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Home , Franz Weidenreich, Julian Huxley, Rudolf Virchow

AMERICAN JOURNAL OF PHYSICAL 56:411-429(1981)

Tales of the Phylogenetic Woods: The and Significance of Evolutionary Trees C. LORING BRACE Museum of Anthropology, University of Michigan, Ann Arbor, Michigan 481 09 KEY WORDS Classification, Phylogeny, Trees ABSTRACT The styles of continuing intellectual traditions can have a major effect on the way in which scientific findings are expressed. Darwin and Huxley, for all their intellectual daring followed the skeptical tactics of the Scottish Enlightenment and avoided the construction of phylogenetic trees, even though they were aware of the evidence on which such could have been con- structed. The romantic evolutionism of Haeckel, Keith, and many subsequent writers in English produced suggested phylogenies on the basis of largely hypothetical forms including Homo “alalus,” “stupidus,” and “Eoanthropus.” The structural aspects of phylogenetic schemes that derive from the French in- tellectual ethos, from catastrophism to and punctuated equilibria, have stressed discrete categorical entities in the tradition of Platonic essentialism and have tended to avoid a consideration of evolutionary dynamics.

Intellectual traditions frequently shape the The Comparative Reception of way in which scientific questions are posed as (Glick [ed.], 1974) admirably documents the well as the procedures undertaken to answer fact that there are indeed different national them. When I attempted to provide a paleoan- styles of thinking when it comes to dealing thropological illustration of this realization with major aspects of science. If this has been some 15 years ago (Brace, 1964), one of the true for the treatment of the interpretation of commentators phrased the rhetorical query, organic evolution in general, it has also been “Since when in science can one base oneself on true for approaches to the study of human arguments of nationality?” (Genoves, 196423). evolution in particular. As this paper will at- This objection was raised in regard to issues tempt to show, this is graphically illustrated relating to the phylogenetic treatment of a par- by the various forms that are offered as human ticular set of human fossils, and, from the phylogenetic trees. perspective of the history of science, a relative- ly minor matter. The proper response, had I NINETEENTH CENTURY ENGLAND been adequately prepared, would have been to Verbal portrayals of human descent in the note the reaction of a figure of unimpeachable form of a tree are present in the very earliest stature to the various ways in which a truly written records and are commonly found in cul- major scientific synthesis was perceived. The tures that lack a system of writing. Further, best example is the reaction of none other than many cultures contain accounts of the kinship to the reception of his theory between and particular members of of evolution by means of . In a the animal kingdom, although these are usual- letter he wrote to the French anthropologist ly expressed in symbolic and totemic form and Armand de Quatrefages, he said, “It is curious rarely, if ever, rendered as identifiable parts of how nationality influences opinion; a week a literal family tree (cf. treatments by Frazer, hardly passes without my hearing of some 1887; Freud, 1950; Levi-Strauss, 1962). While naturalist in who supports my views, there were occasional earlier attempts to por- and often puts an exaggerated value on my tray a more than biblical human antiquity and works; whilst in France I have not heard of a a putative lineal kinship with nonhuman an- single zoologist, except M. Gaudry (and he on- ly partially) who supports my views” (in F. Dar- Delivered at the University of Virginia, Charlottesville. December win [ed.], 1887:299). 11-12, 1980.

