor of modern Homo supiens in abeyance. However, following the announce- ment of the discoveries at Piltdown, Boule lost no time in endorsing these “fossils,” which he said, established the fact that there had not been one, but two “races of man” living in the lower Pleistocene -one being the “Piltdown race,” and the other the “Heidelberg race” (represented by the Mauer jaw). Of these two so- called races, Boule said: “The Piltdown race seems to us the probable ancestor in the direct line of the recent species of man, Homo sapiens; while the Heidelberg race may be considered, until we have further knowledge, as a possible forerunner of Homo neanderthalensis” (Boule, 1913:245-246). I t is interesting to note, however, that Boule was among the few European workers at this time to question if the Piltdown cranial remains and jaw belonged to the same indivi$ual (Boule, 1917).

From the outset Hrdlicka had viewed the Piltdown ‘‘@sils” with considerable skepti- cism (Hrdlicka, 1914:501-509). Aside from the nagging doubt of the reliability of the mid- Tertiary geologic age assigned to the specimens (Woodward, 1913), there was the “insurmountable problem” of equating what was an essentially modern-looking skullcap with an entirely apelike jaw. Had this jaw articulated with an equally primitive skullcap like that of Pithecanthropus, then this would have been an entirely different matter alto- gether. But as it was, this “monstrous hybrid” demanded the existence in Tertiary times of a hominid precursor with a modern high forehead and overall cranial vault. From Hrdlizka’s standpoint this was completely at variance with his understanding of biome- chanics and the evolution of hominid crani- ofacial morphology ie.g., Hrdlixka, 191 1). But while, for obvious reasons, doubting the %tiquity of the cranial fragments, Hrdlicka nevertheless regarded the jaw as a “truly remarkable specimen” and morphologically commensurate with its suggested geological age (see HrdliEka, 1914:501-509, 1930536).

After the First World War, Hrdlixka made a detailed study of the Piltdown “fossils” in an attempt to discredit the association of the jaw and calotte. At the same time he reopened his

Wgnificantly, Hrdlizka’s description of the La Chapelle skeleton both in his 1914 and 1930 monographs were essentially very similar t o those of Bode. In his 1914 draft, however, Hrdlicka noted that many of Boule’s statements on the “inferior characteristics” of La Chapelle had been ”somewhat over-emphasized,” and that most (if not all) of these features could he duplicated in modern skeletal samples. But for reasons which appear to have been entirely political. these comments were deleted from the published text (Spencer, 1979).

HRDLICVKAS NEANDERTHAL PHASE OF MAN 443

study of the European Neandertals. The results of this work were presented in a series of scientific reports published in his journal, the American Journal of Physical Anthropology (Hrdlizka, 1922, 1923a,b, 1924), and culmi- nated with his now classic paper, “The Nean- derthal Phase of Man,” which he prepared as the Huxley Memorial Lecture of 1927.

The Huxley Memorial Lecture By 1926, when Hrdlizka was invited to de-

liver the Huxley Lecture of 1927, he was acutely aware of the fact that he was now almost alone in considering Neandertals an ancestral stage in human evolution. Indeed, the roster of scientists opposed to such an interpretation was impressive and included (among others): Boule (1923), Broom (1918), Burkitt (1921), Elliot Smith (1924), Fleure (1921), Gregory (1927; see also Gregory and McGregor, 1926), Giuffrida-Ruggeri (1918), Hill-Tout (1921, 1924), Lull (1922), MacCurdy (1924), Moir (1926), Morant (1927), Osborn (1916, 1926, 1927), Todd (1914a,b), Wilder (1926), and Woodward (1923). The pre-sapiens theory was now even being espoused by former supporters of a unilineal approach (e.g., Keith, 1912, 1915, 1925; Sollas, 1924). In view of the weight of opinion against him, and the importance of the unilineal approach to his

general conception of evolution and his ideas regarding the peopling of the earth, Hrdlizka decided to use this prestigious occasion to systematically defend his position that Neandertals had not become “extinct without issue,” but had in fact been gradually trans- formed into modern Homo sapiens.

In his defense of a “Neanderthal phase of man,” HrdliEka (1927, see also 1930:319-349) presented his classic definition of Neandertals as “the man of the Mousterian culture” (1927:251) and outlined their geographical distribution (Europe, western Asia, and North Africa) and paleontological and geological context. The latter (Fig. 20) he determined from a consideration of the work by Bayer, Boule, Breuil, MacCurdy, and Peyrony (Spencer, 1979).

From the analysis ofv360 known Paleolithic sites in Europe, Hrdlicka found that as one proceeded from the Pre-Chellean on through the Acheulean to the Mousterian there was a dramatic decrease in the number of open living sites. This suggested to him that during Pre-Chellean and Chellean times the climate had been more temperate-namely that these cultures had flourished during the “Main Interglacial Stage.” The gradual decrease in the number of open sites in the Acheulean he interpreted as a direct response to changing environmental conditions, and for this reason

Fig. 20 Hrdlizka’s version of Pleistocene chronology and its relation to the Neandertd phase of human evolution (based on HrdliEka, 1927).


he considered the Acheulean must have straddled the tail end of the interglacial period and the opening stages of t$e last glacial complex. By comparison, Hrdlicka found that well over two-thirds of the Mousterian sites were confined to either rock shelters or caves. Cn the basis of this evidence he placed the Mousterian tradition on the fluctuating downward curve of the “Late Glacial Period (see Fig. 20). But of greater interest to Hrdlixka were the statistics on sites from the Aurignacian and succeeding periods. Here he found that the number of rock-shelters and cave sites increased, rather than decreased. This finding harmonized completely with his conception of the European environment at the close of the glacial epoch, and served to reinforce his argument for the cultural continuity between the Mousterian and Aurignacian (Hrdlizka, 192 7:256-257).

Equating this evidence with the notion that the Mousterian culture and its author had been “swept away” by the coming of a “distinct and superior species of people,” Hrdliska encountered a number of “inconsistencies” and “difficulties.” Assuming for the moment, however, that the hominids of the Aurignacian period had been “superior” and more “virile,” as Keith and others contended, then the expectation, Hrdlizka felt, would be to find cultural evidence indicating a radical departure from the preceding cultural traditions. But this was not the case. In fact, not only did the Aurig- nacians continue to occupy many of the sites previously occupied by Neandertals, but their cultural artifacts showed quite plainly that the Aurignacian life-style was not significantly removed from that of the Mousterian. To develop this argument Hrdlizka had made a careful study of the various cultural assemblies of the European Paleolithic. From this study he could find no evidence to support the view that these lithic indust,ies were indepenent of one another (Hrdlicka, 1927:258). At the sites of Le Moustier, La VerriAre, and especiaIly the rock-sheIter of Audi in the village of Les Eyzies, Hrdliska said there was palpable evidence of a cultural transition from the upper Mousterian to the lower Aurignacian (Hrdlizka, 1927:258-259). Hrdlixka also noted that just because Neandertals apparently did not produce the cave art characteristic of the Aurignacians, it did not mean that Neandertals had no esthetic appreciation and went on to note other possible expreszions of Mousterian artistic endeavor (Hrdlicka, 1927:262).

