Dental metric assessment of the omo fossils: implications for the phylogenetic position of Australopithecus africanus

The discovery of Australopithecus afarensis has led to new interpretations of hominid phylogeny, some of which reject A. africanus as an ancestor of Homo. Analysis of buccolingual tooth crown dimensions in australopithecines and Homo species by Johanson and White (Science 202:321-330, 1979) revealed that the South African gracile australopithecines are intermediate in size between Laetoli/hadar hominids and South African robust hominids. Homo, on the other hand, displays dimensions similar to those of A. afarensis and smaller than those of other australopithecines. These authors conclude, therefore, that A. africanus is derived in the direction of A. robustus and is not an ancestor of the Homo clade. However, there is a considerable time gap (ca. 800,000 years) between the Laetoli/Hadar specimens and the earliest Homo specimens; "gracile" hominids from Omo fit into this chronological gap and are from the same geographic area. Because the early specimens at Omo have been designated A. afarensis and the later specimens classified as Homo habilis, Omo offers a unique opportunity to test hypotheses concerning hominid evolution, especially regarding the phylogenetic status of A. africanus. Comparisons of mean cheek teeth breadths disclosed the significant (P less than or equal to 0.05) differences between the Omo sample and the Laetoli/Hadar fossils (P4, M2, and M3), the Homo fossils (P3, P4, M1, M2, and M1), and A. africanus (M3). Of the several possible interpretations of these data, it appears that the high degree of similarity between the Omo sample and the South African gracile australopithecine material warrants considering the two as geographical variants of A. africanus. The geographic, chronologic, and metric attributes of the Omo sample argue for its lineal affinity with A. afarensis and Homo. In conclusion, a consideration of hominid postcanine dental metrics provides no basis for removing A. africanus from the ancestry of the Homo lineage.     

Brief communication: new reconstruction of the Taung endocast

Earlier reconstructions of the Taung endocast, from the juvenile type specimen for Australopithecus africanus, were achieved without benefit of the advanced computer technology that is available today and before morphological differences were identified that distinguish endocasts of Paranthropus from those of A. africanus. Here, we reconstruct and measure a relatively complete virtual endocast of Taung and provide a new cranial capacity estimate of 382 cm(3) and a projected adult capacity of 406 cm(3), which are smaller than previous estimates. Linear measurements and ratios were also obtained from an endocast of Sts 5 and five Paranthropus endocasts and compared with those of Taung. A number of previously unrecognized foramina, processes, and canals are identified in the bony material that adheres to the base of the Taung endocast. The newly reconstructed virtual endocast of Taung displays a number of shape features that sort it more closely with gracile than robust australopithecines, including squared-off frontal lobes in dorsal view, and the shape of the tips of its temporal poles. The Taung endocast also shares some features with Paranthropus endocasts, while other characteristics such as small temporal lobes may be due to its juvenile status. Just how much of Taung's unique morphology is due to its juvenile status may eventually be clarified by comparing its endocast with those from other juvenile australopithecines such as the 3.3-million-year-old juvenile from Dikika, Ethiopia.     

Cranial morphometry of early hominids: facial region

We report here on early hominid facial diversity, as part of a more extensive morphometric survey of cranial variability in Pliocene and early Pleistocene Hominidae. Univariate and multivariate techniques are used to summarise variation in facial proportions in South and East African hominids, and later Quaternary groups are included as comparators in order to scale the variation displayed. The results indicate that "robust" australopithecines have longer, broader faces than the "gracile" form, but that all australopithecine species show comparable degrees of facial projection. "Robust" crania are characterised by anteriorly situated, deep malar processes that slope forwards and downwards. Smaller hominid specimens, formally or informally assigned to Homo (H. habilis, KNM-ER 1813, etc.), have individual facial dimensions that usually fall within the range of Australopithecus africanus, but which in combination reveal a significantly different morphological pattern; SK 847 shows similarly hominine facial proportions, which differ significantly from those of A. robustus specimens from Swartkrans. KNM-ER 1470 possesses a facial pattern that is basically hominine, but which in some respects mimics that of "robust" australopithecines. Early specimens referred to H. erectus possess facial proportions that contrast markedly with those of other Villafranchian hominids and which suggest differing masticatory forces, possibly reflecting a shift in dietary niche. Overall the results indicate two broad patterns of facial proportions in Hominidae: one is characteristic of Pliocene/basal Pleistocene forms with opposite polarities represented by A. boisei and H. habilis; the other pattern, which typifies hominids from the later Lower Pleistocene onwards, is first found in specimens widely regarded as early representatives of H. erectus, but which differ in which certain respects from the face of later members of that species.     

