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.     

Taxonomic affinity of the early Homo cranium from Swartkrans,South Africa

A quantitative analysis that employs randomization methods and distance statistics has been undertaken in an attempt to clarify the taxonomic affinities of the partial Homo cranium (SK 847) from Member 1 of the Swartkrans Formation. Although SK 847 has been argued to represent early H. erectus, exact randomization tests reveal that the magnitude of differences between it and two crania that have been attributed to that taxon (KNM-ER 3733 and KNM-WT 15000) is highly unlikely to be encountered in a modern human sample drawn from eastern and southern Africa. Some of the variables that differentiate SK 847 from the two early H. erectus crania (e. g., nasal breadth, frontal breadth, mastoid process size) have been considered to be relevant characters in the definition of that taxon. Just as the significant differences between SK 847 and the two early H. erectus crania make attribution of the Swartkrans specimen to that taxon unlikely, the linkage of SK 847 to KNM-ER 1813, and especially Stw 53, suggests that the Swartkrans cranium may have its closest affinity with H. habilis sensu lato. Differences from KNM-ER 1813, however, hint that the South African fossils may represent a species of early Homo that has not been sampled in the Plio-Pleistocene of eastern Africa. The similarity of SK 847 and Stw 53 may support faunal evidence which suggests that Sterkfontein Member 5 and Swartkrans Member 1 are of similar geochronological age. © 1993 Wiley-Liss, Inc.     

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.     

Brief communication: cutmarks on a plio-pleistocene hominid from Sterkfontein, South Africa

Cutmarks inflicted by a stone tool were observed on the right maxilla of Stw 53, an early hominid partial skull from Sterkfontein "Member 5" (South Africa). The morphology of the marks, their anatomical placement, and the lack of random striae on the specimen all support an interpretation of this linear damage as cutmarks. The location of the marks on the lateral aspect of the zygomatic process of the maxilla is consistent with that expected from slicing through the masseter muscle, presumably to remove the mandible from the cranium. Although radioisotopic dates are not available and relative faunal dating of the deposit from which Stw 53 derives is problematic, the morphology of the hominid skull suggests a Plio-Pleistocene age for the specimen. This therefore constitutes the earliest unambiguous evidence that hominids disarticulated the remains of one another.     

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 natural history of the helicoidal occlusal plane and its evolution in early Homo

In modern man the pitch of the occlusal plane may vary along the tooth-row. When anterior cheek-teeth show a plane sloping upward palatally, whilst that on posterior cheek-teeth slopes upward buccally, there results a twisted or helicoidal occlusal plane (Ackermann). Several hypotheses have been proposed for the structural basis of the helicoidal occlusal plane. Campbell's proposal ('25) has gained widest acceptance, namely that the helicoid results from anteroposterior differences in upper and lower alveolar arch width. In the early 1960s, while studying the Olduvai hominids assigned to Homo habilis, the author noted changing occlusal slopes along the tooth-row and a slight helicoid, although these featues had not been noted in other early hominids. Subsequently, Wallace showed a total absence of the helicoid from South African australopithecines, and its presence in Swartkrans Homo, SK 45 and SK 80. Recent studies confirm the presence of the helicoid in all available specimens of H. habilis, including Stw 53 found at Sterkfontein in 1976. Hence, this trait may distinguish between Australopithecus and early Homo. Measurements of the maxillary arch widths have shown that, whereas in Australopithecus arch widths increase to a maximum at M3, in early Homo maxillary arch widths are greatest at M2. The decline in posterior maxillary arch width is part of a general reduction of that region. Thus despite striking elongation of premolars and M1 in early Homo, M2 and M3 are mesiodistally abbreviated. It is hypothesized that the onset of posterior arch reduction, with the appearance of a helicoid, was a structural and functional concomitant of the transition from the presumed australopithecine ancestor to H. habilis.     

