australopithecus, homo habilis, tool-using and tool-making

South African Archaeological Society

Australopithecus, Homo Habilis, Tool-Using and Tool-MakingAuthor(s): P. V. TobiasSource: The South African Archaeological Bulletin, Vol. 20, No. 80 (Dec., 1965), pp. 167-192Published by: South African Archaeological SocietyStable URL: http://www.jstor.org/stable/3887824 .

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AUSTRALOPITHECUS, HOMO HABILIS, TOOL-USING AND TOOL-MAKING

P. V. TOBIAS, M.B., B.CH., PH.D., F.R.S. (S.AFR.)

Presidential address to South African Archaeological Society, Cape Town, April 1965

In the 6 years which have elapsed since the Darwin Centenary was celebrated in 1959, new discoveries and new observations have largely re- volutionized our knowledge of man's emergence. Major contributions have been made by the palaeo-anthropological discoveries of the Leakeys at Olduvai Gorge; secondly, important consequences flow from the newer information on behaviour patterns of living Primates and especially, of tool-using and tool-making by non-human Primates, such as the great apes and even monkeys.

In this address, I present some of the newer evidence relating to the structure and affinities of early hominids and suggest the possible consequences for hominid phylogeny. Thereafter, I review what is known about the implemental activities of living non-human Primates and try to assess the relevance of such activities for our understanding of the cultural status of the australopithecines. The supposed association of the australopithecines with osteodontokeratic objects and with early stone tools is critically examined. An attempt is made to create a case for associating (a) australopithecines with osteodontokeratic activities, and (b) the new hominine species, Homo habilis, with the Oldowan culture. Finally, the probable nature of two crucial cultural advances

that of the australopithecines over the pongids, and that of Homo habilis over the australopithecines-is tentatively defined.

This provisional synthesis of a small part of the field of human evolution is offered in the full knowledge that tomorrow's new discoveries may well invalidate today's re-thinking about yes- terday's discoveries. And if Dr. and Mrs. Leakey and Jonathan, Richard and Philip Leakey, con- tinue to work at the breathtaking pace they have maintained for the last 6 years, tonight's thoughts may well not survive the clear light of another day. It is a pleasure to acknowledge my deep gratitude to them for having invited my partici- pation in the exciting task of studying and evalu- ating the Olduvai hominids.

The Discovery of the Taung Skull 40th Anniversary

On the 7th of February this year, we celebrated the 40th anniversary of the publication by Pro-

fessor Raymond A. Dart (1925) of his paper in Nature, describing the discovery and the features of the Taung skull. He christened the hitherto- unknown species to which it belonged Australo- pithecus africanus. In contrast with the earlier discoveries of Neandertal man in Europe (1848 and 1856) and of Homo erectus (or Pithecanthropus) in Java (1891), it fell to Dart to make the crucial discovery of a yet earlier and lowlier creature, so primitive in form as initially to leave serious doubts in practically all minds whether it was indeed a hominid (a member of the Hominidae or family of man) or whether it was simply another ape or pongid (a member of the ape- family or Pongidae). The very fact that the con- troversy over its status raged about the little head of the five-years-old Taung child for a quarter of a century testifies to the borderline position it occupied. Man or Ape? Or an intermediate? Could it, with its burgeoning brain, its large though incredibly human teeth, its balanced head bespeaking an upright though imperfectly human posture, be a real "missing link"? Was it the fossilized embodiment of a transitional form, such as Darwin and Haeckel had envisaged in the 19th century?

Dart came out unequivocally in favour of the "missing link" idea. He called the species represented by the creature Australopithecus africanus -the southern ape of Africa-but declared that it was part of a group which had moved deci- sively in a humanoid direction. It had embarked on a series of bodily changes which Teilhard de Chardin called "hominization".

"The Taung child betrayed the existence of a form so thoroughly intermediate between pongid and hominid as to fulfil all the requirements of a "missing link" . . . It provided the first convincing evidence of the earlier existence of a form transitional between non-man and man." Seen in this light, the discovery of 1924 outshines those of 1848 - 1856 and of 1891; in fact, it is no exag- geration to claim that it is the most important single discovery in the study of human evolution.

Dart's perspicacity is the more remarkable when one considers the prevailing notions of his day. He came to Africa at a time when Asia was commonly believed to be the labour-ward of human emergence. Africa was not deemed to be a probable cradle-except in a vague, half-forgotten

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prediction of Charles Darwin. Not only was Dart in the wrong part of the world to find an ancestral prehuman; when he found it, it was the wrong sort of creature! Those in the know believed that the brain was the vanguard and the pace-maker of human evolution-and none pro- pagated this view more indefatigably than Dart's old teacher and fellow Australian, Elliot Smith: dawn-man when found would have an advanced and sapient brain, but large, primitive teeth and jaws. Small wonder that an industrious forger could provide the world with a Piltdown skull showing just this combination of features-and escape undetected for decades. Dart's child from Taung laughed at the long-held dogma: for its teeth and jaws were man-like, but its brain relatively small and very much in the rearguard of evolution! To assert that this child was on the road to manhood, Dart had to defy the doctrines of the day and even the subtle ties of loyalty and affection which bound him to his old teacher.

"A further factor may reveal the full measure of Dart's contribution. He based his claims on a child-skull, whose teeth told of death at about five years of age. Had the first australopithecine discovered been an adult, Dart's task would have been far easier. For the full characteristics of a hominid show themselves only from puberty on- wards; earlier than that milestone, the hallmarks of the species are only dimly adumbrated. Yet Dart read the signs of manhood in the child, whilst to be convinced, the rest of the world waited for an adult.

"The full story of how the tide of opinion changed has been oft-repeated. The discoveries of adults by Broom and, later, Robinson, the Kitchings and Hughes, and above all, the pains- taking anatomical studies on the fossils by Le Gros Clark, gradually confirmed that the australopithecines were indeed on the pathway of hominization. We know today that structurally they fulfilled all the requirements of an ancestral hominid." (Tobias 1964b.)

After four decades, it is instructive to re-read Dart's paper of 1925. He was in no doubt about the significance of the discovery:

"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" (p. 198).

He noted the forward position of the foramen magnum and devised a "head-balancing index" which he found to be intermediate in value between those of apes and men. The implications of this observation were far from being lost on Dart: this is what he made of it -

"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 pro- gression, 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 role 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 weapons of offence. (Descent of Man, p. 81 et seq.)" (Dart 1925, p. 197).

It was only in the fifties and especially in the sixties, that the world came to consider seriously all these functional implications which Dart had limned in those early and youthful years.

THE DISCOVERY OF OTHER AUSTRALOPITHECINES

Meantime, for about a dozen years, most scientists agreed with Sir Arthur Keith's view that, "like most parents, Professor Dart was over- enthusiastic about his offspring", and that the child was closely related to the anthropomorphous apes, gorilla, chimpanzee and orang - utan (Andrews, ca. 1945). When Broom found an adult australopithecine at Sterkfontein in 1936, he stated:

"I think it likely that Australopithecus will prove to lie somewhere near the common ancestor of the chimpanzee, the gorilla and man . . . Not improbably it will be seen to be a little way along the line that branched off to grow up to man. And there seems no doubt that it is the fossil ape nearest to man's ancestor at present known." (Broom 1937.)

When, in 1938, Gregory and Hellman came to South Africa to study the australopithecine fossils -which by that time included a specimen from a third site, Kromdraai, near Sterkfontein- Gregory was able to state in an address to the Associated Scientific and Technical Societies of South Africa:

"Dr. Dart concluded at that time that his form represented a long step in the direction of the human race; and I do not believe, after

168



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the most critical studies that my colleagues and I have been able to make, that any reasonable exception whatever can be taken to that conclusion. It is the missing link no longer missing. It is the structural connecting link between ape and man." (Gregory 1939.)

TABLE 1: DATES OF DISCOVERY OF AUSTRALOPITHECINE FOSSILS

1924 Taung (S. Afr.) 1936 - 1949 Sterkfontein Type Site (S. Afr.) 1938 - 1954 Kromdraai (S. Afr.) 1939 Garusi (E. Afr.) 1947 - 1961 Makapansgat (S. Afr.) 1948 - 1952 Swartkrans (S. Afr.) 1955 - 1959 Olduvai (E. Afr.) 1957 - 1958 Sterkfontein Extension Site (S. Afr.) 1964 Peninj, Lake Natron (E. Afr.)

The immediate post-war period brought two more South African sites to light, namely Maka- pansgat, from which tell-tale fossilized baboons were recovered in 1945 and the first of some 40 australopithecine fragments in 1947, and Swart- krans, which Broom opened up in 1948 in con- junction with the University of California African Expedition (Broom and Robinson 1952).

These years also brought the news that, in February 1939, Kohl-Larsen had discovered a number of hominid skull fragments in the area north-east of Lake Eyasi in Tanganyika: some of these came from the Garusi gully or korongo and included a fragment of maxilla with two premolars. When Kohl-Larsen published a brief account of these finds in 1943, he mentioned that Abel (who had published a detailed study of the Taung skull in 1931) had identified the maxillary fragment and teeth as those of "Menschenaffe"- man-ape, by which he presumably meant australopithecine. In 1948, Hennig referred to this specimen as Praeanthropus; Weinert (1950) subsequently related it to the Javanese Meganthro- pus, calling the African form Meganthropus africanus. This name was retained by Remane (1951) in his detailed study of the teeth. However, in 1952 Teilhard de Chardin and, in 1953, von Koenigswald revived Abel's old suggestion that the Garusi specimen was actually australopithecine. This was convincingly demonstrated in 1953 and 1955 by Robinson. As Robinson wrote in 1953, "The chief importance of the specimen lies in the fact that it proves that the australopithecines were not confined, to southern Africa." Nevertheless, a still more detailed study of the teeth by Senyurek (1955) led him to conclude that, although the Garusi teeth come closer to the australopithecines of South Africa than to other hominids, there were sufficient differences between the Tanganyikan specimen and those of

South Africa to justify keeping it in a separate genus: he therefore proposed to call it Prae- anthropus africanus, after the generic name proposed by Hennig (1948) and the specific name suggested by Weinert (1950). Senyurek's views have gained no support in the ensuing decade, as far as I can ascertain. In neither of two recent international discussions on the taxonomy of fossil man, one at Burg Wartenstein in 1962 and one at Moscow in 1964, was any serious consideration given to retaining the genus Prae- anthropus. My own study of the published des- criptions and of the cast of the Garusi maxilla and premolars has led me to support provisionally Robinson's conclusion that it belongs to an East African australopithecine. Unfortunately, nothing is known about the date of the Garusi deposit from which the specimen was obtained.

Another scanty suggestion of the presence of an australopithecine in East Africa came in 1955, when a very large molar tooth was found at BK II, now known to be stratigraphically located high in Bed II, Olduvai Gorge (Fig. 6). No fewer than four papers have been written about this one tooth (Leakey 1958, Robinson 1960, von Koenigs- wald 1960, Dahlberg 1960). Some have called it a deciduous and others a permanent molar, some an upper and others a lower tooth! However, although the last word has not yet been written on these points of difference, its australopithecine status would now seem to be common cause. What is more, its probable position in the Olduvai sequence would make it one of the latest australopithecines yet found-for it represents a creature which has survived well into the Middle Pleistocene.

Much more satisfactory evidence of the presence of australopithecines in East Africa came with Mary Leakey's dramatic discovery in July 1959 of a beautiful australopithecine skull at FLK in Bed I, Olduvai Gorge (Fig. 2). Origin- ally called Zinjanthropus boisei by Leakey (1959), it has more recently been proposed to lump it into the genus Australopithecus, though as a distinct species, A. boisei (Leakey, Tobias and Napier, 1964) (Table 2).

No fewer than 14 individuals are represented by hominid fossils found at various levels in the Olduvai Gorge between 1955 and 1963. They cover an enormous span of time, the oldest being now well-dated both by potassium-argon deter- minations and by fission track dating (Fleischer et al, 1965) to approximately 1.75 - 2.00 million years. Some of these fossil hominids fit comfortably into the now well-defined category of the australopithecines, which may be regarded as a partially hominized group. Others, as we shall see presently, represent a more markedly hominized lineage than the australopithecines.

169

2



TABLE 2. GENERA AND SPECIES OF AUSTRALOPITHECINAE

Present Name Older Name/s Locality

A ustralopithecus A. africanus Taung africanus A. africanus Makapansgat

A. transvaalensis Sterkfontein Plesianthropus Sterkfontein

transvaalensis Meganthropus Garusi

africanus Praeanthropus Garusi

africanus

Australopithecus Paranthropus Kromdraai robustus robustus

Paranthropus Swartkrans crassidens

P. robustus Swartkrans crassidens

A ustralopithecus Zinjanthropus Olduvai Bed I boisei boisei

Paranthropus Olduvai Bed I boisei

? Peninj

Finally, in January 1964, the newest australopithecine site, lacustrine beds on the Peninj Stream, on the north-western side of Lake Natron in northern Tanganyika, close to the Kenyan- Tanzanian boundary, yielded a superb mandible of australopithecine affinities (Leakey and Leakey, 1964).

Thus there is a lapse of 40 years between the discovery of the first site, Taung, in 1924, and that of the latest site, Peninj, in 1964 (Table 1).