0002-948318115604-0411$05.50 0 1981 ALAN R. LISS, INC. 412 C.L. BRACE cestors (see the treatments in Eiseley, 1958; of the Scottish Enlightenment (Gruber, Greene, 1959; Osborn, 1894), it is clear from 1974:39, 80-81). And, for whatever historical the record that a systematic concern for the reason, one of the most penetrating observers course and forms of human ancestral develop- of the history of science has noted that “Hux- ment did not begin until after the publication ley’s personal creed was a kind of scientific in 1859 of that extraordinary scientific land- Calvinism” (de Beer, 1970:917). mark, , by Charles Not only did they exemplify the application Darwin. of the Protestant Ethic in the realm of science To be sure, despite popular assumptions to (Merton, 1938, esp. pp. 415-419), but, as has the contrary (Wilberforce, 1860:135; Broca, been noted for their British contemporaries in 1862:314),Darwin only alluded to the possibili- the physical sciences (Olson, 1975), the par- ty that his approach could be applied toward ticular style displayed was very much that of an analysis of the human condition barely the Scottish Philosophy of Common Sense. If more than two paragraphs from the end of his the naive extremes of the inductive empiricism epoch-makingwork when he wrote, “Light will of Bacon and Locke were avoided, nonetheless be thrown on the origin of man and his history” there was a faith that the powers of human (1859:488). In subsequent editions, this was reason were sufficient to discover the of expanded only to the extent that he said the world, whose structure was assumed to be “Much light will be thrown. . . .” (cf. 1872504). logical. And if they recoiled from the extremes If this seems in retrospect to be brief to the of skepticism of a David Hume, and feared the point of being cryptic, the implications were related position of “materialism,” at the same not lost on Darwin’s contemporaries despite time they displayed an elaborate caution when the fact that he himself waited a dozen years it came to considering anything that could be before developing the theme in The Descent of regarded as a possible projection of the ideals Man (1871). While Darwin proceeded with whose loci are primarily in the human mind. deliberate caution, others were quicker to Given the nature of the available evidence and follow the path to which he had pointed. Nor the tenor of , it is hardly surprising did these accounts display signs of unseemly that the examplars of British science in the lat- haste. The evidence for human antiquity pre- ter part of the nineteenth century refused to sented by Sir (1863)was the pro- speculate on the possible specifics of a human duct of several decades of meticulous fact col- line of descent. lecting. Likewise the demonstration of the biological affinities of human with particular FRANCE nonhuman form by in If the English were slow to suggest schemes Evidence as to Man’s Place in Nature (1863) for human phylogeny, the French were even was a masterful synthesis based upon an enor- slower. For one thing, even among the few who mous amount of information. accepted the possibility of organic evolution, a Curiously, although Huxley, Lyell, and later Darwinian form of mechanism was explicitly Darwin (1871) dealt with the prehistoric rejected (Gaudry, 1878:250, 257; Topinard, skeletal material known at that time, including 1888:473; Stebbins, 1974:138, 164). Further- the original Neandertal remains, they all avoid- more, the effective founder of the field of an- ed the gambit of arranging them in hypotheti- thropology in France, Pierre Paul Broca, was cal lineages purporting to show the course of an avowed polygenist and vocally unsym- . That they should eschew pathetic to a Darwinian approach (Broca, 1870, such a step seems an odd bit of caution on the 1872; Schiller, 1979:226, 232). part of the authors whose works embody some By this time, of course, the whole field of of the most daring innovation of the entire vertebrate had been in existence nineteenth century. In fact, however, in spite for more than a generation following its crea- of their very different personalities, all three tion by Georges Cuvier by the beginning of the displayed an intellectual style that came century (Simpson, 1961:43, Coleman, 1964:2, relatively directly from the ethos of the Scot- 114).As the field of human paleontology devel- tish Enlightenment. Lyell, as intellectual heir oped in France late in the nineteenth and early of Hutton and Playfair, was born and raised in in the twentieth centuries, it adopted an Scotland (Eiseley, 1959; Wilson [ed.], 1970; outlook that was rooted in the catastrophism Wilson, 1972). Darwin’s introduction to both that characterized its older model (Brace, 1964, natural science fieldwork and biological theory 1966).Because of the quantity of physical evi- was in Edinburg just before the effective end dence for the form of prehistoric human ap- PHYLOGENETIC TREES 413 pearance found in France late in the last and viewed as proceeding separately in the form of early in the present centuries, and because of a “bush (Vallois, 19525‘8-79; see Fig. 9; the interaction of European political and in- Thoma, 1973; Gould, 1976; de Benoist, 1979). tellectual history, an essentially French view- The same analogy has been used by French ar- point has played a very important role in shap- chaeologists to depict the course of prehistoric ing the enduring traditions in the field of paleo- cultural development (Bordes, 1950). It is in- anthropolugy . teresting to recall that Cuvier himself insisted As a consequence of this, much of the field, that “ was a bush, not a ladder” (Eiseley, especially in France, has continued to show an 1958:88), although in that instance he was op- orientation that has been either covertly op- posing the formulations of those who were at- posed to or silent regarding the depiction of tempting to defend an integrated and hierar- schemes that try to arrange the known fossil chical scala naturae. record into an evolutionary sequence. Even Finally one must note that a kind of mystical when ultimate human origins are conceded to evolutionism was promoted in a French con- be “monophyletic,”the depiction of subsequent text even though it had no positive impact on human development is in a “polyphyletic the actual study of the course of human evolu- mode” where the lines of development are tion. This was initiated by in f