Hrdlizka’s belief that the Aurignacian was not an intrusive technology but an indigenous development from the Mousterian was also due in no small measure to the analysis of the Mousterian sequence at La Quina by Henri Martin (e.g., 1911), whom he met in 1912. Unlike many of his contemporaries, notably Boule, Martin felt that the Mousterian had undergone an increase in sophistication from its earlier to later phases (see Spencer, 1979:403).

Yet another aspect of the “Aurignacjan influx” orthodoxy which troubled Hrdlicka was why these people had waited until the height of the glacial period to invade Europe. Hrdlizka (1921b3545) felt that this violated the “laws” governing the movement of human and animal populations. According to Hrdlizka, human populations always tended to move in the “direction of least resistance [climatej” and “in the direction of better material prospects [food]” (HrdliEka, 1927:260).

Finally, and perhaps most significantly, the invasion hypothesis presupposed the existence of not only a large invading population capable of exterminating the resident Nean- dertals, but also the existence of a still lar$er mother population elsewhere. And as Hrdlicka noted, there was no evidence for any such non- Neandertal mother population either in Europe or elsewhere. As for the suggestion that this “invasion” may have been a “peaceful extension,” Hrdlixka contended that this would have led to an “amalgamation” with, rather than the “extinction” of, the European Neandertals (Hrdlizka, 1927:260).

The crux of the entire issue, however, was the skeletal material, for it was essentially on this evidence that the idea of the separateness and discontinuance of NeanFtals had been based. This impression, Hrdlicka claimed, was based on the erroneous supposition that Nean- dertals were an unusually homogenous group which conformed to a “generalized primitive type.” This latter concept had been based on specimens like La Chapelle, Spy 1, and Nean- dertal (Feldhofer Cave). However, when the entire” Neandertal sample was considered, Hrdlicka saw not only extensive variation but also a clear tendency for certain specimens and features to approach closely the pattern of modern Homo sapiens. Considering this point, he wrote:

Here is facing us, evidently, a very noteworthy example of morphological instability, an instability, evidently, of evolutionary nature, lead-

HRDLIEKAS NEANDERTHAL PHASE OF MAN 445

ing from old forms to more modern. . . . Taking the remainder of the skulls, jaws, and bones attributed to the Neanderthal phase, it is seen that both the variability and the number of characters that tend in the direction of later man increase considerably. The Krapina series, by itself, is probably more variable from the evolutionary point of view than would be any similar series fr3m one locality a t the present (Hrdlicka, 1927: 267- 268).

Since it was not possible to arrange the Neandertal fossil rectrd in a reliable chronological sequence Hrdlicka was obliged to base his transformational argument on the “somatological” evidence. This he did by comparing the Neandertal assemblage with a corresponding number of early postglacial hominids. From this comparative study he said it was possible to demonstrate “scales of gradation from forms that stand considerably apart from those of later man (as in Neanderthal, Spy, La Chapelle, Le Moustier) to forms that approach to, or merge with, the modern (many parts of the Krapjna, La Ferrassie, La Quina skeletons)” (Hrdlicka, 1927:269).

Although this would seem to imply that Hrdlizka was advocating the partition of Neandertals into earlier and later forms (namely, what later became known as the “progressive” and “classic” Neandertals), it appears that what he was really intending to convey to his London audience was that each of the fossils under consideration could be regarded as an expression of the range of morphological variation of the Neandertal phenotypic pattern. This is particularly apparent from his description of the Belgian Nean- dertals. “Here the student is confronted with a find in the same terrace and deposits,” Hrdlizka said:

at the same level, and but 6 feet apart of two adult male skeletons from the later Mousterian time. One of these skeletons, No. 1, has a skull the vault of which is a replica of that of the Neanderthal cranium, with typically Neanderthal bones of the skeleton. But this skull is associated with. upper and lower jaw and teeth that may be duplicated today among the lower races. And the skull of the second skeleton is so superior in size, shape, height of the vault, and the

height of the forehead, to No. 1, that the morphological distance between the two is greater than that between No. 2 and some of the Aurignacian crania, such as the Most (Brux) or Brno No. 1 (Brunn) specimens (Hrdlizka, 1927:268).

The full significance of this argument was not fully appreciated until after the appearance of the published description of the Nean- dertal material from Mount Carmel in 1939.

By advancing the idea that Neydertals were a highly variable group, Hrdlicka was thereby able to argue that natural selection had operated on a basic neandertaloid pattern, in which those hominids endowed with the genetic properties favoring their survival, or as he phrased it: “the most fit or able-to-cope- with-the-condition group,” evolved into early modern sapiens, while their less fortunate kin became extinct.

As for the pre-sapiens hypothesis, Hrdlizka astutely pointed out that:

they give us Homo sapiens without showing why, or how, and where he developed his superior make-up. . . . They place Homo sapiens in Africa or Asia without troubling to offer the evidence of his ancient dominion in these regions.. . .If he lived in Europe, coexisting with the Neander- thaler, where are his remains, and why did he not prevail sooner over his inferior cousin? His traces, it will be recalled, never, in Europe or elsewhere, precede or coexist with, but always follow the Mousterian (Hrdlizka, 1927:270).

Summarizing his picture of the evolutionary development of the Mousterian hominids, Hrdlizka said:

During this period man is brought face to face with great changes of environment . . . which call for new adaptations and developments.. . . Such. . . factors must inevitably have brought about, on the one hand, greater mental as well as physical exertion and, on the other hand, an intensification of natural selection, with the survivd of only the more, and perishing of the less, fit. But . . . evolution would certainly differ from region to region, as the sum of the


factors affecting man differed, reach- ing a more advanced grade where the conditions in general proved the most favorable; while to many of the less favored groups disease, famine, and warfare would bring extinction. . . . Here seems to be a relatively simple, natural explanation of the progressive evolution of Neanderthal man, and such an evolution would inevitably carry his most advanced forms to those of primitive Homo supiens (Hrdlizka, 1927:271-272).