Some other claimants for australopithecine status

Apart from fossils from these 8 sites, several other fossils have been claimed as possibly australopithecine. For example, a cranio-facial fragment from northern Chad discovered by Coppens in 1961 was claimed to be australopithecine (Coppens 1961a, 1961b, 1962). In 1963, we invited M. Coppens to visit South Africa and study the original australopithecine material. As a result, he has reached the same conclusion as Leakey and I had reached independently, namely that the Chad fragment represents a more advanced hominid than Australopithecus (pers. comm.)

Outside Africa, the possibility has been raised that the teeth and cranial fragments found at Ubeidiya in the Jordan Valley in Israel may have belonged to an australopithecine (Stekelis et al, 1960). From a preliminary study of these scanty remains, generously placed at my disposal

by Professor M. Stekelis, they are highly likely to have belonged to Homo rather than Austra- lopithecus, although it may be impossible, without the discovery of further material, to attribute them to a particular species of Homo. Chemical dating on these pieces is at present being carried out by Dr. K. P. Oakley.

From Java has come another form of early hominid known as Meganthropus palaeojavanicus. Some three or four mandibular fragments of this form have been found in both the Put- jangan and the Kabuh Beds of the early part of the Middle Pleistocene. Robinson has suggested that this Javanese Meganthropus is simply an australopithecine (1953, 1955). Recently, von Koenigswald and I had the opportunity of com- paring the originals of Meganthropus I and II with original material from Africa (Tobias and von Koenigswald, 1964). We concluded that while Meganthro pus has some strong australopithecine resemblances, it shows several features in which it is somewhat advanced beyond the australopithecine grade (Fig. 3). In this sense, it shows a greater degree of hominization than Australo- pithecius, as does Homo habilis of Africa (Fig. 4). We do not think therefore that Meganthropus is australopithecine, as understood in Africa. Its separate generic status is no longer considered justified either (Le Gros Clark 1955). Like Homo habilis in Africa, I think that Meganthropus will have to be re-classified as a member of the genus Homo; for the time being, however, I should not like to commit myself as to whether it should be classed as a distinct species, Homo palaeojavanicus, or else lumped into Homo erectus (which is the present most widely-accepted name of the group known formerly as Pithecanthropus). Noth- ing approaching a total morphological picture of "Meganthropus" is known; we have no upper jaws or upper teeth; we have no brain-case or estimate of cranial capacity; and we have no knowledge of the limb-bones. It would perhaps be best to defer a definite decision as to the species until more adequate material is discovered.

Another group of Asian fossils has been thought to possess australopithecine status, namely a handful of isolated teeth from China attributed by von Koenigswald to Hemanthropus peii (1957). Simons (1963) has suggested that these teeth may be australopithecine. However, they may represent a more advanced hominid, such as Homo habilis (Tobias, 1965a), but it may be impossible to resolve the problem of their status until more specimens are recovered, including teeth in a mandible or cranium.

In sum, the case for an australopithecine stage being represented in Asia is unproven (Table 3); the only convincing australopithecine sites remain the 8 African sites, 5 in the Republic of South Africa and 3 in the Republic of Tanzania (Fig. 5).

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TABLE 3. PLEISTOCENE HoMINOIDS OF UNCERTAIN TAXONOMIC POSITION

Locality Older Name or Suggested Attribution Taxonomic

Assignation

Chad Tchadanthropus Homo sp. Ubeidiya, ? Australopithecine Homo sp.

Israel J ava Meganthropus Homo sp.

palaeojavanicus (? Homo Paranthropus palaeojavanicus)

palaeojavanicus China Hemanthropus ? Homo sp.

peii (Tobias) ? Australopithecus

(Simons) Kwangsi Gigantopithecus ? Pongid (most

blacki authorities) ? Hominid (Wu)

The Distribution of the Australopithecines in Time

The australopithecines are a purely fossil group although it is of interest to note that there exists a "Committee for the Study of the Hairy, non- human Bipeds", which follows up stories of "yetis"

,- i-0PENINJ OLDUVAI

GARUS I

MAKAPANSGAT

STERKFONTEIN SWARTKRANS &

KROMDRAAI TAUNG

Fig. 5. Distribution of the eight australopithecine localities of Africa.

and "hairies", in the hope of finding somewhere a small, surviving colony of australopithecines, clinging to a precarious foothold as living fossils. For the moment, we are on safe ground in declaring that the australopithecines did not out- live the mid-Pleistocene. It would seem from faunal comparisons, as well as stratigraphic evidence, that Kromdraai in the Transvaal competes with BK II in the upper part of Bed II of the Olduvai Gorge as being the latest deposit from which australopithecine remains have been ob-

tained. Both of these deposits are classified as of mid-Pleistocene age and are characterized by a fauna which is post-Villafranchian. On the terminology proposed by Cooke (1961) and Wells (1962), the Kromdraai faunal deposit would be assigned to the Swartkrans Faunal Stage. That sets a later limit to the known survival of the australopithecines (Table 4).

TABLE 4. DISTRIBUTION OF AUSTRALOPITHECINES IN TIME AND SPACE

(Highly provisional-as at April, 1965) Africa

South East

Middle Kromdraai Olduvai IL Pleistocene (Upper)

Peninj (Lake Natron)

Swartkrans Olduvai 1I (Middle)

? Sterkfontein Extension Site

Lower Olduvai II Pleistocene (Lower)

Makapansgat ? Garusi

Sterkfontein Type Site

Taung Olduvai I

On the other hand, there are two South African competitors, and one East African, for the oldest deposit bearing an australopithecine. New un- published faunal analyses by Dr. Basil Cooke, who has kindly allowed me access to his manu- script, suggest that the available fauna from Sterkfontein Type Site and from Taung permit of two possible interpretations-either that Taung is slightly older than Sterkfontein or vice versa. Either way, both sites belong to the Sterkfontein Faunal Stage, which preceded the Swartkrans Stage. In the Sterkfontein stage only some 30 % of species, or probably fewer, are tentatively identified with living forms; in other words, 70% or more of the species are extinct (Wells 1962).

At present, it is difficult to say whether these two South African sites are older or younger than the oldest East African australopithecine site, FLK in the middle of Bed I at Olduvai. The newer faunal analyses just published by Leakey (1965) certainly show that the Bed I fauna is far more archaic than the earlier analyses by Hopwood (1951) indicated. This re-assessment has been made possible partly by the accretion of better information on ranges of variation within certain species, but mainly by the discovery of new species. In the 1965 Olduvai volume, 152 different mammals are. listed and reference is made to a further 10 new species not yet listed, whereas the 1951 Olduvai volume listed only 51

171



mammals. it is possible on this new faunal evidence to say, with Simpson (1965), that "the oldest of these faunas probably belongs somewhere in the latter part of the Villafranchian . . . that is, sometime towards the end of what is now formally and somewhat arbitrarily designated as early Pleistocene". Such a description would be essentially true as well of Cooke's and Wells's Sterkfontein Faunal Stage from South Africa.

Within any one major geographical zone, faunal comparisons are valuable among the sites in that zone. However, faunal comparisons between the East and South African sites are somewhat viti- ated by the large distance and ecological differences between these two major zones. Any scheme, such as that presented here, must therefore remain highly provisional. It is of interest to note that, following the publication of Volume I of the new Olduvai series, with some of the palaeontological evidence, Howell (1965) has recently commented: ". . . it seems increasingly likely, as I have for some time maintained, that the occurrences of Australopithecus africanus in Southern Africa antedate the earliest hominids known from Olduvai Gorge."

Apart from possibilities of absolute dating, the discovery and excavation of geographically intermediate sites could place correlations between East and South Africa on a more secure footing: herein lies part of the potential importance of Desmond Clark's new site in Malawi (Nyasaland), to be excavated later this year.

How much earlier than the Upper Villafran- chian phase the australopithecines came into being, we do not know, because of the paucity of fossils from the Pliocene. It is of interest and possible relevance that, as Simons (1961, 1964) has pointed out, the jaws of a Mio-Pliocene hominoid, Ramapithecus pun jabicus, are very similar to those of Australopithecus. We have an African relative or representative of Rama- pithecus from Fort Ternan in Kenya, which Lea- key (1961a) has called Kenyapithecus. Perhaps it is to Ramapithecus (including Kenyapithecus) that we should look for the ancestry of the australopithecines. If this proved true, it could provisionally be concluded that the australopithecines, as such, emerged sometime between the Lower Pliocene and the latter part of the Lower Pleistocene. In terms of potassium-argon dates, that gives a time range of about 10 million to just under 2 million years B.P., within which the Australopithecinae emerged. Their extinction may very tentatively be set at somewhat less than one million years ago.* * I do not include Meganthropuis palaeojavanicus,

either from the Putjangan Beds or from the Kabuh Beds, as australopithecine (Tobias and von Koenigswald 1964); otherwise the range would have to be extended up to 2 million years B.P. (Oakley 1964b).

Lower Pleistocene Hominines

For some time, it has proved convenient to classify all the Pleistocene and Recent hominids in two sub-families, the Australopithecinae (or australopithecines) and the Homininae (or hominines) (Chart 1).

CHART 1.

Family: HOMINIDAE

Subfamilies:

I AUSTRALOPITHECINAE HOMININAE

Until recently, the australopithecine chapter was considered largely to precede the opening of the hominine chapter of hominid phylogeny. Thus, while the australopithecines were essentially a Lower Pleistocene manifestation surviving into the Middle Pleistocene, the hominines were considered not to appear before the beginning of the Middle Pleistocene. The earliest hominines recognized until recently were the fossil hominid remains found along with a Djetis Fauna in the Putjangan Beds of Java: and Hooijer (1952) has shown that these beds are most probably Middle Pleistocene. In his new work on the dating of fossil man, Oakley (1964b) does not estimate the age of the Djetis fauna, but ascribes an age of about 500,000 years to the later Trinil fauna (after von Koenigswald 1962). So the Djetis beds would exceed I million years old. At the same time, Oakley makes clear that he no longer regards the dating of tektites in the Upper Trinil Beds of Java as having any valid bearing on the antiquity of Homo erectus. Kurten (1956) has quoted the presence of a subspecies of Hyaena brevirostris in the Djetis Beds as suggesting that these Beds are of late Villafranchian date, that is, Lower Pleistocene, but faunal dating from a single species is beset with many dangers. The differential survival of different species largely rules out such a conclusion, and Hooijer's interpretation of the Djetis Beds as early Middle Pleistocene is now widely accepted.

Until the recent discoveries at Olduvai. Homo erectus in the Djetis Beds was the earliest and morphologically the least hominized member of the Homininae (Table 5). Thus, it could safely be claimed up to a year ago that there was no evidence for the existence of a hominine earlier than the Middle Pleistocene. Again, it could be

172



TABLE 5. GENERA AND SPECIES OF HOMININAE

Present Name Older Name/s Locality

Homo habilis Olduvai Bed I

Homo erectus Pithecanthropus Java Sinanthropus Chou-Kou-Tien

Lantien Atlanthropus N.W. Africa "Homo leakeyi" Olduvai Bed II Telanthropus Swartkrans

Homo sapiens H. s. neander- Homo Europe

thalensis neandertha- Asia lensis

Homo primigenius

Homo Africa rhodesiensis, etc.

H. s. sapiens Homo Europe sapiens Asia

Africa America Australia

said that in the Lower Pleistocene, there was no evidence for the existence of any hominid more advanced than an australopithecine. It was tempt- ing to infer from these two statements that, late in the Lower Pleistocene, some populations of australopithecines underwent further hominizing changes-especially an increase in brain size and a decrease in the size of teeth and jaws-and that these changes were of sufficient magnitude to carry the populations concerned across the next taxonomic frontier into the realm of the hominines.

This viewpoint was based on the assumption that it was but a short step from the most advanced australopithecines to the lowliest Homo erectus. I shall presently question that assumption and try to show that the gap between Australo- pithecus and Homo erectus was bigger than had commonly been assumed.

In 1959, shortly before the discovery of A. (Zinjanthropus) boisei in Bed I at Olduvai, there came to light a left lower third molar embedded in a fragment of mandible and an isolated lower premolar from a site called MK I about 8 feet above the basal lava. This level was appreciably lower in Bed I than the living-floor at FLK I, from which the type-cranium of A. boisei was obtained the same year (Leakey and Leakey 1964, Leakey, Tobias and Napier 1964). In fact, no stratigraphically lower hominid remains have yet been found at Olduvai (Fig. 6). They come from

a layer lower than that for which a K/A date of 1.75 m. years has been obtained. Yet the MK I teeth are on the small side and show a narrowing and elongation not characteristic of australopithecine teeth. Their discovery was, however, soon overshadowed by that of Zinjanthro-

BED

MIDDLE PLEISTOCENE

s Vtlli~~~~~~~~~6ff~raxshn) Ha. erctu

>5 A. specl a

BED

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LN%--- FAUNAL BREAK

Fill LOWER Hominid sD.

Hlomoind sp PLEISTOCENE

w w BED A boisai H. habilis

(type)

F71 ia H. habilis NH.habli

1 75 mn. (t ype)

H. habilis

" 777 17/777 /7 77/77//7/77

OLDUVAI SEQUENCE TO SHOW APPROXIMATE VERTICAL POSITION

OF HOMINID REMAINS

Fig. 6. Schema of the lower part of the Olduvai sequence, showing the approximate vertical positions of hominid remains in Beds I and II.

pus. Nothing was published on the very different teeth of MK I.

