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Part Three

In the thirty-five years following the discovery of thebones of Pithecanthropus the discipline of paleoanthropology grew into a mature program of research marked byan increase in discoveries of fossil hominid specimens,the development of new techniques for analyzing thefossils, the establishment of research centers for humanpaleontology, the teaching of university courses on thissubject, the formulation of new theories to interpret thefossil record, and an increa sing awareness among scholarsthat the discipline of genetics could shed light on theproblems of the origin of man and his history. A majorfeature in this coming of age of paleoanthropology was notonly the discovery of fossil hominids in deposits beyondthe borders of Europe, bu t also the realization that theEuropean, Asiatic, and African fossil specimens nowknown were of varieties that could not be classified withinthe narrow dichotomy of the Neanderthal or Cro- Magnantypes.The realization that ancient man might be discovered inAfrica, Asia, and Australia was initiated by Dubois'sdiscovery in 1890-91. From a fluvial deposit at Trinil inJava Dubois recovered a skullcap and long bones ofa hominid that could not be matched by any specimenin the European fossil record at that time. At the meeting ofthe Royal Dublin Society in 1895 the discoverer stressedthat his Pithecanthropus erectus occupied a phylogeneticposition between the anthropoid and human stems, asHaeckel had conceived of his hypothetical Pithecan-thropus. Immediately differences of interpretation aroseover the question of whether this specimen was an ape,a transitional ape-man form, a Homo sapiens, or at least a

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T90 Part Three / 1890 to 1925 Part Three / 1890 to 1925 191creature more closely akin to man than to the anthropoids. Had Dubois notbeen offended by his colleagues' speculations and by the hesitancy withwhich his fossil man was received by some of them, he might have broughtforth additional specimens which he had discovered in Southeast Asia.Instead he did not reveal the mandibular fragment from Kedung Brubusuntil 1924, although it had been found twenty-six years earlier, and thefour femora that he had recovered from the Trinil beds in 1900 did notcome to scientific attention until 1932. The bones from Wadjak, also inJava, were resurrected in 1921 throug h the intercession of the Americananthropologist, Ales Hrdlicka.

That early man had a wider range of phenotypic variation than could berepresented by the Neanderthal and Cro-Magnon types of fossils found inEurope was supported by new evidence from Asia, Africa, and Australia,and especially by the discovery of the Heidelberg mandible from Germany.The type of hominid represented by Pithecanthropus had not been foundoutside the island of Java during this period of paleoanthropologicalresearch. But the morphological affinities of the European Neanderthalspecimens with the Broken Hill fossil from Northern Rhodesia suggestedthat here was a fossil form with a wider geographical distribution. Itis curious to note that during this period when the fossil record for Europewas being considerably amplified, morphological variation for the Neanderthal specimens was hardly admitted, whereas for the later Cro- Magnonfossils the paleoanthropologists were eager to isolate sub-types andracial entities on the basis of individual specimens. Of the Neanderthalspecimens from La Ferrassie, Le Moustier, La Quina, and La Chapel eaux-Saintes it was the complete male specimen from La Chapelle-auxSaintes that was set up by Boule as the prototype of Neanderthal Man.The other Neanderthal specimens were better or worse examples of thistype of ancient humanity in accordance with the degree of their resemblanceto this ideal form. Among the Cro- Magnon specimens thus far discovered,Verneau claimed the male and female burial at the Grotte des Enfants asindicative of a negroid racial element in this population, just as the specimenfrom Combe Capelle was accorded a sub-specific classification by Klaatsch.With the eskimoid specimen from Chancelade and the Cro- Magnonspecimen proper, the Upper Paleolithic hominids of Europe became dividedfurther into racial sub-types while the Neanderthals tended to be lumpedinto a single phenotype.