Contrary to Hrdlixka’s expectations, the Huxley Lecture was not successful in turning the tide of general scientific opinion (e.g., Elliot Smith, 1928; Osborn, 1930; Hooton, 1930, 1931; MacCurdy, 1937). The pre-sapiens scheme was now firmly entrenckd, and for the next several decades, Hrdlicka’s phylogenetic interpretations were generally either avoided completely or mentioned as a highly unlikely alternative to the pre-sapiens view (e.g., Vallois, 1954).

A CURRENT VIEW OF UPPER PLEISTOCENE HOMINID EVOLUTION IN EUROPE AND

WESTERN ASIA

The years since Hrdlixka’s Huxley Memorial Lecture have witnessed an enormous accumulation of various types of information pertinent to Upper Pleistocene hominid evolution in the Old World. Much of this information has fortunately been in the form of additional fossil hominid remains (see Mann and Trin- kaus, 1974 for a review of all but the most recent finds). However, equally important, at least in some respects, are such factors as a better understanding of the evolutionary process and how it applies to the later phases of hominid evolution, a more accurate picture of chronology and paleoecology of the Late Pleis- tocene, as well as an improved understanding of the cultural complexes of the Middle and Upper Paleolithic in Europe and technologi- cally equivalent periods in other areas of the Old World. While it is neither appropriate nor feasible to discuss all of this in detail, it would seem a fittin contribution to this consideration of Hrdlicka to examine the degree to which his observations, interpretations, and criticism of other perspectives on human evolution are still valid or “measure up” in the light of this knowledge.

%

A rchaeobgy

In his archaeological discussion, Hrdlizka (1927) emphasized the continuity between the Mousterian and the earliest phases of the Up- per Paleolithic and argued, contrary to the vast majority of his colleagues, that the Aurignacian emerged from the Mousterian in Europe. In this regard, he noted that no non- European source area for the Aurignacian had been conclusively demonstrated. Today it is generally accepted that the earliest Upper Paleolithic traditions in Europe, the ChB telperronian in France and the Szeletian in central Europe, developed from indigenous Mousterian (e.g., Bordes, 1968a,b, 1972; Chmielewski, 1972; Movius, 1969; Pradel, 1966; de Sonneville-Bordes, 1972). The Aurig- nacian, however, is still considered by most prehistorians to be an intrusive culture into western Europe (e.g., Bordes, 1968a,b), although the apparent contemporaneity of some Aurignacian and Ch&telperronian levels has led to the suggestion that they might represent adoptive or functional variants of the same tradition (Binford, 1972). In central Europe, although some workers also consider the Aurignacian to be intrusive (Kozlowski, 1979) into this area, a growing number of scholars argue that the roots of the Aurig- nacian are clearly evident in central European Middle Paleolithic traditions (e.g., Hahn, 1973, 1977; Valoch, 1972, 1976). I t is interesting to note that the earliest dates for the Aurignacian in central Europe (Gabori-Cshnk, 1970; Koztowski, 1979) are in excess of 40,000 years B.P., while the earliest in western Europe are around 34,000 years B.P. (Movius, 1960, 1972).

In addition to the distinct possibility (and perhaps even probability in central Europe) that the Aurignacian is an indigenous development in Europe, it is important to note that those who favor an extraneous origin for the Aurignacian in Europe have yet to identify the source area. In the Near East, which has traditionally been considered the most likely source, the Middle Paleolithic-Upper Paleo- lithic transition occurs at roughly the same time as in Europe (Farrand, 1965, 1972). And while there are elements that can be regarded as somewhat Upper Paleolithic-like in pre- Upper Paleolithic cultural complexes in the Near East, the same can also be said for pre- Upper Paleolithic complexes in Europe fe.g.,

V HRDLICKAS NEANDERTHAL PHASE OF MAN 447

Brose and Wolpoff, 1971). Thus, there would appear to be no conclusive evidence to indicate an earlier development of the Upper Paleo- lithic in the Near East than in Europe, which consequently makes it difficult at this time to support the Near East as the source area for the European Aurignacian.

Where then might the source area be? To our knowledge, the only other area where certain technological advances in lithic industries might be occurring significantly earIier than equivalent developments in Europe is southern Africa during the Middle and Late Stone Ages (Klein, 1977). However, the chronology (specifically the early dates for the Middle Stone Age) is not conclusive. Further- more, there is no indication of a clinal spread of these innovations toward Europe, which would be necessary to demonstrate (unequivocally) their influence on the European Paleolithic .

Hence, on the basis of present kTowledye, the following points asserted by Ales Hrdlicka are still strongly defendable: (1) The Aurigna- cian could certainly have developed out of the Mousterian in Europe, and (2) a non-European source area for the Aurignacian has yet to be conclusively established.

In other respects, however, certain of Hrdlizka’s 1927 ideas and arguments have not held up. For example his concept of the nature of Pleistocene glacial phenomena was far too simplistic and his chronology, though not as far off as some, was nevertheless incorrect (cf Musil and Valoch, 1966; Butzer, 1971). From all indications, the earliest Aurignacian, at least in central Europe, is associated with an interstadial period. Consequently his argument that the Aurignacian peoples were probably not invaders because they would be unlikely to “invade” Europe at the height of the “glacial period” would no longer seem valid, if in fact it ever was.

Likewise, Hrdlixka’s definition of Neander- tal as the man of the Mousterian period no longer seems accurate. The simple Mous- terian-Neandertal and Modern Humans/ Upper Paleolithic association pattern, though certainly the norm, clearly does not apply in all cases. For example, at Qafzeh and Skhal in Israel basically modern Homo sapiens-grade hominids are associated with Levalloiso- Mousterian (McCown and Keith, 1939; Vandermeersch, 1977; Wolpoff, 1980), while in Europe the recent discoveries at St. Cesaire in

France (Leveque and Vandermeersch, 1980, 1981) and possibly at Vindija in Yugosolavia (Wolpoff et al., 1981) illustrate the association of Neandertals with early Upper Paleolithic traditions. As was suggested several years ago (Stringer, 19741, continuing to define “Neandertal” on the basis of culture does not appear appropriate.

