Revised age estimates of Australopithecus-bearing deposits at Sterkfontein,South Africa

The Sterkfontein fossil site in South Africa has produced the largest concentration of early hominin fossils from a single locality. Recent reports suggest that Australopithecus from this site is found within a broad paleontological age of between 2.5-3.5 Ma (Partridge [2000] The Cenozoic of Southern Africa, Oxford: Oxford Monographs, p. 100-125; Partridge et al. [2000a], The Cenozoic of Southern Africa, Oxford: Oxford Monographs, p. 129-130; Kuman and Clarke [2000] J Hum Evol 38:827-847). Specifically, the hominin fossil commonly referred to as the "Little Foot" skeleton from Member 2, which is arguably the most complete early hominin skeleton yet discovered, has been magnetostratigraphically dated to 3.30-3.33 Ma (Partridge [2000] The Cenozoic of Southern Africa, Oxford: Oxford Monographs, p. 100-125; Partridge et al. [2000a], The Cenozoic of Southern Africa, Oxford: Oxford Monographs, p. 129-130). More recent claims suggest that hominin fossils from the Jacovec Cavern are even older, being dated to approximately 3.5 Ma. Our interpretation of the fauna, the archeometric results, and the magnetostratigraphy of Sterkfontein indicate that it is unlikely that any Members yet described from Sterkfontein are in excess of 3.04 Ma in age. We estimate that Member 2, including the Little Foot skeleton, is younger than 3.0 Ma, and that Member 4, previously dated to between 2.4-2.8 Ma, is more likely to fall between 1.5-2.5 Ma. Our results suggest that Australopithecus africanus should not be considered as a temporal contemporary of Australopithecus afarensis, Australopithecus bahrelghazali, and Kenyanthropus platyops.     

Vertebrae numbers of the early hominid lumbar spine

General doctrine holds that early hominids possessed a long lumbar spine with six segments. This is mainly based on Robinson's (1972) interpretation of a single partial Australopithecus africanus skeleton, Sts 14, from Sterkfontein, South Africa. As its sixth last presacral vertebra exhibits both thoracic and lumbar characteristics, current definitions of lumbar vertebrae and lumbar ribs are discussed in the present study. A re-analysis of its entire preserved vertebral column and comparison with Stw 431, another partial A. africanus skeleton from Sterkfontein, and the Homo erectus skeleton KNM-WT 15000 from Nariokotome, Kenya, did not provide strong evidence for the presence of six lumbar vertebrae in either of these early hominids. Thus, in Sts 14 the sixth last presacral vertebra has on one side a movable rib. In Stw 431, the corresponding vertebra shows indications for a rib facet. In KNM-WT, 15000 the same element is very fragmentary, but the neighbouring vertebrae do not support the view that it is L1. Although in all three fossils the transitional vertebra at which the articular facets change orientation seems to be at Th11, this is equal to a large percentage of modern humans. Indeed, a modal number of five lumbar vertebrae, as in modern humans, is more compatible with evolutionary principles. For example, six lumbar vertebrae would require repetitive shortening and lengthening not only of the lumbar, but also of the entire precaudal spine. Furthermore, six lumbar vertebrae are claimed to be biomechanically advantageous for early hominid bipedalism, yet an explanation is lacking as to why the lumbar region should have shortened in later humans. All this raises doubts about previous conclusions for the presence of six lumbar vertebrae in early hominids. The most parsimonious explanation is that they did not differ from modern humans in the segmentation of the vertebral column.     

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.     

Possible Brucellosis in an Early Hominin Skeleton from Sterkfontein,South Africa

We report on the paleopathological analysis of the partial skeleton of the late Pliocene hominin species Australopithecus africanus Stw 431 from Sterkfontein, South Africa. A previous study noted the presence of lesions on vertebral bodies diagnosed as spondylosis deformans due to trauma. Instead, we suggest that these lesions are pathological changes due to the initial phases of an infectious disease, brucellosis. The macroscopic, microscopic and radiological appearance of the lytic lesions of the lumbar vertebrae is consistent with brucellosis. The hypothesis of brucellosis (most often associated with the consumption of animal proteins) in a 2.4 to 2.8 million year old hominid has a host of important implications for human evolution. The consumption of meat has been regarded an important factor in supporting, directing or altering human evolution. Perhaps the earliest (up to 2.5 million years ago) paleontological evidence for meat eating consists of cut marks on animal remains and stone tools that could have made these marks. Now with the hypothesis of brucellosis in A. africanus, we may have evidence of occasional meat eating directly linked to a fossil hominin.     

Stw 441/465: a new fragmentary ilium of a small-bodied Australopithecus africanus from Sterkfontein, South Africa

In 1986 and 1987, a hominid left ilium fragment consisting of a spina iliaca anterior superior and crista iliaca was discovered during excavations at Sterkfontein, South Africa. Although the specimen is small it gives valuable hints for muscle insertions and origins at the pelvis of Australopithecus africanus. It indicates that the anatomy of the abdominal muscles and of the mm. glutei medius et minimus of A. africanus was quite different from that of the great apes and more similar to that of modern humans. This has major implications for the interpretation of the bipedalism and locomotor efficiency of the early South African hominids.     

The carbon isotope ecology and diet of Australopithecus africanus at Sterkfontein,South Africa

The stable carbon isotope ratio of fossil tooth enamel carbonate is determined by the photosynthetic systems of plants at the base of the animal's foodweb. In subtropical Africa, grasses and many sedges have C(4)photosynthesis and transmit their characteristically enriched 13C/(12)C ratios (more positive delta13C values) along the foodchain to consumers. We report here a carbon isotope study of ten specimens of Australopithecus africanus from Member 4, Sterkfontein (ca. 2.5 to 2.0Ma), compared with other fossil mammals from the same deposit. This is the most extensive isotopic study of an early hominin species that has been achieved so far. The results show that this hominin was intensively engaged with the savanna foodweb and that the dietary variation between individuals was more pronounced than for any other early hominin or non-human primate species on record. Suggestions that more than one species have been incuded in this taxon are not supported by the isotopic evidence. We conclude that Australopithecus africanus was highly opportunistic and adaptable in its feeding habits.     