Upper Pleistocene

XXXXXXXXXXXMXX Homo neandert halensis

Middle Pleistocene ? xXXXXXxxxxxxxxx 1 Pithecanthropus erectus

Aus trabpithecus africanus Lower Pleistocene

Fig. 9. A version of the polyphyletic formulation favored by French anthropologists,adapted from Vallois 11952:77)with the addition of Australopithecus. All nonmodern fossil hominids are considered to have become extinct without descendants (from Brace et al., 1979:168). 414 C.L. BRACE his Evolution Creatrice (1907) and continued gloating, after having outlived Virchow, when in Teilhard de Chardin, especially his posthum- he felt that Dubois’ discoveries and ous Le PhSnomkne Humain (1955).Lovejoy’s Schwalbe’s interpretations proved that Vir- appraisal of Bergson was written before chow had been wrong (Schwalbe, 1901; Teilhard’s works were published, but it could Haeckel, 1906:108-109). There is more to the serve to characterize them as well. He noted story, however, and the point of adding it is to that the central insight of such a philosophy demonstrate how matters of distinctly nonsci- was “a thing to be reached, not through a con- entific nature can have a bearing on scientific secutive progress of thought guided by the or- assessment. dinary logic available to every man, but Eughne Dubois received his training in Ger- through a sudden leap whereby one rises to a many and in 1880 was an assistant at the in- plane of insight different in its principles from stitute of anatomy at Jena just at the time that the level of mere understanding” (Lovejoy, Haeckel’s influence was on the rise. It was 1936:ll-12). As Medawar observed in regard there that Dubois picked up his interest in to Teilhard, this “stands square in the tradition evolution and the orientation that he was later of (German) Naturphilosophie” (Medawar, to try to apply to the material he collected in 1967:72), but although he regarded much of it Java (Weinert, 1947:33). It is interesting to as “nonsense,”he added, “on further reflection I note that Dubois first referred to the famous see it as a dotty euphoristic kind of nonsense Trinil discovery as “Anthropopithecus,”(l891), very greatly preferable to solemn long-faced the term Haeckel used for the chimpanzee. germanic nonsense” (1967:9).In any event it When he found the femur, he kept the same contributed nothing to interpretations of the generic designation although he changed the hominid fossil record. species from “troglodytes”to erectus (1892).IR his full published report he recognized that he HAECKEL AND GERMAN had more than a chimpanzee, and it was then ROMANTIC EVOLUTIONISM that he used Haeckel’s generic term for The first formal phylogenetic tree purport- “apeman” and called it “Ptihecanthropuserec- ing to depict the course of human evolution tus” (1894). He presented substantially the was the creation of that extraordinary scion of same views at the third International Con- German Naturphilosophie, . The gress of at Leyden in 1895 (Dubois, prehistoric portions of this structure, at least 1896). in its earliest manifestations, were based on Then, as has often been noted, he remained hypothetical constructs. The first of these was silent on the matter for more than 20 years and “Pithecanthropus” which he included without refused to let visitors see his material (Spencer, comment in his system of the mammals in the 1979:415-420). Finally, when he did resume second volume of his Generelle Morphologie consideration of it, he had undergone a radical (1866:CLX). At the same time he also sug- change of mind and regarded his “Pithecan- gested the possibility that the term “Erecta” thropus” not as a transition form but as a giant could be used as a taxonomic designation for extinct gibbon (Dubois, 1932, 1935). Some of the human family. In later discussions he add- the accounts tend to lead the reader to suspect ed the form “alalus”to serve for “speechless that Dubois had become a little “queer” men” while he used “Pithecanthropus”to repre- (Wendt, 1956:299), but there are some other sent “ape-men”(1870:590, 597). By the end of things that may have been involved. For one the century, “alalus” was demoted to become thing, the interpretation that regarded his the species name for this hypothetical fossil as a giant gibbon had in fact been offered “Pithecanthropus” and this in turn was con- by none other than Virchow at that Leyden sidered to be ancestral to Homo sapiens via the Congress in 1895 (Virchow, 1895:746-747). further hypothetical form of Homo “stupidus” Even though Dubois later claimed that he got (1899:35; see Figure 10). the idea from the first edition of Boule’s Les All of this is relatively well known and con- Hommes Fossiles (Boule, 1921:109; Dubois, sidered relevant only insofar as it contributed 1935:583), it would appear that it was the in- the name which the Dutch physician Dubois fluence of Virchow that was of principal impor- later gave to the material he found in Java be- tance (Ackerknecht, 1953903). tween 1890 and 1892. Also well known are the The fact that Dubois switched from the different positions which Haeckel and Virchow Haeckel-orientedviews of his youth to those of took in regard to the interpretation of the Virchow, Haeckel’s long-time rival, brings us original Neandertal skeleton and Haeckel’s back to the nature of their disagreement. An PHYLOGENETIC TREES 415

