Australopithecus Africanus: The man-ape of South Africa

Cranial form in living anthropoids of similar age (after Duckworth) and in the new fossil. For this comparison, the fossil is regarded as having the same calvarial length as the gorilla.

Published in Nature on February 7, 1925, this was the first proper public paper by Raymond Dart announcing his discovery, and the start of the entire Taung story.

A note on illustrations: Because of the poor quality of the copy of the article available to us, we could only reproduce three of the six figures Professor Dart used to illustrate his original article.

"Towards the close of 1924, Miss Josephine Salmons, student demonstrator of anatomy in the University of the Witwatersrand, brought to me the fossilised skull of a cercopithecid monkey which, through her instrumentality, was very generously loaned to the Department for description by its owner, Mr. E.G. Izod, of the Rand Mines Limited. I learned that this valuable fossil had been blasted out of the limestone cliff formation - at a vertical depth of 50 feet and a horizontal depth of 200 feet - at Taungs, which lies 80 miles north of Kimberley on the main line to Rhodesia, in Bechuanaland, by operatives of the Northern Lime Company. Important stratigraphical evidence has been forthcoming recently from this district concerning the succession of stone ages in South Africa (Neville Jones, Jour.

Dioptographic tracing of Australopithecus africanus (right side), x 1/3

 Roy. Anthrop. Inst., 1920), and the feeling was entertained that this lime deposit, like that of broken Hill in Rhodesia, might contain fossil remains of primitive man.

"I immediately consulted Dr. R.B. Young, professor of geology in the University of the Witwatersrand, about the discovery, and he, by a fortunate coincidence, was called down to Taungs almost synchronously to investigate geologically the lime deposits of an adjacent farm. During his visit to Taungs, Prof. Young was enabled, through the courtesy of Mr. A.F. Campbell, general manager of the Northern Lime Company, to inspect the site of the discovery and to select further samples of fossil material for me from the same formation. These included a natural cercopithecid endocranial cast, a second and larger cast, and some rock fragments disclosing portions of bone. Finally, Dr. Gordon D. Laing, senior lecturer in anatomy, obtained news, through his friend Mr. Ridley Hendry, of another primate skull from the same cliff. This cercophithecid skull, the possession of Mr. De Wet, of the Langlaagte Deep Mine, has also been liberally entrusted by him to the Department of scientific investigation.

"The cercopithecid remains placed at our disposal certainly represent more than one species of catarrhine ape. The discovery of Cercopithecidae in this area is not novel, for I have been informed that Mr. S. Haughton has in the press a paper discussing at least one species of baboon from this same spot (Royal Society of South Africa). It is of importance that, outside the famous Fayüm area, primate deposits have been found on the African mainland at Oldaway (Hans Reck, Sitzungsbericht der Gesellsch. Naturforsch. Freunde, 1914), on the shores of Victoria Nyanza. (C.W. Andrews. Ann. Mag. Hist., 1916), and in Bechuanaland, for these discoveries lend promise to the expectation that a tolerably complete story of higher primate evolution in Africa will yet be wrested from our rocks.

"In manipulating the pieces of rock brought back by Prof. Young, I found that the larger natural endocranial cast articulated exactly by

Contour tracings of coronal sections through the widest part of the parietal region of the endocranial casts in Australopithecus …….. and in a gorilla

 its fractured frontal extremity with another piece of rock in which the broken lower and posterior margin of the left side of a mandible was visible. After cleaning the rock mass, the outline of the hinder and lower part of the facial skeleton came into view. Careful development of the solid limestone in which it was embedded finally revealed the almost entire face depicted in the accompanying photographs.

"It was apparent when the larger endocranial cast was first observed that it was specially important, for its size and sulcal pattern revealed sufficient similarity with those of the chimpanzee and gorilla to demonstrate that one was handling in this instance an anthropoid and not a cercopithecid ape. Fossil anthropoids have not hitherto been recorded south of the Fayüm in Egypt, and living anthropoids have not been discovered in recent times south of Lake Kivu region in Belgian Congo, nearly 2 000 miles to the north, as the crow flies.

