The credibility of Homo habilis.

15 THE CREDIBILITY OF HOMO HABILIS

C.B. Stringer, Dept. of Palaeontology, British Museum (Natural History), Cromwell Road, London SW7 5BD, England.

INTRODUCTION The creation of new hominid species was a regular occurrence

in the earlier part of this century (see e.g. Campbell, 1965), but in the last 20 years such events have become rare, and the subject of heated debate in the case of A. afarensis and H. habilis. At a time when major studies of the material referred to H. habilis from Olduvai and Koobi Fora are in preparation, or in press, it seems a hard task to present a significant review about this species which will not shortly be considered insignificant or redundant, particularly when the author has had virtually no contact with the original specimens concerned. But rather than attempt a dry review of the subject based entirely on the original work of others, I wanted to try to introduce some new data and anlyses. Through the co-operation of colleagues I have managed to rework some of their original data, and I have measured available casts of relevant crania wherever possible. This latter activity gave me my first shock when I realised that some casts were more than 5% smaller than the actual fossils for which I had reliable data, so I decided to rely on cranial angles instead of raw data in comparisons of casts and originals.

This paper will first briefly review the history of the species H. habilis from its inception in 1964, then turn to aspects considered important by other workers viz endocranial size and dental characters, then present new analyses of cranial and facial variation in H. habilis specimens, and finally conclude with comments on the credibility of the species. The material allocated to H. habilis from Olduvai must form an essential part of any discussion of the species, while the material from Koobi Fora has become important to the problem of H. habilis because of its excellent preservation and remarkable morphological variation. However for this exploratory paper I do not intend to discuss in any detail the postcrania: material allocated to H. habilis from these sites, nor will I discuss claimed H. habilis material from South Africa (Tobias, 1983) or Ethiopia (Howell, 1978), except where I have been able to study at least a cast the relevant material (SK 847).

HOMO HABILIS AT OLDUVAI From 1959, a series of fossil hominid remains was recovered

from Bed I and lower Bed II at Olduvai Gorge, Tanzania, which clearly differed from those of the Australopithecus (Zinjanthropus) boisei materaa_ identified at the site. In 1964, Leakey, Tobias & Napier named a new species of the genus Homo for these fossils, Homo habilis ("handy man",. ' irp* Ipueq,,) s-Tq€u our6g ,sTrssoJ asaq?;og: offi snuab el{tr Jo sarcads .vieu e perueu :erdeg B sETqoJ, ,â)ipeT ,196T uI .OJTS aLIf f,p perrr-]ueDT::.EUl Tasroq (sndo-rr{lueLurz) sncei{aTdoTp.rfsnv ai_{]._lo asorfl ulo-f,J peJoJJTp

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SITISVH OI^]OH JO AITTISTOSUC SHJST

Stringer: Credibility of Homo habilis 267

Table 1. Homo habilis material (from Leakey, Tobias & Napier, 1964)

Bed I Bed II

TYPE 0.H.7: mandible, upper M*, parietals, hand bones of juvenile

PARATYPES 0.H.4: mandible frag. with 0.H.13: mandible, M and tooth frags. maxillae, cranial

parts of adolescent? 0.H.6- P 3 , M1 or M2 , • 3' cranial frags (and tibia and fibula?)**

0.H.8: hand bones, foot bones, tooth, clavicle of adult***

REFERRED 0.H.14: cranial frags of juvenile.

0.H.16: partial skull and dentition of adult

later renumbered 0.H.45

** tibia and fibula later renumbered 0.H.35

*** tooth later renumbered 0.H.46, clavicle 0.H.48, hand bones not hominid?

The holotype, paratype and referred specimens are listed in Table 1, and the main distinctive features of the species were given as a mean brain size intermediate between that of Australopithecus and H. erectus; maxillae a_nd mandibles in the size range of H. erectus and H. sapiens rather than • Australopithecus; relatively large anterior dentition; a tendency to tuccolingual narrowing of the dentition, especially in the lower premolars ant: molars; external sagittal curvature of the occipital slight as in H. =Lens, but not like that of Australopithecus or H. erectus; various -=_tures of the clavicle, hand bones and foot bones resembling H. sapiens Lens, with some distinctive features such as greater robusticity.

Subsequent research has led to the renumbering of certain of the specimens as trobably representing different individuals (Table 1). The tibia and fLt.ila (now numbered 0.H.35) were found at the site of 0.H.5 ("Zinjanthro- • but Davis (1964) refrained from referring them to the same species a7.:1 Leakey et al (1964) tentatively associated these bones with those of :.H.6 (H. habilis - Table 1). Despite the fact that the remains were

over 200 metres apart, Susman & Stern (1982) suggested that 0.H.7 a7.d 0.H.8 and 0.H.35 might all represent one subadult individual (age • :3-14 years).

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268 Stringer: Credibility of Homo habilis

The erection of this new species for the Olduvai material was not well received on the grounds that while some of the Bed I specimens were not clearly differentiated from A. africanus, the Bed II specimens closely resembled known specimens of H. erectus (in 1964 the time span of Bed I and lower Bed II, now known to be c. 0.3 myr, was unknown but thought to be as much as 1 myr by some workers). Following direct comparison of the Olduvai and Indonesian fossils (Tobias & von Koenigswald, 1964) Tobias modified his view of the Bed II material, suggesting that some of it might represent the same grade of hominid as some of the "Pithecanthropus" material from Java (i.e. H. erectus), while 0.H.16 was perhaps an australopithecine rather than representing H. habilis, to which it had been referred (Table 1). However by 1967, Tobias had apparently re-considered 0.H.13 as an H. habilis specimen (Tobias, 1967), and by 1978 0.8.16 was once again considered to represent this species (Tobias, 1978).

A useful summary of early opinions about the status of H. habilis can be gathered from the comments following an article by Tobias on H. habilis (Tobias 1965). Howell tended to agree with Robinson (1966) that the Bed I material represented an advanced australopithecine, while the Bed II material represented a descendant early form of H. erectus. In fact Howell saw enough cranial and dental resemblances between the Bed II and Indonesian fossils to suggest that H. erectus modjokertensis or H. modjokertensis might eventually prove to be the appropriate name for these fossils (see also Boaz & Howell, 1977; Howell, 1978). Howell disagreed with the view that O.H. 16 might represent an australopithecine, and both he and Napier anticipated Susman & Stern (1982) by noting the excellent match of the 0.H.8 foot with the 0.H.35 tibia and fibula. However they both cautioned that the lack of comparable australopithecine material precluded the use of any of the Olduvai postcranial specimens in taxonomic arguments about the distinctiveness of H. habilis. Robinson (1966) repeated his view that the Bed I and Bed II materials were more like, respectively, Australopithecus and H. erectus than they were like each other. This view was echoed by Simons, Pilbeam & Ettel (1969), Brace et al (1972) and others.