- Anthropomorph~ Anthropini Anthropoiden rfricrnrr c Homo Anthropoiden uhtiou Authropithecus - sapieus aorilla schimpause gina

Pliopithecus antiquus

Plstyrrhinae Dynmopitheor Oynopithtor Mycetes Semuopithecus Atelea Cebus atavus Cercopithecns

Cynocephslida Hnpalida

Lemaravids Lemurogona Rorimiu gmrral*fas Rommk pialbt~ Archipithecus ’ Necrolemures Chirolemure8 . Anaptomorpha Tarsolemures (ChiromY8) Adapida \ 1 (Tarsius)

ant ole mu re^

Lemuravida iU a Prohylemurn [C a r n a LI LI i a]

Archiprimas Proohorieta

Fig. 10. The Primate family tree according to Haeckel(1899:35). older reading of the history of science sug- unintended and relatively inconsequential by- gested that the issue was where they stood in product of a much more important dispute regard to Darwinian evolution. Haeckel (Virchow,1877; Haeckel, 1878; Gasman, 1971). presumably stood for an evolutionary inter- Haeckel, in fact, in the latter part of the nine- pretation of both Neandertal and “Pithecan- teenth century, was busily promoting an ag- thropus,’’ and Virchow opposed it. The real gressive German nationalism of an anti- nature of the HaeckelNirchow opposition, Semitic and anti-Christian (particularly anti- however, was quite different, and the buf- Catholic) nature which may well have been fetings suffered by anthropology, however im- painful to Dubois’ Catholic sensibilities. In portant they may seem to us, were an any case, the movement of which Haeckel was 416 C.L. BRACE

an active part resulted in a gigantic armed con- Haeckel, evolution did not only mean flict that swirled around the edges of Dubois’s the process of change from one own small country. A possible index of species to the next. Evolution for him Dubois’s feelings can be seen in the fact that, was a cosmic force, a manifestation of after was over and he returned to the creative energy of nature. the arena of scholarly publications, he changed (Gasman, 1971:ll) the spelling of his given name from its German form (Eugen) to its French one (Eughe) and What Virchow, the cautious, empirical ceased entirely to use the German language as liberal really feared was not materialism or the medium for his reports. Whatever the mechanistic evolutionism but the elevation of truth may be, it is clear that Dubois’ switch Haeckel’s peculiar brand of evolutionary from the position advocated by Haeckel to mysticism to the status of a state religion (Vir- that defended by Virchow cannot be explained chow, 1877).The subsequent course of German by a consideration of the anatomical and history has shown that Virchow’s fears were paleontological data available at the time. fully justified. As the most careful appraisal If some thought that Haeckel simply has shown, Haeckel displayed “a romantic represented Darwin written in German, the rather than a materialist approach to reality was quite otherwise. As Gasman has . . . The content of the writings of Haeckel and written: the ideas of his followers-their general political, philosophical, scientific, and social orientation - were proto-Nazi in character . . . a Although he considered himself to be prelude to the doctrine of National Socialism” a close follower of Darwin and . . . in- (Gasman, 1971:xiv). voked Darwin’s name in support of his In a major way, this had an impact on the own ideas and theories, there was, in world which has been far more momentous fact, little similarity between them. than French romantic evolutionism. One other Haeckel himself thought of evolution minor and almost unnoticed casualty was the and science as the domain of religion effort to produce a human phylogenetic tree and his work therefore assumed a based on real fossils. This construction was for- character which was wholly foreign to mulated by the Strassburg anatomist, Gustav the spirit of Darwin. Darwin’s em- Schwalbe, and was greeted with considerable piricism, his caution in the face of satisfaction by Haeckel himself (1906:109). speculative theories, his general Schwalbe produced two possible arrangements mechanical conception of the work- of the then-known erectus, , and ings of nature were all in striking con- modern fossils (see Fig. 11, from Schwalbe, trast to Haeckel’s biology. For 1906:14). One could almost be regarded as a

1. Aiiffussiing :

T 10111o s:t pi c n s 3 T toin o+ p ri 111 i g c n i 11 s I’itlrccnritlrropiis.