In the following year, further evidence came to light of a second type of hominid, completely different from A. boisei. Some of the remains were of relatively small teeth found on the same living floor as the massive-toothed A. boisei. Although these isolated teeth came from a later level and a different site from MK I with the oldest specimen, these contemporaries of A. boisei showed the same kinds of departure from the australopithecine dental pattern, as did the MK I teeth. An upper molar showed similar lengthening and a marked narrowing, such that its buccolingual width of 12.3 mm. was smaller than the widths of every one of 67 australopithecine upper first and second molars. Similarly, an upper

173



lateral incisor substantially exceeded in its length- breadth index the most elongated of 13 australopithecine 12's Thirdly, a lower premolar from this living floor at FLK I was both absolutely and relatively narrower than every one of 44 australopithecine lower premolars-the narrowest of which was a good 1I mm. broader than the FLK tooth.

Thus, as contemporary of A. boisei there lived an individual whose teeth differed in size from those of A. boisei by the whole range of all the South African australopithecines. Clearly, this was no mere sex difference: the disparity was so great as to justify regarding the smaller-toothed individuals as belonging to a totally different population from A. boisei. Not only a size difference, but a shape difference, distinguished the second hominid from all known australopithecine teeth, including those of A. boisei. Corresponding dental departures had characterized the earlier teeth from MK I: here, clearly, was another individual of a somewhat earlier period whose tooth size and shape veered in the same direction away from those of australopithecine teeth.

And still nothing was published. The handful of teeth constituted an enigma and a challenge: we sat on them and bided our time. We did not have long to wait.

Later in 1960, there came to light another set of hominid remains from FLK NN I (Leakey 1960). They were found on a living-floor some 14 inches lower than the A. boisei floor (Fig. 6), and that is why for some years we called the specimen pre-Zinjanthropus. The remains com- prised the greater part of a juvenile mandible with teeth, an isolated upper molar, large parts of two juvenile parietal bones and a number of juvenile finger bones. In addition, at least one adult was represented by further hand-bones, foot-bones and probably the clavicle. To our astonishment we found that the teeth-both those in the lower jaw and the isolated upper tooth- showed the selfsame departures from the australopithecine teeth. Thus, the upper molar was absolutely longer than 15 A. africanus MI's and its length-breadth index (102.3) likewise exceeded the top of the A. africanus range. Similarly. the lower teeth showed a marked lengthening, both absolutely in the case of 1,, V and P and relatively in respect of P,, P., M, and MW. Thus. the L/B index of P. exceeded the values for all 23 australopithecine Pu's; that of P, exceeded the indices for all 21 australopithecine P 's; of M, both left and right teeth on Robinson's measurements (though only one of the two on mine) exceeded all 32 australopithecine M, 's; while of M, the index exceeded those for 11 M,'s of A. africanus, but was exceeded by only 1 out of 15 MW's of A. robustus and A. boisei.

The same dental trends which characterized

the earlier MK I individual and the later FLK I specimen also typified the intermediate "pre- Zinjanthropus" individual. All showed departures from the australopithecine features of general dental enlargement and especially dental broaden- ing (Robinson 1956)-departures which were in a hominine direction. The K/A dating suggested that this group of dentally-distinct creatures persisted for at least 100,000 years, from about 1.75 m. to 1.65 m. B.P. Clearly, they must represent a distinctive strain of hominids. This deduction was reinforced when a series of spectacular discoveries in 1963 (Leakey and Leakey 1964) showed that corresponding dental features persisted right through into Bed II times, that is, into the Middle Pleistocene. The same hominizing departures from the australopithecine pattern were present, though in an intensified or even more markedly hominized form (Fig. 7). We could think at this stage in terms of an ancient and long-lasting lineage characterized by increasing degrees of hominization of the teeth.

But we were to learn that the hominizing trend was not confined to the teeth. The parietals, which we could confidently associate with the juvenile mandible, were far bigger than any australopithecine parietals thus far obtained. The sagittal, coronal and lambdoid chords and arcs of these parietals exceed those for all available australopithecine parietals, although they apparently belonged to a juvenile of not more than 11 - 12 years of age. As Leakey (1961b) pointed out, "these parietals suggest that we are dealing with a hominid with a larger brain capacity than Zinjanthroputs". It is interesting to note, too, that in 1962, Robinson apparently accepted these parietals-which he described as "too large for either type of australopithecine" (1962a, p. 105) as evidence that something "more advanced" than the australopithecines was living at their time level (op. cit., pp. 102, 105, 106).

It was clearly tremendously important to estimate the cranial capacity and to do so in a more reliable manner than the rather wild guessing which had earlier conjured up figures such as 700 and 750 - 800 cc for two child crania from Swartkrans and "probably over 800" and "probably over 1,000" for two adult specimens (Broom and Robinson 1952). Not one of these estimates on the robust australopithecines was acceptable, largely owing to the incomplete or crushed state of the specimens. The first of the robust australopithecines to provide a fairly accurate estimate was A. boisei, on which we determined a capacity of 530 cc (Tobias 1963)-surprisingly to some, no larger than that of the gracile australopithecines!

It was not difficult to reconstruct the arch formed bv the two parietal bones, as all four edges of these quadrilateral bones were preserved on one or other side, and some partly on both.

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A reconstruction by A. R. Hughes and myself in Johannesburg differed by not more than a milli- metre or two from one made independently by Leakey and myself in Nairobi (Fig. 8). A partial

~~~~....#......

Fig. 8. The biparietai calvariai arch reconstructed from the parietai fragments of the type specimen of Hlomo habilis, together with the contained partial endocraniai cait.

endocranial cast was made from each of the two reconstructions and its volume was determined by displacement of water. Five readings on the first cast ranged from 362 to 364 cc and gave a mean of 363.6 cc; while five readings on the second ranged from 362 to 365 cc with a mean of 363.4 cc. The virtual identity of the two sets of readings gave us confidence that the reconstructions were not very inaccurate.

The next problem was to determine what proportion of the brain size lies under the two parietals: we made a series of total and partial endocasts of australopithecines and Homo erectus specimens and, in each case, compared the two values. A relatively narrow range of readings was obtained-from 50.22 % to 56.54%?S. It was thus possible to compute the probable total volume of the pre-Zinjanthropus endocranium (Fig. 9): the figures ranged from 642.7 cc to 723.6 cc (Tobias 1 964a). It is not surprising that

even the smaller of the two values exceeded the largest australopithecine capacity, namely the estimate of 600 cc* for the Taung child if it had survived to adulthood. Thus, the estimated cranial capacity merely confirmed what had been obvious from the parietals-that they were part of a larger cranium than that of any known australopithecine. For practical purposes we selected the middle value of 680 cc. (Table 6). Clearly, this represented an advance on the australopithecine brain-size-and thus a further advance in a hominizing direction. In other words, the same creature which showed hominizing tendencies in its teeth also showed a more hominized brain-size.

so27 ~ 7243c

. .~~~~~~5 . 646c

Fig. 9. Estimates of the total endocranial volume from the biparietal partial endocranial volume - type specimen of Homo habilis.

Meantime, our colleagues in London, John Napier, Peter Davis and Michael Day, had shown how very hominine the limb-bones were (Napier 1962, Davis, Day and Napier 1964). Hence, although we cannot say whether the limb-bones showed an advance on those of Australopithecus, for lack of adequate remains of the latter, it is possible to conclude that as much as we can determine of the total morphological pattern of the hominid is markedly hominine in aspect, and, where adequate comparative materials are available, shows an advance on the australopithecine pattern. The teeth had already shown that at least some features of this hominine trend persisted for a considerable time.

In the face of all this evidence, we felt justified in concluding that we were dealing here with a new taxon. It was a taxon which was more hominized than Australopithecus, but not as hominized as Homo erectus. Was there room for a new species between these two stages in hominization? That is, if we compared the most man- like of the australopithecine species, namely A. a;ricanus, with the lowliest species of Homo then recognized, namely H. erectus, was there sufficient morphological distance between them to allow of an additional species being interposed?

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TABLE 6. CRANIAL CAPACITY OF EARLY HOMINIDS

Taxon Capacity Reference

Australopithecine Sterkfontein 1 435 Schepers 1946 Sterkfontein 5 480 Broom, Robinson

& Schepers 1950 Sterkfontein 7 480-520 Broom, Robinson

& Schepers 1950 Sterkfontein 8 530 Broom &

Robinson 1948 Makapansgat 38/39 480 Dart 1962 Taung-as child 500-520 Dart

Schepers 1946 -as an adult Le Gros Clark

(estimated) 570-600-624* 1947 Le Gros Clark

1955 Dart 1956

Olduvai 5 530 Tobias 1963 (Zinjanthropus)

Homo habilis Olduvai 7 680 Tobias 1964a Homo erectus "Pithecanthropus" I 935 Weidenreich 1943 "Pithecanthropus" II 775 Weidenreich 1943 "Pithecanthropus" III ca. 880 Weidenreich 1943 Choukoutien II 1030 Weidenreich 1943 Choukoutien III 915 Weidenreich 1943 Choukoutien X 1225 Weidenreich 1943 Choukoutien XI 1015 Weidenreich 1943 Choukoutien XII 1030 Weidenreich 1943 Olduvai 9 1000 Tobias 1964c

("Chellean Man")

When we examined this question more closely, we found that the two species had been assumed to be closer to each other than, indeed, the facts would allow. In the formulations of Le Gros Clark (1955) and of Arambourg (1958), for instance, it seems to have been taken for granted that A. africanus graded insensibly into H. erectus. Nevertheless, in tooth size and shape and in cranial size and capacity, there remains a sizeable gap between the two species ranges, despite fairly good samples (Table 7). In tooth size and shape, for instance, there is a much closer fit between A. africanus and A. robustus, than there is between A. africanus and H. erectus (Figs. 10 and 11). The patterns in the latter two species are widely divergent, as the graphs demonstrate. In brain size, too, there is a gap between the two groups of fossils (Fig. 12). And, of course, in time there is a gap-for A. africanus is essentially a Lower Pleistocene form and H. erectus a

*Later re-estimated at 562 cc. (Tobias (1965) Nature, 208, 206).

TABLE 7. NUMBERS OF AUSTRALOPITHECINE TEETH AVAILABLE FOR STUDY (1965)

Taung ...... ..... .... 24 Sterkfontein ... ...... 162 Kromdraai ...... ...... 39 Swartkrans ...... ...... 311 Makapansgat .. ...... 55 Garusi ......2 .... ..... 2 Peninj (Natron) ..... 16 Olduvai ... ..... ...... ?38

Total ...... 647

Middle Pleistocene form. We concluded that on the presently available samples of fossils in the two groups, there is sufficient morphological distance between them to accommodate an additional species. Of course, future discoveries of australopithecines and of H. erectus fossils may narrow the distance between the two species; if that happens, the taxonomic position of the new hominid may have to be re-considered. Meantime, however, on the present data, the erection of a new species seems the most reasonable taxonomic

,7- - BUCCOLINGUAL BREADTHS OF

MANDIBULAR TEETH

16--

6- ARBUSTUS

52- - l t 3 ? l 1

Fig. 10. Sample ranges of buccolingual breadths of mandibular teeth in A. africanus and A. robustus.

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BUCCOLINGUAL BREADTHS OF

MANDIBULAR TEETH

...~

~~~~~~~~~~..... o ..~~~~~~~~~~~~~~~~~~......