A number of more recent discoveries from Olduvai have been assigned to H. habilis (Tobias, 1978), the most complete being the cranium of 0.8.24 from Bed I (Leakey et al, 1971). This crushed and distorted specimen was skilfully reconstructed by Clarke to produce a cranium which was con-sidered to be similar to 0.8.13 from Bed II, with an initial estimate of endocranial volume of c. 560 ml, later increased to c. 600 ml (Tobias, 1972; Holloway, 1983b). Following suggestions (Anon, 1971) that the specimen did not represent the genus Homo as defined by Leakey at al (1964) because of its low endocranial capacity and "dished" face, Tobias (1972) defended its inclusion in the genus, while refraining from actuall: assigning it to H. habilis. Tobias' caution in 1972 no doubt reflected the poor preservation of the specimen and the doubts of Leakey at al (1971) that, while it was possible that 0.8.13 and 24 represented females, and 0.8.7 and 0.8.16 males of H. habilis, occipital differences between the two presumed sexes perhaps indicated further taxonomic variation. However Tobias (1978) later accepted 0.H.24 as H. habilis, while others remained doubtful (Howell, 1978; Walker, 1981). '(TB6T 'ra)iTel4 ,BL6T ,TTelroH)

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CLASSIFICATION OF THE KOOBI FORA HOMINIDS Work at Koobi Fora, Kenya, from 1968 has provided a large

number of specimens of relevance to the status of Homo habilis (Leakey et al, 1978). Initial descriptions of the material, where a basic classification of the first specimens into two genera, viz. Australo-pithecus and Homo was proposed (Leakey, 1972), were criticised by Robinson (1972) on the grounds that there was nomenclatural confusion between Australopithecus and Paranthropus. He proposed that the South African and East African forms should instead both be divided into two genera, Paranthropus and Homo, and that the gracile forms of both areas probably represented H. africanus. Further discoveries greatly enlarged and complicated the picture with the appearance of large and small-brained "gracile" hominids with apparent affinities variously to H. erectus, H. habilis and A. africanus (Walker, 1976, 1981). Walker (1976) discussed the classification of the large brained fossil KNM-ER 1470 and tested the specimen using indices and other characters which distinguish Australopithecus from Homo. Overall the specimen appeared more like Australopithecus than Homo using these criteria, and while he did not actually classify the specimen as an australopithecine, he certainly urged caution about classifying it as anything else. In a later paper (Walker, 1981) he appeared to prefer classifying specimens such as 1CNM-ER 1470 with 0.8.7 and 16 as Australopithecus habilis, while assigning the small- brained gracile cranium KNM-ER 1813 to a late surviving form of A. africanus along with 0.8.13 and 24. The model favoured by Walker appeared to be that A. africanus gave rise to A. habilis through body size increase, and this species was then probably ancestral to H. erectus (represented by speci-mens such as KNM-ER 3733 and SK 847). Wood provided a longer review of the East African material (Wood, 1978) and also suggested the possibility of a third hominid lineage in East Africa,' apart from H. habilis/H. erectus and A. boisei. Although differing from Walker in preferring to classify specimens such as KNM-ER 1470 as Homo, he also regarded the habilis-erectus groups as representing a Homo lineage and noted the distinctive-ness of KNM-ER 1813 from this lineage. But rather than regarding this latter specimen as a small-toothed australopithecine, he regarded it as probably distinct, without naming it.

Many other workers have been prepared to place all the non-erectus gracile hominids of Koobi-Fora in one taxon, usually H. habilis, and view varia-tion between specimens such as KNM-ER 1470 and KNM-ER 1813 as related to sexual dimorphism and anagenetic change (e.g. Howell, 1978; Wolpoff,1980; White et al, 1981; Cronin et al, 1981; Tobias, 1983). Opinions vary about the classification of individual specimens such as KNM-ER 1805, regarded as H. habilis by some workers (and possibly abnormal, White et al, 1981) or as H. erectus by others (Howell, 1978; Wolpoff, 1980). Recent work on the cranial base by Dean & Wood (1982) has demonstrated that specimens such as KNM-ER 1470, 1813, and 0.H.24 share features in common with KNM-ER 3733 which favour their inclusion in Homo, rather than A. africanus. KNN-ER 1805 is also unlike A. africanus but the evidence of the basicranium is compatible with its inclusion in either Homo or A. boisei.

H. ergaster Differences between the Olduvai H. habilis specimens and pue suarlTsads srTTqerl .H re^npTo eql uaaA?aq sesuefaJJTc

tafsebre .H

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gracile hominids from Koobi Fora led Groves & Maza.k (1975) to reaffirm the specific identity of the Olduvai material, but also to recognise a new species of Homo, H. ergaster. The holotype of this new species was the mandible KNM-ER 992. They accepted that H. habilis could be clearly distinguished from A. africanus (termed by them H. africanus) in features such as lower premolar shape and endocranial capacity, while H. ergaster was probably distinct from both in having smaller molars and premolars, with reduced numbers of premolar roots. H. ergaster was also characterised by a rather thick and massive mandible, and a large endocranial capacity compared to A. africanus. Specimens in the hypodigm included KNM-ER numbers 730, 820 and, less certainly, 1805, while the resemblances between H. ergaster and the "Telanthropus" mandible (SK 15) from Swartkrans were noted. The authors also speculated that earlier specimens from Koobi Fora, such as KNM-ER 1470, may have been ancestral to H. ergaster.

The main problem with this new species was that it was not properly differentiated from specimens assigned to H. erectus and this appears difficult to achieve for the holotype KNM-ER 992. Partly because of this, the new species name has been ignored, or considered premature (Wood, 1978'. However the naming of H. ergaster was significant because if a new species were to be created for any of the gracile hominid material from East Africa, and the taxon included the ergaster holotype, this name would be available for the new species. The relevance of this to the classifica-tion of specimens such as KNM-ER 1813, which has been linked to the KNM-ER 992 mandible (Leakey et al, 1978), will be disucssed later.

ENDOCRANIAL VOLUME Since the days of Keith's "cerebral Rubicon", brain size (as

reflected by endocranial volume) has loomed large in studies of human evolution. Although absolute brain size is obviously significant in differentiating modern humans from apes, the relationship between brain size and body size becomes more important in assessing fossil hominids (Holloway, 1983b). For this paper I do not intend to use absolute endo-cranial volume as an important taxonomic character for the recognition of the genus Homo, as I feel that body size variation is still too poorly known in early Homo individuals to allow an appropriate adjustment for relative brain size. Instead I will examine variation in endocranial volume to identify whether one or more groups may be represented in a widely-defined Homo habilis taxon, encompassing specimens such as KNM-ER 1805, 1813 and 1470, as well as 0.H.7 and 13.