Molar microwear and dietary reconstructions of fossil cercopithecoidea from the Plio-Pleistocene deposits of South Africa

The South African Plio-Pleistocene cave deposits have yielded a diverse cercopithecoid fauna. In this study, the possible dietary proclivities of these extinct species are examined using details of molar microwear. Although sample sizes are often small, wear patterns suggest possible temporal changes in the diets of Parapapio jonesi from Makapansgat to Sterkfontein, of Papio robinsoni from Sterkfontein to Swartkrans, and Cercopithecoides williamsi from Makapansgat to Sterkfontein to Swartkrans. However, there does not appear to have been a significant change in the dietary habits of Parapapio broomi over time. The microwear patterns of the two temporally successive congeners, Theropithecus darti and T. oswaldi show no significant differences from one another. The sympatric congeners, Parapapio broomi and Pp. jonesi, have microwear signatures that differ significantly at Makapansgat (Members 3 and 4) but not at Sterkfontein (Member 4). Finally, the microwear analyses suggest that the extinct cercopithecoid species did not necessarily have diets similar to those of their closest living relatives.     

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.     

Description d’un bassin fragmentaire de Paranthropus robustus du site Plio-Pléistocène de Drimolen (Afrique du Sud)

The Plio-Pleistocene site of Drimolen (Gauteng Province, South Africa) has yielded a fragmentary adult pelvis, DNH 43, composed of a robust, right innominate bone (DNH 43B) associated with the sacrum and the posterior arch of the last lumbar vertebra (DNH 43 A). Comparisons with other Plio-Pleistocene hominids (AL 288.1, Sts 14, Sts 65, Stw 431, TM 1605, SK 50, SK 3155b) pelves have been made which indicate that this specimen belongs to Paranthropus robustus. The ilium of the Drimolen hominid is narrowed just above the acetabulum. The small auricular surface is set far away from the acetabulum. The sacrum is curved, the upper lateral angles are clearly expressed. Some anatomical features reveal a lumbar lordosis in DNH 43. This lumbar lordosis is specific of australopithecines because different from extant human.     

Reappraisal of the taxonomic status of the cranium Stw 53 from the Plio/Pleistocene of Sterkfontein,in South Africa

A fossil skull, Stw 53, from the Plio/Pleistocene of Sterkfontein, in South Africa, has been referred toHomo habilis Leakey, Napier, andTobias, 1964. Reappraisal of its putative hominine affinity reveals a closer resemblance toAustralopithecus africanus Dart, 1925. The skull, as reconstructed, is too small forH. habilis; with no indication of brain expansion overA. africanus; has a facial angle outside the hominine range, but identical with that ofA. africanus; and whose teeth are not elongated but display buccolingual expansion. Although it was found in the same strata (Member 5) as stone tools, there is no causal connection. It has been dated faunistically at 2–1.5 my BP, but due to an unconformity it is suggested that it could be older. In spite of its late date, Stw 53 shows no intermediate characters which could support a trend towardsH. habilis orA. robustus Broom, 1938. It may, therefore, represent a relict population ofA. africanus.     

Carnivora from the Plio-Pleistocene hominin site of Drimolen, Gauteng, South Africa

Drimolen is one of the newest and most productive hominin sites in South Africa, and is dated on faunal grounds between 2.0 Ma to 1.5 Ma. This paper provides the first overview of the Carnivora from Drimolen, updating the previously published preliminary faunal list, and describing all currently prepared craniodental and postcranial material. The Drimolen specimens are described in comparison with other modern and fossil South African carnivore material. The carnivores cover a range of taxa including hyaenids, felids, canids and herpestids. Most notable amongst these are the sabretooth Dinofelis aff. piveteaui craniodental and postcranial remains, which are described in detail, and a Chasmaporthetes nitidula cranium. The genus Chasmaporthetes is found at three other sites in the area - Sterkfontein, Swartkrans and Coopers D. There are two models for the geographic origin of Dinofelis piveteaui, in that it may have arisen in either eastern or southern Africa. These possibilities are discussed in the light of the new South African Dinofelis material, as the Drimolen material appears to represent a more primitive form with affinities with D. piveteaui. Fossil leopard material from Kromdraai B and Drimolen is also discussed, as the metapodia assigned to P. pardus from these two sites are very small, but lie within the variation of modern leopards. Such size differences in fossil postcrania may have implications for the niches that these animals may have occupied in the past.     