Fig.~

~~~~~~~~~~~~~~~+///

A.P3as

N

1

2 3 ~~~~~~~~~~~~~. . . 4 1 2 3

Fi.11. Sample ranges of buccolingual breadths of mandibular teeth in A. africanues and H. erectus.

procedure to adopt. It is, too, a compromise between various extreme viewpoints which have been expressed: such as, that the new hominid

Fig. 12. Ranges of cranial capacity in samples of Australo- pithecus (based on 6 specimens of A. africanus and 1 of A4. boisei) and of Homo erectus.

should be regarded as a new species of Australo- pithecus or a new genus between Australopithe- cus and Homo or a new subspecies of Homo erectus. The latter classification would claim a greater measure of hominization for the new hominid than we have so far dared to suggest.

Accordingly, we have proposed a new species, Homo habilis, to accommodate the Bed I fossils which were in the hominine tradition. The name "habilis" means "handy, skilful, able" and was suggested to us by Professor R. A. Dart.

Before coining the new name and creating the new species, we had subjected the specimens to study, measurement and comparison for at least four years. During all that time, we were re-

peatedly urged by our colleagues to name the new fossils or at least to indicate where their affinities lay. It was not until January 1964- about 5 years after the discovery of the MK I habiline and 4 years after that of the three habi- lines from FLK I and FLKNN I-that we finally decided upon the creation of a new species, when the conclusions we had reached from a study of the earlier specimens were seen to be borne out and even strengthened by the batch of new specimens discovered late in 1963. Homo habilis made its debut through the pages of Nature in April 1964 (Leakey, Tobias and Napier 1964).

We at first included the Bed II hominines as paratypes of H. habilis, but from further study by von Koenigswald and myself (1964), we have concluded that the Bed fl specimens have taken the hominizing trends so much further as to have crossed the next taxonomic barrier: we now regard them as being more closely related to Homo erectus (Fig. 7). Our decision to place the Bed I hominines into the genus Homo was based primarily on the anatomical and functional considerations mentioned; but we felt the decision was supported by the strong suggestive evidence that Homo habilis made the Oldowan stone tools. This brings us to a consideration of the culture of Lower Pleistocene hominids.

Cultural capacity There are several sides to this question. Firstly,

what indirect inferences may be made from the bodily structure of the Australopithecinae? Secondly, what direct inferences can be made from the associated objects found with the australopithecines? Thirdly, what deductions may be made about australopithecine behaviour from a study of the behaviour patterns of other Pri- mates?

The questions are closely interrelated: When Dart first suggested that the australopithecines were capable of violent manual activities (1926, 1929) and of using and fabricating tools of bone, horn and teeth (1957b), relatively little was known of the extent to which other higher Primates were capable of tool-using and even, to an extent, of tool-making. Since that time, more and more information has accumulated on such primate activities, and proportionately, the resistance to attributing tool-using and tool-making activities to the australopithecines has lessened. Perhaps it may be useful to review briefly some of the recent information on the implemental activities of non-human Primates.

The implemental activities of monkeys and apes

The capacity for tool-using is widespread, not only in the higher Primates, the great apes, but also in lower primates. Numerous reports exist of the use of objects as fighting tools as well

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3



as for non-agonistic purposes: these reports relate to behaviour both in captivity and in the wild. For example, Bolwig (1961) reported on the tool-using activities of a captive olive baboon from the Sudan: these included purposeful throwing of sticks, the raking in of food beyond arm's reach and the breaking of sticks to make a "ladder" with which to reach titbits hanging overhead. The remarkable feats of that clever little New World monkey, the capuchin, have been known at least since 1882, when G. J. Romanes reported the use by a Brown Capuchin of the flat bottom of a dish to crack walnuts (cited by Weiner 1963). Kluver (1937) had likewise reported on the capuchin's versatility and quickness in the use of a variety of differently- shaped sticks, as well as wire, rope, cardboard, boxes and other objects for obtaining food out of reach. Osman Hill (1960) has summarized the available evidence in the volume of his Primates series which deals with the Cebidae. A further example is the use by a male capuchin in the London Zoo of a large marrow-bone to crack open Brazil nuts and almonds, when it was no longer able to do so with its teeth-which it had used when young (Vevers and Weiner 1963). Re- cently, we were able to confirm such behaviour by a capuchin in the Johannesburg Zoo; in the company of Dr. M. Lyall-Watson, zoologist to the Zoo and staff members and science students of the Anatomy Department, I observed two capuchins trying to open walnuts with their teeth; one succeeded but the other failed. The second animal then laid the walnut on the floor of the cage, picked up a stone and cracked the walnut open after a few well-aimed blows.

Kortlandt and Kooij (1963) have most recently reviewed what they describe as "Protohominid Behaviour in Primates". Analysing a number of reports on agonistic throwing, scooping, clubbing and stabbing by New and Old World monkeys and gibbons, they find these activities best developed in ground-living genera, such as baboons, and types which live predominantly in more open and diversified habitats, including rocky landscapes. Such behaviour is less characteristic of more arboreal genera which live mainly in closed canopy forests: the only obvious exception to this rule being the capuchin. They concluded that, as is true of the great apes, a semi-terrestrial life and a more or less open habitat in the wild tend to promote the evolution of the genetic basis for the ability to perform aimed agonistic throwing, and/or for the actual inclination to do so in captivity-to which generalization the capuchin remains the exception.

Conversely, however, as Weiner (1963) has nointed out, the tool-using abilities of a tree- living form like the capuchin emphasize the point

that a bipedal terrestrial mode of life per se is not a necessary pre-adaptation for the emergence of tool-using activities. He adds, though, that it must have been crucial for the change from tool- using to tool-making. I shall later question the latter view of Weiner, on the basis of observations on higher Primates.

Abundant observations have now accumulated, not only on tool-using by the great apes, but even on tool-fabricating. As Le Gros Clark stated, in his recent Raymond Dart Lecture (March 1965), "it has now become clear that they (modern large apes with their small brains) are capable of more elaborate patterns of behaviour than had hitherto been suspected".

The classical studies on chimpanzee behaviour by Kohler (1924), Yerkes (1948) and others, have in recent years been supplemented by a host of new studies. The facilities provided by the Wit- watersrand University's Uganda Gorilla Research Unit enabled Donisthorpe (1958), Osborn (1963) and others to study the mountain gorilla, while Carl Akeley's Africa provided the venue for the comprehensive studies on the mountain gorilla by G. B. Schaller (1963). At the same time, the field studies of Kortlandt (1962) and of Jane Goodall (1963a, 1963b, 1964) have thrown much new light on the behaviour of chimpanzees. These field researches have been supplemented by the zoo observations summarized by Kortlandt and Kooij (1963), the experiments of the Hayes' (1952) with Viki, and of Mrs. N. Kohts (1935) with her child's gorilla contemporary and playmate, and the new exploration by H. F. Khroustov (1964) or the highest frontier of implemental activity in chimpanzees.

From this wealth of new information, we may select certain key observations bearing on tool- using and tool-making or, if one prefers it, tool- modifying activities by the great apes. Tool- iising activities-such as agonistic throwing, clubbing and stabbing--characterize all three of the great apes, chimpanzee, gorilla and orang-utan, as Kortlandt and Kooij (1963) have well summarized recently. Non-agonistic uses by anthropomorphous apes include isolated instances of branches being used by gorillas and one orang- utan as raking tools for reaching fruit or other objects (Kortlandt and Kooij 1963); Beatty (1951) observed a chimpanzee pick up a rock and break open a dried palm nut; Merfield (1956) watched a group of chimpanzees poking sticks into an underground bees' nest and then licking off the honey; while Kortlandt and Kooij (1963) have reported the use of leaves as "medicated pads" by a chimpanzee and of small sticks and fruit as "toilet" aids by another chimpanzee. Goodall saw chimpanzees use natural objects as tools on many occasions. For instance, she saw chimpanzees use sticks to feed on different species of

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ants (1964): the nests were opened, a stick was thrust in, left for a moment, and then withdrawn with the ant-mass on the end. The delicate morsel was licked off with the lips. Termite "fishing", however, provided her with one of her crucial observations. In those months (November to January, approximately), when the termites extend their passages to the surface of the nest, chimpanzees plucked stalks and small twigs, pushed them into the openings on the surface of the termitarium, withdrew the tools and picked off the insects with their lips. They would move away from the termite heap to pluck grass stalks, carry back one or even several, and settle down for an hour or two's peaceful "fishing". "Some- times tools were carefully prepared: leaves were stripped from stems or twigs with the hand or lips, and long strips were sometimes pulled from a piece of grass which was too wide" (Goodall 1964). As the straw became bent at the end, the chimpanzee would break off the bent pieces until the tool was too short for further use. In other words, we have here evidence not only for the use of natural objects, but also for the modification of natural objects to render them more suitable for the purpose to which they are put.

Another form of rudimentary tool-fabrication consisted in the use of leaves as drinking tools. Chimpanzees would drink water from natural bowls until the level was too low for the water to be reached with the lips. Thereafter, leaves were stripped from twigs and chewed briefly, giving a crumpled surface. Then the leaf-mass, held between the index and middle fingers, was pushed into the bowl, withdrawn and the water sucked out of it. The process was repeated, until the bowl was empty or until the chimpanzee lost interest (Goodall 1964).

Cultural Primates By modifying natural objects in this way, the

chimpanzee may be said to have reached the first crude beginnings of tool-making. Goodall (1963b) has suggested that "It is unlikely that this practice of fishing for termites is an inborn behaviour pattern. Among higher primates, behaviour is found to depend more and more on learned techniques and less and less on 'instincts'. It seems almost certain that this method of eating termites is a social tradition, passed from ape to ape by watching and imitation. As such, it must be regarded as a crude and primitive culture." (op. cit., p. 308.)

Kortlandt and Kooij (1963) would agree with this view. According to the laboratory experiments and zoo experience they have reviewed, the behaviour of the great apes is formed largely by maternal education, social traditions and other environmental factors, rather than by innate fac-

tors as in monkeys. "This applies," they point out, "to locomotion, nest building, food choice, sexual behaviour, social intercourse, etc., and, to some extent, even to maternal care." (op. cit., p. 61). Therefore, according to these workers, the great apes and man might be characterized as "cultural Primates", while the gibbons and monkeys would be classified as "instinctual Primates", provided that we may use the term "culture" for a non-verbalized system of social traditions. (Un- fortunately, there exists no appropriate term to designate non-human, speechless culture.) (op. cit., p. 62.)

In other words, these primatologists align the apes with man because their behaviour patterns are largely learned, rather than genetic or instinctual. Of course, there is no sharp dividing line. There must be a smooth transition from the instinctual to the learned patterns of behaviour, and even in modern man there is often doubt as to the degree to which he has thrown off instinctual patterns and substituted learned ones for them. However, Kortlandt's distinction between the cultural Primates and the instinctual Primates is a useful one and is in line with other recent evidence which is tending to show a much closer relationship between the great apes (especially the African ones) and man. These new evidences include data on the chromosomes and on the serum proteins of the apes. To such evidence, as well as that of comparative anatomy, it seems, must now be added ethological evidence, data drawn from the field of animal behaviour.

The limits of implemental activities by apes

Further evidence of tool-making activities among chimpanzees has been provided by the experiments of H. F. Khroustov in Moscow, as reported to the VIIth International Congress of Anthropological and Ethnological Sciences held in that city in August 1964. In an attempt to explore the limits to which implemental activity could extend in apes, he devised an elegant series of experiments on chimpanzees. A food-bait was placed in a metal tube, of such diameter and length that the chimpanzee's fingers and lips could not reach the bait. Then, to get at the bait, the chimpanzee was faced with a series of situations of increasing complexity and which made increasing demands of tool-using, tool- modification and tool-making for their solution. The series of experiments may be summarized as follows:-

(1) A straight smooth rod was provided, that is, an implement fully suitable for the action required by the experiment. The chimpanzee had no diffi- culty in getting at the bait.

(2) The same rod was provided, but with a cross-piece at one end. The chimpanzee did not

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immediately find the correct solution. The idea that the stick had to be inserted into the tube had, apparently, fixed itself in the animal's mind: but no more than that, for it at first grasped the stick by the free end and attempted to insert the end with the cross-piece into the metal tube. Later, it tried the other end with success.

In both these preliminary experiments, success could be achieved without any modification of the tool.