While the discovery of Pithecanthropus provided paleoanthropologistswith a hominid fossil that seemed similar in some of its physical featuresto the anthropoid apes, it was recognized nevertheless as being a hominidvery different from Haeckel's Pithecanthropus alalus, that medley of simianand human physical conditions. Scientists continued to be embarrassedfor the lack of a convincing missing link. Then, among the discoveries ofthis period, came a series of remarkable finds from Piltdown and SheffieldPark in Sussex between 1908 and 1915. Here was a fossil to jar the anti-

evolutionists out of their complacency and scepticism about the possibilityof discovering missing links. Piltdown Man, or Eoanthropus dawsoni,sported a modern-like skullcap and a simian mandible. Was he not the truerepresentative of that transitional form between man and ape? This happydiscovery seemed at first to answer many of the needs of those paleoanthropologists harassed by embarassing questions. It was remarkable thatthe different anatomical portions of this fossil turned up in deposits atPiltdown and Sheffield Park in an order that paralleled chronologically thesuccession of questions asked about the specimen by interested scholars.In the light of our present knowledge that Piltdown was a fraud, it has beensuggested that the hoax was a sincere attempt on the part of its manufacturer to hasten the day when human evolution could no longer be amatter of doubt. Interestingly, the Eoanthropus specimen had outlived itsservice some quarter of a century before its fraudulency was ascertained,and the fossil was frequently described in the textbooks and scientificpapers of the early twenties of this century as an aberrant type of hominidwhose very nature precluded its serious consideration in the constructionof man's phylogen etic tree.

How were the differences in the physical type of early man and thegreat geographical dispersions of some of these types to be explained interms of human phylogeny? To answer this question the early twentiethcentury paleoanthropologists needed to know two things: the relative agesof the specimens that they considered the prototypes of the different kindsof early men, and the loci of human evolution from whence these typesbegan their adaptive radiation.The dating by geological stratification and faunal association of thedeposits at Trinil and Heidelberg demonstrated that the antiquity of mancould be extended as far back as the Mid- Pleistocene. Dubois's datingof the Pithecanthropus bones to the Early Pleistocene or Pliocene wasrejected by the members of the Selenka expedition who visited Trinil in1906, but reports from other regions of discoveries of Tertiary Man werestill current. However, during this period the claims for Tertiary Man werenot as enthusiastically received as they had been in the earlier part of thesecond half of the nineteenth century. Ameghino's assertions of a SouthAmerican origin for man, which were based on his studies of osteologicalseries of New World monkeys, were appreciated only by a limited numberof scholars in Europe. By 1927 the revelation that a supposed earlyAmerican hominid progenitor, Hesperopithecus haroldcooh was nothingbut the tooth of an extinct peccary did much to undermine the prestige ofclaims for the fossils of Tertiary Man, especially in the New World. Supportfor this notion was stronger in the field of prehistoric archaeology, where itwas asserted that pre- Paleolithic tools called "eo\iths" were to be found inTertiary deposits in western Europe and India. The eolithic controversyof the 1890s had no sooner subsided when Moir revived interest in theso-called rostrocariante tools from the Pliocene beds of East Anglia. The

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eolithic and rostrocarinate controversies became less intense when itwas realized that such "tools" could be produced by natural agents,for example water activity working with an abrasive agent on stones. Itcame to be recognized that eoliths were not confined to Tertiary deposits,that they were found in obvious pre-hominid deposits, and that they werenever found in regions where the materials of which they were composedwere not already present.

The venerable idea that man had originated somewhere in Asia nowseemed confirmed by the discovery of Pithecanthropus. This thesis foundfurther support when in the Siwalik Hills of northern India the fossils ofMiocene and Pliocene apes were uncovered by Pilgrim. Some of thesefossils were immediately recognized as possessing morphological featuressimilar to those represented in the hominid stem, various scholars favoringone fossil anthropoid over another as nearest to man. Although fossilanthropoids from the Tertiary were being found in Egypt at this time, theidea that Africa might be the birthplace of man, as Darwin had earliersuggested, was overshadowed by opinions that favored Asia as the placeof human beginnings, as seen in the writings of Richard Swan Lull.

