the nature and status of the florisbad skull as revealed by its nonmetrical features

THE NATURE AND STATUS OF THE FLORISBAD SKULL AS REVEALED BY ITS NON-

METRICAL FEATURES

ALEXANDER GALLOWAY Department of Anatomy, University of the Witwatersrand, Johannesburg,

South Africa

ONE PLATE

INTRODUCTION

In 1932, Professor Dreyer unearthed at Florisbad, 20 miles north of Bloemfontein in the Orange Free State, the most important human fossil found in Southern Africa since the discovery of Rhodesian Man. The circumstances relating to this find are outlined in Drennan’s ( ’37) analysis of this skull and its endocranial cast.

The first diagnosis of this skull was made by Dreyer in 1935 when he described it “as a very primitive form related to Homo sapiens,” and he tentatively placed it in a subgenus ‘ Africanthropus. ’ Kappers, at the same time, commenting on the endocranial cast, approximated it to fossil representatives of Homo sapiens, but his interpretation, as Dreyer ( ’36) has pointed out, was vitiated by an incorrect orientation of the cast. In this latter paper, Dreyer concludes that “the Floris- bad skull belongs to the prehistoric South African race-the Bushman, of which he is a very early and very primitive repre- sentative.” In this conclusion Dreyer appears tacitly to ac- cept Kappers’ view that the specimen is one within the limits of Homo sapiens.

On the other hand, Drennan in a brief communication in 1935, and in a more detailed study presented to the Royal Society, South Africa, this year has argued in favor of the view that

1

AYEDICAN JOUENAL OF PHYSICAL ANTEROPOLOGP. VOL. XXIII. NO. 1 JULY-SEPTEYBER, 1937

2 ALEXANDER GALLOWAY

Florisbad Man is best interpreted as an African variant of the Neanderthal race. He adds as an after thought but without detailed demonstration that “in Florisbad Man, Rhodesian Man, and the Cape Flats Australoid we have a closely related phylogenetic sequence linking the Homo primigenius to the Homo sapiens type.’’

Both these diagnoses are correct as far as they go and may be regarded as elucidating different facets of this most com- plicated skull. The outstanding features of the skull are its excessive absolute length (210 mm. 4 ) and its thickness of bone, which seem to Drennan “to cry out for a Neanderthal interpretation.” Fossil skulls with such general features are al- ready known in South Africa and it is surprising that Drennan turned his eyes to Europe for his diagnosis and by so doing missed a possible diagnosis lying at hand under his eyes.

“he skull consists of the greater part of the vault and part of the right facial skeleton. As Drennan has analyzed it in detail metrically but has paid only scant attention to non- metrical features I will confine myself to the latter.

NON-METRICAL FEATURES OF THE FLORISBAD SKULL

The contour of the vault is a low even convexity (pl. 1, A) and is characterized by a metopic ridge in the frontal region, and an interparietal groove, the only one among South African skulls to equal that of the original Boskop skull. The contour of the forehead passes on to the supraorbital torus without the intervention of an ophryonic groove.

Cun- ningham ( ’09, pp. 291-296) classified the supraorbital torus of primates, including man, in three types :

The supraorbital margin extends in a continuous and arch-like manner from the fronto-malar suture on the outer side to the fronto-maxillary suture on the inner side. It is divided into a long outer part and a short inner portion by the supraorbital notch. The outer portion is a projecting and sharply defbed edge; the inner part, quite distinct from the supraciliary arch, sweeps downwards immediately in front of the trochlear pit, and although much less prominent than the outer part is yet quite obvious.

The nature of the supraorbital torus is important.

Type I.

THE FLORISBAD SKULL 3

The supraciliary ridge or arcw supraciliaris is a semilunar smooth elevation which lies above the inner part of the inner portion of the supraorbital margin. Its inner end curves downwards into the glabella, and is separated from the corresponding eminence of the opposite side, by .a narrow, shallow median depression; its outer end fades away as it approaches the trigonurn supra-orbitale.

The trigortum supra-orbitale lies to the outer side of the supraciliary eminence. It is a triangular depressed field, with its apex a t the fronto-malar suture, which is included between the outer parts of the margo supra-orbitalis and the anterior prominent part of the temporal ridge.