Fig. 11. Two possible arrangements of the known hominid fossils by Gustav Schwalbe. “Homoprimigenius”was the term he used for Neandertal (from Schwalbe, 1906:14). PHYLOGENETIC TREES 417

cladogram, and the other, for which he cism from Keith (l910,1911b),the type of con- declared his preference, was a simple phyletic struct which it represented disappeared from line. serious scientific consideration. Initially, Schwalbe’s formulation was adopted with some approval in England ROMANTICISM AND SIR ARTHUR KEITH (Sollas, 1908; Keith, 1911 a,b). By the outbreak When the late Lord Snow spoke of “the two of the First World War, however, Keith had a cultures,” he provided evidence for the fact complete change of (1915, 1925, 1931 that the paradigm shift from the Age of a,b), and for the next half-century almost no Enlightenment to that of Romanticism had recognizable vestige of Schwalbe’s well- not been as all-encompassing as Lovejoy’s considered formulation was to be found. Fur- treatment would lead one to believe (Snow, ther, Hrdlixka, in his splendid summary of the 1963; Lovejoy, 1936). Snow himself was at known human fossil record, completely mis- home in both a literary and a scientific milieu, represented Schwalbe’sposition and its signifi- although he was aware that this was not true cance (Hrdlixka, 1930:327).Hrdlixka’s curious for most intellectuals. In the middle of the treatment of Schwalbe’s views, which were ac- nineteenth century, the gulf between the two tually quite similar to his own, may have been realms, while appreciable, was not quite so un- due to his antipathy for things German bridgeable. Charles and Emma Darwin, for ex- (Spencer, 1979:15, 18, 769-770), although this ample, enjoyed the company of Thomas and can be no more than informed speculation. Jane Carlyle. Conversely, the historian of The last pre-World War I phylogenetic philosophy, George Henry Lewes, was given scheme associated with German romantic an advance copy of Darwin’s Origin to review evolutionism was the wondrous concoction by for Blackwood’s (Barzun, 1958:34). After the the Breslau anatomist, (see “Oxford Movement” and the conversion of Fig. 12 from Klaatsch, 1910:567; Wegner, John Henry Newman in England (1845, see 1910:120). As with the phylogenetic trees of OConnell, 1969; Weatherby, 1973)and the end Haeckel, the organizing principle was basically of the Scottish Enlightenment (Hook, 1975), hypothetical, and known fossil and modern however, the gap grew to such an extent that specimens were only added after the fact. It communication between the sides dwindled to had few defenders, and after a blast of criti- the point where some of those who attempted

ffeanderfd Aurignac

,- A.y,.