Table 2 presents data on variation, of endocranial volume in living and fossil hominoid samples, determined in most cases by the use of whole endocranial casts. In common with previous estimates, it is evident fr:n these data that a coefficient of variation of about 10% is a reasonani= figure for a hominoid species, with the largest variation found in G - - - (figures as high as 13% are sometimes reported for this ape - Tobias, Wood, 1976). Turning now to a broadly defined sample of H. habilis Fig. 1 presents determined and estimated values for the sample taken fr:n Holloway (1983b) and my own estimates for incomplete specimens. The -.--__ quoted vary greatly in reliability, but the minimum and maximum value: are from whole endocast determinations, rated as the most reliable -ny

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africanus, A. robustus and A. boisei (Johanson & White, 1979; White at al, 1981). Looking at the early Homo samples as a whole it is evident that variation in this group is generally comparable with that found in australopithecine samples (Tobias, 1978; White et al, 1981).

Martin (1983) suggested criteria for establishing the existence of more than one species in a dental sample using characteristics of the range, mean and standard deviation of tooth measurements, compared with one- species samples of living hominoids. The methods would identify an unusually variable sample as probably representing at least two species, although some samples actually consisting of two species might not always be variable enough for this to show from the criteria used. Applying these criteria to the posterior dentitions of the early Homo group of White et al, (1981), only the Ml (m-d length) and M3 (b-1 breadth) values immediately suggest the presence of more than one species in the sample, although the M2 (m-d and b-1), M3 (b-1) and P 3 (b-1) dimensions are also quite variable. Since the White at al (1981) samples appear to contain some individuals otherwise assignable to African H. erectus, the size variation in the sample is perhaps not surprising and indicates that this approach is not

Figure 1. Coefficients of variation for endocranial volumes of early Homo crania. C.V. values are uncorrected for small sample sizes, so are probably underestimates. Most values were obtained from Holloway (pers. comm., 1982, 1983b) but see text for discussion of my estimates for O.H. 16, 1590 and 3732.

C.V. = 12.4

KNM-ER

1

1

10.9

9.5

1 6.0

1805

582 ml

OH 16

667 ml

3732

(700ml)

1590

(750m1)

1813 OH 24 OH 13 OH 7 1470

510 ml

590 ml 650 ml 687 ml 752 ml

475

525

575

625

675

725

775

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likely to produce significant data to separate possible subgroups in the early Homo sample - if two groups do exist, they are similar in most dental dimensions.

An alternative approach is to examine dental proportions, since these have been used to distinguish H. habilis from australopithecines (Tobias, 1965, 1978) and to distinguish specimens within the early Homo group (Groves & Mazik, 1975). In Table 3 I have presented data on premolars and first molars of living and fossil hominoids. Looked at in terms of primitive and derived characters, it is probable that broad premolars and M1 (i.e. a small L/B index) are plesiomorphous for hominoids, as is a relatively small P4/large M1 (low index value). For M1, which is distinctively narrow in many of the early Homo specimens (especially O.H.7), the polarity of this character is unclear, and a fairly high value of the index (c.110) may have been the plesiomorphous condition, based on living and fossil hominoid values. If this is so, it is the A. afarensis specimens which seem especially derived in their broad M 1 , while the Homo specimens are closer to the assumed plesiomorphous condition.

For the fossil groups considered, the early Homo sample does appear derived in the narrow P 4 and M 1 , but relatively primitive in P4 and P4/M 1

indices. Of the individual specimens listed, KNM-ER 992 and the Olduvai hominids are most contrasted in P 4 and P 4/M1 indices, while the rest of the individual specimens are rather similar, save for the (plesiomorphous?) broad P4 in KNM-ER 1805 and the (apomorphous?) large P 4 in KNM-ER 1802. The similarity of KNM-ER 992 and 1802 in the P 4/M i index may look odd to anyone who is familiar with these specimens, placing them both as more derived in the direction of A. boisei than is A. africanus, but examina-tion of the tooth areas suggests that for the former specimen it is the very small M1 size which is the significant factor, whereas for KNM-ER 1802 the size, shape and crown form of the P 4 does indeed resemble that of some South African australopithecines.

Overall the data of Table 3 confirm the narrowness of P4 and M1 as probable synapomorphies for the early Homo specimens, while suggesting that the P4, ml and P4 /M1 indices overall remain plesiomorphous. It is somewhat ironic that the holotype mandible of H. habilis (O.H.7) should be extreme for the group in its narrow P 4 and M1 , partly explaining Tobias' emphasis on this character (Tobias, 1965, 1966) and Groves & Maz5k's (1975) differentiation of H. habilis and H. ergaster. The tooth shape data, then, confirms the presence of relatively narrow premolars and molars in the Homo sample, but does not suggest any clear division of the material beyond the differences already mentioned.

Dental morphology In a study of the morphology of mandibular molars, Wood &

Abbott (1983), and Wood, Abbott & Graham (1983) examined variation in crown areas, cusp areas, crown shape, crown profile and cusp numbers and patterns. They establish criteria which characterised the four main taxonomic groups studied, South African gracile hominids (SAFGRA), South African robust hominids (SAFROB), East African gracile hominids (EAFHOM) and East African robust hominids (EAFROB). They then applied these criteria

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274 Stringer! Credibility of Homo habilis

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in order to classify a number of "unknown" specimens in the two geographical areas into one of the two local groups (e.g. in South Africa, the Taung Mi was classified as SAFROB using relative cusp area and fissure pattern, while crown area and additional cusps of the tooth could not clearly allocate it to either the SAFROB and SAFGRA groups. Similarly KNM-ER 1302 was predominantly classified as EAFHOM rather than EAFROB using these criteria).

In order to examine variation in these characters in the early Homo group I was allowed access to the original data used in the study and as an experiment I extended the classification criteria used for two members of the EAFHOM group, and KNM-ER 1802, to include SAFGRA as an additional category. The revised classification of these three lower dentitions from the EAFHOM sample is shown in Table 4, excluding the criteria of crown profile, which I did not examine.

It is evident from this table that in tooth area all the specimens con-sidered more closely resemble the South African australopithecine sample than the East African Homo sample to which two of them have been allocated (and in fact, for M 2 , KNM-ER 1802 and O.H. 16 resemble South African robust australopithecines even more closely:). There is less differentiation for the cusp and fissure patterns, while for cusp areas the classification of M1 and M2 in KNM-ER 1802 and O.H. 7 is divided between the australopithe-cine and Homo groups. Thus morphological characters suggest variation in the direction of A. africanus in some of the larger early Homo specimens, but an overlap with A. africanus in both morphology and size of teeth was

Table 4: Classification of lower molars of KNM-ER 1802, 0.H.7 and O.H. 16 using criteria of Wood & Abbott (1983) and Wood, Abbott & Graham (1983). GRA = South African gracile hominid group, HUM = East African gracile hominid group, NA = not available, - = no discrimination.