The Fossil Hominids Some time ago, Howells (1974) suggested

that a clearer understanding of the evolutionary relationship between archaic Homo sapiens (Neandertals in Europe and the cir- cum-Mediterranean region) and early modern Homo sapiens might be attainable by considering the problem from a more regionalized “populational” approach rather than on a worldwide basis. Concentrating on the Nean- dertal/early modern relationship, there would appear to be three generalized regions where such an approach is possible: western Europe (essentially France, Belgium, England, Italy, and the Iberian Peninsula), south-central Eur- ope (basically the Pannonian Basin and its surrounding highlands - composed primarily of northern Yugoslavia, eastern Austria, western Rumania, and parts of Czechos- lovakia), and the Near East (the Levant and surrounding inland countries, like Iraq and Iran). Other areas, such as eastern Europe (primarily consisting of European Russia) and North Africa, were also inhabited by similar hominid forms during the Late Pleistocene, but there are not enough pertinent specimens from these regions to make this type of analvsis Dossible.

Western Europe. For a number of reasons (see Brace, 1964), the study of Neanderthals and European Late Pleistocene hominid evolution in general has been concentrated primarily on material from this region. As a consequence of this, the sites and hominid fossil remains from western Europe are the most well-known (e.g., La Chapelle, La Ferrassie, Gibraltar, Cr6- Magnon, Combe Capelle, and Grimaldi), and the conceptions of the course and nature of European Late Pleistocene hominid evolution have historically been based primarily on western European evidence.

For the most part, the Neandertals from this area conform to the generalized descriptions given by most authorities as characteristic for the taxon H p o sapiens neanderthalensis (e.g., Hrdlicka, 1930; Vallois,

F. SPENCER AND F.H. SMITH 448

1954; Howell, 1957; Howells, 1973, 1974; Brace and Montagu, 1977; Trinkaus and Howells, 1979; Wolpoff, 1980), and are often referred to as “classic” Neandertals (e.g., Howell, 1952). Significant features include a continuous, projecting supraorbital torus; a relatively low, broad cranial vault, generally exhibiting occipital bunning and lambdoidal flattening; a large, prognathic face; lack of a canine fossa; very broad nasal aperture and anterior palate; large anterior teeth, generally exhibiting heavy attrition; a mandible lacking a mental eminence and trigone, but exhibiting a receding symphysis and retromolar space; and certain postcranial characteris$cs (to be discussed later). Of course, as Hrdlicka noted, and as more recent workers have emphasized (Brose and Wolpoff, 1971; Brace and Montague, 1977; Wolpoff, 1980), there is a considerable degree of variability in the expression of these features. Several specimens exhibit certain features that approach the modern H o m o sapiens condition to a greater degree than is usual for western European Neandertals. For example, rather vertical symphyses and incipient mental eminences are noted in Monte Circeo 2 and La Ferrassie 1, while very little occipital bunning is ex- hibited in specimens like Saccopastore 1 and Le Moustier.

The chronological problem that faced Hrd- lixka in 1927 still frustrates the analysis of western European Wurm Neandertals. Presently the majority of these remains (including those from La Chapelle, La Ferrassie, La Quina, and Hortus) are correlated to Wurm I1 (Heim, 1976a,b; de Lumley, 1976; Vandermeersch, 1976), placing them temporally between approximately 55,000 and 37,000 years B.P. However, for various reasons, the temporal position of many of the specimens (particularly those found early in this century) within this period cannot be determined conclusively, which severely hampers the search for systematic patterns of change over time during Wurm 11. Only a meager number of Late Pleistocene specimens seem to be pre-Wurm I1 (see Wol- poff, 1980), and there appear to be very few systematic differences between the pre-Wurm I1 and Wurm I1 Neandertal samples in western Europe. There is, however, clear evidence of anterior dental reduction from the pre- Wurm I1 to Wurm I1 Neandertal samples (Frayer, 1978; Wolpoff, 1980); and reduction of Wurm I1 anterior teeth is particularly evident at the late Wurm I1 site of Hortus (de Lumley,

1973, 1976), where the anterior teeth are consistently smaller than the rest of the Wurm I1 sample. Except for this tendency for dental reduction, the only other existence of “progressive” features in the Wurm I1 western Euro- pean sample consists of occurrences of traits such as those mentioned earlier, which are generally isolated in specimens otherwise exhibiting “typically” Neandertal morphology.

The recent discovery of a Neandertal in Chh telperronian context at Saint Cesaire in France (L&v&que and Vandermeersch, 1980, 1981) deserves special consideration a t this point. Although the preliminary considerations of the morphology of this specimen em- phasize that it is virtually identical to earlier (Wurm 11) Neandertals (Levbque and Vander- meersch, 1980, 1981; ApSimon, 1980), limited examination of the specimen by one of us (F.H.S.) in Paris in 1978 left the impression of a strong similarity to the Vindija Neandertals in certain features. Others (Wolpoff, personal communication) have come to the same conclusion. I t is interesting to note that the only other hominid remains definitely attributable to the Chiitelperronian (teeth from Arcy-sur-Cure) also exhibit Neandertal affinities (Wolpoff, 1980).7 While more information is needed to be certain, it would appear that the latest Neandertals in western Europe, Saint Cesaire and possibly Hortus, do indicate some pattern of morphological change toward the direction of modern H o m o sapiens. However, if the age of the Saint Cesaire specimen is comparable to the few available radiocarbon dates for the Chatel- perronian at other sites (like Arcy-sur-Cure), the specimen would date about 31,000 to 33,000 years B.P., which would make it rather late for a Neandertal. Some support for this date comes from the approximate 35,000-year- B.P. date associated with the aparently Neander- tal La Quina H27 temporal (Vallois, 1969). We shall return to the possible significance of this later.

Except for the above mentioned Chiitel- perronian specimens, the earliest significant Upper Paleolithic remains known from western Europe are associated with the Mid- dle Aurignacian. Thus, a critical (albeit short) temporal gap still remains between the latest Neandertals and earliest moderns in western Europe, which also complicates evaluation of

’The robust skeleton from Combe Capelle, recovered by Klaatsch and Hauser 1191Ol under somewhat dubious circumstances, cannot he definitely considered to be Chhtelperronian.

V HRDLICKAS NEANDERTHAL PHASE OF MAN 449

the nature of the relationship between these two hominid types. The earliest Homo sapiens sapiens specimens include the famous remains from CrG-Magnon and Grotte des Enfants (Billy, 1976), which while quite robust are nevertheless unquestionably modern Homo sapiens in total morphological pattern. The robustness itself, however, especially in the occipital region, is reminiscent of Neandertal morphology, as are other aspects of their morphology (see Wolpoff, 1980). The impression that there is virtually no difference between these specimens and modern Euro- peans (see Vallois, 1954) is far from the truth. A significant amount of evolutionary change is documented from the early to the late Upper Paleolithic samples in western Europe (Billy, 1976; see also Frayer, 1978). Thus, the western European early Upper Paleolithic material is demonstrably intermediate in the expression of certain features and in overall robusticity between Neandertals and later Europeans.