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.     

Faunal assemblage seriation of southern African Pliocene and Pleistocene fossil deposits

Fossil assemblages from the Pliocene and Pleistocene of southern Africa were seriated in order to give a better idea of their relative chronology. Time-sensitive mammals were selected for calculation of the Faunal Resemblance Index among 17 site units. On the basis of a logistical seriation and subsequent site analysis, the following sequence of sites was deemed most probable: Makapansgat Member 3, Makapansgat Member 4, Taung Dart deposits, Sterkfontein Member 4 and Taung Hrdli?ka deposits, Sterkfontein Member 5 (in part) and Kromdraai B, Kromdraai A and Swartkrans Member 1, Swartkrans Member 2, Swartkrans Member 3, Plovers Lake, Cornelia, Elandsfontein Main Site, Cave of Hearths Acheulian levels, Florisbad and Equus Cave and Klasies River Mouth. © 1995 Wiley-Liss, Inc.     

Computed tomography and enamel thickness of maxillary molars of Plio-Pleistocene hominids from Sterkfontein, Swartkrans, and Kromdraai (South Africa): An exploratory study.

This paper is one in a series which explores the possibility of using the non-destructive CT technique to identify patterns in tooth enamel distribution and structure of hominid molars from Plio-Pleistocene sites in South Africa, notably Swartkrans, Sterkfontein, and Kromdraai. Whereas previous investigators have emphasised gross differences in absolute and relative or average enamel thickness between hominid taxa, the present study highlights differences in enamel thickness over functionally significant regions of the crown. Differences in the distribution of enamel in A. robustus, A. africanus, and Homo sp. are identified through the use of bivariate and multivariate analyses, and are interpreted in terms of dietary regimes.     

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.     

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.     

Beyond leopards: tooth marks and the contribution of multiple carnivore taxa to the accumulation of the Swartkrans Member 3 fossil assemblage

The ca. 1.0 myr old fauna from Swartkrans Member 3 (South Africa) preserves abundant indication of carnivore activity in the form of tooth marks (including pits) on many bone surfaces. This direct paleontological evidence is used to test a recent suggestion that leopards, regardless of prey body size, may have been almost solely responsible for the accumulation of the majority of bones in multiple deposits (including Swartkrans Member 3) from various Sterkfontein Valley cave sites. Our results falsify that hypothesis and corroborate an earlier hypothesis that, while the carcasses of smaller animals may have been deposited in Swartkrans by leopards, other kinds of carnivores (and hominids) were mostly responsible for the deposition of large animal remains. These results demonstrate the importance of choosing appropriate classes of actualistic data for constructing taphonomic inferences of assemblage formation. In addition, they stress that an all-encompassing model of assemblage formation for the hominid-bearing deposits of the Sterkfontein Valley is inadequate and that each must be evaluated individually using not just analogical reasoning but also incorporating empirical data generated in the preserved fossil samples.     

Brief communication: evidence bearing on the status of Homo habilis at Olduvai Gorge

Students of the early hominin career have debated the status of Homo habilis since its discovery in 1960. Today discussion centers on which specimens should be included in the species and what constitutes the holotype. Recent reviews of early Homo suggest that the Olduvai Hominid 8 foot may sample Paranthropus while the OH 7 skull bones, mandible, and hand sample H. habilis. Moreover, some suggest that while H. habilis in Middle Bed I at Olduvai is craniodentally Homo-like, the postcranial skeleton of H. habilis is more like that of Australopithecus. Evidence presented here indicates not only that OH 7 and OH 8 represent H. habilis but also that they come from a single individual. The association of OH 35 with OH 7 and OH 8 is less certain. Morphological, pathological, and taphonomic evidence favors the inclusion of OH 35 in the holotype. However, stratigraphic evidence suggests that OH 35 and OH 8 are not coterminous. With or without OH 35, the holotype of H. habilis ranks as one of the most complete early hominin skeletons and the most complete and functionally informative specimen of early Homo.