This type of eyebrow region does not appear to be distinctive of, nor indeed more frequently present in, any one race more than another. It occurs in most, if not in all, recent races . . .

Type 11. I n type I1 are included those skulls which exhibit that condition of the supraorbital region which Schwalbe has described as being peculiar to recent man, and also to the mandrill and other species of apes. The supraciliary projection has coalesced with the part of the supraorbital margin which lies to the inner side of the supraorbital notch. On the outer side of the notch the eminence extends outwards with a varying degree of prominence and for a varying dis- tance towards the trigonurn supraorbitalis. From the latter i t is separated by a faint groove which ascends obliquely upwards and outwards from the supraorbital notch. The trigonum supraorbitale, which varies in its extent according to the degree of development of the arcus supraciliaris is, as a rule, depressed and flattened. This form of the supraorbital region occurs in all races, past and present, with the excep- tion of the Neanderthal race, and is undoubtedly the condition which is most distinctive of man.

Type 111. In this form of the supraorbital region all the three elements which enter into its formation become fused together so as to constitute a continuous arch, the torus supraorbitalis, which bounds the orbital opening above and forms a varying amount of the forepart of the roof of the orbital cavity. This arch extends from the glabella to the fronto- malar suture, and in its typical condition it shows no trace of the constituent elements of which it is composed, so completelv have these become blended the one with the other. Of such a nature is the supraorbital region in the adult chimpanzee, in the gorilla, and in a large number, if not the majority, of the lower apes of the old world . . . .


The supraorbital region in the Neanderthal, Spy, and Krapina remains presents features which place it within type 111. As Schwalbe has shown, the different elements have become blended in a strongly projecting torus supraorbitalis in which there is little or no indication of its composite char- acter.

Type 111, which is characteristic of the Neanderthal race, may be found in Homo sapiens, but types I and I1 are distinctive of Homo sapiens and are not found among Neander- thal types.

The supraorbital torus of the Florisbad skull consists of two parts (pl. 1, B). The part medial to the supraorbital notch is massive and boldly convex in its vertical plane. I t incorporates the medial portion of the superior orbital margin and its medial down-flowing between the orbits seems to carry with it the glabellar prominence. Laterally, however, it fades out along the medial margin of the well-defined sulcus supraorbitalis, which runs upward and laterally from the supraorbital notch. Tbe lateral part of the torus consists of the thickened, round, everted lateral portion of the superior orbital margin, and is definitely limited medially by the sulcus supraorbitalis. As such, it conforms to Cunningham’s type 11, a type distinct from the Neanderthal torus.

In Neanderthal Man there is always an ophryonic groove. Sollas ( ’08, p. 284) remarks that the glabellar fossa is strongly marked in the Neanderthal, and barely existent in the Aus- tralian skull. “It is, indeed, largely owing to the presence of its associated fossa that the torus in the Neanderthal skulls, as in the anthropoid apes, stands out in such high relief.’’ This ophryonic groove (i.e., the glabellar fossa of Sollas) is even more exaggerated in Homo palestinus. I n Florisbad Man, as in the modern Australian skull, the forehead slopes upward and backward directly from the supraorbital torus withiut any trace of an ophryonic groove.

Sollas (ibid.) also points out that in the Australian skull the nasal bones


are pinched in as it were, and pressed under the glabella, so that the nasion is situated at the point of abrupt contrary flexure. It is to this feature that the male Australian skull in general owes much of its ferocious appearance. I n the Neanderthal type as represented by the Gibraltar skull, the nasal bones are very broad at the upper end, and instead of terminating at the point of contrary flexure are continued upwards with gently flowing outline in the curve of the glabella itself, so that the nasion is situated some four or five mms. above the bottom of the trough which marks the passage from the nasal to the glabellar region.

In the Florisbad skull the situation of nasion is identical with that in the Australian skull.

The orbits of Neanderthal Man are described as ‘large round, widely opened’ (Sollas ibid., p. 336), a description cor- roborated by Boule ( ’23, p. 204). The plane of the floor of the Neanderthal orbit is only slightly sunk below the plane of the inferior orbital margin; the medial part of this iu- ferior margin is bevelled off, but the lateral portion always presents a sharp edge. In the Florisbad skull the orbital width greatly exceeds the orbital height, the floor is well depressed below the level of the inferior margin, the lateral portion of which margin is so widely bevelled off that the contours of the floor of the orbit andsthe body of the malar bone at the infero-lateral angle form a wide even convexity.