Central Europe. Although several Upper Pleistocene fossil hominid specimens in South- Central Europe were ong the earliest such discoveries made (e.g.Fpka, i,n 1880; Mladez, beginning in 1881; Predmosti, in 1894-1895; Krapina, beginning in 1899; and Ochoz, in 1905) and certainly consitute some of the most important fossil hominid samples in Europe, they have received proportionately little detailed attention in comparison to western European and western Asian specimens - except by central European scholars (e.g., Jelinek, 1969, 1976; Vlxek, 1$67, 1969). I t is interesting to note that Hrdlicka was quick to recognize the tremendous importance of this material, especially that from Krapina and P:edmosti (Hrdlizka, 1912a, 1914, 1927, 1930), to the question of the Neander- talimodern Homo sapiens relationship in Europe. Recently, interest in and analysis of this material by noncentral European workers have increased significantly (e.g., Smith, 1976a,b, n.d.; Smith and Ranyard, 1980; Frayer, 1978; Wolpoff, 1979, 1980; Wolpoff et al., 1981).

While many of the sites in this region were initially excavated around the turn of the century, and thereby suffer from the same problems of imprecise methodology that plague contemporaneously excavated sites in western Europe, several of the more important sites have been recently reexcavated using much more acceptable techniques of excavation and documentation. Furthermore, reanalysis of stratigraphy and other aspects of earlier-

excavated sites has been carried out in many instances. Thus, while problems certainly remain, a reasonably good stratigraphic/ chronologic framework is available - at least for the most productive hominid fossil- producing regions in Czechoslovakia (Musil and Valoch, 1966) and northern Yugoslavia (Malez, 1978).

In the context of this framework, the Nean- dertal remains from South-Central Europe can be confidently divided into two temporal samples (see Smith, n.d.). The early sample consists of the remains from Krapina, GBnovce, Ochoz, and probably Subalyuk. The time span represented extends from the Riss/Wiirm Interglacial through the Early Wurm Stadial (roughly equivalent to Wurm I in western Europe). The late sample is comprised of the sp imens from Vindija

encompassed within the time range of the Lower Wurm Stadial (Wurm 11).

The early Neandertals from this region, when considered as a group, clearly exhibit all of the characteristic morphological features of the taxon Homo sapiens neanderthalensis; and despite the fact that many of them have often been considered more “progressive” or more “generalized (i.e., less extreme) in their expression of Neandertal anatomical characteristics than western European “classic” Neandertals, there is no feature or complex of features which supports this assertion. In every aspect of cranial, dental, and postcranial anatomy where comparisons are possible, the early South-Central European Neandertals exhibit the same morphological pattern” as their counterparts in western Europe (Vlcek, 1969; Smith, 197613, n.d.; Wolpoff, 1980). For example, the Neandertals from Krapina have often been portrayed as being more “progressive” than thswestern European Neandertals. In fact, Hrdlicka frequently cited the Krapina remains as an example of Neandertals exhibiting features which presented a “considerable variation and that of a rather progressive tendency” (1930:215). I t is clear, however, that he did not consider them any more “progressive; as a group than the western material (Hrdlicka, 1914, 1927, 1930). Recently, a series of detailed studies on various aspects of the Iirapina samples (Smith, 1976b, 1978; Trinkaus, 1978; Wolpoff, 1979) have demonstrated conclusively that the Krapina hominids do not differ from western European Neandertals to any significant extent.

(level G3), Khlna, and 2 ipka, all of which are


The late Neandertal sample from South-Cen- tral Europe exhibits basically the same total morphological pattern as the early sample; ut as the descriptions of Vindija, Kfilna, and

kieka (Wolpoff et al., 1981; Jelinek, 1967; Vlcek, 1969) demonstrate, there are a number of features in which this late group ap- proaches the early Homo sapiens sapiens condition to a consistently greater extent than the early group does. This fact has been pointed out repeatedly by centralvEuropean scholars (e.g., Jelinek, 1969; Vlcek, 1958, 1969), but these assertions were based on the

ther sparse Czec oslovakian material from

discovery of the Vindija hominids (Malez et al., 1980) has solidified this position. At Vin- dija additional specimens preserving the same anatomical regions (supraorbital tori , mandibles, and maxillae) represented in the Czechoslovak sample have been recovered, thus markedly increasing sample sizes. The Vindija specimens have confirmed the same general “progressiveness” in these areas previously observed in the Czechoslovak specimens, indicating that the entire “population” of late Neandertals in South-Central Europe were more evolved than the early sample in the direction of early modern Homo sapiens. In the supraorbital region, for example, it has been documented elsewhere (Smith and Ran- yard, 1980; Wolpoff et al., 1981) that the

indija supraorbital tori (as well as that of x ala) exhibit both metric diminution and certain morphological changes from the earlier Krapina sample. The latter basically concerns relatively greater midorbit reduction in the late sample, which can be considered an incipient stage in the emergence of distinct supraorbital trigone and superciliary arch segments of the supraorbital area. Addition- ally, the mandibular remains from Vindija exhibit (as a group) relatively vertical symphyses (all less than 90”); d two of the

give indications of at least incipient mental eminences and trigones. There is also evidence of some degree of reduction in certain significant facial characteristics in the late group. For example, the Khlna and both Vin- dija maxillae exhibit rather narrow nasal apertures compared to the western European sample. Furthermore, the Vindija (but not Kfilna) specimens have rather shallow palates and reduced alveolar heights. ,As0 the Khlna and perhaps the Vindija maxillae possess very shallow canine fossae. For further details on

!$ ipka, Kfilna, and !3 ala. The relatively recent

Vindija specimens, as well as r ipka (probably),

these finds see Wolpoff et al. (1981) and Smith (n.d.).

The early modern Homo sapiens (early Upper Paleolithic) sample from South-Central Europe exhibits several characteristics that indicate a close morphological connection with late South-Central European Neandertals and suggest an intermediate position between Ne- andertals and later European modern hominid populations. The material that constitutes this sample is dated to older than circa 25,000 years n.P. and includes specimens from Pyedmosti, Zlaty, Kfig, MladeE, Brno, Velika PeCina, Dolni V&tonice, and Pavlov. Hrdlizka clearly recognized that the morphology of stme of these specimens, specifically the Predmosti and Brno crania, was more similar to Neandertals than that of most western Euro ean Upper Paleolithic specimens (Hrdlska, 1912, 1914, 1927, 1930). There is, however, no record of his examining the complete Mladex sample, which indicates this connection even more distinctly. This morphological “connection” is illustrated by a number of features (see Smith, n.d.; Smith and Ranyard, 1980; Wolpoff, 1980 €or more details).

