New hominid fossils from the Swartkrans formation (1979-1986 excavations): craniodental specimens

Seventy-two individually numbered hominid craniodental fossils from recent excavations at Swartkrans are described. All derive from in situ decalcified breccia and/or unconsolidated sediments. A total of 20 specimens, representing 13 to 16 individuals derive from Member 1 "Lower Bank," two teeth derive from sediments along the Member 1-2 Interface, 38 fossils representing 19 to 24 individuals come from Member 2, and 12 teeth representing 9 to 11 individuals derive from Member 3. All but four of the specimens are attributable at the generic level; one specimen from Member 1 "Lower Bank" and five specimens from Member 2 are attributed to Homo, while the others represent Paranthropus. The proportional representation of Homo in the Swartkrans Formation is markedly higher in Member 2 (c. 33%) than in the Member 1 "Lower Bank" (c. 8%) and Member 1 "Hanging Remnant" (c. 5%) samples.     

New first metatarsal (SKX 5017) from Swartkrans and the gait of Paranthropus robustus

A new complete hallucal metatarsal (SKX 5017) was recovered from the "lower bank" of Member 1 at Swartkrans (ca. 1.8 m.y. BP). The new metatarsal is attributed to Paranthropus robustus, the predominant hominid found in Member 1 (greater than 95% of hominid individuals). SKX 5017 is similar to Olduvai Hominid 8-H from bed I, Olduvai (ca. 1.76 m.y. BP), and both resemble humans most closely among extant hominoids. The base, shaft, and head of SKX 5017 suggest human-like foot posture and a human-like range of extension (= dorsiflexion) at the hallucal metatarsophalangeal joint, while at the same time the distal articular surface indicates that a human-like toe-off mechanism was absent in Paranthropus. The fossil evidence suggests that Homo habilis and Paranthropus may have attained a similar grade of bipedality at roughly 1.8 m.y. BP.     

Stratigraphy, artefact industries and hominid associations for Sterkfontein, member 5

A revised stratigraphy for the early hominid site of Sterkfontein (Gauteng Province, South Africa) reveals a complex distribution of infills in the main excavation area between 2.8 and 1.4 m.y.a, as well as deposits dating to the mid to late Pleistocene. New research now shows that the Member 4 australopithecine breccia (2.8-2.6 Ma) extends further west than was previously thought, while a late phase of Member 4 is recognized in a southern area. The artefact-bearing breccias were defined sedimentologically as Member 5, but one supposed part of these younger breccias, the StW 53 infill, lacks in situ stone tools, although it does appear to post-date 2.6 Ma when artefacts first appear in the archaeological record. The StW 53 hominid, previously referred to Homo habilis, is here argued to be Australopithecus. The first artefact-bearing breccia of Member 5 is the Oldowan Infill, estimated at 2-1.7 Ma. It occupies a restricted distribution in Member 5 east and contains an expedient, flake-based tool industry associated with a few fossils of Paranthropos robustus. An enlarged cave opening subsequently admitted one or more Early Acheulean infills associated in Member 5 west with Homo ergaster. The artefacts attest to a larger site accumulation between ca. 1.7 and 1.4 Ma, with more intensive use of quartzite over quartz and a subtle but important shift to large flakes and heavier-duty tools. The available information on palaeoenvironments is summarized, showing an overall change from tropical to sub-tropical gallery forest, forest fringe and woodland conditions in Member 4 to more open woodland and grassland habitats in the later units, but with suggestions of a wet localized topography in the Paranthropus -bearing Oldowan Infill.     

Cladistic analysis of early Homo crania from Swartkrans and Sterkfontein, South Africa