AREA CUSPS CUSP AREA FISSURE

1802 M1 R GRA HOM? HOM HOM

M1 L GRA HOM? HOM HOM

M2R GRA - GRA -

M2 L GRA - GRA -

Q.H. 7 MLR GRA - GRA NA

M1 L GRA - GRA NA

M2 L

GRA - HON HOM

0.H.16 M1 R GRA - - GRA

M2R GRA - HOM HOM

M3R GRA - HOM NA

M3L GRA - HOM NA

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276 Stringer; Credibility of Homo habilis

not excluded by the original describers of H. habilis. What is needed nex -: is for this useful work on molar morphology to be extended to A. afarensia and modern hominoid samples in order to test resemblances to the other groups analysed and to begin to establish probable plesiomorphous and apomorphous conditions.

MIDSAGITTAL CRANIAL ANGLES Cranial angles measured between various points in mid-sagittal

section have a long history of use (Howells, 1973; Krukoff, 1978) and abuse (Baker, 1974) in anthropology. Here I intend to use them, as I have previously (Stringer, 1978; Stringer & Trinkaus, 1981) to examine variaticn in crania with the aim of identifying polarities in values of cranial angles between different hominoid species. The values of the cranial angles used here are dependent entirely on the relative position of four ostec-metric points on the cranium (prosthion, nasion, bregma and basion, Howells, 1973; Stringer, 1978), and as such may be affected by various ontogenetic, functional and allometric considerations. Nevertheless I hope to demonstrate below that a consistent pattern of cranial angles can be recognised in most adult living hominoids regardless of overall size, allowing recognition of plesiomorphous and apomorphous hominoid conditions in cranial angle values, against which the fossil crania of the early Homo group can be assessed.

Three midsagittal cranial angles have been calculated for the facial tri-angle (between nasion, basion and prosthion) consisting of nasion angle (NAA), basion angle (BAA) and prosthion angle (PRA) and for the anterior cranial triangle (between nasion, bregma and basion) consisting of nasion-bregma angle (NBA), bregma angle (BRA) and basion-bregma angle (BBA). Samples of male and female ape crania of G. gorilla, P. pygmaeus, P. troglodytes and Symphalangus syndactylus were measured to provide a comparative base, and the calculated cranial angles were compared with those of modern H. sapiens (data from Howells, 1973) as in Table 5.

From Table 5 it is evident that two main patterns of cranial angles can be discerned in living hominoids. One is the pattern found in ape crania, characterised by high NAA, low PRA, low NBA, high BRA and low BBA values. The other pattern, found in H. sapiens crania, is the opposite of the condition found in the apes in each case, and can therefore be assumed to be the derived condition as against the shared primitive condition of the ape crania. For the purposes of this paper the data for Pan will be used to represent the (plesiomorphous) ape condition. Variation in BAA values will not be discussed here since clear polarities cannot be identified.

The mean values for the Pan and Homo sample are shown diagrammatically in Fig. 2, with the figures scaled to the same basion-nasion length to eliminate size differences in this visual comparison. The contrast between the two is immediately apparent, with Pan showing a relatively large and prognathic facial triangle (high NAA, low PRA), and a small and low anterior cranial triangle (low NBA and BBA, high BRA). Values for the fossil sample were compared with those obtained for Pan and H. sapiens. Original Middle and tipper Pleistocene crania were measured for the analysis, but the Plio-Pleistocene crania could only be studied as casts, /sJsec sp perpnfs eq

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and in the case of specimens such as KNM-ER 1470 and D.H. 24 casts of reconstructions were used, which must be taken into account in any conclusions reached. It was also necessary to reconstruct the position of some of the osteometric points in some specimens, and where the osteometric point nasion lay near glabella, the measurement was actually taken 5 mm below glabella, to provide a more comparable data point.

Figure 3 shows the cranial angles calculated for Pan, modern H. sapiens, a Neanderthal sample (Stringer & Trinkaus, 1981) and the Pilo-Pleistocene specimens. Where polarity could be clearly determined on the basis of the modern hominoid data (Table 5), an arrow has been added to indicate the derived condition, which is in all cases found in H. sapiens and to a lesser extent, the Neanderthals. Crania KNM-ER 3733 and 3883, assigned here to H. cf. erectus, are generally further from modern human values than the Neanderthals, lying in an approximately intermediate position between H. sapiens and Pan for NAA, PRA, BRA and NBA values. These crania will be used here as a guide to the probable plesiomorphous Homo condi-tion in these angles, against which the other Plio-Pleistocene specimens can be tested.

Figure 3 shows that all three australopithecine crania (Ste 5 - A. africanus; Sr 48 - A.robustus (Paranthropus); KMM-ER 406 - A. boisei (Paranthropus)) are close to the mean values for Pan in NAA, PRA and BBA angles and can be considered to be primitive in these values. But where SK 48 and KNM-ER 406 are also primitive in BRA and NBA values, Sts 5

Figure 2. Cranial angles of sample of Pan troglodytes (n = 16) and modern H. sapiens (n = 1652, Howells, 1973), scaled to same basion-nasion length.

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appears to share the distinctive derived Homo condition for these angles. Examining the data for the early Homo specimens, it is evident that KNM-ER 1813 appears most derived in NAA and PRA values, and 0.H. 24 in BBA, BRA and NBA values, while KNM-ER 1470 tends to occupy an intermediate position between the other two crania.

Drawing out the values of the three early Homo crania together with KNM-ER 3733, scaled to the same basion-nasion length (Fig. 4), shows interesting similarities and contrasts. Now it can be seen that while KNM-ER 1470 and 1813 closely resemble KNM-ER 3733 in cranial angles, 0.H.24 is distinctly more primitive (i.e. like Pan and Australopithecus) in the facial triangle, and quite aberrant (i.e. ultra derived even compared with H. sapiens) in the anterior cranial triangle. While the facial triangle data agree with the appearance of the specimen in indicating a high and primitive degree of total facial prognathism, the anterior cranial triangle data are peculiar, giving values for BBA and BRA which are more extreme than any modern human population. This seems an unlikely combination, and suggests that a large amount of distortion remains in the cranial vault of the O.H. 24 reconstruction, as was recognised by Leakey et al (1971) and Tobias (1972).

Figure 3. Cranial angles for Pan sample, modern H. sapiens and fossil hominids. Where polarity can be established, an arrow indicates the inferred derived condition. Neanderthal data from Stringer and Trinkaus (1981). NAA = nasion angle, PRA = prosthion angle, BAA = basion angle, BBA = basion-bregma angle, BRA = bregma angle, NBA = nasion bregma angle.