For example, the supraorbital superstruc- tures, although essentially modern in form, are generally considerably more salient and robust than later Homo sapiens sapiens populations. Furthermore, the division between the supraorbital trigone and superciliary arch is not always distinct, as in later populations. Some specimens, such as Mladec 5, almost exhibit a supraorbital torus. I t is argued elsewhere in more detail (Smith and Ranyard, 1980) that the supraorbital region in South-Central European hominids represents a morphological continuum from early Nean- dertals, through late Neandertals, to early modern Homo supiens. In addition, hominids which make up this early modern Homo supiens sample are intermediate in tooth size between Wurm Neandertals and later Upper Paleolithic-associated specimens (Frayer, 1978; Smith, 1976133, contributing to a distinct pattern of dental reduction traceable from RissiWurm Neandertals through Mesolithic and Neolithic Europeans (Brace, 1979; Frayer, 1978). Nasal apertures tend to be only slightly narrower than late South-Central European Neandertals, and canine fossae are not extensively more excavated. Prognathism, general facial robustness, and rugosity are generally more developed in the early modern Homo supiens sample than in later European

HRDLI~KA’S NEANDERTHAL PHASE OF MAN 451

Upper Paleolithic “populations.” Occipital buns, though somewhat different from and generally less developed than those of Nean- dertals, are common in the early Upper Paleo- lithic sample and are most logically considered remnants of a Neandertal ancestry. These structures are much less frequent in later populations.

While these and other features indicate a good deal of morphological similarity between early Upper Paleolithic and late Neandertal specimens from South-Central Europe, the total morphological pattern of early Upper Pale- olithic hominids unequivocally places them in the taxon Homo sapitns sapiens. No specimen, not even Mladec 5 or 6, possesses a complex of features that would warrant its classification as a Neandertal. Stringer (1974, 1978) has illustrated this quite nicely by means of multivariate analysis of the pertinent material (see also Brauer, 1980; Trinkaus and Howells, 1979). Although the multivariate evidence has had the effect of enforcing the idea of a qualitative difference between Neandertal and early modern Homo sapiens morphology, an idea which has its roots in the original replacement concepts (cf. Boule, 1914, 1923; see also Vallois, 1954), it is important to note that neither Stringer’s nor Brauer’s analyses can include the late Ne- andertal remains from South-Central Europe, due to their fragmentary condition. Therefore at present, it would appear that impressions from such studies are not directly applicable to the Late Pleistocene hominid sequence in this area of Europe.*

Western Asia. The Near East has been a focal point for the study of Upper Pleistocene hominid evolution ever since the discovery of the Mount Carmel hominids (McCown and Keith, 1939). Historically, the early chronological interpretation of these sites (e.g., the contemporaneity of the Skhiil and Tab- hominids and their Riss/Wiirm equivalent age) contributed to the impression that modern Homo sapiens had evolved somewhere to the “east” of Europe. Recent stratigraphic, faunal, and archaeological analyses (Jeligek et al., 1973) have shown that the sites are not contemporary and certainly not RisslWiirm in age.

Presently, all workers agree that the Near East was populated by populations of Homo sapiens neanderthalensis during the period circa 60,000 (or earlier) to 46,000 years B.P. Specimens attributed to this taxon come from the sites of Tabiin, Shanidar, Amud, and Zut- tiyeh and exhibit only rather minor regional

differences from European Neandertals (Stewart, 1977; Trinkaus and Howells, 1979; Wolpoff, 1980). The SkhQl hominids are considered by most scholars to be early modern Homo sapiens, although certainly exhibiting a number of features very reminiscent of Near East Neandertals (Wolpoff, 1980). In fact, the most ,well-known cranium (Skhd 5 ) , which is generally represented as “typical” of the Skhiil hominids, is one of the most modern-looking of the specimens in the sample. Other specimens (e.g., Skhiil 4 and 9) are somewhat more Neandertal-like. The Skhnl hominids are clearly more recent than the Neandertal specimens listed above (Smith, 1977; Trinkaus and Howells, 1979; Wolpoff, 1980) and would appear to date between 31,000 and 35,000 years B.P.

The Qafzeh hominids, the first of which were discovered in the 1930s, have only recently been systematically described by Vander- meersch (1977). They are very similar to the Skhtil hominids in total morphological pattern, as well as in the expression of considerable variation (see Wolpoff, 1980), and would appear to represent the same evolutionary stage of development. These hominids, however, have been claimed to be anywhere from 44,000 to around 70,000 years old. But even in the detailed geological analyses of Farrand (1972, 1979), who favored an early date, there is nothing that would preclude an age of between 30,000 and 40,000 years for these hominids - a range which is supported by certain archaeological considerations and some admittedly questionable amino-acid racemization dates (Bada and Masters Helf- man, 1976), as well as morphology. Thus at the present time, there appears to be little reason for considering the Qafzeh hominids to date significantly earlier than those of Skhtil. Unfortunately, there are no hominid remains attributed to the early Upper Paleolithic of western Asia.

Postcranial evidence. Although supposedly primitive postcranial morphology constituted one of the major arguments for a separate Neandertal lineage (Boule, 1911-1913, 1914, 1923; Vallois, 1954), more recent analyses have shown that while Neandertal postcrania are characterized by extreme rugosity and robusticity, virtually all aspects of Neandertal

SBrauer’s (1980) complete analysis of the Hahnofersand frontal (dated at ca. 36,000 years B.P. ) indicate it to be demonstrably in termediate between Neandertals and modern Homo sapiens in a number of features.


postcranial morphology are within the range of modern human variability (e.g., Trinkaus, 1976b, 1978; Stoner and Trinkaus, 1981). Exceptions are (1) the unusually elongated and thinned superior pubic ramus of Nean- dertals (Stewart, 1960; Trinkaus, 1976a; Smith, 1976b), which possibly relates to par- turition, and (2) the unusual pattern of the axillary border of the scapula, which exhibits a dorsal rather than a ventral axillary groove (the latter being typical of modern hominids). Trinkaus (1977) relates this to different muscular demands, noting that transitional morphologies are found in both Neandertals and modern Homo sapiens and demonstrate that this morphology is the norm in “transitional’’ groups like European Upper Paleoli- thic and Skhd hominids.