The phylogenetic relationships of early Pleistocene Homo crania from the South African sites of Swartkrans and Sterkfontein were investigated through cladistic analyses of 99 morphological characters. The Swartkrans Member 1 specimen SK 847 and the Stw 53 cranium from Sterkfontein Member 5A were treated as separate operational taxonomic units (OTUs), distinct from the three species of early Homo-H. erectus, H. habilis, and H. rudolfensis-that are recognized from the Plio-Pleistocene deposits of East Africa. The cladistic analyses differed in the treatment of the South African OTUs (separate Swartkrans and Sterkfontein OTUs vs. a single Swartkrans+Sterkfontein OTU). PAUP 4.0 was used to construct cladograms and address hypotheses about relationships. In the analysis that treated the South African specimens as a single OTU, the position of that OTU was stable as a separate branch on the Homo clade between H. rudolfensis and [H. habilis+(H. erectus+H. sapiens)]. When SK 847 and Stw 53 were treated as separate OTUs, the majority of most parsimonious trees indicated that they were positioned in similar positions as the combined South African Homo OTU; that is, as separate branches between H. rudolfensis and [H. habilis+(H. erectus+H. sapiens)], with the Swartkrans OTU generally occupying a more derived position. The position of the Sterkfontein OTU was more stable than that of the Swartkrans OTU, which was found in several other positions among the minimum length trees. Running the analyses with only those characters preserved by SK 847 and Stw 53 resulted in similar topologies for minimum length trees, although the positions of Stw 53, SK 847, and H. habilis exchanged places in some trees. In no case was an exclusive sister relationship between either South African OTU and a particular species of Homo supported statistically. Both South African OTUs differ from H. habilis in the fewest number of cladistic characters.     

Radius of Paranthropus robustus from member 1, Swartkrans formation, South Africa

Recently recovered hominid postcrania from Member 1, Swartkrans Formation include the proximal and distal ends of a right radius attributed to a single individual of Paranthropus robustus. These fossils are essentially similar to Australopithecus afarensis, A. africanus, and P. boisei homologues. The head manifests an ape-like circumferentia articularis, and the distal end has prominent medial, dorsal, and lateral tubercles and a well developed brachioradialis crest, features also commonly exhibited by extant great apes. The volar set of the P. robustus radiocarpal joint, like that of Australopithecus homologues, more closely resembles the neutral condition exhibited by Homo than the greater flexion evinced by living apes. Compared with fossil and recent specimens of Homo, the configuration of the P. robustus radial head suggests enhanced stability against medial displacement during pronation and supination; the strong crest for the attachment of brachioradialis may attest to enhanced forearm flexor capability. In addition, this crest and the prominent dorsal tubercles may indicate enhanced hand extensor and, therefore, hand flexor capabilities. The differences in radial morphology between Paranthropus and Homo may relate to significant behavioral differences between these two synchronic taxa.     

Early hominid brain evolution: a new look at old endocasts

Early hominid brain morphology is reassessed from endocasts of Australopithecus africanus and three species of Paranthropus, and new endocast reconstructions and cranial capacities are reported for four key specimens from the Paranthropus clade. The brain morphology of Australopithecus africanus appears more human like than that of Paranthropus in terms of overall frontal and temporal lobe shape. These new data do not support the proposal that increased encephalization is a shared feature between Paranthropus and early Homo. Our findings are consistent with the hypothesis that Australopithecus africanus could have been ancestral to Homo, and have implications for assessing the tempo and mode of early hominid neurological and cognitive evolution.     

A new pelvic fragment from swartkrans and the relationship between the robust and gracile australopithecines

A recently discovered hominid pelvic fragment from Swartkrans (SK 3155) is described in detail with particular reference to the relationship of the two presently recognized forms of australopithecines in South Africa. Results of this examination and metrical analysis indicate that the acetabulum and iliac blade of the early hominids are similar to Homo sapiens except for a unique pattern of traits: a relatively small sacral articular surface, a relatively small acetabulum, a relatively large iliac fossa, and wide lateral splaying of the iliac blades. The new Swartkrans fossil expresses these traits more strongly than does the gracile australopithecine (Sts 14) and is therefore somewhat less similar to Homo sapiens but it is very unlike any pongid.     

New hominid fossils from the Swartkrans formation (1979-1986 excavations): postcranial specimens

New postcranial fossils of Paranthropus robustus and Homo cf. erectus were recovered from Swartkrans from 1979 through 1986. These fossils are from Members 1, 2, and 3. The new fossils are described here along with their morphological affinities. Fossils that are assigned to Paranthropus indicate that the South African "robust" australopithecines engaged in tool behavior and were essentially terrestrial bipeds at around 1.8 Myr BP. The manual dexterity and bipedal locomotion of Paranthropus may have equaled that of Homo habilis in East Africa at approximately the same time.     