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In order to display the overall results more clearly, I have plotted out values for three of the angles whose polarity is most clear in figures 5 and 6, adding data from the Middle Pleistocene crania from Broken Hill, Petralona and Bodo to provide further information on variation in the genus Homo. Figure 5 provides an overall measure of total facial prognathism and shows the plesiomorphous characteristics of the australopithecine crania, together with the KNM-ER 1470 and O.H. 24 reconstructions. KNM-ER 1813 is clearly much more Homo-like in these characters. Figure 6 shows a comparison of the two cranial angles at nasion, and here the spread of specimens reflects total facial prognathism (NAA values) and relative cranial height (NBA values). The ratio of the two angles is also a measure of the position of basion in relation to nasion. The robust australopithecine crania remain within the Pan range, while KNM-ER 3733 now appears more primitive than KNM-ER 1470 and 1813 by virtue of its relatively low cranial height. O.H. 24 and Sts 5 are distinct in their combination of (plesiomorphous) high NAA values and (derived) high NBA values. The significance of the cranial angle data will be discussed further in the concluding remarks.

FACIAL SHAPE Variation in the facial form of the Plio-Pleistocene hominids

has recently received much and deserved attention (Rak, 1983; Kimbel et al 1984). A series of facial, cranial and mandibular characteristics linking A. africanus, A. robustus and A. boisei has been identified and associated with trends in the expansion of the postcanine tooth row, an

Figure 4. Cranial angles of O.H. 24, KNM-ER 1470, 1813, and 3733, scaled to same basion-nasion dimension. Compare with figure 2.

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Stringer; Credibility of Homo habilis 281

anteriorly positioned infraorbital region and origin of the masseter muscle, and the predominance of the anterior temporalis fibres. Primitive characters linking A. afarensis and early Homo have been noted, lending support to the phylogeny proposed by White et al (1981) which identifies a distinct australopithecine clade for A. africanus, A. robustus and A. boisei, from which A. afarensis is excluded on the basis of its primitive morphology. A. afarensis, not A. africanus, is identified as the probable last common ancestor of the Homo and australopithecine clades.

My own work on the Neanderthal face (e.g. Stringer & Trinkaus, 1981) has been much concerned with defining its unique characters, and in many ways the Neanderthal cranio-facial morphology is the exact antithesis of that described by Rak (1983) for the 'robust' australopithecines. In Neander-thals it is the anterior teeth which are relatively enlarged, while the premolars are small relative to molars (Table 3). The middle of the Neanderthal face is dominated by a large and projecting nasal opening, while the zygomaxillary region is swept back, associated with posterior placement of the masseter origin and the ascending ramus of the mandible. Since this distinctive Neanderthal morphology can be readily measured using simple transverse facial angles, I decided to calculate these same angles for the Plio-Pleistocene sample in order to investigate variation in these characters.

Figure 5. Scatter diagrams of values for prosthion angle (PRA) and nasion angle (NAA). Bars indicate 2 standard deviations for Pan sample (n = 16) and 34 subsets of modern H. sapiens (Howells, 1973).

70

60

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70 80 90 100

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Two transverse facial angles can be used to measure the projection of the base of the nose (subspinale) in front of the zygomaxillare points (subspinale angle, SSA), and the projection of the origin of the nose (nasion) in front of the frontomalare points (nasiofrontal angle, NFA). As before, a sample of modern hominoid crania was measured in order to assess the probable primitive hominoid condition. A position of 'nasion' inferior to glabella was taken where necessary, and where subspinale could not be identified, the central point of a line projected between the lateral nasal borders was used instead (Howells, 1973). The values obtained for samples of modern apes were compared with those for modern H. sapiens (Howells, 1973), as shown in Table 6.

Two somewhat different patterns can be identified. One, found in the apes, is characterised by a low SSA value and relatively high NFA value, while the opposite condition is found in H. sapiens. Thus, while apes are generally characterized by a flattened upper face and projecting lower face, humans have a more projecting upper face and flatter lower face (Neanderthals are exceptional for the latter character). Data for Miocene fossil hominoids are not available, but examination of casts of Sivapithe-cus and "Ouranopithecus" specimens suggests that values for these specimens were probably similar to those of living apes, which show the assumed primitive pattern.

Figure 6. Scatter diagram of values for nasion-bregma angle (NBA) and nasion angle (NAA). Bars indicate 2 standard deviations for Pan sample (n = 16) and 34 subsets of modern

70 80 90 100

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Data for fossil hominids were obtained using casts of Plio-Pleistocene crania and original specimens for Middle and Upper Pleistocene crania. In some cases (e.g. Sts 17 and 71, and SK 847) data had to be obtained by duplicating measurements from the midline, and once again it should be noted that casts of the existing reconstructions of specimens such as O.H. 5, O.H. 24 and KNM-ER 1470 were used. The peculiar and superiorly positioned zygomaxillary regions of some australopithecine specimens also led to measurement difficulties for SSA values.

The values obtained for the fossil hominid specimens are displayed in Fig. 7, together with the means of the Pan and H. sapiens samples. Two main 'trends' are apparent, and the Pan and H. sapiens data now appear quite similar to each other in comparison with some of the fossil specimens. A large number of fossil specimens retain the primitively flat upper face, while only KNM-ER 1813 and SK 847 closely resemble H. sapiens in showing a lower NFA value. In SSA values, all the living hominoids (including H. sapiens) are relatively similar when compared with the high values for this angle found in the 'robust' australopithecines

Table 6: Data on transverse facial angles of modern hominoids, with standard deviations. Recent data from Howells (1973) with overall standard deviations calculated from means of 34 modern subsets of the data

Species Sample Subspinale angle

SSA Nasiofrontal angle

NFA

H. sapiens d 834 128.8 5.0 142.1 4.3

9 818 128.7 5.1 142.8 4.3

combined 1652 128.7 4.4 142.4 2.5

P. troglodytes d 8 110.0 6.3 155.8 5.0

9 8 114.7 4.1 154.7 4.5

combined 16 112.3 6.8 155.2 4.6

G. gorilla d 15 104.9 4.8 159.4 6.5

9 14 105.3 3.1 157.0 4.3

combined 29 105.1 4.0 158.2 5.6

P. pygmaeus d 16 108.4 5.2 163.6 5.7

9 12 117.2 5.7 161.6 3.7

combined 28 112.3 7.0 162.7 4.9

S. syndactylus d 4 88.4 3.1 150.3 5.1

9 5 87.6 5.3 149.5 4.7

combined 9 88.0 4.2 149.9 4.6

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284

Stringer: Credibility of Homo habilis

while KNM-ER 1470, and to a lesser extent Sts 71 and O.H. 24, approach the highly derived condition found in the latter group. The KNM-ER 3733 and 3883 crania, usually assigned to H. erectus, seem more primitive than KNM-ER 1813 and SK 847 in NFA values, while the A. africanus specimens show variation in SSA, from values like H. sapiens to those approaching KNM-ER 1470 and the 'robust' australopithecines.