THE EVIDENCE FOR A NEANDERTAL PHASE OF MAN

The archeological and morphological information presented above indicate, in our opinion, that a transition between Neander- tals and modern Homo sapiens is even more defendable than in HrdliEka’s day. There are demonstrable morphological continua in the Late Pleistocene hominids of both South- Central Europe (early Neandertals-late Neandertals-early moderns) and the Near East (Neandertals - SkhWQafzeh group) which do not exhibit the extensive morphological breaks we would expect with replacement senso stricto. Late Neandertals in South- Central Europe, for example, make excellent intermediates between the earlier, more “typical” Neandertals and the early modern Homo sapiens sample. The latter sample exhibits numerous well-developed Neandertal- reminiscent features, which occur in rather high frequencies and decrease in both frequency and similarity to Neandertals in later samples. Clearly, the morphological gradient between Neandertals and modern humans which Hrdlizka recognized in 1927 has been supported by subsequent discoveries. Like Hrdlizka, we believe that the most logical (although not the only) interpretation of this gradient is that it represents a phylogenetic continuum between Neandertals and modern humans in these regions.

In western Europe, a temporal/morphological continuum in Late Pleistocene hominids is less clear, and it appears that Neandertals may have survived later in this region than the other two. Furthermore, on the basis of

present evidence, the earliest Upper Paleo- lithic in western Europe seems to appear more recently than in central Europe. All of this may indicate that the appearance of modern Homo sapiens in western Europe was influenced by gene flow from central Europe, but these indications may also be the result of the chronological and other problems noted earlier.

Even if it should be demonstrated that western European Neandertals didFot evolve into modern Homo supiens, Hrdlicka’s arguments are not invalidated. I t is evident that, although Hrdlixka is generally considered to be a unilinealist in the same mold as Schwalbe (see Vallois, 1954, 1958), this is not completely accurate; his views are somewhat more complex (Spencer, 1979). As his writing continuously emphasizes, HrdliEka did not conceive of all Neandertal populations as evolving to modem Homo supiens. Further- more, while not attempting a regional approach, largely because the fossil record was not complete enough at that time to allow such an analysis, it is clear that Hrdlizka recognized regional geography as an integral factor in the evolutionary process (e.g., Hrdlicka 1912a:3-5, 1927:271-272). At any rate, in order to defend the “Neandertal phase of man” in Hrdlizka’s sense, it is not necessary to demonstrate that all Neandertal populations everywhere were ancestral to European modern Homo supiens but only that at least some were. In the light of the above discussion, we obviously believe this can be demonstrated.

THE PRE-SAPIENS AND OTHER RE PLACEMENT HYPOTHESES

In his classic presentation of the pre-sapiens theory, Vallois (1954) noted two major points as evidence favoring this perspective: (1) the lack of morphological evidence of a transition between Neandertals and modern humans (1954:116-119), and (2) the existence of fossil specimens proving the existence of a pre-sapiens lineage (1954:120-125). Having demonstrated that evidence does exist for a transition, we now turn to the question of the existence of a pre-sapiens lineage. Like Boule, Vallois’s pre-sapiens lineage was European and consisted of the specimens from Fontechevade and Swanscombe. More recent analyses (e.g., Brace, 1964; Corruccini, 1975; Trinkaus, 1973; Wolpoff, 1980) have shown that none of these specimens differ significantly from other contemporaneous

V

HRDLICKAS NEANDERTHAL PHASE OF MAN 453

specimens. A more recently suggested European pre-sapiens specimen, the Vertesszollos occipital (Thoma, 1969), has also been demonstrated to be indistinguishable from Homo erectus (Wolpoff, 1971). Thus, there is no convincing evidence to support the existence of a European pre-sapiens lineage.

Outside of Europe, the initial claim of pre- sapiens specimens was registered by L.S.B. Leakey for remains from the Kenyan sites of Kanam and Kanjera discovered in 1932. These have subsequently been excluded as suitable pre-sapiens specimens for differing reasons (see Coon, 1962; Wolpoff, 1980). Also, the possibility of a very early appeareance of modern humans in the Near East was once popular but is now hard to defend, at least in the strict pre-sapienist sense (see previous discussion). More recently, the Niah juvenile (Borneo) has been suggested to indicate the presence of a very modern group in this region at 40,000 years B.P. Apart from the problem of inferring this in a juvenile specimen, there is some question regarding its stratigraphic context, which is difficult to resolve at this point (Kennedy, 1979). Furthermore, no other indications of this early occurrence of modern humans have been recovered from any other East Asian site. Additionally, we do not consider a 40,000-year date to be significantly earlier than the earliest appearance of modern Homo sapiens in Europe or the Near East.

Presently, the best case for a very early date for anatomically modern humans comes from South Africa, specifically the site of Border Cave (Beaumont et al., 1978; Rightmire, 1979). Among other remains, a very modern-looking cranium and separate mandible have been recovered from this site, and are said to come from levels dating in excess of 80,000 years B.P. (see Butzer, 1978). There are, however, contextual problems with these specimens; and the existence of modern humans at so early a date has not yet been conclusively demonstrated at other sites in southern Africa, with the possible exception of the Klasies River Mouth sites (see Klein, 1977). Otherwise, there is a reasonably clear morphological sequence in South Africa (Rightmire, 1976, 1978) in which evolutionary grades appear equivalent to those contemporary hominids in other areas of the Old World (see Wolpoff, 1980). Thus, while we view the possibility with great interest, we feel that considerably more substantiation is necessary before the idea that South Africa is the area of

origin for modern Homo sapiens, in a classic monocentric sense (Beaumont et al., 1978; Protsch, 1975), can be accepted. In fact, as the evidence now stands, although clade differences certainly exist on a broad geographic basis, grade differences appear to be minimal at any particular time during the Middle and Late Pleistocene of the Old World (Coon, 1962; Wolpoff, 1980).