During this period the major theories of human phylogeny becamedefined. Many interpretations of the fossil human record appearing before1925 persisted to the present, but with some striking modifications. Sometenets that once seemed unassailable have had to be abandoned. Thuswhile the thesis of a catarrhine origin for the human stem was not entirelyrejected, the discovery of fossil primates from India and Egypt lent muchsupport to the hypothesis of a pongid ancestry for man. Gregory was atthis time the recognized American authority on the comparative anatomyof the higher primates. On the basis of similarities of dental form betweenthe fossil apes and the fossil men then known he championed Dryopitheeusas nearest to the human stem. Pilgrim also accepted an ape ancestry forman, and he saw osteological and dental features in the fossil ape Sivapitheeus that led him to hail it as the Miocene progenitor. Indeed, he placedSivapithecus in the family Hominidae with man, the other large anthropoidsfalling under the family Simiidae. In reaction to these concepts of ananthropoid human ancestor, several anatomists argued that the humanline had its origins in primate forms much lower on the taxonomic scale.Thus Wood-Jones suggested that the distinctively human line originated atthe beginning of the Tertiary without ever having been directly connectedwith the anthropoid stem. Rather, man had evolved from a tarsioid stoc kwhose only living representative today is the small arboreal Tarsius ofMadagascar. Wood-Jones admitted that no fossil form of such a humanancestor was yet known, bu t he based his thesis on data from comparativeanatomy and embryology. Tate Regan, on the other hand, placed Tarsiusand Homo at the ends of two lines that began to diverge in Cretaceoustimes. Others students of primate phylogency asserted that the nonhumanprogenitor of man was to be found in the lemuroid line or among the

Part Three /1890 to 1925

platyrrhines of South America, theories that never found the same degreeof acceptance that was accorded the anthropoid ape and tarsioid hypotheses. The writings of Boule kept alive the notion that man's closestrelatives in the phylogenetic tree were to be sought among the catarrhines.Finally, there is the bizarre notion of the German prehistorian HermannKlaatsch, who believed that the human species is older than either theanthropoid or the infra-anthropoid lines, the apes being degenerate formsof humanity. Klaatsch maintained that the human stem arose from a nowextinct being which was already human rather than simian, and that afterit had reached the evolutionary stage represented by Pithecanthropusit ramified into various primate forms which continued to evolve as theliving races of man as well as the living kinds of great apes. Klaatschreflected certain eighteenth century ideas that apes were degenerateforms of humanity, founding his interpretation on the premise that thehuman body is more primitive in its structure than that of any of the apes,the latter being specialized deviants pointing in a direction of evolutiondifferent from that of the more generalized catarrhine and human stocks.Klaatsch was the chief representative of the Polyphyletic School duringthis period.By the early years of the present century the fossil hominid record hadbeen amplified to such a degree that it was being asked if Homo sapienscould be descended from all of the earlier men of the Pleistocene thus fardiscovered, or was our species descended from some or perhaps fromnone of these fossils, arising as still another line whose members might yetbe undiscovered? If Homo sapiens could be shown to have followed aseparate evolutionary line, the antiquity of this line could be determinedindependently of the estimates of the antiquity of the lines for Pithecanthropus, Neanderthal, and related fossil hominid forms. During this periodthere appeared to be some evidence that Homo sapiens was more ancientthan formerly supposed on the basis of putative Homo sapiens fossil speci-mens from sites that could be dated as earlier than Upper Pleistocene.Eoanthropus had already been interp reted as a hominid of great antiquity;its Homo sapiens type of cranium was the principle evidence for thisdecision. Furthermore there was the hesitancy to admit the implications ofwhat man's brute ancestry might have been, an intellectual inhibition thatwas no less foreign to the sensitivities of dedicated paleoanthropologistsof this period than it was to their non-scientificallyoriented contemporaries.A separate human line afforded that psychological buffer to the humanego which still found it distasteful to admit the meanness of man's origins.But the presence of certain sapiens-like hominid fossils in geologicaldeposits then described as Second Interglacial or earlier was the mostsignificant factor in the query as to whether Homo sapiens might not haveevolved earlier than the followers of the Unilinear School had supposed.If the sapiens line was separate from that which included the Neanderthaland Pithecanthropus forms, then these more primitive hominids could be

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195Part Three / 1WIO to 1925 Dubois / Place of Pithecanthropus in the Genealogic;d TreeI

related only indirectly to modern man. Theirs were phylogenetic lines thatwithered, while the sapiens line flourished and survived. Sir Arthur Keithwas the leading voice in support of this "Presapiens School" and the firstto draw up a phylogenetic tree to represent the branching off of the Homosapiens from those stems leading to other fossil hominids. Keith differs in hisearly works from later adherents of the Presapiens School, for he did not setapart particular fossil specimens as representative of direct human ancestors of modern man. Also he relegated Eoanthropus, in spite of its Homosapiens skullcap, to a separate evolutionary stem that had become extinct.