0.H.24 falls in a rather extreme position in Fig. 7, but taking the varia-tion of the Pan and H. sapiens samples into account, it could belong to the same group as KNM-ER 1470 or Sts 5, but is much less likely to belong to a population also containing KNM-ER 1813 and SK 847. The supposedly 'dished' form of the O.H. 24 face has been discussed before by Tobias (1972), but he in fact did not consider the specimen to possess a trans-versely dished face in the manner of the 'robust' australopithecines. This is certainly true for the lower face, since the SSA value does not approach the 180° values found in 'robust' australopithecines, but the upper face appears extraordinarily flat even compared with australopithecine speci-mens. Once again some caution must be used in interpreting data such as these, where heavily reconstructed specimens are measured. The results of the analysis of facial angles will be discussed further in the next section.

THE CREDIBILITY OF HOMO HABILIS The two main arguments against the existence of H. habilis

have centred on the supposed lack of "morphological space" between A.

Figure 7. Scatter diagram of values for nasiofrontal angle (NFA) and subspinale angle (SSA). Bars indicate 2 standard deviations for Pan sample (n = 16) and 34 subsets of modern H. sapiens (Howells, 1973).

100 120 140 L 160 "" 11.`" 180 FLAT L.FACE

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africanus and E. erectus for such a species, and the sheer variation in specimens assigned to the species. Regarding the first argument, which was based on an assumed evolutionary connection between A. africanus and H. erectus, it now appears invalid when viewed against the present uncertainty about the phylogenetic position of A. africanus. Regarding the second argumeht, I hope the preceding analyses of material from Olduvai and Koobi Fora have shown that while it is difficult to separate the early Homo specimens on dental grounds, the cranial, facial and endo-cranial data suggest a wide morphological and metrical variation. The problem remaining is whether this variation is sufficient to warrant the recognition of more than one species for the material generally assigned to H. habilis, and, if so, how the individual specimens should be allocated.

As we have seen, the dental distinctions drawn by Leakey et al (1964) do generally distinguish the early Homo sample by its relatively narrow teeth (Table 3), but it is the upper dentition which seems most derived for this character. For the lower dentition, many specimens seem merely primitive in P4 and M1 shape, although O.H. 7 does appear exceptional. However, it must be remembered that the early Homo sample of Table 3 is dominated by immature or young adult specimens with little interstitial wear, and hence relatively long teeth. It is fairly easy to match, or even exceed, most of the early Homo shape index values amongst South African australopithecine specimens when unworn specimens are compared, although this is more difficult for the premolar indices of O.H. 7 and KNM-ER 1813, where A. boisei or A. afarensis values are more likely to be comparable.

There is little basis for dividing up the early Homo sample using characters such as those presented in Tables 3 and 4, apart from separating off KNM-ER 992 for its broad and relatively large P 4 . The low P4 shape index for this fossil appears highly derived, most resembling specimens such as O.H. 22, Ternifine and late Pleistocene Homo, but the high P 4 /M1

index is atypical. It has been suggested by several workers that this mandible does not belong in the same group as O.H. 7, but while Groves & Mazak (1975) separated it from all the Olduvai specimens and made it the holotype of H. ergaster, Leakey et al (1978) grouped it with KNM-ER 1813, and by implication, with O.H. 13, O.H. 24 and A. africanus. I believe the cranial and dental resemblances between O.H. 13 and KNM-ER 1813 noted by Walker (1981) certainly indicate that these specimens should be classified together, in which case the O.H. 13 mandible becomes representative of the mandibular morphology expected for KNM-ER 1813. Unfortunately, as Groves & Mazik (1975) note, the O.H. 13 mandible is unlike KNM-ER 992 in several respects, making it more difficult to add this Koobi Fora mandible to the group. An alternative possibility is that KNM-ER 992 belongs with specimens such as KNM-ER 3733 and 3883 (H.cf. erectus), although the high P 4/M 1 index would again be unusual, as already indicated. Additionally, the ascending ramus of KNM-ER 992 looks rather low to match specimens such as KNM-ER 3733, but variation in samples such as Ternifine range from one specimen like the Koobi Fora mandible to another with an ascending ramus too large to match the KNM-ER 3733 cranium. Unfortunately I have to con-clude this discussion on KNM-ER 992 by admitting that I cannot decide whether it belongs with the group containing KNM-ER 1813 and O.H. 13, or

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with H. cf. erectus. However, the characteristics of the recently dis-covered WT 15000 mandible of H. cf. erectus should be of value in resolving this question.

Regarding the mandible KNM-ER 1802, dental data presented here show it is essentially like other early Homo specimens in tooth shape indices examined, but resembles South African australopithecines in certain aspects of tooth size and morphology, as do specimens such as O.H. 7 and O.H. 16. In addition, according to data presented by Chamberlain & Wood (in press), the cross-sectional area of the mandible below Ml is very large, falling above values for all A. afarensis mandibles except AL-333-32, all early Homo specimens (unless KNM-ER 1483 is included) and all four A. africanus mandibles measured. However, it falls within the ranges for A. robustus and A. boisei in this character. Additionally it lacks the supposed Homo-like form of the inner contour of the mandible discussed by Robinson (1966) and Tobias (1974), and instead resembles Australopithecus. The premolar crowns are large and rhomboidal, and as Abbott & Wood (in press) have discussed, they have "molarised" roots like those of australopithecines, particularly 'robust' specimens. Finally, even allowing for the subadult/young adult age of the individual, the anterior origin of the ascending ramus and development of an extramolar sulcus, apparent even at the M2 position, are further indications of australopithecine affinities, using the criteria of White (1977). Putting these data together suggests that this specimen might have belonged to a population with a masticatory emphasis on vertical occlusal forces. This provides an interesting parallel with the anteriorly-placed zygomaxillary region (and inferred masseter origin) of KNM-ER 1470 (see Fig. 7), since these two specimens have generally been assumed to be closely related on other grounds (Leakey et al 1978; Wood, 1978).