Growth changes in measurements of upper facial positioning

Growth changes in the position of the midline upper face are examined for samples of Pan troglodytes, Gorilla gorilla, and modern humans. Horizontal and vertical distances between nasion and the anterior end of the cribriform plate are plotted against stage of dental development. Kendall's nonparametric correlations between facial positioning and stage of dental development are tested for significance. In African apes, the upper face becomes more projecting and positioned higher relative to the anterior cranial base. The extent of this horizontal and vertical separation reflects primarily facial size. In modern humans, the upper face becomes more projecting but is relatively stable in its vertical position. Comparison of Pan and modern human crania in the youngest dental age category indicates that the upper face of modern humans is positioned lower early in postnatal life. The position of the upper face (glabella) relative to the anterior and posterior cranial base is presented for several fossil hominid crania. The fossil crania are similar to Pan and modern humans in facial projection relative to the anterior cranial base. However, glabella is positioned low in the fossil crania. Total facial projection (relative to hormion) for Sts 5 is similar to the mean for Gorilla. Fossil Homo and robust australopithecine crania display very projecting upper faces. We suggest that the upper face of Homo is projecting due to the length of the anterior cranial fossa, while robust australopithecines possess a thick frontal bone.     

Hominins do not share a common postnatal facial ontogenetic shape trajectory

This paper examines the hypothesis raised by recent studies that postnatal trajectories of shape change in the facial skeleton are parallel between, at least, chimpanzees, modern humans and also fossil hominins, specifically australopithecines and possibly Neanderthals. In contrast, other studies point to divergences in postnatal shape trajectories within diverse groups of primates. As such there is some debate regarding the relative contributions of pre and postnatal ontogeny to adult morphological differences. This paper presents a series of geometric morphometric studies of the ontogeny of facial shape in hominins with the specific aim of resolving these issues. The results indicate that many differences in facial shape between hominins are established prenatally, however highly significant divergences of postnatal facial ontogeny are found among living hominins. Our studies point to possible differences between the shape ontogeny of the Australopithecus africanus face and that of African apes on the one hand and humans on the other. However, sampling experiments indicate that the small sample size of available specimens of A. africanus does not permit any conclusions to be drawn regarding comparative shape ontogeny of the face.     

Dental development in South African australopithecines. Part I: Problems of pattern and chronology

It is well known that humans take about twice as long as apes to mature. The traditional view that such delayed maturation was already present in australopithecines has been called into question during the past several years. We have approached this problem by looking at patterns of dental development in gracile and robust australopithecines from South Africa and comparing them to patterns found in extant humans and apes. We have employed both 2 and 3 dimensional computed tomography in our research. The dental growth patterns in these two australopithecine morphs differ, particularly in M1/I1 development. The robust australopithecines are more humanlike and the gracile australopithecines more apelike in this feature (“humanlike” and “apelike” are not used in any taxonomic sense). Pattern and chronology of dental development must be considered separately. Several major problem areas for future research are identified, most of which revolve around the issue of intra- versus interspecific variation.     

Protective buttressing of the hominin face

When humans fight hand‐to‐hand the face is usually the primary target and the bones that suffer the highest rates of fracture are the parts of the skull that exhibit the greatest increase in robusticity during the evolution of basal hominins. These bones are also the most sexually dimorphic parts of the skull in both australopiths and humans. In this review, we suggest that many of the facial features that characterize early hominins evolved to protect the face from injury during fighting with fists. Specifically, the trend towards a more orthognathic face; the bunodont form and expansion of the postcanine teeth; the increased robusticity of the orbit; the increased robusticity of the masticatory system, including the mandibular corpus and condyle, zygoma, and anterior pillars of the maxilla; and the enlarged jaw adductor musculature are traits that may represent protective buttressing of the face. If the protective buttressing hypothesis is correct, the primary differences in the face of robust versus gracile australopiths may be more a function of differences in mating system than differences in diet as is generally assumed. In this scenario, the evolution of reduced facial robusticity in Homo is associated with the evolution of reduced strength of the upper body and, therefore, with reduced striking power. The protective buttressing hypothesis provides a functional explanation for the puzzling observation that although humans do not fight by biting our species exhibits pronounced sexual dimorphism in the strength and power of the jaw and neck musculature. The protective buttressing hypothesis is also consistent with observations that modern humans can accurately assess a male's strength and fighting ability from facial shape and voice quality.     