(3) In the third stage, some modification of the tool was necessary for achievement of the final result. The rod was furnished with two cross-pieces, one at either end. After unsuccess- ful attempts to use it directly as an implement, the animal broke off one cross-piece: however, the earlier associations were not sufficiently fixed in its mind and it then attempted to thrust into the tube the other end with intact cross-piece! Only after the failure of such attempts did the animal find the correct way to use the rod. In other words, it seems that the connection had not been established at this point between the changes which it wrought on the implement, and the possibility of using it as an implement. To quote Khroustov, "The destructive actions of the anthropoid on the one hand and its attempts to perform implemental action on the other, remained separate operations not united into a chain. Only by establishing the connection between these operations and their organization into a single chain do the destructive actions receive definite direction, turning into a directed treatment of the implement and thus forming a link in the whole chain of activity. Further experiments led the chimpanzee to hit upon the connection that had escaped it before." It was given a tree branch with a side-shoot: it broke off the shoot and so gained the bait with the branch. When it was again given the rod with two cross-pieces, after some hesitation and several abortive trial-and-error actions, the connection seems suddenly to have been established, for it grasped the rod, immediately broke off both cross- pieces and the string binding them, and then used the free rod as an implement.

It seems that this stage fixed in the mind of the animal the idea of treating an object to adapt it for an action which the animal tried to perform with its help.

Up until now the elements of the object were clearly manifest to the chimpanzee; but in ensuing experiments, the features of the implement were progressively obscured.

(4) The chimpanzee was provided with a long, thin, rectangular plate. In this case, the material given was not structurally divided into a ready implement and elements which hindered its use. Now it was necessary for the animal to break the object: this it did in a variety of experiments,

until one of the resulting fragments suggested the form of the original rod-shaped implement.

(5) In the next experiment, a circular disc of the same sort of wood was provided: thus, nothing was manifest of the form of the necessary implement. The animal was required to transform the disc into certain parameters, not on the basis of any indications on the disc, but solely on the basis of information about them fixed in the mind of the ape. The chimpanzee proved equal to this task.

(6) It was noticed that the chimpanzee before breaking the disc turned it so that the grain lines of the wood ran away from its body. Hence, it was subsequently given discs with imitation grains running across the true grain. At first, the chimpanzee began to operate according to the illusory grain; finding the disc more difficult to break, it changed the direction of its actions and was successful.

(7) The disguise was made more complete: grooves were carved on the disc at right angles to the grain of the wood. Both surfaces of the wood were then covered in thick paper, with the grooves forming a clearly discernible relief; furthermore, black lines were drawn between the furrows perpendicular to the grain. Again, the illusory grain misled the chimpanzee; but after repeated trials-and-errors, it changed the direction and had success, assisted by its teeth making a number of consecutive dents along the true grain.

Khroustov was able to conclude that "active transformation of material having a preliminary neutral form, even despite its false outer appear- ances, into an implement with definite parameters, is a fully attainable stage of activity for a chimpanzee". In Khroustov's opinion, this is the level of implemental activity characteristic of the australopithecines. "Its coming into being signifies not the appearance of the human principle, but a high stage of its functional preparation, which is a necessary prerequisite for the transition to the primary forms of labour, as an activity pecu- liar to human beings." We shall return to this aspect when we discuss the implemental activities of the australopithecines.

(8) Finally, once the chimpanzee had mastered the full manufacturing of the implement under the conditions just outlined, further experiments were devised to find the limits of his implemental activity. A similar series of objects was given to the animal and he had to make the same implement. Only this time, the rods and discs were of tough oak wood, which the animal was unable to break with its natural equipment of hands, feet and teeth. A stone intermediary, in the form of a Chellean hand-axe was provided, to assist the chimpanzee break the tough wood. Numerous and persistent attempts were made by

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the chimpanzee to break the wood, but none of them made any attempt to treat the unyielding material with the stone implement, or with anything other than their natural or bodily equipment. For the first time in the whole series of experiments, an attempt was made by demon- stration to teach the chimpanzees to use the stone tool. Even then, no attempt was made to copy the actions of the experimenter and use the stone implement, despite the most strenuous and pro- longed efforts of the chimpanzee after the demon- stration to overcome the resistance of the material. It seems that the decisive step of using a tool to make a tool was impossible for the apes under the experimental conditions.

Whether an ape could overcome this barrier under other conditions can be determined only by further experiment. Meantime, Khroustov concludes that "The limited scope of using the natural organs as means for making adaptational changes of implements tied down the develop- mental activity of man's ancestors to definite boundaries and possibilities. When these were exhausted, the immediate forebears of man by force of necessity were brought to the threshold of the frontier, on the surmounting of which depended their survival"-that is, the use of objects as means for transforming materials into implements-and so the frontier itself: "the creation of artificial means for producing implements."

Thus, on a somewhat narrower definition of human cultural behaviour than that of Kortlandt and Kooij, Khroustov draws a line between the apes and man and aligns the australopithecines in this regard with the apes.

Wherever the line is to be drawn and indeed, whether it is at all possible to draw a line, it is clear that the level of implemental activity which the living great apes can attain in the wild, in captivity and under experimental conditions, far exceeds what had earlier been thought. These new bodies of information culled by specia- lists in primate behaviour have scarcely made their impact yet upon the interpretation of archaeological problems and of australopithecine cultural potentialities. That is why I make no apology for having dwelt on them at such length here: for they must alter the climate of opinion in palaeo-anthropology and thev must influence the way in which *we consider the claims made for the australopithecines. In fact, the accretion of this new information in the last decade has been Parallelled by a steadv decline in the resistance to claims made about implemental activity of the australopithecines.

IMPLEMENTAL ACTIVITY OF THE AUSTRALOPITHE-

CINES : ANATOMICAL CONSIDERATIONS

Even if the Australopithecinae showed no greater degree of implemental activity than living

great apes, it is clear trom the foregoing that a considerable range of cultural activities would have been within their capacity. Nobody, to my knowledge, has suggested that ape-men were less capable than apes of implemental activity! Much indirect evidence however, points to the possibility that australopithecines were culturally advanced as compared with apes. The first line of evidence comes from the bodily structure of australopithecines.

(a) Teeth: The absence of large canine teeth strongly suggests that the australopithecines must have used alternative mechanisms to solve those sorts of problems for which apes use their teeth.

(b) Brain size: The mean cranial capacity of the australopithecines is 508 cc., that of the large- bodied gorilla 498 cc. and that of the smaller chimpanzee 411 cc. Thus, the mean brain-size of the australopithecines is absolutely larger, despite its much smaller body, than that of the great apes. Jerison (1963) has suggested a method of dividing brain-size into two components: one of which depends on body size, whilst the other comprises "excess" nerve cells available for increasing the range of adjustments. In very broad terms, the extra nerve cells are a measure of the degree of development of the portion of the brain related to intelligence. When Jerison's for- mulae are applied to apes and to australopithecines, it is seen that while African great apes have some 3.4 to 3.6 thousand million of extra nerve cells, the australopithecines, on various estimates of their body weight, would have had some 3.9 to 4.9 thousand millions of extra nerve cells, i.e. on the average about 1,000 m. extra nerve cells more than the apes (Table 8). Since Homo habilis is estimated to be only 1,000 m. more than that again, and the lower limit of Homo erectlus is about 11 thousand million extra nerve cells more than the upper limit of the australopithecines (Fig. 13), the potential significance of the jump of 1,000 m. from ape to Aus- tralopithecus is seen to be appreciable. It is a reasonable assumption that such a jump would have given the australopithecines some advantage over the apes in cultural potential, although it is always well for us to remember, with Dobzhan- sky (1963), that "We cannot administer I.Q. tests to our remote ancestors, and can only engage in nebulous speculations about their cultural capacities". But, he adds, "it is flying in the face of evidence to overlook this increase of the brain- size as an indication of an important adaptive trend in human evolution" (op. cit., p. 312).

Selection for larger brains, as Bielicki (1964) has pointed out, was not strong at the cultural level characterised by tool-using (with little role played by tool-making), for the anthropomorphous apes could manage this with relatively small brains. But these selective pressures must have

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TABLE 8. ESTIMATES OF "EXTRA NEURONES" IN HoMINOIDS

(After the method of Jerison 1963)

Estimates of total Estimates of extra neurones neurones

(in thousands of (in thousands of Brain Size millions) millions)

African great apes 320-685 (752)* 4.3-5.7 3.4-3.6 Australopithecines 435-600 4.6-5.7 3.9-4.9 Homo habilis 680 (middle est.) 6.2 5.3-5.5

643 (lowest est.) 6.0 5.1-5.3 724 (highest est.) 6.5 5.6-5.8

Homo erectus 775-1225 6.7-9.3 5.8-8.4 Homo sapiens ? 1276- + 1455 + 9.4- + 10.3 + 8.4- + 9.2

* Includes a single large male zoo specimen.

become very powerful after the invention of tool- making: in the time-interval between the australopithecines and Neandertal man, the braincase expanded almost threefold. Later, again, it seems that the culture-brain feedback slackened con- siderably.

The interpretation offered by Bielicki (1964) is that, in the Middle Pleistocene, progress in technology was rather closely dependent on the evolution of the brain. "Man's tool-kit could increase, so to speak, only as much as the growth of

genetically determined mental capacities permitted" (and conversely). "But this dependence gradually weakens, and with the emergence of anatomically modern man the evolution of brain has apparently reached some 'critical point', at which there has been accumulated a 'surplus intelligence', a vast-and at that time, only partly utilized- reservoir of potential capacities for acquiring and enriching culture; as the result of this, further very dramatic cultural advance became possible, without at the same time invoking the necessity

EXTRA" NEURONES OF HOMINOID BRAINS

HOMO SAPIENS

HOMO ERECTUS

HOMO HABILIS

AUSTRALOPI THECINES

AFRICAN GREAT APES

U i -4~~I

3 4 S 6 7 8 9

Extra" . neuronts (in thousands of millions)

Fig. 13. Estimates of "extra nerve-cells" (in thousands of millions) in hominoids-following the method of Jerison (1963).

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of an equally rapid improvement of man's mental endowment. In other words, cultural advance has been largely freed from the rigid control of gene- tical determinants and has acquired, temporarily at least, the character of a 'self-propelled' process." (op. cit., pp. 2-3.)

(c) Bipedalism-and the importance of sitting upright

The australopithecines were essentially bipedal; they did not depend upon their hands in locomotion to the same extent as the apes and presumably therefore were more capable of manipula- tion. Did this freeing of the hand for essentially manual activities release a greater potential for cultural development?

To quote Washburn and Howell (1960), "Dar- win and many workers after him stressed development of bipedal locomotion as a factor in diffe- rentiating man from ape. This process freed the hands, made possible the use and manufacture of tools, and led to reduction in the size of the teeth and the facial skeleton. The australopithecines in general represent such a stage in human evolution." (op. cit., p. 35.) But they modify this view by suggesting a page or two further on, "Perhaps, as Darwin suggested, tool use is both the cause and effect of hominid bipedalism, and the evolution of erect posture occurred simul- taneously with the earliest use of tools" (op. cit., p. 37).

These views remind us that there must have been a subtle reciprocal feedback relation between culture and its genetic bases, between physical and cultural evolution (Dobzhansky 1963, Bielicki 1964). Did the genetic basis of culture appear before culture? is a question we often hear asked. But Dobzhansky points out that this is a wrong way to ask the question; culture and its genetic basis developed, and are developing, together. Bielicki (1964), following Washburn (1960), says, "It seems, for instance, that feedback between bipedalism and tool-using . . . was at work only in Pliocene proto-hominid apes, and faded out at the beginning of the Pleistocene, since in the Australopithecinae the process of acquiring bipedal gait had already been com- pleted and further evolution of culture did not demand any further change in the mode of locomotion".

The views of some workers are, however, at variance with this suggested reciprocal feedback and are couched in terms of a more rigid dependence of implemental activities upon bipedalism and erect posture. Thus, as quoted earlier, Weiner (1963) has suggested that a bipedal mode of life, though not essential for the emergence of tool- using activities, "must have been crucial for the change from tool-using to tool-making" (p. 117). Robinson (1962a) goes so far as to state, "Effec- tive tool-using could only have become possible

after erect posture had been acquired". On the evidence from ape studies cited earlier,

the conclusion is inescapable that upright posture and bipedal gait were not a necessary prerequisite either to effective tool-using, or even to the beginnings of tool-making. The fact that chimpanzees, which are not bipedal, can make tools both in the wild and under experimental conditions, proves that bipedalism is no sine qua non of tool-making, let alone tool-using. It suggests, too, that tool-making has a longer history in Primate phylogeny than has bipedalism.