The 0.H. 7 holotype mandible of H. habilis can be distinguished more readily from australopithecines in some aspects of dental shape, but less so on size or morphology (Tables 3,4). Like KNM-ER 1802 it possesses a relatively narrow P4 , but the premolars of 0.H. 7 are absolutely and relatively smaller (Table 3) and differ in morphology and root form. How-ever the O.H. 7 mandible was probably also large in corpus area, and displays an anteriorly placed ascending ramus with a developed extramolar sulcus. However comparisons with KNM-ER 1802 must be tempered by the juvenile status of 0.H. 7, which affects not only tooth wear but relative position of the ascending ramus origin. From a comparative perspective, the O.H. 7 ascending ramus certainly originates anteriorly compared with many later Homo specimens at an equivalent developmental stage, but is positioned somewhat more posteriorly than that of MLD 2, a juvenile A. africanus mandible. The parietal bones of 0.H. 7 do indicate a large endo-cranial capacity, probably approaching 700 ml, and I agree with Holloway about similarities with the parietal arch of KNM-ER 1470. Thus despite dental differences between O.H. 7 and KNM-ER 1802 I am inclined to associate them both with KNM-ER 1470 as Homo habilis.

Regarding the crania studied here it must again be stressed that casts of reconstructed specimens were used, and for KNM-ER 1470 this may have particularly influenced measures of facial prognathism, while for 0.H. 24 vz 'H'O f,oJ arrl{,o 'ursrq1eu60-rd 1erceJ Jo ssrnseaur eneq .{eur srql olrr ug-r/ig>r roJ pue ,pesn a.re&

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a number of other aspects of overall cranial form must be treated with ;teat caution. Nevertheless the conclusions regarding KNM-ER 1813, and to a lesser extent, KNM-ER 1470 seem clear. They both show derived Homo-like features in the reduced NAA of the facial triangle, and in the three angles of the anterior cranial triangle (Figs. 3-6). KNM-ER 1470, however, shows a lower and more primitive PRA value which probably cannot be entirely attributed to reconstruction, and quite distinctive transverse facial angles compared to the Homo-like values shown by KNM-ER 1813, none of which appear to be due to reconstruction (Fig. 7). While differences between KNM-ER 1470 and 1813 in NFA values follow the pattern found in dimorphic modern hominoids (male values larger i.e. flatter upper face), the SSA values show the opposite pattern (SSA values in the great apes are generally smaller in males, probably related to the larger size of the anterior dentition). Only if it were argued that Plio-Pleistocene hominids had an atypical pattern of dimorphism in lower facial projection could the differences between KNM-ER 1470 and 1813 be readily attributable to sexual dimorphism, with the former specimen as a male individual. Alternatively, if we accept that variation in endocranial volume between these two specimens rules out the possibility that they are conspecific, the SSA value of KNM-ER 1470 appears derived in the direction of the australopithecine Glade, combined with a primitively high NFA value, while KNM-ER 1813 instead appears relatively primitive in its lower SSA value, and derived in the low NFA value in the direction of Homo.

It is certainly possible that the 0.H. 24 facial triangle values are affected by distortion, as those of the anterior cranial triangle certainly appear to be. But, as reconstructed, O.R. 24 is primitive in measures of facial prognathism (Figs. 3-6) and resembles Sts 5 in this respect, as does its combination of high NAA and NBA values (Fig. 6). In transverse facial angles (Fig. 7) it is characterised by a very large NFA value and a fairly high and derived SSA value similar to values for KNM-ER 3733 and A. africanus specimens. While variation between 0.H. 24 and KNM-ER 1470 might be containable in a population range similar to the variation shown by living hominoids, the implied pattern of dimorphism, assuming 0.H. 24 to be female and KNM-ER 1470 to be male, is again contrasted with that of the great apes (Table 6). Additionally, while on the basis of their cranial angles, it is possible to accept O.H. 24 and KNM-ER 1470, or KNM-ER 1470 and 1813, as belonging to one population, it is much more difficult to accept all three as conspecific. I am inclined, therefore, to initially group O.H. 24 with KNM-ER 1470 rather than with KNM-ER 1813, on the basis of their shared higher SSA values, and their more primitive PRA and NFA values. The alternative would be to classify O.H. 24 as distinct from either, and it is true that in its present state it is the most similar overall to A. africanus of the three crania studied here. However, the cranial shape and endocranial volume of the specimen seem to ally it primarily with KNM-ER 1470 rather than with A. africanus, so provisionally it can be assigned, like KNM-ER 1470, to H. habilis.

From the preceding remarks it is evident that the degree of distinction accorded to the differences between KNM-ER 1470 and KNM-ER 1813 is critical to the delineation of the species H. habilis. Figure 8 displays a division of the early Homo material into two groups, based on an initial

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separation of the two specimens on the grounds of endocranial volume. Specimens which can be grouped with KNM-ER 1470 on various grounds, including chronostratigraphic considerations where necessary, are placed in group 1, and those linked with KNM-ER 1813 in group 2. The marked over-all similarities between KNM-ER 1813 and O.H. 13 in preserved parts make their classification together one of the most secure starting points for assessing other material. Cranial similarities between KNM-ER 1813 and 0.H. 16, for example in frontal bone, supraorbital torus and occipital morphology, can further extend this grouping, giving an indication of the expected level of sexual dimorphism. The problematic KNM-ER 1805 cranium could be grouped here or with H. cf. erectus on chronostratigraphic grounds, but given the endocranial size and sagittal cresting of the specimen, it seems reasonable to classify it with O.H. 16, which, as reconstructed, has a somewhat larger endocranial volume but temporal lines which also converge markedly posterior to bregma. Apparent facial resemblances between KNM-ER 1805 and 1470, which might instead suggest its classification with the latter specimen must be viewed with caution until new reconstructions of the face of the specimen (Walker, 1981) can be studied.

Regarding body size estimates for the early Homo groups, it is evident from similarities in overall cranial dimensions that specimens such as KNM-ER 1470 were likely to be similar to smaller individuals of H. cf. erectus in body size, and a comparable conclusion can be reached from the size of the sub-KBS postcranial material assigned to early Homo. Thus, males of group 1 were probably similar to H. cf. erectus (and some modern humans) in body size, while specimens such as O.H. 24 (group 1) and KNM-ER 1813 (group 2) were probably smaller-bodied, in the size range of A. africanus.

Referring again to Fig. 8 it is evident that the most secure taxonomic assignments are the grouping of KNM-ER 1470 with 1590, 1802 and 3732, and the grouping of KNM-ER 1813 with 0.H. 13 and 16. Both subsets can be distinguished from A. africanus, with the group 2 specimens in fact being more clearly delineated than the group 1 set which shows more retained primitive characters and a masticatory system apparently paralleling or sharing features with those of australopithecines. Nevertheless derived Homo features are also present in group 1 and with an extension of varia-tion to include O.H. 7, the specific designation becomes H. habilis. The group 2 subset of KNM-ER 1813, O.H. 13 and 0.H. 16 displays more Homo-like characters, particularly in an external cranial morphology recalling H. cf. erectus (Cronin et'al, 1981; Hublin, 1983). However there is little evidence to support assignment of any of these specimens to the same taxon as Indonesian H. erectus (or H. modjokertensis, Howell, 1978). There are large differences in cranial robusticity and endocranial size between specimens such as KNM-ER 1813 and O.H. 13, and specimens such as Sangiran 4, and this contrast extends to differences in dental proportions, where the Sangiran material more closely resembles the Zhoukoudian sample and other later Pleistocene hominids.