The facial aspect of the maxilla of Neanderthal Man is characterized by the complete absence of any infraorbital excavation, so that the plane of the malar bone and that of the body of the maxilla are almost confluent, while the nasal process of the maxilla protrudes so markedly anteriorly that its plane faces almost laterally.

In the Florisbad skull the plane of the malar lies at almost a right angle to that of the body of the maxilla, so extensive is the infraorbital excavation. This excavation combined with the bevelling of the infero-lateral angle of the orbit causes the bone flanking the zygomatic-maxillary suture laterally to be protuberant, while in Neanderthal Man this region recedes evenly in a postero-lateral direction. Thus the main plane


of the malar in Florisbad Man falls laterally whereas in the Neanderthal group it is antero-lateral. Further the Neander- thal malar is an insignificant thing compared with that of the Florisbad. The plane of the nasal process in the Florisbad skull faces as much anteriorly as laterally. In marked con- trast to this process in Neanderthal Man, it does not show a marked projection in front of the infero-medial angle of the orbit.

From the endocranial cast it is apparent that the vascular grooves on the interior of the vault were very prominent and that the crista frontalis interna is massive with a broad base and a sharp posterior border.

In its non-metrical features then, the Florisbad skull has little, if any, Neanderthal affinities, while the nature of its supraorbital torus, malar bone and maxilla link it definitely with Homo sapiens.

THE ENDOCRANIAL CAST OF T H E FLORISBAD SKULL

Drennan has sought further support for his interpretation of the Florisbad skull in the features of its endocranial cast. Here, as in his study of the skull fragments, he has pointed out facts of undoubted significance, yet it cannot be said that he has exhausted the possibilities of interpretation of this specimen.

The late Sir Grafton Elliot Smith and his pupil Professor Shellshear in a long series of studies have traced the evolution of the brain from ape to man, as shown both by endocranial casts from fossil skulls and by brains of primitive living races. Their work has made clear the steps by which the great expansion of the association areas of the pre-frontal and parieto-temporal region of the brain, which distinguishes the human from the simian type of cerebral organization, has been brought about. They have shown that in the parieto- temporal region, expansion first takes place in the posterior portion of the middle temporal convolution, and in the supra- marginal area which surrounds the posterior extremity of the great lateral or Sylvian fissure. In the pre-frontal region,


it is the inferior frontal and fronto-orbital areas lying just above the orbital margin which first undergo expansion. From these centers the process of expansion gradually extends throughout the association areas. Further, the sequence in which the various territories undergo this expansion in the course of evolution is the same as that in which they attain full functional development in the growth .of the brain of the modern human child.

As a result of these studies by Elliot Smith and Shellshear, it is possible by an examination of the relative expansion of the different portions of a human brain or endocranial cast to assess with reasonable accuracy its position in the ascend- ing scale of brain development.

The Florisbad cast unfortunately comprises only the frontal lobes and the anterior median portion of the parietal lobes. From these, however, a great deal of information can be gathered.

At first glance, the frontal portion of the cast presents a most peculiar appearance, due to its extreme breadth and flatness, which misled Kappers into thinking the cast to be incomplete inferiorly. Drennan has shown, however, that a nearly equal degree of broadening and flattening of the frontal lobes may be found within the normal limits of Homo sapiens. The endocranial cast of Dean Swift has a frontal region as broad as, and but little higher than, that of Florisbad Man.

This flattening gives to the inferior frontal area a prominence which is hardly warranted by its degree of expansion. Meiring ('36, p. 969) has in fact demonstrated the lesser degree of development of this region in the Florisbad cast as compared with that of the Boskop skull M. R. 1. Hence, while Drennan has noted that, apart from its lesser height, the contour of this region in the Florisbad cast agrees with that of Neanderthal casts, more elaborate comparison reveals that the inferior frontal expansion of the Neanderthal specimens is considerably greater, being comparable in degree with that of the Boskop type. The expansion of this area in the Floris- bad specimen is of the same order as is seen in the Rhodesian

8 ALEXANDER QALLOWAY

endocranial cast. By this criterion alone, then, the Florisbad type proves to be less advanced than either the Neanderthal or the Boskop type.