More important, bipedalism is not the only circumstance under which a creature's hands are freed for such activities as tool-using and tool- making: all that an animal needs to do is to sit upright, for its hands to be available for implemental and other activities. In fact, most of the implemental activities of man and of the great apes are carried out in a sitting position. It is in this position that our bodies find greater stability than in the standing position. Stability, it is suggested, is a most important structural and functional consideration in the development of manual skills. Sitting upright is widespread throughout the Primates (except the Prosimii); it characterises arboreal genera such as the capuchin monkey and terrestrial ones such as the baboon. It is in the sitting position that a great part of the manipulative activities of Jane Good- all's chimpanzees are carried out, as her films and photographs clearly show. Sitting is an activity which frees the hands for implemental and other activities as surely as bipedalism does. This means that the postural basis for tool-using and tool-making is widespread in the Primate order, although bipedalism is not.

If sitting upright is an adequate postural basis for implemental activities in the Primates, what extra element did bipedalism add? Bipedalism seems clearly to have catalysed the implemental process, for now a creature could use or make tools not only whilst sitting down, but also whilst standing. A larger proportion of the life of a Primate could now be occupied with manipulative activities, since standing and walking need not have interrupted the process. A second possible catalysing aspect of bipedalism is that the hands, freed from a major rale in standing and walking, must have undergone morphological changes in a hominizing direction. Let us take a closer look at this aspect.

(d) The hand Structurally, the hands of the great apes are

very different from those of man, especially in the relative proportion of thumb and fingers. Yet it is important to remember, with Napier (1960), that great apes are capable of a "precision grip". Because of the structural difference from the hand of man, the nature of the precision grip in apes

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is different from that in man; but, as in man, it depends upon the interaction of fingers and thumb, as Napier has demonstrated (1960). Thus, small objects are picked up by apes from a flat surface in one of several different ways - (i) between the pulp of the thumb and the side of the middle or terminal phalanx of the index finger, (ii) between the side of the thumb and the side of the middle phalanx, and (iii) between the dorsum of the thumb and the dorsum of the terminal phalanx. A fourth method was to pick up a small object between the middle phalanges of the index and middle fingers (op. cit., pp. 652-653). The power grip is produced in chimpanzees by flexion of the fingers alone. The manual mechanisms of the apes, it seems, are adequate to the task of rudimentary tool- making. Thus, not only the postural but also the manual mechanisms of the great apes provide an adequate anatomical basis for the observed implemental activities.

Just how hominized the hand of Australopithe- clls was we cannot yet say, but even if no more hominized than that of a pongid, it is clear that Australopithecus like the pongids must have possessed the manual basis for tool-making of a sort. As for H. habilis, Napier (1962) has shown that it almost certainly possessed at least two out of three "essential osteological correlates" of the modern human precision grip: as compared with the hands of adult living great apes, this is a partially hominized hand. It would not be un- reasonable on general grounds to suggest that the structure of the australopithecine hand was probably not as hominized as that of H. habilis -which shows more hominization than Australo- pithecus in at least two other directions (teeth and brain-size). However, this argument cannot be pressed in view of the mosaic character of the evolutionary process. Thus, the feet of H. habilis are manifestly more hominized (nearer to those of modern man) than the hand or the teeth. Only more specimens can answer the problem of what sort of hand Auistralopithecus possessed.

(e) Dental crowding

As indirect evidence bearing on implemental activities of Australopithecus, Oppenheimer (1964) has cited the crowding of the front teeth shown in several australopithecine mandibles. Crowding arises, according to Oppenheimer, when the functional demands upon the jaws are significantly diminished. He dismisses the possibility of a major change in dietary habits as leading to reduction of the jaws. Rather he concludes that it was a behavioural revolution, such as the use of tools for butchering animals, cutting up and pounding roots and vegetable food, peeling tough- rinded fruits, preparing skins and the like-thus

significantly reducing the r6le of the masticatory apparatus. Weapons likewise took the place of teeth in defence and the dynamics of dominance, according to this writer. Hence, on this view, we are led to infer his dependence upon tools from the very crowding of the teeth in his mouth, no less than from the reduced size of the canines.

The foregoing formidable array of facts points to a battery of well-developed implemental activities of Australopithecus. What does the direct evidence indicate? Associated objects have led to two main lines of thinking: (1) osteodontokeratic activities; and (2) stone tool-making activities of the pebble tool or Oldowan pattern.

1. OSTEODONTOKERATIC ACTIVITIES OF AUSTRALO- PITHECUS

At the Third Pan-African Congress of Pre- history in 1955 at Livingstone, Dart (1957b) made his first public announcement about what he called the Makapansgat australopithecine osteodontokeratic culture. Subsequently, he elaborated this theme in the Report of the Smithsonian Institute for 1955 (Dart 1956a) and in a series of articles (1956b, 1957a, 1957b, 1958, 1959a, 1959b, 1959c, 1960a, 1960b, 1961a, 1961b, 1961c, 1962a, 1962b), as well as in a monograph (1957c). The facts and Dart's claims are too well known for me to need to review them. Only a few points need be mentioned here:-

(1) The analysis of many thousands of bones from the Makapansgat deposit has shown definite evidence of selection of bones, not only from different parts of the body, but of different parts of the same bone. Thus, the ratio of proximal to distal humeral fragments is 33:336, while the ratio of humeri to femora is 518:100.

(2) Tooth-marks of hyaena and porcupine are absent from all but a handful of the many thousands of bones.

(3) Eighty per cent. of over 50 baboon skulls from Taung (21), Sterkfontein (22) and Maka- pansgat (15) show signs of damage by localized violence.

(4) Large concentrations of ungulate humeri show damage inflicted before fossilization on the epicondyles, some of which fit the occasional doubly-indented fracture depressions of the baboon crania.

(5) Many of the bone flakes show signs of differential wear and tear along one edge or at one end, but not on the other.

(6) A number of special cases include horn- cores and smaller long bones rammed and lodged up the marrow cavities of broken larger bones.

(7) In several instances, small bone and even stone flakes have been wedged between the con- dyles of long bones.

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(8) Many of the long bones show evidences of having been broken by a kind of spiral tor- sional stress.

(9) Some fragments suggest that stalactites and/ or stalagmites were broken off, presumably for use, and some were further fractured.

(10) There is some suggestion of stone-collecting habits: a small number of quartz and quartzite pebbles and fragments have been found in the breccia.

(11) So far, the 41 hominid fragments identified are all of Australopithecus. We have no evidence for any other hominid having lived in or near the Makapansgat Limeworks caves during the time that the Limeworks deposit was accumulating.

Comparable masses of osteodontokeratic objects have not been reported from Taung or Sterk- fontein, but in the former case, the deposit seems to partake more of the nature of a fissure than of a cave deposit, while in the latter, it cannot be said that more than a fraction of the breccia has been thoroughly searched for broken bone fragments, other than taxonomically identifiable parts. In the Swartkrans cave, where we have large numbers of australopithecine remains, as weil as a few pieces associated with Homo crectus, associated faunal remains are conspicuous by their paucity. This would be in keeping with the vegetarian habits which Robinson (1962b) claims characterised the larger-toothed australopithecines he calls Paranthropus; it would also be in keeping with the australopithecine in that instance having been the hunted rather than the hunter, australopithecine remains having perhaps provided a major item in the diet of the early Transvaal H. erectus (formerly called Telanthro- pus).

Thus, the mere absence of hitherto detected osteodontokeratic objects from other South Afri- can sites does not weaken Dart's claims based upon Makapansgat.

At this stage in our knowledge, it would seem that Dart's hypothesis is the only one which can explain a great many lines of evidence.

Would this kind of activity be in keeping with what might have been inferred from the animal studies quoted earlier? To answer this, we have to consider the mechanism which might have been used to break the bones in the typical patterns described by Dart. On the basis of a suggestion and a series of experiments by Trevor Jones, Dart (1959a) claimed that one bone was used to strike another bone in order to fracture it and make the so-called "cannon bone scoops". If this were correct, it would represent precisely the stage of tool-fabrication which Khroustov's chimpanzee had not been able to master. But, when Dart reported on these matters at the 4th

Pan-African Congress on Prehistory at Leopold- ville in 1959, J. Desmond Clark suggested that the same effect could be achieved by Australo- pithecus simply by bashing his bones up against the rock sides and floor of the cave, in much the same way as chimpanzees today bash pieces of stick on the floor (Clark 1962). This seems a plausible enough explanation of the mechanism and, if correct, would imply that Australopithecus had gone no further than could the experimental chimpanzees in Moscow, at least as far as this kind of bone tool-making was concerned.

In sum, I reiterate that Dart's overall hypothesis on the osteodontokeratic activities of Aus- tralopithecinae is the most reasonable and plausible explanation of the otherwise almost inexplic- able mountains of bones, with their selected, fractured and patterned characteristics at Maka- pansgat. Furthermore, this kind of activity is entirely within the somatic possibilities which might be inferred on a priori grounds from behaviour studies on anthropomorphous apes and from anatomical studies of the degree of hominization of the australopithecines themselves. Diffi- culties do remain, it is true; one does not feel that one can go all the way with Dart in accepting all that he has claimed for the osteo- dontokeratics. In fact, I am not sure if it is justified to ascribe the epithet Osteodontokeratic Culture to the implemental activities of the australopithecines. Is this exactly what the prehistoric archaeologists mean by a culture? Perhaps this becomes a mere matter of definition, but I cannot help feeling that the phrase "osteodontokeratic activities" would more fittingly describe what in fact seems to be deducible.

2. WAS AUSTRALOPITHECUS THE MAKER OF THE OLDOWAN "PEBBLE-TOOLS"?

It has long been known that pebble-tools were made in South Africa during the period of the australopithecines. Partly because of this con- temporaneity and partly because of the discovery of a fragmentary australopithecine maxilla and a few possible pebble-tools in the same layer near the top of the Makapansgat deposit, partly too because of the a priori considerations of ape behaviour, the idea has grown up in the last decade or so that Australopithecus was responsible for the pebble tools. This view has been espoused by Oakley since 1951 (1951, 1964b), by J. D. Clark (1963, 1964), by Washburn (1959) and Washburn and Howell (1960), and others. On the other hand, Mason (1957, 1961, 1962) has doubted whether the australopithecines were stone tool-makers, and so have Robinson (1957, 1962a), von Koenigswald (1961), Inskeep (1959) and others.

The lithicultural evidence was not convincing from Makapansgat; but in 1956, Brain discovered

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4



stone implements in what has since come to be known as the Sterkfontein Extension Site. Further implements were recovered by Robinson the following year, in the breccia of the Extension Site, along with some hominid fragments identified by Robinson as australopithecine (1957, 1962a).

But what seemed at first to be the most convincing evidence for the association of australopithecines with early stone tools came from Olduvai, where, in 1959, Dr. and Mrs. Leakey found the cranium of A. (Zinjanthropus) boisei on a living-floor along with broken bones and crude artefacts of the Oldowan Culture. A. boisei was at once hailed as the maker of the stone implements (Leakey 1959). This evidence has clearly influenced some of the above-quoted views in favour of the australopithecine authorship of the Oldowan Culture.

Subsequently, however, teeth of Homo habilis were found on the same living-floor and Leakey (1961b) then stated, "If I am right in believing that the juvenile from F.L.K.N.N.I. is not an australopithecine, but a very remote and truly primitive ancestor of Homo, then it is possible (and I stress the fact here that I only use the word 'possible' at this stage of the inquiry) that it was this branch of the Hominidae that also made the Oldowan tools at the site F.L.K.I. where Zinjanthropus boisei was found". (op. cit., p. 418.)

The fact that Leakey changed his mind about the authorship of the Oldowan tools has been levelled at him as a criticism (Oakley 1964a). Yet, Leakey's second thoughts would seem to be not only legitimate and beyond reproach, but scientifically correct. At the time when A. boisei was the most advanced creature known alongside the tools, it was reasonable to suggest an association between them. But once a more advanced hominid is identified, no matter what label we apply to it, it is surely more reasonable to attribute the making of the stone tools to the more advanced hominid.

In his 1962 paper in the S. African Archaeo- logical Bulletin, Robinson was at pains to demonstrate that the australopithecines were not capable of tool-making of the level seen in the Sterk- fontein industry. He stated:-

"If it can be proved that australopithecines occur in direct association with a stone industry over a significant period of time when no evidence whatever exists of the presence at that time level of anything more advanced than the australopithecines, then there will be a sound case for regarding the australopithecines as tool-makers. In fact there is evidence throughout the entire australopithecine period either proving or suggesting the presence of a more advanced hominid." (Robinson 1962a, p. 102.) In the light of his recent comments (Robinson