The pre-neandertal hypothesis is probably the most popular perspective on the phylogenetic position of Neandertals at the present time. According to this viewpoint, modern Homo sapiens evolved out of “progressive” Neandertal or other archaic Homo sapiens populations outside of Europe and subsequently influenced the origin of modern Homo sapiens in Europe. This influence is generally considered to have been derived from the Near East and to have taken one of two forms: (1) an influx of fully modern Homo sapiens into Europe, probably with some assimilation of Neandertal genes into the invader’s gene pool, or (2) extensive gene flow, with “progressive” genes being introduced into Europe, but without a significant influx of modern peoples themselves. As has been demonstrated, the available evidence provides virtually no support for the first possibility. Regarding the second proposition, it is our considered opinion that, because paleontological models are presently inadequate to deal effectively with gene flow, it is simply not possible at this juncture to judge what the precise effects gene flow (without major population movements) may have had on the transition from Neandertals to modern Homo sapiens in Europe (see Weiss and Maruyama, 1976). From all indications, there were no impassable geographical barriers preventing populational contacts within Europe and between Europe and any adjacent region; thus, it is quite likely that gene flow may have played an important role. On the other hand, there is no a priori reason why the transition in South-Central Europe, for example, would have necessarily been influenced by the influx of “progressive” alleles t o any greater extent than the transition in other regions, including the Near East, was influenced by the influx of “progressive” alleles from South-Central Europe. Similarly, while it is tempting to suggest that the origins of modern Homo sapiens in western Europe was strongly influenced by gene flow from central Europe, such an assertion requires further clarification of the nature of Neandertaliearly


modern Homo sapiens relationships in the former area of Europe.

In light of the evidence presently available, we believe that HrdliEka’s major point of criticism regarding the replacement hypothesis continues to be valid. There is still no conclusive evidence for the existence of a pre- sapiens lineage separate from and contemporary with archaic Homo sapiens. This is true both within Europe and in the remainder of the Old World. Furthermore, though southern Africa may prove to be an exception, we do not see convincing chronological evidence that modern Homo sapiens appears significantly earlier in any portion of the world than in Europe. Thus the source area for modern Homo sapiens necessary for any strict replacement interpretation regarding the origin of modern Europeans has yet to be identified.g

MONOCENTRISM AND POLYCENTRISM

There are some important differences in how Alex Hrdliska viewed his Neandertal phase of man and what the existing fossil record indicates, Hrdlixka (1921b, 1927, 1930) was clearly espousing a strict monocentric perspective on the origin of Homo sapiens. In his opinion Europe had been the “cradle and nursery” of early modern Homo sapiens. In our opinion, the existing human fossil record does not support this or any other strictly monocentric concept of the origin of modern Homo sapiens. To the contrary, it appears to support the idea of several, regionally somewhat distinct and apparently contemporary, origins of modern Homo sapiens-grade hominids - ana- logous to the ideas presented by Weidenreich (1947, 1949) and Coon (1962). However, there are certain theoretical problems which make it very difficult to accept the idea of a strict polycentric origin for modern Homo sapiens (see Howells, 1976).

Wolpoff (1980:298-300) has suggested that the changes necessary for the archaic-to-modern Homo sapiens transition are essentially the same throughout the Old World and “can be understood and documented in terms of worldwide changes in selection acting on already differential local populations of archaic Homo sapiens.” Furthermore, he notes that the “explanation for these developments relates to the consequences of the last series of cultural and technological changes that were truly worldwide (before the industrial age): the appearance of Middle Paleolithic industries and the development of the prepared-core tech-

nique” (Wolpoff, 1980:299). Basically, his argument is that these innovations changed the orientation toward the use of the anterior teeth for nonmasticatory and paramasticatory purposes. The resulting anterior dental reduction and decreased degrees of stress generated in the face and cranial vault in turn brought about the morphological changes marking the emergence of a modern Homo sapiens grade worldwide. This is a more complex restatement of Hrdlixka’s (1911) suggestion that changes in “masticatory function” was an integral function in later patterns of human cranial evolution. This idea was subsequently developed by Brace (196213, see also 1979) and later espoused in various forms by several other workers (e.g., Brose and Wolpoff, 1971; Frayer, 1978; Smith, 1976b, 1978).

SUMMARY AND CONCLUDING REMARKS

In this pa er we have taken the position that

phase in human evolution is strongly supportable. I t is contended that the existence of a Ne- andertal phase is supported by the following evidence: (1) the presence of a morphological continuum in appropriate chronological frame- works for both Europe and the Near East between archaic Homo sapiens (Neandertals) and early modern Homo sapiens; (2) the lack of either any evidence for a pre-sapiens lineage in Europe or the unequivocal appearance of modern Homo sapiens anywhere in the Old World at any appreciably earlier time than in Europe; (3) the indications that the Aurignacian assemblage can be logically considered an indigenous European phenomenon.

Furthermore, we have hopefully shown that Hrdlizka’s concept was not a simple intel- lectual outgrowth of earlier unilineal schemes but in fact was more complex, involving the idea of differential survival of Neandertal groups. A defense of Hrdlizka’s position, therefore, does not require proving that all Nean- dertal “populations” everywhere evolved into modern Homo sapiens. In the light of the available evidence from both central Europe and the Near East we feel that Hrdlizka’s proposition is much more supportable now than it was in 1927.

Alex Hrdli P ka’s basic idea of a Neandertal

gEven if a very early presence of modern Homo sapiens is demonstrated in southern Africa or elsewhere, this does not necessarily mean that such a phenomenon would have had an effect on Late Pleistocene hominid succession in Europe. In a strict replacementist scheme (i.e., population replacement), it would also be necessary to document the movement of such populations toward and into Europe as well as their morphological similarity to European modern Homo sapiens.

H R D L ~ K A ’ S NEANDERTHAL PHASE OF MAN 455

We trust that the importance of the Nean- dertal phase to Hrdlizka’s general conception of the peopling of the world and his beliefs concerning the antiquity of man in the New World is evident. His ideas on these and other issues are often considered as if they were totally independent segments of his thinking, which is clearly not the case. When one appreciates this fact, it is that much easier to understand why he doggedly defended some of the things he did (e.g., his insistence that “Sinanthropus” was a Neandertal). Furthermore, Hrdlixka is often criticized for such concepts as his “morphological dating.” As we demonstrated, this was a necessary and pragmatic solution to the obstacle of imprecise methodology which often made the question unanswerable on strictly geological grounds. Besides, it made sense in terms of his concept of the evolutionary process (which was quite accurate for its day); and it worked well, in his experience, with those hominid remains whose antiquity was demonstrable by other means. What is more, an analysis of the evidence will show he was invariably right!

To what extent the anticipated discoveries of the next 50 .gears will change our assessment of Hrdlicka’s Neandertal phase of human evolution remains to be seen. But whatever the outcome we are confident that future generations of American physical anthropologists will continue to regard his contributions with respect and admiration.

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Documents

The significance of Aleš Hrdlička's “Neanderthal phase of man”: A historical and current assessment