Mechanical and spatial determinants of Paranthropus facial form

It is well documented in the anthropological literature that the distinctive morphology of the “robust” hominid facial skeleton reflects its dietary specialization. Rak (1983) has provided the most comprehensive evaluation of Paranthropus facial morphology and this important study concluded that bone strain generated during mastication was responsible for the scaling of measures of facial height and breadth. The present study evaluated Rak's analysis by examining the relationship between bizygomatic breadth and facial height in an ontogenetic series of Pan and Gorilla crania. Results of this analysis indicate that facial height and breadth dimensions were not mechanically scaled in the “robust” australopithecines. Structural analysis of African ape facial maturation was also used to examine alternative spatial methods of malar elongation in Paranthropus. It is concluded that the increased height of the malar region in these specimens is not related to either vertical expansion of the posterior facial skeleton or to expansion of the temporal fossa. Malar elongation is, however, consistent with a derived pattern of facial growth in crania possessing a thickened hard palate. © 1994 Wiley-Liss, Inc.     

Femoral lengths and stature in Plio-Pleistocene hominids

This study reports the femoral lengths of 31 Plio-Pleistocene hominids dated between 3.1 and 0.7 million years ago, and uses those lengths to estimate stature by way of the femur-stature ratio reported by Feldesman et al. (Am. J. Phys. Anthropol. 78:219-220, 1989). By this method the average female Australopithecus afarensis is 105 cm and the average male is 151 cm. The respective values are 115 and 138 cm for A. africanus. As defined by Howell (In VJ Maglio and HBS Cooke (eds): The Evolution of African Mammals. Cambridge: Harvard University Press, 1978) and Johanson et al. (Kirtlandia 28:1-14, 1978), Homo habilis is a sexually dimorphic species, with females standing 118 cm and males 157 cm. Such apparently strong dimorphism may be due to the possibility that there are actually two species of nonrobust hominids between 2 and 1.7 m.y.a. The estimate for the female Australopithecus boisei is 124 cm and for the male, 137 cm, but these estimates are especially difficult to be certain of because there are no femora that can be positively identified as male A. boisei. Australopithecus robustus is estimated to be 110 cm (female) and 132 cm (male). African Homo erectus stood 160 cm (female) and 180 cm (male). From these estimates several generalizations are apparent. First, there is apparently strong sexual dimorphism in stature in A. afarensis and H. habilis, but less in the other species. Second, the "robust" australopithecines were relatively small statured. Third, it is apparently not true that humans have been getting progressively taller throughout their evolutionary history. Some individuals were as tall as modern humans 3 m.y.a., by 2 m.y.a. one individual stood about 173 cm, and by 1.7 m.y.a. a stature of 180+ cm was not uncommon.     

Alleged synapomorphy of the M1/I1 eruption pattern in robust australopithecines and Homo: evidence from high-resolution computed tomography

Ever since Broom and Robinson (1951) published their claim that the eruption pattern of permanent incisors in robust australopithecines was most similar to that of modern man and different from that of gracile australopithecines and apes, the accuracy of this observation has been the subject of periodic debate (e.g., Wallace: Ph.D. thesis, 1972; Dean: Am. J. Phys. Anthropol. 67:251-257, 1985; Grine: Am. J. Phys. Anthropol. 72:353-359, 1987). Part of the problem is that the developing incisors in one of the specimens most crucial to this argument (SK61) are difficult to visualize clearly by conventional radiographic techniques because of the heavy mineralization in the fossil. This study reanalyzes SK 61 by high-resolution computed tomography in order to contribute to the final resolution of its incisor development. Grine's (op. cit.) assessment of the incisors as the deciduous ones, not the permanent ones, is fully confirmed. This fact, in conjunction with the observation that permanent incisor root formation had only just commenced in this specimen, further weakens the argument of M1/I1 eruption pattern synapomorphy between Homo and robust australopithecines.     