1965), it is interesting to note that as his first

example of this more advanced hominid, he went on to quote the large parietals in the 'pre-Zinj' level of Olduvai, i.e. those of Homo habilis!

Again, elsewhere in the same work, Robinson states, "stone tools can only be attributed to australopithecines with any certainty if it is known that nothing more advanced was living at their time level. This is not the case . . ." and he goes on to cite "the apparently much earlier presence at Olduvai of a hominid with parietal bones too large for either type of australopithecine" (op. cit., p. 105.)

Seen against the background of these comments by Robinson, the procedure of preferring Homo habilis to A. boisei as the probable fabricator of the Oldowan tools is perfectly reasonable. When one takes all the evidence bearing on the relation between Australopithecinae and stone tools into account, it becomes not only reasonable, but the hypothesis which meets more of the facts than any alternative.

Let us critically examine the evidence, a sum- mary of which was presented at the Moscow Congress in August, 1964 (Tobias 1964c) (Table 9).

TABLE 9. ASSOCIATIONS OF EARLY HOMINID FOSSILS AND OF EARLY STONE IMPLEMENTS

Presence Presence of australo- of more Presence

pithecine advanced of stone fossils hominids tools

Site / Deposit Sterkfontein

Type Site Yes Sterkfontein

Extension Site Yes ?Yes Yes (Middle Breccia)

Taung Yes Makapansgat Yes ? Swartkrans Yes Yes Yes Kromdraai Yes Garusi Yes Peninj Yes Olduvai Bed I

MKI - Yes Yes FLKNNI Yes Yes FLKI Yes Yes Yes

Olduvai Bed II MNKII (Lower) Yes Yes MNKII (Upper) ?Yes BKII Yes LLK Yes Yes

Olduvai: Bed I:

Three Oldowan living-floors or levels have yielded identifiable hominid remains; at two of these levels the only hominid found has been H. habilis. On one floor, H. habilis and A. boisei have been found together. Another Oldowan level in Bed I has yielded one hominid toe

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phalanx, but we are at present unable to assign this to A. boisei or H. habilis.

Olduvai: Bed II: According to Mary Leakey, developed

Oldowan and early Acheulean remains occur at various levels in Bed II and here, too, an advanced hominine has been found, as well as some probable australopithecine remains. Thus, hominine remains have been found in at least four stone implement-bearing levels or sites in Beds I and II, while remains of Australopithecus occur at or near two of these levels as well. That is to say, at every level or site where stone tools occur alongside hominid skeletal remains, these remains include a more advanced hominid, irrespective of whether, in addition, australopithecine remains are present. Even apart from its morphological advancement, this would suggest that the hominine rather than Australopithecus was the confirmed tool-maker.

Taung: Australopithecus present; stone tools absent;

hominine absent.

Makapansgat: A ustralopithecus present; stone tools absent if

we exclude the handful of specimens claimed by van Riet Lowe (Brain et. al. 1955), since these have failed to convince most prehistorians; more advanced hominid absent.

Sterkfontein Type Site: Australopithecus present; stone tools absent;

more advanced hominid absent.

Swartkrans: A. robustus crassidens present; stone tools have

been reported from this site (Brain 1958); the more advanced hominid present is that known formerly as Telanthropus and now designated Homo erectus.

Sterkfontein Extension Site: Stone tools definitely present. The so-called

middle breccia from this site has yielded 6 fragments of hominid (Robinson 1957, 1958, 1962a), all of which Robinson has attributed to Australo- pithecus. A closer examination of the published accounts and measurements of these fragments has suggested to me that, while some of them may justifiably be attributed to Australopithecus, they include several isolated teeth which seem to belong to a more advanced hominid. Unfortu- nately, no illustrations of these 6 crucial fragments have been published. Measurements have been published for only three of the teeth (Robin- son 1962a), and two of these-the lower canine and the upper first molar-fall outside the A.

af ricanus range of variation for several metrical features reflecting size and shape, but can be comfortably accommodated within the H. habilis range. Furthermore, apart from the teeth referred to in the three papers by Robinson, it seems that 2 further fragments were recovered which have not, to my knowledge, been published: part of the crown of a left upper molar, probably M2, and the lingual half of a left P3. The latter is clearly labelled in the Transvaal Museum card index of specimens as "Homo erectus (Telanthro- pus)?". It is concluded that there is a trace, however slight, of the presence of a more advanced hominid in the middle breccia of the Sterkfontein Extension Site, as well as an australopithecine remnant or two and stone implements.

Kromdraai: A. robustus present; as far as is known, no

stone tools; no trace of a more advanced hominid.

Garusi: Australopithecine present, apparently as a sur-

face find; faunal and cultural associations uncertain.

Peninj: Australopithecine present, but no cultural or

hominine remains from the jaw site. However, scattered stone artef acts and two early Acheu- lean sites have been found in nearly contemporary strata of the Peninj beds, but 3 Km and 9 Km removed from the mandible site proper (Isaac 1965).

Mason (1962) has rightly warned against specu- lating on the identity of the tool-maker in terms of negative evidence; while Oakley (1964a) has reasoned that "Only in certain circumstances would the evidence that they (the australopithecines) were tool-makers be found. For example, they might be expected to occur at an occupation- site such as Sterkfontein Extension site, but not in Karst-hole debris such as that composing the breccia at Sterkfontein Main Site". This seems to be a somewhat circular argument, since the Extension Site is presumably described as an occupation site for the simple reason that tools are found there; while this status is denied to the Type Site presumably because none has been found there! In any event, the two sites are continuous with each other and really comprise a single cave deposit.

Irrespective of the nature of the deposits it is surely of some significance that: -

(1) At every australopithecine site at which stone tools occur in close association with hominid remains, there is evidence of the sym- patric co-existence of a more advanced hominid;

(2) at all sites where we find the co-presence of Australopithecus and a more advanced hominid, there too we find stone tools;

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(3) at every site where stone tools are f ound, along with associated hominid remains, these hominid remains include a more advanced hominid, whether or not australopithecine remains are present in addition;

(4) at every early site which has yielded a more advanced hominid, stone tools are present.

Unless we are to resort to a series of special pleas, the most reasonable hypothesis to explain these data is that Aristralopitheclis was not the maker of the stone implements, but that more advanced hominids almost certainly were.

This conclusion is in no way intended to pre- judge the issue of how many kinds of more advanced hominids were involved; present evidence suggests that the earlier hominine was of the species Homo habilis, whereas the later one was of the species Homo erectus.

The evidence of these cultural-hominid associations, coupled with the a priori considerations advanced by Robinson and Mason and those which we have inferred from morphological and comparative evidence, all conspire to leave little doubt that the earliest and most primitive stone implements found in Africa belonging to the Oldowan Culture, were made by the hominid we have called Homo habilis. He seems to have been a maker of stone implements to a set and regular pattern, which, moreover, shows some changes over a long period of time. Such behaviour characterizes the other species of Homo, namely crectus and sapiens; it has nowhere been convincingly demonstrated to be characteristic of A ustra/opithecius.

CONCLUSIONS

We have seen that structurally the australopithecines are more hominized than the great apes: they have, on the average, somewhat bigger brains in somewhat smaller bodies, so that their endowment of extra nerve cells exceeds that of the great apes by about one (American) billion on the average. Furthermore, they possess the general Primate structural basis for implemental activities, namely the ability to sit upright and probably to manipulate both powerfully and to an extent precisely; but in addition, they possess the trait of bipedalism, although, as Chopra (1961, 1962) has shown, they were imperfectly adjusted skeletally to plantigrade, bipedal stance and gait. But then, as Straus (1962) has suggested, even modern man has not effected a perfect transition, as witness his liability to frequent breakdown of the structural mechanism, by the development of flat feet, hernias, slipped discs, and so on. Never- theless, the greater degree of bipedalism which the anatomical facts permit us to attribute to Australopithecus must have potentiated his implemental activities.

Our analysis has led us to conclude that his

techniques included tool-making, and that osteo- dontokeratics provided his major cultural outlet. This does not preclude his having tinkered with stone, but we find no convincing evidence that Australopitheclus was responsible for the first distinctive stone culture, the Oldowan. The makers of these Oldowan tools must have mastered one intricate conceptual and technological mechanism: the ability to use a tool to make a tool. The osteodontokeratic objects, it seems to me, can be adequately explained without postulating that other tools were used in their fabrication. In other words, we do not need to postulate that the australopithecines had gone beyond the stage of using their natural organs or the floor and walls of their caves for making their osteodontokeratic objects. If there is no reason to suppose that this highest implemental frontier had been crossed in their bone implemental activities, it is a fortiori highly unlikely that it would have been crossed in lithicultural activities. This indirect reasoning, therefore, strengthens the conclusion reached from skeletal-cultural associations, that Auistralopithecuis was not a cultural stone tool- maker.

We may validly enquire: wherein lies the cultural advance of australopithecines over the great apes if they could not or did not cross the highest implemental frontier of the apes? I suggest that the answer resides in the frequency of implemental patterns of behaviour. In apes, tool-using and tool-making are infrequent, sporadic; it cannot be said that the apes' way of life is built around such implemental activities. Survival does not depend on implemental means, but rather on formidable natural defence mechanisms and on the sheltered forest habitat of the living great apes.

The australopithecines, on the other hand, lived in a habitat providing little natural protection and they had no natural weapons of offence or defence like large canines. Their implemental activities had come to loom very largely in their pattern of adjustment. Indeed, it would not be too much to claim that their very survival depended on implemental activities. This, it is suggested, is the great step forward of the australopithecines over the apes.* They learned to exploit a mental and manipulative capacity, a

* Since writing the above, I have come across an interestingly relevant passage in the paper entitled "Ecology and the Protohominids" by Bartho- lomew and Birdsell (1953): "Rather than to say that man is unique in being the "tool-using" animal, it is more accurate to say that man is the only mammal which is continuously dependent on tools for survival. This dependence on the learned use of tools indicates a movement into a previously unexploited dimension of behaviour . . ."

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cultural potentiality, which even apes possess. And they exploited it so effectively that they became dependent on it for survival. Cultural capacity was the greatest evolutionary asset of the australopithecines: and it was on this aspect of their form and function that selection operated with the greatest vigour.

Small wonder then that, sooner or later, some among the numerous Africa-wide populations of australopithecines did acquire the mental capacity to overleap the highest implemental frontier of Khroustov. Sooner or later, some of them did acquire the right quantity and/or quality of brain to be able to use a tool to make a tool. Initially, perhaps, this would have been an isolated flash in the pan, but if it conferred a sufficient selective advantage, the capacity for it would have spread -with a definite feedback interplay between the genetic basis for learning such behaviour and the performing of this more intricate type of implemental activity. This new development clearly did not require a very much bigger brain than that possessed by Australopithecus - perhaps another (American) billion nerve cells were sufficient.

Now a new major breakthrough occurred. Stone tool-making of the complexity of an Oldo- wan culture became feasible: new, virtually limit- less possibilities opened up. A new kind of man was born-Homo habilis-with a new set of implemental capacities, achievements and frontiers.

The recognition of Homo habilis brings three important consequences in its wake.

Firstly, the new primitive species of Homo provides us, for the first time, with a knowledge of the makers of the Oldowan Culture.

Secondly, the finding of a Homo alongside an Australopithecus in the earlier Pleistocene must modify somewhat our conception of hominid phylogeny. If both lines stemmed from a common ancestry, and there is scarcely room for doubt that they did, then the division into two or more lineages must have occurred earlier than had been imagined. Previously, it was still possible to claim that the lineage of Homo stemmed off from an australopithecine lineage within the Pleistocene and even fairly late in the Lower Pleistocene. Now, we have to push these hominine origins back, at least into the later Pliocene, or the very beginning of the Pleistocene (Fig. 14).

Thirdly, there previously remained a large morphological gap between Australopithecus africanus and Homo erectus, and in several respects ranges of variation of the two species did not overlap even slightly, despite fairly large samples. Homo habilis has now provided us with a morphological intermediate, of which the range of variation for various morphological traits undoubtedly overlaps with those of both Australopithecus africanus and Homo erectus.

pien sapien

w

-J 0.

W A b . WHho b itins 0

Fig. 14. The stages of hominization-the darker the shading. the nearer to modern man. A - hypothetical ancestral australopithecine of the Plio-

cene. A.a. - Australopithecus africanus. A.b. - Australopithecus boisei. A.r. - Australopithecus robustus.

It may thus be claimed that, from every point of view, its position in space and time, its mor- phology and inferred behaviour, Homo habilis bridges the last remaining major gap in the Pleis- tocene part of the story of human evolution.