"All fossil anthropoids found hitherto have been known only from mandibular or maxillary fragments, so far as crania are concerned, and so the general appearance of the types they represented has been unknown; consequently, a condition of affairs where virtually the whole face and lower jaw, replete with teeth, together with the major portion of the brain pattern, have been preserved, constitutes a specimen of unusual value in fossil anthropoid discovery. Here, as in Homo rhodesiensis, Southern Africa has provided documents of higher primate evolution that are amongst the most complete extant.

"Apart from this evidential completeness, the specimen is of importance because it exhibits an extinct race of apes intermediate between living anthropoids and man.

"In the first place, the whole cranium displays humanoid rather than anthropoid lineaments. It is markedly dolichocephalic and leptoprosopic, and manifests in a striking degree the harmonious relation of calvaria to face emphasised by Pruner-Bey. As Topinard says, "A cranium elongated from before backwards, and at the same time elevated, is already in harmony by itself, but if the face, on the other hand, is elongated from above downwards, and narrows, the harmony is complete." I have assessed roughly the difference in the relationship of the glabella-gnathion facial length to the glabella-inion calvarial length in recent African anthropoids of an age comparable with that of this specimen (depicted in Duckworth's "Anthropology and Morphology," second edition, vol. i), and find that, if the glabela-inion length be regarded in all three as 100, then the glabella-gnathion length in the young chimpanzee is approximately 88, in the young gorilla 80, and in this fossil 70, which proportion suitably demonstrates the enhanced relationship of the cerebral length to facial length in the fossil.

"The glabella is tolerably pronounced, but any traces of the salient supra-orbital ridges, which are present even in immature living anthropoids, are here entirely absent. Thus the relatively increased glabella-inion measurement is due to brain and not to bone. Allowing 4 mm. for the bone thickness in the inion region, that measurement in the fossil is 127mm.; i.e. 4mm. less than the same measurement in an adult chimpanzee in the Anatomy Museum at the University of the Witwatersrand. The orbits are not in any sense detached from the forehead, which rises steadily from their margins in a fashion amazingly human. The inter-orbital width is very small (13 mm.) and the ethmoids are not blown out laterally as in modern African anthropoids. This lack of ethmoidal expansion causes the lacrimal fossae to face posteriorly and to lie relatively far back in the orbits, as in man. The orbits, instead of being subquadrate as in anthropoids, are almost circular, furnishing an orbital index of 100, which is well within the range of human variation (Topinard, "Anthropology"). The malars, zygomatic arches, maxillae, and mandible all betray a delicate and humanoid character. The facial prognathism is relatively slight, the gnathic index of Flower giving a value of 109, which is scarcely greater than that of certain Bushmen (Strandloopers) examined by Shrubsall. The nasal bones are not prolonged below the level of the lower orbital margins, as in anthropoids, but end above these, as in man, and are incompletely fused together in their lower half. Their maximum length (17 mm.) is not so great as that of the nasals in Eoanthropus dawsoni. They are depressed in the median line, as in the chimpanzee, in their lower half, but it seems probable that this depression has occurred post-mortem, for the upper half of each bone is arched forwards (Fig 1). The nasal aperture is small and is just wider than it is high (17 mm. x 16 mm.). There is no nasal spine, the floor of the nasal cavity being continuous with the anterior aspect of the alveolar portions of the maxillae, after the fashion of the chimpanzee and of certain New Caledonians and negroes (Topinard, loc. cit.).