A large male hominin cranium from Sterkfontein, South Africa, and the status ofAustralopithecus africanus

Stw 505 is the most complete hominin cranium discovered in Sterkfontein Member 4 since Broom's excavations. It was found in situ in Member 4 breccia in 1989 and is larger, on the whole, than any other cranium from Sterkfontein that has comparable parts. Displacement due to breakage, as well as plastic deformation, has affected Stw 505 in several areas, especially the face and the vault. Diagnosticmorphology is nevertheless abundant in the specimen. In several areas-the distinct anterior pillar, the straight inferior border of the zygoma, the pattern of cresting on the naso-alveolar clivus, the basal aspect of the temporal bone-Stw 505 closely matches the morphology of specimens of Australopithecus africanus and is distinct from other hominins. Some isolated characters overlap with other groups, mainly early Homo and/or A. robustus. However, only the hypodigm of A. africanus can accommodate the entire suite of morphology.In some cases, Stw 505 introduces more variation into the Sterkfontein sample. For example, prominent superciliary eminences occupy the medial portions of the supraorbital region and flow medially into a strongly protruding glabellar mound. These characteristics are probably attributable to sexual dimorphism. In many respects, Stw 505 highlights similarities between A. africanus and early Homo. Comparison with other species suggests that males of A. africanus do not show derived features of A. robustus that are not also present in females, and that cranial differences between A. afarensis and A. africanus have, if anything, been understated.     

Craniofacial diversity in earlyHomo and the affinities of the Sterkfontein Valley

The Sterkfontein Valley specimens SK 847 (Swartkrans Member 1) and Stw 53 (Sterkfontein Member 5) provide important evidence of earlyHomo in southern Africa. However, specific identity has been disputed, with that of SK 847 especially contentious. Opinions differ markedly as to whether the specimens are conspecific or not, whether they should be referred to East African earlyHomo species, or whether they represent new species. Morphometric analysis of facial dimensions reveals contrasting affinities for the two South African fossils, and so does not support claims for their conspecifity. Stw 53 is very like smaller East African crania referred toH. habilis, whereas SK 847 has a distinctive facial pattern. In some respects it resembles early AfricanH. erectus (=H. ergaster), but with a markedly more projecting mid-face, prominent zygomatic and unexpanded frontal region, all of which militate against inclusion in that species. The taxonomic implications of these contrasting facial affinities are briefly discussed.     

The hunters and the hunted revisited

The dietary niches of extinct animals, including hominids and predators, may be constrained using stable carbon isotope ratios in fossil tooth enamel.(13)C/(12)C ratios of many of the primates abundant in the faunal assemblages of Members 1 and 2 at Swartkrans, including cercopithecoids and Australopithecus (Paranthropus) robustus, and a range of other possible prey species, have been reported previously. Resulting suggestions of a mixed, or omnivorous, diet for A. robustus raise questions about niche overlap with coeval, larger brained Homo. Here we present(13)C/(12)C data from Homo and several large predators including Panthera pardus, Dinofelis sp., Megantereon cultridens and Chasmoporthetes nitidula in Member 1, and P. pardus and P. leo in Member 2, in order to compare the two hominid species and to determine likely predators of the various primates and other macrovertebrates. Results for three Homo cf. ergaster individuals are indistinguishable from those of A. robustus, showing that proportions of C(3)- and C(4)-based foods in their diets did not differ. P. pardus, Megantereon and Crocuta are shown to be likely predators of the hominids and Papio baboons in Member 1, while the Dinofelis individual concentrated on prey which consumed C(4)grasses. The hunting hyaenid C. nitidula preyed on either mixed feeders or on a range of animals across the spectrum of C(3)and C(4)variation. The data from Members 1 and 2 confirm a shift in leopard diets towards animals that consumed C(4)grasses.     

A cranial base of Australopithecus robustus from the hanging remnant of Swartkrans, South Africa