A quantitative and qualitative reanalysis of the endocast from the juvenile Paranthropus specimen l338y-6 from Omo, Ethiopia

Based on an analysis of its endocast, Holloway (1981 Am J Phys Anthropol 53:109-118) attributed the juvenile Omo L338y-6 specimen to Australopithecus africanus (i.e., gracile australopithecines) rather than to Paranthropus (Australopithecus) boisei (robust australopithecines) favored by other workers (Rak and Howell [1978] Am J Phys Anthropol 48:345-366). Holloway's attribution was based on the specimen's (1) low cranial capacity, (2) gracile-like meningeal vessels, (3) gracile-like cerebellar hemispheres, and (4) absence of an enlarged occipital/marginal (O/M) sinus system. Recent work, however, has shown that criteria 1 and 2 are not useful for sorting gracile from robust australopithecines (Culotta [1999] Science 284:1109-1111; Falk [1993] Am J Phys Anthropol 92:81-98). In this paper, we test criterion 3 by quantifying the endocranial cerebellar and occipital morphology reproduced on the Omo L338y-6 endocast, and comparing it to seven endocasts from South and East African early hominids. Our preliminary results show that metric analysis of this specimen cannot be used to sort it preferentially with either robust or gracile australopithecines. Finally, we demonstrate that, contrary to previous reports, the Omo L338y-6 endocast reproduces an enlarged left occipital sinus (criterion 4). This observation is consistent with the original attribution of the Omo specimen to robust australopithecines (Rak and Howell [1978] Am J Phys Anthropol 48:345-366). Furthermore, if Omo L338y-6 was a robust australopithecine, this discovery extends the occurrence of an enlarged O/M sinus system to one of the earliest known paranthropines. Am J Phys Anthropol 110:399-406, 1999.     

Petite bodies of the “robust” australopithecines

The “robust” australopithecines are often depicted as having large and powerfully built bodies to match their massive masticatory apparatus, but until 1988 the sample of postcranial remains attributed with certainty to this group was very limited. Almost nothing was known about the body of the East African “robust” australopithecine because taxonomic attribution of the postcrania was so uncertain. The body of the South African “robust” australopithecine had to be reconstructed from about a dozen isolated fragments of postcrania. Now a partial skeleton is attributed with confidence to the East African “robust” group along with several isolated bones. The South African sample has more than tripled. Analyses of this vastly expanded sample reveal that a large portion of postcrania attributed to “robust” australopithecines from Swartkrans Member 1 (35%) are from extraordinarily small-bodied individuals similar in size to a modern Pygmy weighing as little as 28 kg. These small elements include parts from the forelimb, spine, and hindlimb. About 22% of these Swartkrans 1 “robust” australopithecines are about the same size as a modern human weighing about 43 kgs and about 43% are larger than this standard but less than or equal to a 54 kg modern human. Approximately the same pattern is true for the Swartkrans 2 hominids, but taxonomic attribution is less certain. All of the Member 3 specimens are similar in size to the 45 kg standard. The partial skeleton of the East African “robust” australopithecine (KNM-ER 1500) has hindlimb joints that would correspond to a modern human of 34 kgs although the actual weight may be 5 to 10 kgs greater judging from shaft robusticity and forelimb size. The largest postcranial element attributed with some certainty to the East African “robust” australopithecine group (the talus, KNM-ER 1464) is about the same overall size as a modern human of 54 kgs, although its tibial facet is slightly smaller. Although many previous studies have hinted at the possibility that “robust” australopithecines had relatively small bodies, the new fossils provide substantial evidence that these creatures ranged from quite small to only moderate in body size relative to modern humans. These were the petite-bodied vegetarian cousins of our ancestors. Sexual dimorphism in body size appears to be greater than that in modern humans, similar to that in Pan, and less than that in Gorilla or Pongo, although such comparisons are of limited value given the small samples, poorly known body proportions, time averaging, and many other problems.     

How big were early hominids?