ACKNOWLEDGEMENTS

I am deeply indebted to the Wenner-Gren Foundation for Anthropological Research, the Boise Fund, the South African Council for Scien- tific and Industrial Research, and the Research Committee of the University of the Witwaters- rand. My warmest thanks are extended to Dr. and Mrs. L. S. B. Leakey, Professor R. A. Dart. Dr. G. H. R. von Koenigswald, Dr. V. FitzSimmons, Dr. C. K. Brain, Miss J. Soussi. Mrs. R. W. Levine, Mr. A. R. Hughes and Mr. L. P. Morley.

REFERENCES

Abel, W. (1931). Kritische Untersuchungen tiber Australopithecus africanus Dart. Morph. Jahr- buch, 65 (4), 539-640.

Andrews, R. C. (n.d.). Meet your ancestors: A biography of primitive man. London: John Long.

Arambourg, C. (1958). Les stades evolutifs de

189



1'humanite. The Leech, 28 (3, 4, 5), 106-111. Bartholomew, G. A. and Birdsell, J. B. (1953):

Ecology and the protohominids. Amer. Anthrop., 55, 481-498.

Beatty, H. (1951). J. Mammal., 32, 118 (Quoted by Goodall, 1964).

Bielicki, T. (1964. Evolution of the intensity of feedbacks between physical and cultural evolution from man's emergence to present times. UNESCO/SS/Race/24, 10th July, 1964, 3 pp.

Bolwig, N. (1961). An intelligent tool-using baboon. S. Afr. J. Sci., 57, (6), 147-152.

Brain, C. K. (1958). The Transvaal ape-man bearing deposits. Transvaal Mus. Mem. No. 11, Pretoria.

Brain, C. K., Dart, R. A. and Lowe, C. van Riet (1955). Kafuan stone artefacts in the post- australopithecine breccia at Makapansgat. Nature, 175, 16.

Broom, R. (1937). On Australopithecus and its affinities. Early Man: International Sympo- sium. Acad. Nat. Sci. Phil., 285-292.

Broom, R. and Robinson, J. T. (1952). Swartkrans ape-man Paranthropus crassidens. Transvaal Mus. Mem. No. 6, Pretoria.

Broom, R. and Schepers, G. W. H. (1946). The South African fossil ape-men, the Australo- pithecinae. Transvaal Mus. Mem. No. 2, Pretoria.

Chopra, S. R. K. (1961). The angle of pelvic torsion in the primates. Zeits. Morph. u Anthrop., 51 (3), 268-274.

Chopra, S. R. K. (1962). The innominate bone of the Australopithecinae and the problem of erect posture. Bibl. primat., 1, 93-102.

Clark, J. D. (1962). Discussion on paper by R. A. Dart (1962a), q.v.

Clark, J. D. (1963). The evolution of culture in Africa. Amer. Naturalist, 97 (892), 15-28.

Clark, J. D. (1964). The prehistoric origins of African culture. J. Afr. Hist., 5 (2), 161-183.

Clark, W. E. Le Gros (1955). The fossil evidence for human evolution. Chicago: UniversityPress.

Clark, W. E. Le Gros (1965). "There is a trans- cendence from Science to Science." Second Ray- mond Dart Lecture. Johannesburg, Witwaters- rand University Press.

Cooke, H. B. S. (1961). The Pleistocene sequence in South Africa and problems of correlation. Paper presented at INQUA meeting, Warsaw, 1961.

Coppens, Y. (1961a). D6couverte d'un Australopi- thecine dans le Villafranchian du Tchad. Comp- tes rend. seances Acad. Sci., 252, 3851-3852.

Coppens, Y. (1961b). Un Australopithecine au Sahara (Nord-Tchad). Bull. S.P.F., 58 (11 and 12), 756.

Coppens, Y. (1962). Decouverte d'un Australopi- thecine dans le Villafranchian du Tchad. Note preliminaire. Editions du C.N.R.S., 455-459.

Dahlberg, A. A. (1960). The Olduvai giant homi-

nid tooth. Nature, 188, 962. Dart, R. A. (1925). Australopithecus africanus:

the man-ape of South Africa. Nature, 115, 195-199.

Dart, R. A. (1926). Taungs and its significance. Nat. Hist., 26, 315-327.

Dart, R. A. (1929). A note on the Taungs skull. S. Afr. J. Sci., 26, 648-658.

Dart, R. A. (1956a). Cultural status of the South African man-apes. Smithsonian Inst. Report for 1955, 317-338.

Dart, R. A. (1956b). The myth of the bone- accumulating hyena. Amer. Anthrop., 58 (1), 40-62.

Dart, R. A. (1957a). An australopithecine object from Makapansgat. Nature, 179, 693-695.

Dart, R. A. (1957b). The Makapansgat Australo- pithecine Osteodontokeratic Culture. 3rd Pan- African Congress of Prehist., Livingstone, 1955, 161-171. London: Chatto and Windus.

Dart, R. A. (1957c). The osteodontokeratic culture of Australopithecus Prometheus. Transvaal Mus. Mem. No. 10, Pretoria.

Dart, R. A. (1958a). Bone tools and porcupine gnawing. Amer. Anthrop., 60 (4), 715-724.

Dart, R. A. (1958b). The minimal bone-breccia content of Makapansgat and the australopithecine predatory habit. Amer. Anthrop., 60, 923- 931.

Dart, R. A. (1959a). Cannon-bone scoops and daggers. S. Afr. J. Sci., 55 (3), 79-82.

Dart, R. A. (1959b). Osteodontokeratic ripping tools and pulp scoops for teething and eden- tulous australopithecines. J. Dent. Ass. S. Afr., 14 (5), 164-178.

Dart, R. A. (1959c). Further light on australopithecine humeral and femoral weapons. Amer. J. Phys. Anthrop., 17 (2), 87-94.

Dart, R. A. (1960a). The persistence of some tools and utensils found first in the Makapansgat grey breccia. S. Afr. J. Sci., 56 (3), 71-74.

Dart, R. A. (1960b). The bone tool-manufacturing ability of Australopithecus Prometheus. Amer. Anthrop., 62 (1), 134-143.

Dart, R. A. (1961a). An australopithecine scoop made from a right australopithecine upper arm bone. Nature, 191, 372-373.

Dart, R. A. (1961b). Further information about how Australopithecus made bone tools and utensils. S. Afr. J. Sci., 57 (5), 127-134.

Dart, R. A. (1961c). The place of the antelope cannon-bones (or metapodials) in australopithecine economy. Zeits. wissens. Zool., 165, 108-121.

Dart, R. A. (1962a). The continuity and origin- ality of australopithecine osteodontokeratic culture. 4th Pan-African Congress of Prehist., Leopoldville, 1961. Ann. Mus. Roy. Afr. Cent., Ser. in 8?, Sciences humaines, No. 40.

Dart, R. A. (1962b). Substitution of stone tools for bone tools at Makapansgat. Nature, 196, 314-316.

190



Davis, P. R., Day, M. H. and Napier, J. R. (1964). Hominid fossils from Bed I, Olduvai Gorge, Tanganyika. Nature, 201, 967-970.

De Chardin, T. (1952). Quoted by Senyurek (1955) p. 2.

Dobzhansky, T. (1963). Cultural direction of human evolution-a summation. Hum. Biol., 35, (3), 311-316.

Donisthorpe, J. (1958). A pilot study of the mountain gorilla (Gorilla gorilla Beringei) in South West Uganda, February to September, 1957. S. Afr. J. Sci., 54 (8), 195-217.

Fleischer, R. L., Price, P. B., Walker, R. M. and Leakey, L. S. B. (1965). Fission track dating of Bed I, Olduvai Gorge. Science (publ. pending).

Goodall, J. (1963a). Feeding behaviour of wild chimpanzees. Symp. Zool. Soc. Lond., No. 10, 39-47.

Goodall, J. (1963b). My life among wild chimpanzees. Nat. Geogr. Mag., 124 (2), 272-308.

Goodall, J. (1964). Tool-using and aimed throwing in a community of free-living chimpanzees. Nature, 201, 1264-1266.

Gregory, W. K. (1939). The bearing of Dr. Broom's and Dr. Dart's discoveries on the origin of man. Ann. Proc., 1938-1939, Asso- ciated Sci. and Tech. Socs. of S. Afr., 45.

Hayes, C. (1952). The ape in our house. London: Gollancz.

Hennig, E. (1948). Quartarfaunen und urgeschichte Ostafrikas. Naturwiss. Rdsch. Jahrg., 1, Heft 5, 212-217.

Hooijer, D. A. (1952). Fossil mammals and the Plio-Pleistocene boundary in Java. Proc. Ko- ninkl. Nederl. Akad. Wetensch., Amsterdam. Ser. B., 55 (4), 436-443.

Hopwood, A. T. (1951). The Olduvai Fauna. In Leakey (1965), Olduvai Gorge, 1951-1961, 20-30.

Howell, F. C. (1965). Footnote to Comment on Olduvai hominids. Current Anthrop, 6, 399-401.

Inskeep, R. R. (1959). Central and West Africa. Encyclopaedia Britannica 1959, 328-330.

Isaac, G. (1965). The Peninj beds-an early Middle Pleistocene formation west of Lake Natron. Quaternaria (pending).

Jerison, H. (1963). Interpreting the evolution of the brain. Human Biol., 35, 263-291.

Kluver, H. (1937). Re-examination of implement- using behaviour in a cebus monkey after an interval of three years. Acta Psychol., 2, 347- 397.

Khroustov, H. F. (1964). Formation and highest frontier of the implemental activity of anthropoids. VIIth Int. Congr. Anthrop. Ethnol. Sci., Moscow, August, 1964.

Koenigswald, G. H. R. von (1952). Quoted by Senyuirek (1955).

Koenigswald, G. H. R. von (1957). Remarks on Gigantopithecus and other hominoid remains from Southern China. Proc. Koninkl. Ned.

Akad. Wetensch., Ser. B, 60, 153-159. Koenigswald, G. H. R. von (1960). Remarks on

a fossil human molar from Olduvai, East Africa. Proc. Koninkl. Ned. Akad. Wetensch., Ser. B, 63, 20-25.

Koenigswald, G. H. R. von (1961). Australopithe- cus und das Problem der Gerolkulturen. Deuts. Gesells. Anthrop., Tubingen, April, 1961, 139- 152.

Koenigswald, G. H. R. von (1962). Das absolute Alter des Pithecanthropus erectus Dubois. In "Evolution und Hominisation", Ed. G. Kurth, Stuttgart: Gustav Fischer Verlag, 112-119.

Kohler, W. (1924). The Mentality of Apes. Eng- lish Edition: New York, Vintage Books, 1959.

Kohl-Larsen, L. (1943). Die menschenfunde des Sudserengeti in der Zeit vom. 10 Januar 1939 bis. 28 Februar 1939. In "Auf des Spuren des Vbrmenschen", Vol. II, 379-381.

Kohts, N. (1935). Infant ape and human child (Instincts, emotions, play, habits). Scient. Mem. Mus. Darwinianum, Moscow.

Kortlandt, A. (1962). Chimpanzees in the wild. Scient. Amer., 206, 128-138.

Kortlandt, A., and Kooij, M. (1963). Proto- hominid behaviour in Primates. Symp. Zool. Soc. Lond., No. 10, 61-87.

Kurten, B. (1956). The status and affinities of Hyaena sinensis Owen and Hyaena ultima Mat- sumoto. Amer. Mus. Novit., No. 1764, 48 pp. (Quoted by Oakley, 1964b).

Leakey, L. S. B. (1958). A giant child among the giant animals of Olduvai? A huge fossil milk molar which suggests that Chellean Man in Tanganyika may have been gigantic. Ill. London News, 1103-1106.

Leakey, L. S. B. (1960). Recent discoveries at Olduvai Gorge. Nature, 188, 1050-1052.

Leakey, L. S. B. (1961a). A new lower Pliocene fossil primate from Kenya. Ann. & Mag. Nat. Hist., Ser. 13, iv, 689-696.

Leakey, L. S. B. (1961b). The juvenile mandible from Olduvai. Nature, 191, 417-418.

Leakey, L. S. B. (1965). Olduvai Gorge 1951-1961. Vol. I: A preliminary report on the geology and fauna. Cambridge: University Press.

Leakey, L. S. B. and Leakey, M. D. (1964). Recent discoveries of fossil hominids in Tanganyika: at Olduvai and near Lake Natron. Nature, 202, 5-7.

Leakev. L. S. B., Tobias, P. V. and Napier, J. R. (1964). A new species of the genus Homo from Olduvai Gorge. Nature, 202, 7-9.

Mason, R. J. (1957). Occurrence of stone artefacts with Australopithecus at Sterkfontein, Part II. Nature, 180, 521-524.

Mason, R. J. (1961). The earliest tool-makers in South Africa. S. Afr. J. Sci., 57, (1), 13-16.

Mason, R. J. (1962). Australopithecines and artefacts at Sterkfontein, Part II. The Sterkfontein stone artefacts and their maker. S. Afr. Arch. Bull., 17 (66), 109-125.

191



Merfield, F. G. (1956). Gorillas were my neigh- bours. London: Longmans, Green & Co.

Napier, J. R. (1960). Studies of the hands of living primates. Proc. Zool. Soc. Lond., 134 (4), 647-657.

Napier, J. R. (1962). Fossil hand-bones from Olduvai Gorge. Nature, 196, 409-411.

Oakley, K. P. (1951). A definition of man. Science News, Penguin Books, 20, 69-81.

Oakley, K. P. (1964a). Letter to the Editor of Discovery on "The Evolution of Man". Dis- covery, Aug. 1964, p. 49.

Oakley, K. P. (1'964b). Frameworks for Dating Fossil Man. London: Weidenfeld and Nichol- son.

Oppenheimer, A. (1964). Tool use and crowded teeth in Australopithecinae. Current A nthrop., 5, (5), 419-421.

Osborn, R. (1963). Observations on the behaviour of the mountain gorilla. Symp. Zool. Soc. Lond., No. 10, 29-37.

Osman Hill, W. C. (1960). Primates, Vols. 4 and 5, Cebidae, Parts A and B. Edinburgh: Uni- versity Press.

Remane, A. (1951). Die Zahne des Meganthropus africanus. Zeits. Morph. u Anthrop., 42, 311- 329.

Robinson, J. T. (1953). Meganthropus, australopithecines and hominids. Amer. bourn. Phys. Anthrop., 11 (1), 1-38.

Robinson, J. T. (1955). Further remarks on the relationship between "Meganthro pus" and aus- tralonithecines. Amer. Journ. Phys. A nthrop., 13 (3), 429-446.

Robinson, J. T. (1956). The dentition of the Australopithecinae. Ann. Transvaal Mus., No. 9, Pretoria.

Robinson, J. T. (1957). Occurrence of stone artefacts with Australopithecus at Sterkfontein. Nature, 180, 521-524.

Robinson, J. T. (1958). The Sterkfontein tool- maker. The Leech, 28 (3, 4, 5), 94-100.

Robinson, J. T. (1960). An alternative interpretation of the supposed giant deciduous hominid tooth from Olduvai. Nature, 185, 407-408.

Robinson, J. T. (1962a). Australopithecines and artefacts at Sterkfontein, Part l: Sterkfontein stratigraphy and the significance of the Exten- sion Site. S. Afr. Arch. Bull., 17 (66), 87-107.

Robinson, J. T. (1962b). The origin and adaptive radiation of the australopithecines. In "Evolu- tion und Hominisation", Ed. G. Kurth. Stutt- gart: Gustav Fischer Verlag.

Robinson, J. T. (1965). Homo 'habilis' and the australopithecines. Nature, 205, 121-124.

Schaller, G. B. (1963). The Mountain Gorilla, Ecology and Behaviour. Chicago: University Press.

Senytirek, M. (1955). A note on the teeth of Meganthro pus africanus Weinert from Tangan- yika territory. Belleten, 19 (73), 1-55.

Simons, E. L. (1961). The phyletic position of Ramapithecus. Postilla Yale Peabody Museum, 57, 1-9.

Simons, E. L. (1963). Some fallacies in the study of hominid phylogeny. Science, 141, 879-889.

Simonm, E. L. (1964). On the mandible of Rama- pithecus. Proc. Nat. Acad. Sci., 51, 528-535.

Simpson, G. G. (1965). Foreword to Leakey, L. S. B. (1965): Olduvcai Gorge, 1951-1961., Vol. I.

Stekelis, M., Picard, L., Schulman, N. and Haas, G. (1960). Villafranchian deposits near Ubeidiya in the Central Jordan Valley (Preliminary report). Bull. Res. Council of Israel, 9G, 175- 184.

Straus, W. L. (1962). Fossil evidence of the evolution of the erect, bipedal posture. Clin. Ortho p., 25, 9-19.

Tobias, P. V. (196). Cranial capacity of Zinj- anthropus and other australopithecines. Nature, 197, 743-746.

Tobias, P. V. (1964a). The Olduvai Bed I hominine with special reference to its cranial capacity. Nature, 202, 3-4.

Tobias, P. V. (1964b). Dart and Taung forty years after. S. Afr. J. Sci., 60 (11), 325-329.

Tobias, P. V. (1964c). The early hominid remains from Tanganyika: Australopithecus and Homo. Proc. VIIth Itit. Con gr. Anthrop. Ethnol. Sci., Moscow, August. 1964.

Tobias, P. V. (1965a). Early man in East Africa. Science, 149, 22-33.

Tobias, P. V. (1965b). New discoveries in Tan- ganyika: their bearing on hominid evolution. Curr. Anthrop, 6, 391-399 and 406-411.

Tobias, P. V. and Koenigswald, G. H. R. von (1964). A comparison between the Olduvai hominines and those of Java and some implications for hominid phylogeny. Nature, 204, 515-518.

Vevers, G. M. and Weiner, J. S. (1963). Use of a tool by a captive capuchin monkey (Cebus opella). Symp. Zool. Soc. Lond., No. 10, 115- 117.

Washburn, S. L. (1959). Speculation on the in- terrelations of the history of tools and biologi- cal evolution. Human Biol., 31, 21-31.

Washburn, S. L. and Howell, F. C. (1960). Human Evolution and Culture. In 'Evolution after Darwin", Vol. II: The Evolution of Man. Chicago: University Press, 33-56.

Weiner, J. S. (1963). See Vevers and Weiner (1963).

Weinert, H. (1950). Uber die neuen Vor- und Fruhmenschenfunde aus Afrika, Java, China und Frankreich. Zeits. f. Morph. it Anthrop., 42, 113-148.

Wells, L. H. (1962). Pleistocene faunas and the distribution of mammals in Southern Africa. Ann. Cape Prov. Mus., No. II, 37-40.

Yerkes, R. M. (1948). Chimpanzees. (4th Ed.) New Haven: Yale University Press.

192



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