SKW 18, a partial hominin cranium recovered from the site of Swartkrans, South Africa, in 1968 is described. It is derived from ex situ breccia of the Hanging Remnant of Member 1, dated to approximately 1.5-1.8 Mya. Although partially encased in breccia, it was refit to the facial fragment SK 52 (Clarke 1977 The Cranium of the Swartkrans Hominid SK 847 and Its Relevance to Human Origins, Ph.D. dissertation, University of the Witwatersrand, Johannesburg), producing the composite cranium SKW 18/SK 52. Subsequent preparation revealed the most complete cranial base attributable to the species Australopithecus robustus. SKW 18 suffered weathering and slight postdepositional distortion, but retains considerable anatomical detail. The composite cranium most likely represents a large, subadult male, based on the incomplete fusion of the spheno-occipital synchondrosis; unerupted third molar; pronounced development of muscular insertions; and large teeth. Cranial base measures of SKW 18 expand the range of values previously recorded for A. robustus. SKW 18 provides information on anatomical features not previously visible in this taxon, and expands our knowledge of morphological variability recognizable in the cranial base. Morphological heterogeneity in the development of the prevertebral and nuchal muscular insertions is likely the result of sexual dimorphism in A. robustus, while differences in cranial base angles and the development of the occipital/marginal sinus drainage system cannot be attributed to size dimorphism.     

Dental microwear and diets of African early Homo

Conventional wisdom ties the origin and early evolution of the genus Homo to environmental changes that occurred near the end of the Pliocene. The basic idea is that changing habitats led to new diets emphasizing savanna resources, such as herd mammals or underground storage organs. Fossil teeth provide the most direct evidence available for evaluating this theory. In this paper, we present a comprehensive study of dental microwear in Plio-Pleistocene Homo from Africa. We examined all available cheek teeth from Ethiopia, Kenya, Tanzania, Malawi, and South Africa and found 18 that preserved antemortem microwear. Microwear features were measured and compared for these specimens and a baseline series of five extant primate species (Cebus apella, Gorilla gorilla, Lophocebus albigena, Pan troglodytes, and Papio ursinus) and two protohistoric human foraging groups (Aleut and Arikara) with documented differences in diet and subsistence strategies. Results confirmed that dental microwear reflects diet, such that hard-object specialists tend to have more large microwear pits, whereas tough food eaters usually have more striations and smaller microwear features. Early Homo specimens clustered with baseline groups that do not prefer fracture resistant foods. Still, Homo erectus and individuals from Swartkrans Member 1 had more small pits than Homo habilis and specimens from Sterkfontein Member 5C. These results suggest that none of the early Homo groups specialized on very hard or tough foods, but that H. erectus and Swartkrans Member 1 individuals ate, at least occasionally, more brittle or tough items than other fossil hominins studied.     

The dental developmental status of six East African juvenile fossil hominids

Radiographs of five juvenile fossil hominids from Koobi Fora, Kenya are described and presented together with measurements and observations made on the original speciments. Data are also presented for a single specimen from Olduvai Gorge, Tanzania. Four of these specimens are attributed to Paranthropus boisei (KNM ER 812, 1477 1820 and OH 30), and are all of remarkably similar dental developmental status. Conventional age estimates for these specimens of Paranthropus based on the first permanent molar, indicate an age at death of around 2·2 to 3 years. Perikymata counts on permanent lower central incisors of these specimens also indicate an age at death between 2·5 and 3 years. Two specimens attributed to early Homo (KNM ER 820 and 1507), are dentally more mature than specimens of Paranthropus boisei described here being closer to 5 years of age. Differences between the spacing and distribution of perikymata on the surfaces of incisor teeth are now apparent between Homo, Australopithecus. Paranthropus boisei and Paranthropus robustus: these are described in this paper. Details of the dental developmental patterns of these hominids are also discussed in the light of recent publications that have presented data about hominid eruption sequences and fossil hominid growth periods.     

Inferring hominoid and early hominid phylogeny using craniodental characters: the role of fossil taxa