The recent discovery of new postcranial fossils, particularly associated body parts, of several Plio-Pleistocene hominids provides a new opportunity to assess body size in human evolution.¹ Body size plays a central role in the biology of animals because of its relationship to brain size, feeding behavior, habitat preference, social behavior, and much more. Unfortunately, the prediction of body weight from fossils is inherently inaccurate because skeletal size does not reflect body size exactly and because the fossils are from species having body proportions for which there are no analogues among modern species. The approach here is to find the relationship between body size and skeletal size in ape and human specimens of known body weight at death and to apply this knowledge to the hominid fossils, using a variety of statistical methods, knowledge of the associated partial skeletons of the of early hominids, formulae derived from a modern human sample, and, finally, common sense. The following modal weights for males and females emerge: Australopithecus afarensis, 45 and 29 kg; A. africanus, 41 and 30 kg; A. robustus, 40 and 32 kg; A. boisei, 49 and 34 kg; H. habilis, 52 and 32 kg. The best known African early H. erectus were much larger with weights ranging from 55 kg on up. These estimates imply that (1) in the earliest hominid species and the “robust” australopithecines body sizes remained small relative to modern standards, but between 2.0 and 1.7 m.y.a. there was a rapid increase to essentially modern body size with the appearance of Homo erectus; (2) the earliest species had a degree of body size sexual dimorphism well above that seen in modern humans but below that seen in modern gorillas and orangs which implies (along with other evidence) a social organization characterized by kin-related, multi-male groups with females who were not kin-related; (3) relative brain sizes increased through time; (4) there were two divergent trends in relative cheek-tooth size—a steady increase through time from A. afarensis to A. africanus to the “robust” australopithecines, and a decrease beginning with H. habilis to H. erectus to H. sapiens.     

Cranium of a juvenile Australopithecus boisei from the lower Omo Basin,Ethiopia

A partial cranium of a juvenile Australopithecus boisei, recovered from the Shungura Formation in the lower Omo basin, southern Ethiopia, and dated at 2.1 m.y. B.P. , is described anatomically and compared to young and adult australopithecines, modern Homo sapiens, chimpanzees, and gorillas. A resemblance to the gracile Australopithecus is observed but is attributed mainly to the generalized appearance of the Omo specimen resulting from its young individual age. An attempt is made to reconstruct part of the ontogenetic process of A. boisei. This process is compared to the developmental changes exhibited by the African great apes and modern man and is found to combine characteristics of both.     

Effects of size and locomotor adaptations on the hominid pelvis: evaluation of australopithecine bipedality with a new multivariate method

Three pelves and eight innominate bones belonging to the fossil species, Australopithecus africanus, Australopithecus robustus, Homo erectus, and Homo sapiens, have been studied biometrically and compared with those of recent humans and apes. A new method of logarithmic factorial analysis suppresses both the size effects and the size reference on pelvic proportions. In combination with principal component analysis it allows specializations to be dissociated from allometrical variations. Some morphological differences on the hominid pelvis prove to be mainly allometric. However, the pelvic morphology of australopithecines is clearly differentiated from that of the genus Homo (including H. erectus, OH 28, KNMER 3227). A. africanus (Sts 14, MLD 7, AL 288) is nearer the humans than is A. robustus (SK 50, SK 3155), which appears to be more specialized in the australopithecine lineage. The pelvic morphology of A. africanus, as integrated with the articular pelvic-femoral link, appears to be biometrically equivalent to that of humans.     

Functional morphology of the asterionic region in extant hominoids and fossil hominids

Asterionic sutural patterns in Plio-Pleistocene hominid crania have never been examined in detail. We present an analysis of this anatomical region in Australopithecus and Homo and relate different sutural patterns to functional changes in the masticatory apparatus. The great apes and A. afarensis share the common adult higher primate sutural pattern referred to as the "asterionic notch," which develops in response to the hypertrophy of posterior temporalis muscle fibers and the consequent formation of compound temporal/nuchal crests. This sutural configuration also appears to be present on the early Homo cranium KNM-ER 1805. In contrast, adult male A. boisei crania exhibit a unique pattern where the temporal squama overlaps the parietal which, in turn, overlaps the par mastoidea and the upper scale of the occipital bone. We relate this arrangement to the need to reinforce the rear of a thin-walled braincase against the net tensile forces exerted by the temporalis and nuchal muscles. The common juvenile hominoid edge-to-edge asterionic articulation is maintained in adult A. africanus, A. robustus, female A. boisei, and most Homo crania. We discuss the latter pattern in regard to anterior temporalis hypertrophy in A. africanus, A. robustus, and A. boisei and to craniofacial paedomorphosis in Homo.     