"In the second place, the dentition is humanoid rather than anthropoid. The specimen is juvenile, for the first permanent molar tooth only has erupted in both jaws on both sides of the face; i.e. it corresponds anatomically with a human child of six years of age. Observations upon the milk dentition of living primates are few, and only one molar tooth of the deciduous dentition in one fossil anthropoid is known (Gregory, "The Original and Evolution of the Human Dentition," 1920). Hence the data for the necessary comparisons are meagre, but certain striking features of the milk dentition of this creature may be mentioned. The tips of the canine teeth transgress very slightly (0.5-0.75 mm.) the general margin of the teeth in each jaw, i.e. very little more than does the human milk canine. There is no diastema whatever between the premolars and canines on either side of the lower jaw, such as is present in the deciduous dentition of living anthropoids; but the canines in this jaw come, as in the human jaw, into alignment with the incisors (Gregory, loc. cit.). There is a diastema (2 mm. on the right side, and 3 mm. on the left side) between the canines and lateral incisors of the upper jaw; but seeing, first, that the incisors are narrow, and, secondly, that diastemata (1 mm.-1.5 mm.) occur between the central incisors of the upper jaw and between the medial and lateral incisors of both sides in the lower jaw, and, thirdly, that some separation of the milk teeth takes place even in mankind (Tomes, "Dental Anatomy," seventh edition) during the establishment of the permanent dentition, it is evident that the diastemata which occur in the upper jaw are small. The lower canines, nevertheless, show wearing facets both for the upper canines and for the upper lateral incisors.

"The incisors as a group are irregular in size, tend to overlap one another, and are almost vertical, as in man; they are not symmetrical and well spaced, and do not project forwards markedly, as in anthropoids. The upper lateral incisors do project forwards to some extent and perhaps also do the upper central incisors very slightly, but the lateral lower incisors betray no evidence of forward projection, and the central lower incisors are not even vertical as in most races of mankind, but are directed slightly backwards, as sometimes occurs in man. Owing to these remarkably human characters displayed by the deciduous dentition, when contour tracings of the upper jaw are made, it is found that the jaw and the teeth, as a whole, take up a parabolic arrangement comparable only with that presented by mankind amongst the higher primates. These facts, together with the more minute anatomy of the teeth, will be illustrated and discussed in the memoir which is in the process of elaboration concerning the fossil remains.

"In the third place, the mandible itself is humanoid rather than anthropoid. Its ramus is, on the whole, short and slender as compared with that of anthropoids, but the bone itself its more massive than that of a human being of the same age. Its symphyseal region is virtually complete and reveals anteriorly a more vertical outline than is found in anthropoids or even in the jaw of Piltdown man. The anterior symphyseal surface is scarcely less vertical than that of Heidelberg man. The posterior symphyseal surface in living anthropoids differs from that of modern man in possessing a pronounced posterior prolongation of the lower border, which joins together the two halves of the mandible, and so forms the well-known simian shelf and above it a deep genial impression for the attachment of the tongue musculature. In this character, Eoanthropus dawsoni scarcely differs from the anthropoids, especially the chimpanzee; but this new fossil betrays no evidence of such a shelf, the lower border of the mandible having been massive and rounded after the fashion of the mandible of Homo heidelbergensis.

"That hominid characters were not restricted to the face in this extinct primate group is borne out by the relatively forward situation of the foramen magnum. The position of the basion can be assessed within a few millimetres of error, because a portion of the right exoccipital is present alongside the cast of the basal aspect of the cerebellum. Its position is such that the basi-prosthion measurement is 89 mm., while the basi-inion measurement is at least 54 mm. This relationship may be expressed in the form of a "head balancing" index of 60.7. The same index in a baboon provides a value of 41.3, in an adult chimpanzee 50.7, in Rhodesian man 83.7, in a dolichocephalic European 90.9, and in a brachycephalic European 105.8. It is significant that this index, which indicates in a measure the poise of the skull upon the vertebral column, points to the assumption by this fossil group of an attitude appreciably more erect than that of modern anthropoids. The improved poise of the head, and the better posture of the whole body framework which accompanied this alteration in the angle at which its dominant member was supported, is of great significance. It means that a greater reliance was being placed by this group upon the feet as organs of progression, and that the hands were being freed from their more primitive function of accessory organs of locomotion. Bipedal animals, their hands were assuming a higher evolutionary rôle not only as delicate tactual, examining organs which were adding copiously to the animal's knowledge of its physical environment, but also as instruments of the growing intelligence in carrying out more elaborate, purposeful, and skilled movements, and as organs of offence and defence. The latter is rendered the more probable, in view, first, of their failure to develop massive canines and hideous features, and, secondly of the fact that even living baboons and anthropoid apes can and do use sticks and stones as implements and as weapon of offence ("Descent of Man," p. 81 et seq.).