\\\‘\I Schimpanse

\v. Tasmanier Fig. 12. The hypothetical scheme of human, ape, fossil, and hypothetical forms suggested by Klaatsch (1910:567). 418 C.L. BRACE to provide bridges produced serious misrepre- Autobiography (1950) he noted his youthful sentations instead. One such was Sir Arthur enthusiasm for Thomas Carlyle, and it is clear Keith. from a careful reading of his writings that Through his numerous books and articles, Carlyle continued to remain the dominant in- Keith earned a reputation in England as one of fluence in his mode of thinking (cf. Keith, the foremost students of human evolution. He 1931b:27).In Darwin’s appraisal of Carlyle, he was a self-proclaimed follower of Darwin recount,ed on one occasion his pleasure in (1950:562),and, upon his retirement from the listening to Carlyle’sdiscourse (Litchfield [ed.], Royal College of Surgeons in , he spent 1915:11:21),and on another that “I never met a the rest of his life in a house on the grounds of man with a mind so ill adapted for scientific the Darwin estate at Down -in the eyes of the research (F. Darwin [ed.], 1887:1:64). In some public, supposedly a continuing symbol of the respects, a bit, of that judgment could be ap- Darwinian spirit. Some years ago (1964)when I plied to Keith himself. Although Sir William noted that Keith and the majority of the stu- Bragg, in his presidential address before the dents of the human fossil record displayed a re- British Association for the Advancement of luctance to interpret the evidence from the per- Science in 1928, chided the outgoing president, spective of Darwinian evolutionary principles, Sir Arthur Keith, for his materialism, it was clear that they had bowed to the in- (Spencer, 197 9: 74 - 75), Keith’s “materialism” fluence of the non-Darwinian ethos which had far more in common with the of prevailed in France after the First World War. Haeckel (and for that matter Bergson) than In the case of Keith, however, I had not real- with Darwinian mechanism. Keith rejected the ized the extent to which the ground was role of chance in evolution in a manner quite already prepared for his defection. akin to Paley’s “natural theology” of more than Keith specifically mentioned the fact that it a century earlier (Paley, 1802).“I could as easi- was the influence of French paleontology ly believe t.he theory of the Trinity as one which led to the change in his views (Keith, which maintains that living developing pro- 1946:141; 1950:318-319). The phylogenetic toplasm, by mere throws of chance, brought trees which then served as the frontispieces for the human eye into existence. The essence of his most infhential work (1915, 1925) had a living protoplasm is its purposiveness” (Keith, distinctly French cast to them as Vallois in ef- 1946:217).Then in a passage that could easily fect noticed nearly 30 years ago (Vallois, 1952; have come straight from Haeckel he added, “I and see Fig. 13). With changes in dates and have just affirmed that there are evolutionary some other details, some of the main points of processes inherent in living things and Keith‘s formulation continue to be apparent in therefore in Nature - trends of change which the schemes of his immediate (Hooton, 1931; are akin to human purpose and human policy” 1946:413; see Fig. 14; Leakey and Goodall, (1946:218).There is little of the cautious em- 1969:108; L.S.B. Leakey, 1965, and inter- pirical Darwin in his writings, and it is in- preted in Cole, 1975:255; see Fig. 15) and more teresting how little of Darwinian mechanism is remote (RE. Leakey and Lewin, 1977:84-85; to be found in the schemes of his English and, Kennedy, 1980) intellectual heirs. In his via Hooton, his American successors. scheme building, however, Keith and his followers did not go to quite the skeptical ex- CLADISTICS tremes of the “evolution buissonante” formula- In order to avoid the manifest subjectivity of tions which the French tend to prefer (see Fig. the past, and in an effort to make assessments 9). Instead they tended to retain some of the of evolutionary relationship more objectively speculative continuity that was first apparent testable, many students of evolution have re- in the diagrams of Haeckel. cently engaged in the practice of constructing It is interesting to note, in this regard, that diagrams with twigs and branches called clad- Keith retained a warm admiration for Haeckel ograms. These are offered to depict phylogen- (Keith, 1935). Further, Keith’s ideas concern- etic relationships. Although there is more than ing racial purity, and his enthusiasm for war a little difficulty in rendering time on a and prejudice as devices for promoting racial cladogram (Delson et al., 1977:265), and this development (1931b) are startlingly close to was not even considered when Julian Huxley Haeckel and subsequent Nazi racial policy. first proposed the term “”(Huxley, 1959), Like Haeckel, Keith‘s self-declaredenthusiasm the sharing of “derived”-that is, modified or for Darwin (cf. 1950:562)masks a more basic non-primitive-traits, which is the basis for ad- commitment to romantic mysticism. In his jacent placement of terminal twigs, is assumed PHYLOGENETIC TREES 419

RECENT PLElSTOCENt 4000 ft. zoo,ooo Yrs. -4----- I PLIOCENE SOOOft. ZS0,oOO Yrs . I -&-- - - - . I I 1 MIOCENE 3000 ft. t50,OOO Yrs. I I I I I 1----- I I I OL~GOCENE 12000 ft. 600,000 yrs. I I I I I I I I .g-- - - .. I EOC~NE lZ000 ft. 600 000 Yrs 1 I I 420 C.L. BRACE

Fig. 14 The refinement of Keith‘s polyphyletic diagram by Hooton (1946:413) to be based on recency of descent from a com- Indeed cladograms based on shared trait mon ancestor (Cain and Harrison, 1960; Mayr, states are easy to draw up, and in many in- 1974). According to some advocates, “trees stances they may constitute satisfactory should always be based on cladograms” and depictions of evolutionary relationships (see discussions of evolutionary dynamics should Fig. 16),but there are some things which they follow after the construction of the trees (Tat- do not handle very well, and a number of tersall and Eldredge, 1977:205).Although the scholars have issued caveats with varying proponents of this approach would prefer to degrees of stringency (Mayr, 1974; Van Valen, label it “phylogenetic ” (Eldredge, 1978; Gingerich, 1979).The fact that cladistics 1979),the term “cladistics” is a lot easier to get has trouble with time is even occasionally of- straight and is preferable for a number of fered as a virtue of the approach as shown in reasons (Mayr, 1974:95). the phrase that it represents “phylogency PHYLOGENETIC TREES 421