Above the inferior frontal territory there is perceptible, despite the unusual shape of the frontal region, an area of depression corresponding to the superior and middle frontal convolutions. Thus, in this cast, as in those of Rhodesian Man, Neanderthal Man and Boskop Man, the upper portion of the pre-frontal region was lagging behind the lower in its evolution. This is confirmed by Meiring’s demonstration of the simple and coarse modelling of these convolutions. At the anterior extremity of the cast, however, the upper part of the fronto-orbital territory constitutes a distinct expanded boss. This is not to be seen in the Rhodesian and Neanderthal casts, in which the orbital margin of the frontal lobe recedes in an even curve as it passes outward from the mid-line. The presence of this boss imparts to the Florisbad cast a suggestion of that fullness of the frontal poles which Elliot Smith noted in the Boskop and Kanjera casts, and regarded as diagnostic of Homo sapiens. In a frontal lobe of such unusual form, and mayhap nearing the extreme limit of normal variation of the type, this feature should not be unduly stressed.

Further, in this region Dreyer has pointed out to me the existence of a depressed convex triangular area medial to this boss, which is not seen in either Neanderthal Man or Homo sapiens, but which in Rhodesian Man is represented by a small flat depressed triangular area. Dreyer suspects this to be a definitely primitive feature, reminiscent of the modelling of the gyrus rectus in the baboon.

The post-coronal portion of the cast, i.e., the area behind the coronal vascular ridge, is flattened and slopes downward and backward. It is true, as Elliot Smith has pointed out, that even in Homo sapiens a distinct flattening is frequently found in this region over the pre-Rolandic and post-Rolandic motor and sensory areas. The downward slope is, however, a feature of great importance, which Drennan has rightly

THE FLOBISBAD SKULL 9

emphasized as indicating the primitive status of this brain. It shows that the supra-marginal area of expansion cannot have been continued upward into the parietal area as it is in both Neanderthal and Boskop Man. Thus the development of this region cannot have passed beyond the stage shown by Sinanthropus and Rhodesian Man.

The evidence of both the frontal and the parietal lobes, therefore, indicate clearly that the brain of Florisbad Man was considerably different from and less specialized than, that of Neanderthal Man. It presents a much closer parallel to the more generalized stage of human cerebral evolution represented by Rhodesian Man. Its chief differences from that of Rhodesian Man lie in its greater size and in the cofiguratioii of the frontal region, differences which suggest the condition characteristic of Homo sapiens. Thus interpreted, there is nothing to negative the possibility of the Florisbad endocranial cast being the precursor of the Boskop type, the latter being only distiguished by a greater degree of expansion in the parietal and inferior pre-frontal regions.

DISCUGGION

To explain the Florisbad skull it has been necessary to compare it with Neanderthal and Rhodesian Man on the one hand, and with the Boskop and the Australian aboriginal skull on the other. The skull of Florisbad Man agrees metrically with that of N.eandertha1 Man, as Drennan has shown, but there is no agreement non-metrically. Since the non-metrical features of the calvaria are the outward expression of the underlying brain and since the endocranial cast of Florisbad Man is more generalized than that of the earlier Neanderthal Man, there can be no question of the phylogenetic origin of Florisbad Man from Neanderthal Man.

Rhodesian Man is just as dissimiliar in norma facialis from Florisbad Man as is the Neanderthal type, but the agreement between the endocranial casts is so close as to warrant the claim that the Rhodesian type must be very closely related to the ancestor of the Florisbad type.


I n table 1, I have set out these non-metrical features which the Florisbad fragments display and the table demonstrates to what extent these features recur in the Australian skull and in the Boskop phpical type.

Trigonism and a well-marked interparietal groove constitute the salient features of the vault of the Florisbad skull, which is elongated and of great thickness. The skull of the Boskop physical type shows all these features added to which is the pentagonoid vault and post-coronal flattening. The vault of the Australian skull, however, differs from the Florisbad skull in having neither trigonism nor an interparietal groove although it is characteristically pentagonoid and flattened post-coronally.