Basicranial anatomy of Plio-Pleistocene hominids from East and South Africa

The results of a metrical analysis of the basicranium of 19 Plio-Pleistocene fossil hominid crania are presented. The sample includes crania attributed to Australopithecus africanus, Australopithecus boisei, and robustus, and Homo erectus as well as crania whose attribution is still under discussion. These results confirm significant differences between the cranial base patterns of the "gracile" and "robust" australopithecines and the three crania attributed to Homo erectus have a pattern which resembles that of modern humans. None of the crania examined from East Africa sites have base patterns which resemble that of the "gracile" australopithecines. The crania KNM-ER 407 and 732 have patterns which are compatible with them being smaller-bodied females of Australopithecus boisei; KNM-ER 1470 and 1813 have base patterns which most closely resemble that of Homo erectus. The cranial base pattern of KNM-ER 1805 is compatible with its inclusion in either Australopithecus boisei or Homo. When account is taken of the immaturity of Taung, the evidence of its cranial base pattern suggests that if it had reached adulthood it would have resembled the "gracile" australopithecine crania from Sterkfontein and Makapansgat.     

A reappraisal of variation in hominid mandibular corpus dimensions

The relationship between breadth and height of the mandibular corpus has been investigated in a sample of 77 hominid mandibles. An interspecific allometric increase in robusticity with size occurs between four taxonomic subgroups of Australopithecus, but subgroups of Homo vary in robusticity while differing little in size. Within taxonomic subgroups, variation in breadth is not significantly related to variation in height among the “gracile” australapithecines; however, it is isometrically related to height in the “robust” australopithecines and bears an allometric relationship to height in Homo. Thus, robusticity, in conjunction with size, may provide a useful indicator of the taxonomic affinities of hominid mandibles.     

What lies beneath? An evaluation of lower molar trigonid crest patterns based on both dentine and enamel expression

The nearly ubiquitous presence of a continuous crest connecting the protoconid and metaconid of the lower molars (often referred to as the middle trigonid crest), is one of several dental traits that distinguish Homo neanderthalensis from Homo sapiens. This study examined variation in trigonid crest patterns on the enamel and dentine surfaces to (1) evaluate the concordance between the morphology of trigonid crests at the inner dentine and the outer enamel surfaces; (2) examine their developmental origin(s); and (3) examine trait polarity through comparison with Australopithecus africanus and Pan. The sample included 73 H. neanderthalensis, 67 contemporary H. sapiens, 5 A. africanus, and 24 Pan lower molars. Results indicate general agreement in the morphology observed on the dentine and enamel surfaces. All but one H. neanderthalensis molar shows some trigonid crest development, whereas trigonid crests occur in low frequency in contemporary humans. Pan and A. africanus both also show high frequencies of a continuous trigonid crest. However, the origin of the trigonid crest differs among groups. H. neanderthalensis uniquely possesses a 'middle' trigonid crest that originates from the mesial accessory ridge of one or both cusps. Based on our results we suggest that presence of a continuous middle trigonid crest at the dentine surface is primitive and the lack of any trigonid crest is derived. Genetic drift may explain the high frequency of trigonid crests in H.neanderthalensis. However, H. neanderthalensis still appears to be derived relative to Pan and A. africanus in its high frequency of the mesial-mesial trigonid crestconfiguration.     

How large were the australopithecines?

Stature of the African early hominids is estimated from most of the available fragments of fossil long bones by means of regression analysis. The average height of the South African gracile australopithecines is predicted to be 145.1 cm (4′9″) where n = 4 and of the South African robust forms, 152.7 cm (5′) where n = 3. The East African early hominids are somewhat taller (x = 163.0 cm or 5′4″, where n = 7). Variability in stature is high even within the same site which is probably a reflection of fairly strong sexual dimorphism in body size. Evidence is presented which suggests that at least in one form of early hominid the size proportions of fore- and hindlimbs are different than in modern man. There is also evidence that average stature may have increased through time. The most significant of these findings is that the two forms of early hominids in South Africa are possibly more similar in stature than is usually cited. This does not imply necessarily that the two forms did not differ significantly in robustness or weight.