Upper Pleisrocene

kUXkXXXXXXXXXXXXXX Middle Pleislocene

ZiriianthroDus‘i boisei Lower Pleistocene \

Pliocene

I Ker?yapithecus

Miocene ProconsulI I Fig. 15. The family tree preferred by the late L.S.B. Leakey. The only hominid fossil accepted as a possible human ancestor is his own “Homo habilis” (adapted from Leakey, 1965115 as depicted in Brace et al.. 1979169). without true time dimension” (Delson et al., Aside from the problem of dealing with time, 1977:265). Having eliminated a concern for which surely is crucial for any study of evolu- time, the authors go on to decry the attempt to tion (Van Valen, 1978).there are twoother mat- depict ancestor-descendant relationships ters that cladistics has trouble handling. One which they declare to be ”untestable.” One of these is the problem of differing rates and must grant, of course, that the paleontologist times of change for different traits within a cannot test his problems in the laboratory like single line or between related lines, i.e., mosaic a modern chemist, but there are more than a evolution (Mayr, 1974).In the case of hominid few who do not regard this as sufficient reason evolution, for example, the timing of changes for declaring that paleontology ceases to be a in size (Holloway,1980a,b) and tooth size science when it deals with matters that are uni- (Brace, 1979a,b,c) is quite different, with the que to its domain, namely time and relation- result that there are still major questions in ships. Even the cladist who has noted that “the regard to the cladistic affinities of more than a probability of recognizing ancestors in a fossil few important early hominids. sample is zero” goes on to suggest that the Another problem that clouds the practice of likelihood of studying the nature of ancestry is cladistic analysis is the matter of continuing “quite substantial” (Vrba, 1980:77). change after a given branch point. As Mayr 422 C.L. BRACE entirely. In fact the extreme exponents of cladistics reject any concern for evolutionary dynamics as a matter of principle (Gaffney, 1979:88). It is also clear from the data of paleoan- thropology that gradualism does occur despite the bald but unsupported statement to the con- trary by Gould and Eldredge (1977:135). Hominid brain size doubled over a span of 1.5 million years until it effectively reached its modern size some time before the onset of the last glaciation. Subsequently it evidently has displayed the expected equilibrium, but at the same time human tooth size has reduced by a full 50% in a broad belt running from Europe to Japan (Brace, 197913, and in preparation). Although this dental change may seem like a sudden event in the full span of evolutionary time, it was not accompanied by , it did not take place in isolation, nor was it par- Fig. 16. An example of a plausible hominid cladogram ticularly peripheral. Furthermore, the pre- (Eldredge and Tattersall, 1975235). ceding expansion in brain size took place gradually over a suitably long period of evolu- tionary time and evidently was shared by the has noted (1971:16, 17), cladistics ignores the entire hominid population. distinction between evolutionary parallelism Those who have been principally behind and evolutionary convergence, but one could developing the picture of punctuated go further and observe that it cannot handle equilibria, whether or not they are proponents phyletic divergence either (cf. Eldredge and of cladistics, have placed it in opposition to the Cracraft, 1980:73-74). It is perfectly possible models of evolutionary gradualism which they for forms to diverge after a branch point and have traced to the world views and political then later reconverge to produce a misleading philosophies of the initial proposers, principal- picture of phyletic affinity. Turner’s depiction ly Darwin, Lyell, and their English contem- of a possible connection between the poraries (Gould, 1977). Others of course have Hoabinhian people of the Southeast Asian noted that Darwin’s views were very much a Mesolithic and the Jomon people of Japan may product of his background. Karl Marx, writing well be a case in point (Turner, 1979). to Engels, observed, “It is remarkable how Cladistic analysis in fact assumes that the Darwin recognizes among beasts and plants main significant steps in evolution take place his English society . . .” (Schmidt, 196237). at branch points and that what happens subse- Commenting on the continuing favor with quently is of considerably less importance. In- which Darwinian ideas are held, Gould and evitably those who favor the use of cladograms Eldredge have noted, “The general preference to illustrate phylogenetic relationships also that so many of us hold for gradualism is a prefer to regard evolution as primarily a pro- metaphysical stance embedded in the history cess of fits and starts. This is embodied in the of western cultures: It is not a high-order em- term “punctuated equilibria” to describe the pirical observation induced from the objective course of organic change (Eldredge and Gould, study of nature” (19773145). 1972; Hecht, et al., 1974). The promoters of The possibility must be considered, this approach in fact seem to have taken one of however, that there may be just as much of Simpson’s three major modes -quantum evo- social conditioning inherent in the recent en- lution (Simpson, 1944:198, 206; 1953: thusiasm for punctuated equilibria as opposed 389-393) -and elevated it to a position of prin- to gradualism. Not only are many of the sup- cipal importance. As has been abundantly porters products of an affluent post-World demonstrated (cf. Vrba, 1980),there are many War I1 milieu in which instant gratification instances where species appear suddenly in the was accepted as due, but as the originators of fossil record and enjoy long periods of ap- the view have noted, “it may also not be irrele- parent stasis before vanishing. Whether or not vant to our personal preferences that one of us the initial shaping was done in a small, isolated learned his Marxism literally at his daddy’s and peripheral population is another matter knee” (Eldredge and Gould, 1977:146). PHYLOGENETIC TREES 423

Finally, one can suggest that the origins of dedicated to Weidenreich in memoriam, the paleontological traditions in the scholar- stressed the long-continued geographic isola- ship of nineteenth-century France may also be tion of separate developing hominid lines of some consequence. The current in modern (Coon, 1962). Another redrew his diagram to paleontology that accepts the discontinuous remove all of the connecting lines that in- nature of the stratigraphic record as indicative dicated contact between geographically adj a- that the history of organic life has been discon- cent populations and then labeled it the tinuous (Gould, 1965),the emphasis on sudden, “Polyphyletic or Candelabra School” (Howells, dramatic change (Gould, 1974, 1978a,b), and 1967:241). In fact, however, the polyphyletic the stress on speciation events in small, stress on regional isolation was more a reflec- isolated groups remote from the area of con- tion of the expectations of the readers than it sideration followed by sweeping take-overs, all was inherent either in Weidenreichs diagrams are remarkably similar to the outlook that or his discussion. characterized nineteenth-century French In their appraisal of the dynamics of hominid paleontology. A century ago this contributed evolution, both Weidenreich and Dobzhansky to the French rejection of the idea of evolution (1944) noted that, once the level of the genus by means of natural selection. Its reemergence Homo had been reached, the geographic isola- today in the writings of a generation of paleon- tion that would lead to local speciation would tologists who have not looked carefully into no longer occur (Weidenreich, 1943b:253, the traditions of the field in which they have 1946b:414). flow across population boun- been trained could well lead to its being daries then would be sufficient so that the designated “neocatastrophism” (Brace, genus would contain a single albeit regionally 1978:983). In view of what we can suspect is differentiated species. If, as Dobzhansky the lurking French connection, it would be ap- noted, modern Homo sapiens constitutes a propriate here to recall what the French would “single polytypic species” (Dobzhansky, say in this regard Plus Fa change, plus c’est le 1944:265), the expectation was implicit that m6me chose. the same thing should have been the case for the previous history of the genus Homo. This THE HOMINID PHYLETIC PICTURE was the basis for the suggestion that, prior to If the interpretive styles of a century ago are the appearance of what we recognize as still with us in several guises, in addition a “modern”form, the human line passed through quantity of fossil material has been ac- a grade of organization (cf. Huxley, 1958) cumulated that was simply undreamed of by characterized by larger jaws and teeth and a the empirically oriented thinkers who framed skeleton displaying greater robustness and the first evolutionary syntheses. Given what more pronounced muscle markings. The first we now possess, the possibility of building such prehistoric skeleton to be recognized was various kinds of descent “scenarios”is almost the one discovered in the Neanderthal in 1856, inevitable. As has already been noted, Gustav and there are some scholars who felt that this Schwalbe made the first attempt to deal with was reason enough to attach that name to the the known hominid fossils early in this century stage through which human form had passed from what would now be regarded as the before reaching its modern configuration perspective of “evolutionary systematics” (see (Hrdlixka, 1927, 1930; Weidenreich, 1928, Fig. 11).Although this was largely forgotten 1943a; Brace, 1964). with the eclipse of German views that general- This does not mean, as some have assumed, ly occurred following World War I, the ap- that the European Neandertals gave rise to the proach was not entirely without subsequent modern Chinese - or Africans or Indonesians. development. Over 40 years later and Instead it means that one would expect to find bolstered by many more fossils, Schwalbe’s that the local predecessors of Chinese, student and, briefly, successor at Strassburg, Africans, and Indonesians -and any other Franz Weidenreich, produced a sophisticated modern population where continuity in situ elaboration (Fig. 17, from Weidenreich, can be expected extending back to the last in- 1947:201; and see also Weidenreich, 1946a:24). terglacial - would display a degree of By this time, however, the paradigm which robustness in teeth, jaws, skeletons, and mus- governed interpretation in , cle markings comparable to that observed in despite some lip-service to Darwin, was largely the European . The idea of a that of the French tradition. As a consequence, polytypic Neanderthal Stage as the antecedent in the few instances where Weidenreich’s work to modern forms of Homo sapiens has been has been mentioned, it has been seriously criticized for many reasons (Howells, 1974, misrepresented. One major work, actually 1976; Thoma, 1973, 1975; Santa Luca, 1978), 424 C.L. BRACE