Recent discoveries of new fossil hominid species have been accompanied by several phylogenetic hypotheses. All of these hypotheses are based on a consideration of hominid craniodental morphology. However, Collard and Wood (2000) suggested that cladograms derived from craniodental data are inconsistent with the prevailing hypothesis of ape phylogeny based on molecular data. The implication of their study is that craniodental characters are unreliable indicators of phylogeny in hominoids and fossil hominids but, notably, their analysis did not include extinct species. We report here on a cladistic analysis designed to test whether the inclusion of fossil taxa affects the ability of morphological characters to recover the molecular ape phylogeny. In the process of doing so, the study tests both Collard and Wood's (2000) hypothesis of character reliability, and the several recently proposed hypotheses of early hominid phylogeny. One hundred and ninety-eight craniodental characters were examined, including 109 traits that traditionally have been of interest in prior studies of hominoid and early hominid phylogeny, and 89 craniometric traits that represent size-corrected linear dimensions measured between standard cranial landmarks. The characters were partitioned into two data sets. One set contained all of the characters, and the other omitted the craniometric characters. Six parsimony analyses were performed; each data set was analyzed three times, once using an ingroup that consisted only of extant hominoids, a second time using an ingroup of extant hominoids and extinct early hominids, and a third time excluding Kenyanthropus platyops. Results suggest that the inclusion of fossil taxa can play a significant role in phylogenetic analysis. Analyses that examined only extant taxa produced most parsimonious cladograms that were inconsistent with the ape molecular tree. In contrast, analyses that included fossil hominids were consistent with that tree. This consistency refutes the basis for the hypothesis that craniodental characters are unreliable for reconstructing phylogenetic relationships. Regarding early hominids, the relationships of Sahelanthropus tchadensis and Ardipithecus ramidus were relatively unstable. However, there is tentative support for the hypotheses that S. tchadensis is the sister taxon of all other hominids. There is support for the hypothesis that A. anamensis is the sister taxon of all hominids except S. tchadensis and Ar. ramidus. There is no compelling support for the hypothesis that Kenyanthropus platyops shares especially close affinities with Homo rudolfensis. Rather, K. platyops is nested within the Homo + Paranthropus + Australopithecus africanus clade. If K. platyops is a valid species, these relationships suggest that Homo and Paranthropus are likely to have diverged from other hominids much earlier than previously supposed. There is no support for the hypothesis that A. garhi is either the sister taxon or direct ancestor of the genus Homo. Phylogenetic relationships indicate that Australopithecus is paraphyletic. Thus, A. anamensis and A. garhi should be allocated to new genera.     

Species attribution of the Swartkrans member 1 first metacarpals: SK84 and SKX 5020

Susman (Am. J. Phys. Anthropol. 75:277-278, 79:451-474; Science 240:781-784; In FE Grine (ed): Evolutionary History of the "Robust" Australopithecines. New York: Aldine de Gruyter, pp. 149-172) has attributed the morphologically similar SK 84 and SKX 5020 hominid first metacarpals to Homo erectus and Australopithecus robustus, respectively, and has inferred that both species exhibited derived pollical morphologies, indicating refined precision grips. Consideration of the structure of his taphonomic arguments indicates that there are no adequate nonmorphological reasons to attribute these specimens securely to one or the other of the craniodentally represented species at Swartkrans. His morphological arguments fail to note any significant differences between the two specimens. Only the contrast in size between the small SK 84 and large SKX 5020 bones might warrant a species distinction; yet comparison of their length ratio to distributions of modern human first metacarpal length ratios indicates that it is not possible to reject conclusively the null hypothesis that they are conspecific. Therefore, early hominid adaptive scenarios based on a derived Homo-like manual functional morphology in A. robustus remain without a secure paleontological basis.     

Affinities of the Swartkrans 847 hominid cranium

Further evidence of the presence of a second hominid species at the Swartkrans locality was obtained in 1969 when the SK.847 specimen was discovered by us to represent the same individual as the SK.80 maxilla. The SK.847 specimen had previously been regarded as robust australopithecine, whereas the latter was first attributed to Telanthropus capensis and subsequently to a species of the genus Homo. Recent criticism of our interpretation of these remains has not evaluated and analyzed critically the primary fossil evidence. Instead it relies on a strict adherence to an as yet unsubstantiated hypothesis that posits only a single hominid species at any point in space and time in the Cenozoic history of Hominidae.     

Variation in the Habiline Crania – Must it be Taxonomic?

The problem of whether the hominid fossil sample of habiline specimens is comprised of more than one species has received much attention in paleoanthropology. The core of this debate has significant implications about when and how variation must be explained by taxonomy. In this paper, we examine the problem of whether the observed variation in habiline sample must be interpreted to reflect species differences. We test the null hypothesis of no difference by examining the degree of variability in habiline sample in comparison with other single-species early hominid fossil samples from Sterkfontein and Swartkrans (Sterkfontein is earlier than the habiline sample; Swartkrans may be within the habiline time span). We use the standard error test for this analysis, a sampling statistic based on the standard error of the slope of regressions between pairs of specimens that relates all of the homologous measurements each pair shares. We show that the null hypothesis for the habiline sample cannot be rejected. The similarities of specimen pairs within the habiline sample are not more than those observed between the specimens in the two australopithecine samples we analyzed.