The Florisbad forehead is characterized by a supraorbital torus of type I1 Cunningham, the absence of an ophryonic groove and a markedly receding forehead, features which are very closely paralleled in the Australian skull. I n the Boskop skull, on the other hand, although its supraorbital region is type I1 Cunningham, the lateral supraorbital portion is re- duced relative to the medial part, while there is always a definite ophryonic groove associated with a low vertical forehead. The vertical ,forehead is an expression of the pedomorphic nature of the skull, while the presence of the ophryonic groove indicates the more generalized nature of the forehead of the Boskop skull (Weidenreich, '37).

The Florisbad calvaria, therefore, may be summed up as the combination of a Boskop vault with an Australian forehead.

Comparisons of the facial regions similarly reveal how the orbits in all three types are identical in shape, direction of axis and nature of margin. The Boskop orbit differs in that the bevelling of the infero-lateral angle forms a definite shelf. The nasal process of the maxilla of none of them is projecting while the plane of its facial surface faces as much anteriorly as laterally. Whereas the nasal bones of the Florisbad skull and the Australian are geroiito morphically raised into a ridge or an arch, those of the Boskop type are pedomorphically flattened.

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Both the Florisbad and the Boskop skull are equally characterized in norma interna by a salient massive crista frontalis interna and well-defined deep canal-like impressions for the middle meningeal artery and the sigmoid portion of the transverse sinus.

As in the vault, so in the face of the Florisbad skull, the features are those of Homo sapiens and their nearest counter- parts are those of the later Boskop and Australian types. While, on the one hand, some features are common only to Florisbad and Australian and, on the other hand, are common to Florisbad and Boskop, there are so many features which are common to all three that two-thirds of the Florisbad features are encountered in the modern Australian skull and two-thirds in the Boskop. Meantime, the endocranial cast of the Florisbad skull is of the generalized type and resembles that of .Rhodesian Man. These similarities pass the limits of coincidence and place the Florisbad type somewhere intermediate between the Rhodesian type and later types.

To suggest a common ancestry for the smaller-skulled modern Australian and the large-headed Boskop South African types may seem hazardous. Strange as it may appear, the Wadjak skull, a type of great cranial capacity and absolute calvarial measurements-yet so similar in its general mor- phology to the Australian aboriginal that it is accepted by practically all anthropologists as proto-hustralian-has features of such a nature as to declare its affinity with the South African Boskop type and to narrow more than ap- preciably the gap between African and Australian human stocks.

The close resemblance of the Wadjak skull and the Boskop type are exhibited by the non-metrical features. I ts vault is pentagonoid shaped and shows an interparietal groove and some postcoronal flattening. The cranial contour in norma lateralis reveals an almost orthognathous face with subnasal prognathism ; the glabellar region and supraciliary ridges are prominent. While the low sharply receding forehead, in which the ophrponic groove is absent, is not Boskopoid, the rest of the


contour with its flat vault whose vertex is in front of porion, the sloping parieto-occipital region terminating in an almost foetal occipital boss, the sigmoid nuchal plane, is emphatically of that order.

The supra-asterionic area is slightly concave, the mastoid process is small, the supramastoid crest turns sharply upward at right angles, the temporal squama is bulged out as in the Fish Hoek skull, and the inferior frontal eminence is well marked-all Boskop characteristics.

Again the face seems small in relation to the calvarial dimen- sions but is broad and massively constructed and is reminiscent of the Fish Hoek skull. The orbits are low and rectangular and their ‘transverse’ axes pass obliquely downward and backward. They are surmounted by a supraorbital region which resembles closely that of the Cape Flats skull, which I have shown to be a Boskop variant (’37). The malar bone faces laterally and it has an upward tilt; the interorbital region is wide and its contained maxillary nasal processes face more anteriorly than laterally and are separated by narrow and only slightly ridged nasal bones. As Keith ( ’29) has shown, the palate also is reminiscent of extinct African types.

It will be seen, therefore, that the Wadjak skull shares too many features in common with the Boskop physical type for them not to be genetically related.