"Lastly, there remains a consideration of the endocranial cast which was responsible for the discovery of the face. The cast comprises the right cerebral and cerebellar hemispheres (both of which fortunately meet the median line throughout their entire dorsal length) and the anterior portion of the left cerebral hemisphere. The remainder of the cranial cavity seems to have been empty, for the left face of the cast is clothed with a picturesque lime crystal deposit; the vacuity in the left half of the cranial cavity was probably responsible for the fragmentation of the specimen during the blasting. The cranial capacity of the specimen may best be appreciated by the statement that the length of the cavity could not have been less than 114 mm., which is 3 mm. greater than that of an adult chimpanzee in the Museum of the Anatomy Department in the University of the Witwatersrand, and only 14 mm. less than the greatest length of the cast of the endocranium of a gorilla chosen for casting on account of its great size.

"Few data are available concerning the expansion of brain matter which takes place in the living anthropoid brain between the time of eruption of the first permanent molars and the time of their becoming adult. So far as man is concerned, Owen ("Anantomy of Vertebrates," vol. iii) tells us that "The brain has advanced to near its term of size at about ten years, but it does not usually obtain its full development till between twenty and thirty years of age." R. Boyd (1860) discovered an increase in weight of nearly 250 grams in the brains of male human beings after they had reached the age of seven years. It is therefore reasonable to believe that the adult forms typified by our present specimen possessed brains which were larger than that of this juvenile specimen, and equalled, if they did not actually supersede, that of the gorilla in absolute size.

"Whatever the total dimensions of the adult brain may have been, there are not lacking evidences that the brain in this group of fossil forms was distinctive in type and was an instrument of greater intelligence than that of living anthropoids. The face of the endocranial cast is scarred unfortunately in several places (cross-hatched in the dioptographic tracing - see Fig.5). It is evident that the relative proportion of cerebral to cerebellar matter in this brain was greater than in the gorilla's. The brain does not show that general pre- and post-Rolandic flattening characteristic of the living anthropoids, but presents a rounded and well filled-out contour, which points to a symmetrical and balanced development of the faculties of associative memory and intelligent activity. The pithecoid type of parallel sulcus is preserved, but the sulcus lunatus has been thrust backwards towards the occipital pole by a pronounced general bulging of the parieto-temporo-occipital association areas.

To emphasize this matter, I have reproduced (Fig. 6) superimposed coronal contour tracings taken at the widest part of the parietal region in the gorilla endocranial cast and in this fossil. Nothing could illustrate better the mental gap that exists between living anthropoid apes and the group of creatures which the fossil represents than the flattened atrophic appearance of the parietal region of the brain (which lies between the visual field on one hand, and the tactile and auditory fields on the other) in the former and its surgent vertical and dorso-lateral expansion in the latter. The expansion in this area of the brain is the more significant in that it explains the posterior humanoid situation of the sulcus lunatus. It indicates (together with the narrow interorbital interval and human characters of the orbit) the fact that this group of beings, having acquired the faculty of stereoscopic vision, had profited beyond living anthropoids by setting aside a relatively much larger area of the cerebral cortex to serve as a storehouse of information concerning their objective environment as its details were simultaneously revealed to the senses of vision and touch, and also of hearing. They possessed to a degree unappreciated by living anthropoids the use of their hands and ears and the consequent faculty of associating with the colour, form, and general appearance of objects, their weight, texture, resilience, and flexibility, as well as the significance of sounds emitted by them. In other words, their eyes saw, their ears heard, and their hands handled objects with greater meaning and to fuller purpose than the corresponding organs in recent apes. They had laid down the foundations of that discriminative knowledge of the appearance, feeling, and sound of things that was a necessary milestone in the acquisition of articulate speech.