Thus, if the difference in endocranial volume between KNM-ER 1470 and 1813 is a valid criterion for separating them taxonomically, as Wood (in press) (sse-rd ur) pooM se '^Trecrrrrouoxel uoql 6ur1p-redes ,roJ uor.re?T-rc prr.A p sr tTBT pue OLTT US_WNII uae^qoq eurnTo^ TeTue-rropua uT ocue.r"l+fp

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also believes, the group to which KNM-ER 1813 belongs is probably neither Australopithecus, H. habilis, H. erectus nor H. modjokertensis. Allocation of the KNM-ER 992 mandible becomes critical here, but as indicated previously, I can go no further than to suggest that the appropriate species name for KNM-ER 1813 may be H. ergaster, or a new presently

unnamed species of the genus Homo. Regarding the relationships of these different groups, it is evident that both group 1 and 2 specimens share synapomorphies with members of the genus Homo, but on the characters assessed here, group 2 specimens appear closer to H. cf. erectus (Africa) and Homo sapiens. Group 1 specimens retain a more primitive cranial morphology in some respects, but also exhibit some characters found in australopithecines which may not be primitive for hominids. Such characters suggest that group 1 specimens were either derived from an ancestor sharing masticatory specialisations with australopithecines (excluding A. afarensis), or had developed such characters by parallel evolution. A gradualistic evolutionary explanation for the different characters shown by specimens such as KNM-ER 1470, O.H. 24 and KNM-ER 1813 is also possible (Cronin et al, 1981) , where the apparently earlier specimens retain more characters from an A. africanus-like ancestor, and the supposedly later specimens (such as KNM-ER 1813) show more characters of H. cf. erectus, but the endocranial volume data contradict such a scheme, since the early segMent of the sample has large values, approaching those of H. cf. erectus, while the late segment has small values, more like A. africanus. Even the argument that the early segment is dominated by male individuals, and the later segment by females seems insufficient to account for this situation.

SUMMARY AND CONCLUSIONS Thus, the data presented here provide little support for the

classification of KNM-ER 1813 as A. africanus, as long as the small endo-cranial volume of the specimen is not accorded undue significance. The same conclusion can also be reached using the criteria suggested by Walker (1976) for assessing the classification of KNM-ER 1470. It appears to represent a small-bodied sister species to H. cf. erectus and H. sapiens. As reconstructed, KNM-ER 1470 also shows Homo characters, but is both somewhat more primitive, and somewhat more like A. africanus and Paranthro-pus in maxillary shape. This is also true of O.H. 24, but this heavily reconstructed specimen is probably still too distorted to yield far-reaching conclusions. Combining KNM-ER 1470, 1802, and O.H. 24 as a group reveals the presence of more specific australopithecine characters than is sometimes recognised for H. habilis, where a large absolute endocranial volume and Homo-like postcranium have usually been emphasised. If, as this paper suggests, there were at least three Plio-Pleistocene species of 'early Homo', we must now consider the possibility that a Pliocene radia-tion of Australopithecus and Paranthropus was followed by a radiation of Homo-like forms. Given the existence of distinct species of australopithe-cine grade at opposite ends of the African continent, we must even consider the possibility that Homo-like forms might have evolved in parallel in different areas of the continent. In which case, of course, they should not all be called Homo.

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291

ACKNOWLEDGEMENTS I would like to thank B. Wood, R. Holloway, P. Andrews, M. Day,

A. Chamberlain and G. Wilson for their help or advice; K. Hebb for assist-ance in collecting comparative data on modern hominoids, and R. Kruszynski and the B.M.(N.H.) photographic studio for preparation of figures.

REFERENCES

Key references

Howell, F.C. (1978). Hominidae. In Evolution of African Mammals, ed. V.J. Maglio & H.B.S. Cooke, pp. 154-248. Cambridge: Harvard University Press.

Kimbel, W.H., White, T.D. & Johanson, D.C. (1984). Cranial morphology of Australopithecus afarensis: a comparative study based on a composite reconstruction of the adult skull. Am. J. Phys. Anthrop. 64, 337-88.

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

Tobias, P.V. (1965). New discoveries in Tanganyika: their bearing on hominid evolution. Current Anthropology, 6, 391-411 (including discussion).

Tobias, P.V. (1978). Position et Ale des australopithecin6s dans la phylogene'se humaine, avec etude particulire de Homo habilis et des theories controversees avancees 'a propos des premiers hominides fossiles de Hadar et de Laetoli. In Les origines humaines at les epoques de l'intelligence, E. Bone at al., pp. 38-77. Paris: Masson.

Wood, B.A. (1978). Classification and phylogeny of East African hominids. In Recent advances in Primatology. Volume 3. Evolution, ed. D.J. Chivers & K.A. Joysey, pp. 351-72. London: Academic Press.

Main references Abbott, S.A. & Wood, B.A. (in press). Mandibular premolar root form and

evolution in the Hominoidea. J. Anat. Anon (1971). Confusion over fossil man. Nature, 232, 294-5. Baker, J.R. (1974). Race. London: Oxford University Press. Boaz, N.T. & Howell, F.C. (1977). A gracile hominid cranium from Upper

Member G of the Shungura formation, Ethiopia. Am. J. Phys. Anthrop., 46, 93-108.

Brace, C.L., Mahler, P.E. & Rosen, R.B. (1972). Tooth measurements and the rejection of the taxon "Homo habilis". Yrbk. Phys. Anthrop. /6, 50-68.

Campbell ; B,G. (1965). The nomenclature of the Hominidae. Occ. Pap. Roy. Anthrop. Inst. 22.

Chamberlain, A.T. & Wood, B.A. (in press). A reappraisal of variation in hominid mandibular corpus dimensions. Am. J. Phys. Anthrop.

Cronin, J.E., Boaz, N.T., Stringer, C.B. & Rak, Y. (1981). Tempo and mode in hominid evolution. Nature, 292, 113-22.

Davis, P.R. (1964). Hominid fossils from Bed I, Olduvai Gorge, Tanganyika. Nature, 201, 967-70.

Day, M.H., Leakey, R.E.F., Walker, A.C. & Wood, B.A. (1976). New hominids

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The credibility of Homo habilis.

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