In his analysis of the Wadjak skull, Keith (’29, p. 446) says “I have placed Wadjak Man as an offshoot from the stem which afterwards diverged into Australian and Negro types. I have placed him thus because of his many resem- blances to the older and more primitive Rhodesian Man on the one hand, and to the Australoid type on the other.” Through- out all Keith’s African studies there is the repeated suggestion of the close relationship between the Negro ancestral stock and the Boskop type. This foregoing comparison of the Wadjak skull with the Boskop type and Keith’s statements show the closeness of the ancestral relations between African and Australian types.


It is now generally accepted that Rhodesian Man, while superficially resembling Neanderthal Man, differs from him fundamentally and that he is more definitely one of the earliest types of Homo sapiens with such affinities to the modern Australian that Keith and Dubois even regard him as proto- Australian. Dubois ( ’37, p. 1) categorically states “Judging from the pulp cavities of the teeth, from the straight and rather slender limb bones, accompanying the skull, from the peculiar nuchal plane of the occipital bone, and above all from the low cranial capacity, this certainly is no Homo w a d e r - thalertis, but a Homo sapiens and a proto-Australian.”

Keith, more cautiously, in his final chapter to the Antiquity of Man (’29) finds that the nearest approach to the type of man who is most likely to serve as a common ancestor for both African and European is the modern Australian in that he retains more characteristics of the common ancestors than any other living race (p. 713), and later on insists that Rho- desian Man, while not the direct ancestor of any living race foreshadows many of the features of the modern type, par- ticularly of the Australian aborigine (p. 721).

This evidence and that of the Sinanthropic Lake Eyassi skull described by Leakey (’36) reveals that the primitive types of Africa and the Orient are so closely related that the Eyassi form and Sinanthropus itself may well be variants of an early generalized type of which Rhodesian Man and Solo Man represent their respective modern recent descendants. Both of these are later types and are termed proto-Australian. Closely akin to this so-called proto-Australian type is the Florisbad skull, showing as it does, a close similarity in endocranial features in Rhodesian Man on the one hand, and on the other, a collection of skeletal features which are almost exactly paralleled by the modern Australian skull. The Floris- bad type seems, therefore, to be an intermediate type of early Homo sapiens, such as could be the ancestor of the modern Australian, the gap between the two being in part bridged by the features of the Wadjak skull.


His purely African position is clear. The endocranial cast shows features linking him closely to the generalized human stem, while the features of the vault and face bones indicate his position as ancestral to the Boskop type. Extinct Boskop and living Australian represent the pedomorphic and gerontomorphic evolution of this early generalized type.

More modern than Neanderthal Man and arising from some Beparate but closely-related stock nearer in cerebral development to Rhodesian Man, Florisbad Man could serve equally well as the ancestor of Boskop Man and the proto-Australian stock of Wadjak. The Boskop and Australian types represent the pedomorphic and gerontomorphic specializations of this earlier, generalized ‘and quasi-sapient and specifically African fossil type.

LJTERATURE CITED

BOULE, M. 1923 Fossil men, elements of human palaeontology. Edinburgh: Olivier and Boyd.

CUNNINGHAM, D. J. 1909 The evolution of the eyebrow region of the forehead, with special reference to the excessive supra-orbital development in the Neanderthal race. Trans. Roy. Soc. Edin., XLVI, 263311.

DRENNAN, M. R. 1935 The Florisbad skull. S. Afr. J. Sci., XXXII, 601-602. 1937 The Florisbad skull and brain cast. Trans. Roy. Soe. 8. Afr.

A human skull from Florisbad, Orange Free State, with a Proc. Acad.

1936 The endocranial cast of the Florisbad skull-a correction.

DUBOIS, E. 1937 On the fossil human skulls recently discovered in Java, and

GALLOFAY, A. 1937 Man i n Africa in the light of recent discoveries. Pres.

The characteristics of the skull of the Boskop physical type.

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S O U S , W. J.

THE FLORISBAD SKULL ALEXANDER GALLOWAY

I’I,ATE 1

A, lateral view; B, front view of a restoration of the Florisbad skull.

17

AMERICAN JOURNAL OF P l i Y S l C A L AGTHROPOILM3Y. YOL. S S I I I . KO. 1

the nature and status of the florisbad skull as revealed by its nonmetrical features

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