There is, therefore, an ultra-simian quality of the brain depicted in this immature endocranial cast which harmonies with the ultra-simian features revealed by the entire cranial topography and corroborates the various inferences drawn therefrom. The two thousand miles of territory which separate this creature from its nearest living anthropoid cousins is indirect testimony to its increased intelligence and mastery of its environment. It is manifest that we are in the presence here of a pre-human stock, neither chimpanzee nor gorilla, which possesses a series of differential characters not encountered hitherto in any anthropoid stock. This complex of characters exhibited is such that it cannot be interpreted as belonging to a form ancestral to any living anthropoid. For this reason, we may be equally confident that there can be no question here of a primitive anthropoid stock such as has been recovered from the Egyptian Fayüm. Fossil anthropoids, varieties of Dryopithecus, have been retrieved in many parts of Europe, Northern Africa, and Northern India, but the present specimen, despite its youth, cannot be confused with anthropoids having the dryopithecid dentition. Other fossil anthropoids from the Siwalik hills in India (Miocene and Pliocene) are known which, according to certain observers, may be ancestral to modern anthropoids and even to man.

Whether our present fossil is to be correlated with the discoveries made in India is not yet apparent; that question can only be solved by a careful comparison of the permanent molar teeth from both localities. It is obvious, meanwhile, that it represents a fossil group distinctly advanced beyond living anthropoids in those two dominantly human characters of facial and dental recession on one hand, and improved quality of the brain on the other. Unlike Pithecanthropus, it does not represent an ape-like man, a caricature of precocious hominid failure, but a creature well advanced beyond modern anthropoids in just those characters, facial and cerebral, which are to be anticipated in an extinct link between man and his simian ancestor. At the same time, it is equally evident that a creature with anthropoid brain capacity, and lacking the distinctive, localised temporal expansions which appear to be concomitant with and necessary to articulate man, is no true man. It is therefore logically regarded as a man-like ape. I propose tentatively, then, that a new family of Homo-simiadae be created for the reception of the group of individuals which it represents, and that the first known species of the group be designated Australopithecus africanus, in commemoration, first, of the extreme southern and unexpected horizon of its discovery, and secondly, of the continent in which so many new and important discoveries connected with the early history of man have recently been made, thus vindicating the Darwinian claim that Africa would prove to be the cradle of mankind.

It will appear to many a remarkable fact that an ultra-simian and pre-human stock should be discovered, in the first place, at this extreme southern point in Africa, and secondly, in Bechuanaland, for one does not associate with the present climatic conditions obtaining on the eastern fringe of the Kalahari desert an environment favourable to higher primate life. It is generally believed by geologists (vide A.W. Rogers, "Post-Cretaceous Climates of South Africa," South African Journal of Science, vol. xix, 1922) that the climate had fluctuated within exceedingly narrow limits in this country since Cretaceous times. We must therefore conclude that it was only the enhanced cerebral powers possessed by this group which made their existence possible in this untoward environment.

In anticipating the discovery of the true links between the apes and man in tropical countries, there has been a tendency to overlook the fact that in the luxuriant forests of the tropical belts, Nature was supplying with profligate and lavish hand an easy and sluggish solution, by adaptive specialisation, of the problem of existence in creatures so well equipped mentally as living anthropoids are. For the production of man a different apprenticeship was needed to sharpen the wits and quicken the higher manifestations of intellect - a more open veldt country where competition was keener between swiftness and stealth, and where adroitness of thinking and movement played a preponderating rôle in the preservation of the species. Darwin has said, "no country in the world abounds in a greater degree with dangerous beasts than Southern Africa," and, in my opinion, Southern Africa, by providing a vast open country with occasional wooded belts and a relative scarcity of water, together with a fierce and bitter mammalian competition, furnished a laboratory such as was essential to this penultimate phase of human evolution.

In Southern Africa, where the climatic conditions appear to have fluctuated little since Cretaceous times, and where ample dolomitic formations have provided innumerable refuges during life, and burial-places after death, for our troglodytic forefathers, we may confidently anticipate many complementary discoveries concerning this period in our evolution.

In conclusion, I desire to place on record my indebtedness to Miss Salmons, Prof. Young, and Mr. Campbell, without those aid the discovery would not have been made; to Mr. Len Richardson for providing the photographs; to Dr. Laing and my laboratory staff for their willing assistance; and particularly to Mr. H. Le Helloco, student demonstrator in the Anatomy Department, who has prepared the illustrations for this preliminary statement.

- Dart, R. "Australopithecus africanus: The man-ape of South Africa", Nature, Vol. 115, No. 2884, February 7, 1925

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