Fig. 17 This famous diagram is more scenario than tree, and has more implications than the simple designation of “polyphyletic” would lead one to believe (Weidenreich, 1947:201)

but it still seems more useful than the alter- (Kibish) to Florisbad (Day, 1969; Rightmire, natives available which, at best, are largely 1978) or anything in between (Robbins, 1972) cladistic in nature. As Mayr has put it, “It that would contradict such a view. And in results in a great deal of loss of information to , Mapa and the recently discovered Da Li ignore the adaptive component of evolution ex- (examined at the Institute of Vertebrate pressed by the concept of grade. . . .” (Mayr, Paleontology and Paleoanthropology in Beij- 1974:107). ing, 1980, courtesy of Prof. Wu Xin-zhi)make In Europe itself, where the evidence col- splendid Neanderthal Stage precursors of the lected is more abundant and better known modern Chinese. Finally, the evidence from In- than anywhere else, the most thorough recent donesia also is consistent with a picture of in analyses have shown that the transition of situ continuity from early Middle Pleistocene Neanderthal to modern tooth and face form oc- erectus to Solo and possibly via Wadjak to curs gradually and without any break (Brace, modern Australian form. 1979b; Smith and Ranyard, 1980), and it is At the Neanderthal level, the quantity of clear that there is no reason why such a transi- material available from widely dispersed areas tion could not be documented for the creates the very real problem of whether a postcranial skeleton as well (Trinkaus, 1976, single term is adequate to encompass it all. In 1977). In Africa, the evidence is scrappier and spite of the efforts of pheneticists and cladists the dating much more tentative, but there is to reduce to a rigidly defined and nothing inherent in the material from Omo presumably objective science, the element of PHYLOGENETIC TREES 425 “art,” as Simpson has noted (Simpson, would have to be consigned to separate 1961:110, 227), will continue to remain in the species. All of this remains unresolved even practice of the proverbial “competent though there can be little doubt concerning the systematist” (Carter, 1951:118-119). probable status of the Hadar and Laetoli Earlier in the course of hominid evolution, specimens as representing the condition there is another problem which is just as dif- ancestral to all subsequent hominids. ficult to deal with. This involves the question At a slightly more recent level of time, the of how to designate continuous change same sorts of problems (and more) becloud the through time. Of course, one solution is simply status of what has been called “habilis.”Using to deny that the hominid phyletic line gradual- the most numerous data available, namely, ly evolved from one species into another, which tooth measurements, a kind of is precisely the tack taken by one recent “stratophenetic” (cf. Gingerich, 1976, 1979) ar- survey (Stanley, 1979:80-82). Otherwise, if full rangement has been made which then is used cladistic logic were followed (Eldredge and to defend a tree in which “habilis”is regarded Cracraft, 1980:114, 244), we would still belong as the direct descendant of “afarensis”and the to Australopithecus africanus if indeed the last immediate precursor of erectus (Johanson and splitting event in hominid evolution occurred White, 1979:327-328). A similar scheme has when africanus and boisei differentiated. been defended using multivariate distance Most paleoanthropologists, however, accept measures (McHenry and Corruccini, an in situ transition from Australopithecus to 1980:1104). Homo at the generic level and then from erec- While this may very well be correct, it does tus to sapiens at the specific level. The prob- not tell us when to change the generic or even lem, then, is the classic one of successive the specific names. Further, it is just possible species-where to draw the lines, and what to that the legitimate H. erectus and A. africanus call the lineage and when. When we only had a specimens may have been lumped from dif- few points in time represented, it was easy ferent sites and time periods to make up the enough to give them names without regard to “habilis” group in the fist place (Brace et al., whether the entities named were discrete 1973).Part of the difficulty has been caused by events of points on a continuum - the “piers”of the prolonged delay in the proper comparative Sollas’s “ruined bridge which once continuous- treatment of the original material on which the ly connected the kingdom of man with the rest proposed was based. of the animal world (Sollas, 1908:337). While Even Holloway’s recent suggestion that the the gaps remained, they provided room either braincase of the type specimen, OH 7, war- for the presence of a presumed “God,” or rants the designation Homo can be challenged justification for the specific essentialism on solid quantitative grounds (Holloway, preferred by the cladists. 1980b:273;Wolpoff, in press). His further com- Now, however, the accumulation of recent ment that final taxonomic placement will have discoveries has confirmed Sollas’s assump- to await the appraisal of the dentition raises tions. While this is gratifying to those with another major problem, even though the one Darwinian expectations, it has created the ter- comparative and quantitative treatment of the minological problems alluded to above. At the dentition published has shown that the teeth earliest level, “afarensis”has been proposed as of the type specimen cannot be distinguished specifically distinct from africanus because of from Australopithecus africanus (Brace et al., its possession of a series of features that are 1973). clearly more primitive (Johanson and White, But the problem is that the brain and the 1979)- “plesiomorphic”in the cladistic lexicon. teeth may very well be following separate But the differences in these traits are no evolutionary courses. This certainly was true greater than those by which modern human late in the Pleistocene where brain sue did not “races” are distinguished, and they indicate at change at all but the dentition reduced by up to most that africanus had simply been pursuing 50%(Brace, 1979a,b,c).Mosaic evolution is not the same adaptive strategy for a longer period only difficult for cladists to handle, it is dif- of time. If the same logic were rigorously ap- ficult for the evolutionary systematist to plied to the recent hominid fossil record, then represent in the form of a tree. At this point we the Neanderthals would once again warrant a are back to the “art” in Simpson’s taxonomic separate specific designation from sapiens. practice. Even the complex schemes offered Even the Skhiil specimens and possibly such (cf. Fig. 17; Brace, 1979c:105)are cluttered and early Upper Paleolithic groups as Predmost difficult. One solution is to indicate the core of 426 C.L. BRACE the tree by points based on the data from one were to have similar quantities of information evolving dimension and then subjectively for other traits, many would present pictures depict a trunk that is wide enough to include a that are quite different. Both schematic number of the detailed schemata that have simplicity (Fig. 18) and indications of con- been proposed. This is done in Figure 18 (from siderable complexity (Fig. 17) have their utili- Brace et al., 1979:173).Whether this can ac- ty. No one has a monopoly on “truth.” comodate the resolution of the brain size issue Although it is legitimate to feel that many and the data from other dimensions that will have helped increase our understanding of the eventually be considered is impossible to dynamics by which humanity emerged, it is predict at the present time. At least it provides just because we can never know the whole of a framework for the various contending that “truththat a multiplicity of schemes will hypotheses that are currently being con- continue to be constructed and will continue to sidered. be of value.’ It is evident from the variations that are pro- duced by the use of the different measures or ACKNOWLEDGMENTS their combinations that such simple quan- titative procedures do not give us a single The text of this paper is a revision of the presentation made at Charlottesville, Virginia Platonic truth. Even the summed use of metrics representing the considerable amount on December 11, 1980 at the meeting com- memorating the fiftieth anniversary of the of teeth available to us - however satisfying founding of the American Association of some of us may think this to be - only gives us a picture of evolution in one dimension. If we Physical Anthropologists. I am grateful to Drs. Frank Spencer and Noel T. Boaz for in- viting me to be on the program at Charlot- tesville and for arranging for the publication, in the American Journal of Physical An- I,lod ern s thropology, of a paper which would otherwise have been rejected because of its failure to Neanderthals comply with standard editorial policy.

‘In the past, I have often closed my comments with a hit of doggerel from the pen of I. Doolittle Wright. His efforts, however, have been limited to the use of the limerick form, and given the serious nature of H erecfu: the present occasion, it occurs to me that a more appropriate mode of expression would he preferable. Since I started the current paper with I. observations concerning the debt of Darwin and his contemporaries to the ethos of the Scottish Enlightenment, it seemed only fitting to record my own levels of skepticism in suitable Scottish form. Although, as the name shows, he is not a proper Scot, nevertheless Wright’s cousin I. Wright Drivell has dabbled in the use of favorite stanza of Robert Rurns. In response to Wright’s simple certainties, \* A afri c anus Drivell. acting the veritable killjoy, replied to his cousin crossly with Y‘ his “Owed to Trees” As we survey the path we’ve trod, Of knowledge gained by labored plod 3r7. mL Now each aspiring learned clod, Will try to see, How he can he a hit like God And make a tree.

Fig. 18. The dots within this tree are based on actual The record shows that those who strived, datapoints, sums of mean cross-sectional areas for the teeth Produced results that look contrived in the dental arches of the populations named. The lowest Are twigs then based on traits derived, dot is based on the material from Had= and Laetoli. The dot To make a clade? Do branches show the route arrived, between africanus and boisei is from the material at To reach a grade? Swartkrans and Kromdraai. The earliest dot in the sapiens section is from the Krapina Neanderthals. The classic However much they try to please, Neanderthals of western Europe are just above that, and the The schemes expand by twos and threes; two dots at the top display the range between Australian The viewer then who thinks he sees, aborigines and modern Europeans. (From Brace et al., Can only fail, To tell the forest from the trees, 1979:173, based on data recorded in Brace, 1979a,b). -To no avail. PHYLOGENETIC TREES 427

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