The Presapiens School arose in part because of the dissatisfaction ofmany early twentieth century scholars with the tenets of the Polyphyleticand Unilinear Schools which had been inherited by the discipline fromthe writings of the previous century. Nor was the Presapiens School theonly result of this need to assess the data in a new way in the light of thecurrent amplification of the hominid fossil record. The Unilinear Schoolhas sometimes been called the Neanderthal School because of its tenetthat the Neanderthal specimens marked a necessary link in the evolutionarysequence of types that led to Homo sapiens. This Neanderthalism, orpan-Neanderthal theory, was favored by Dubois, wh o placed between hisPithecanthropus and modern man the Neanderthal specimens thenknown. Grafton Elliot Smith agreed that Homo sapiens shows a numberof neanderthaloid traits and that our species of the Late Pleistocenefollowed in time the European Neanderthals. But Smith differed from hiscolleagues in the Unilinear School by his interpretation of the popularbelief that while all living races today are members of the same genus andspecies, certain races are more primitive than others. Smith saw theseprimitive groups as the less evolved representatives of the Homo sapiensbranch that departed from the main human stem at a period in thePlesistocene prior to the continuation of the Neanderthals into their blindalley of evolution, marked by the "classic" or extremely robust Neanderthalsfound in western Europe and in the Broken Hill site of Northern Rhodesia.Nor did Smith accept Pithecanthropus as the ancestor of NeanderthalMan. This Javanese fossil appeared to him too specialized to be on thatmain human stem leading from the Miocene apes up to the "Nordicracial group" of modern man. It was primarily Smith's thesis that Homosapiens became divorced from the Neanderthals and evolved separately atsome period of the Third Interglacial that became the basic premise of the"Preneanderthal SchooL" The supporters of this theory differ from thePresapiens adherents mainly by placing the origin of the Homo sapiensline later in time-namely, within the period of the Third Interglacialthan do their colleagues, who would place this time of species formationat the Second Interglacial or earlier.

By the first quarter of the present century these four ways of interpretingthe fossil record had been defined. The results of prehistoric researchessuggested that Europe could not be the cradle of human culture any more

than it was the home of man's biological progenitors. If only a portion ofthe prehistoric cultural patterns of the Old World were within the Europeanarea, then hominid progenitors might also be forthcoming from Africaand Asia. Yet in 1924 when the first Austra/opithecus was discovered,very few scientists were aware of its significant role in future interpretationsof the story of human evolution.

Th e Place of Pithecanthropus in theGenealogical TreeMarie Eugene Francois Thomas Dubois1896

the accompa nying diagr am, repreIn the report on the scit'ntific meet sellting the t'volution of the Ol ding of the Royal Dublin Socit'ty \Vodd apes frOln a hypothrtic;ll comon November 20, in Nature of mon ancestor, whom I call ProcerDecember ;), 189:1, it is statrd thatI placed Pithecanthropus in the copithecus.In Prof. Cunningham"s tree,genealogical tree, drawn by Prof. figured in Nature of December 5, p.Cunningham, below the point of 116, he rt'gards the left branch as alldivarication of the Anthropoid apes human, the right one as entirelyfrom the human line. This indeed 1 simian, an d he placed Pithecanthrodid. But this statement could be pus midway betwern recent ~ l a nmisleading as to my real views on an d the point of divarication. Nowthe grnealogy of Pithecanthropus, 1 could find no place for the fossilsuch as I stated thrIll already on p.

Jayanese form, which I consider as38 of Illy original memoir ("J'ithe- intermediate between ~ I a n an d Ancanthrof!uS erect l iS, Eine TIIenschen- thropoid apt's, many of the branchesahnliche Cebt'rgansform aus Java," of that tree. only in the third chirfBatavia, lWH), and more fully at the lint', tht' main stem, \Try nt'ar to tht'last mt'eting of the Anthropological point of diyaricatioll.Institute of Creat Britain an d Ire Owing to tht' same eircums(;lnces,land, on Novembn 2J. which indirectly prevented me fromI t Illay not be superfluous to explaining my ow n yiews on theexplain my views here by means of"The Place of 'l'ithecanthrofJUs' in the Genealogical Tree" Nature. Vol. 5:;,pp. 245-47. 1896.