The view from the Lincoln Cave: mid- to late Pleistocene fossil deposits from Sterkfontein hominid site, South Africa

The Lincoln-Fault cave system lies adjacent to the Sterkfontein Cave system in the Cradle of Humankind World Heritage Site, Gauteng Province, South Africa. Lincoln Cave contains a mid- to late Pleistocene fossiliferous deposit which has been dated using uranium series methods to between 252,600+/-35,600 and 115,300+/-7,700 years old. Although speleologists presumed that there was no connection between the Lincoln Cave and Sterkfontein Cave systems, results of excavations conducted in 1997 suggest a link between the deposits. Detailed comparisons of artifacts, fauna, hominid material, and a statistical correspondence analysis (CA) of the macromammalian fauna in the deposits strongly support this hypothesis. The recovery of Early Acheulean-type artifacts from the Lincoln Cave suggests that older artifacts eroded out of Sterkfontein Member 5 West and were redeposited into the younger Lincoln Cave deposits. The close physical proximity of these deposits, and the nature of the material recovered from them, indicates that the material was probably redeposited via a link between the two cave systems. Although faunal mixing is present, it is possible to say that large carnivorans become more scarce at Sterkfontein during the mid- to late Pleistocene, while small canids and felids appear to become more abundant, indicating that large and small carnivorans probably varied their use of the site through time. This may also reflect an increasing presence of humans in the Sterkfontein area during the mid- to late Pleistocene.     

Sr/Ca and early hominin diets revisited: new data from modern and fossil tooth enamel

A previous study of strontium/calcium (Sr/Ca) ratios in Paranthropus suggested that it consumed more animal foods than was previously believed. However, that study looked at Sr/Ca in fossil bone, which is known to be highly susceptible to diagenesis. Enamel, in contrast, is resistant to post-mortem alteration making it a more appropriate material for Sr/Ca analysis of Plio-Pleistocene fossils. Yet, we know virtually nothing about Sr/Ca in the enamel of modern African mammals, much less fossil taxa. To address this gap, we studied Sr/Ca in tooth enamel from modern mammals in the greater Kruger National Park, South Africa, as well as fossil fauna from the Sterkfontein Valley. Grazing herbivores have the highest Sr/Ca, followed by browsers and carnivores in both modern and fossil fauna. This similarity in ecological Sr/Ca patterning between modern and fossil fauna shows that diagenesis has not obscured the primary dietary signals. Australopithecus has significantly higher Sr/Ca than Paranthropus, and higher Sr/Ca than fossil papionins, browsers, and carnivores. Paranthropus has lower Sr/Ca than grazers, but its Sr/Ca is higher or equal to that of fossil papionins, browsers, and carnivores. Thus, Sr/Ca for both hominins is relatively high, and provides no direct evidence for omnivory in either taxon. The consumption of underground resources or insects are among the possible explanations for the highly elevated Sr/Ca in Australopithecus.     

The alpha taxonomy of Australopithecus at Sterkfontein: The postcranial evidence

The possibility that the fossils attributed to Australopithecus africanus represent more than a single species is of significance because of the pivotal role that A. africanus has played in discussions about hominin evolution. The A. africanus hypodigm that is currently widely recognized evinces considerable variation in a number of craniodental characters, and this has led to speculation that more than one australopith taxon may be represented among the specimens from Sterkfontein. Although crania, mandibles and teeth have dominated these taxonomic discussions, the Sterkfontein postcranial remains also have been invoked. While several workers have proposed that some of the craniodental remains from Sterkfontein can be partitioned into two groups, there is a notable lack of agreement among them as to their actual sorting. Most of the craniodental observations that have been put forward in support of arguments for taxonomic heterogeneity of the Sterkfontein australopith assemblage have been subjective and anecdotal in nature. So too, the postcranial evidence that has been cited in support of more than one australopith species at Sterkfontein has been largely subjective, and limited to a small number of elements. The results of quantitative statistical analyses of the craniodental and postcranial fossils that have been undertaken to date are not necessarily consistent with the hypothesis of taxonomic heterogeneity.     

Palaeomagnetic analysis of the Sterkfontein palaeocave deposits: implications for the age of the hominin fossils and stone tool industries

Palaeomagnetic analysis was conducted on speleothems from Members 1-5 at Sterkfontein Cave, South Africa. Palaeomagnetic analysis of siltstone and speleothem from the bulk of Member 4 indicate a reversed magnetic polarity that dates the deposits and its Australopithecus africanus fossils to between 2.58 and ∼2.16 Ma. Further confirmation of this age comes in the form of two short normal polarity events correlated to the Rèunion (∼2.16 Ma) and Huckleberry Ridge (∼2.05 Ma) events in speleothem capping the bulk of Member 4 and coeval with deposition of the final phase of Member 4, including A. africanus fossil Sts 5. At ∼2.16-2.05 Ma, Sts 5 is the youngest representative of A. africanus yet discovered. Palaeomagnetic analysis of the Silberberg Grotto deposits identifies a single short geomagnetic field event in flowstone overlying the StW 573 Australopithecus fossil, which is suggested to represent the Rèunion event at ∼2.16 Ma. This further supports the uranium lead age estimates of 2.3-2.2 Ma for the StW 573 fossil. Based on a reversed polarity for the deposits below the skeleton it cannot be older than 2.58 Ma. If StW 573 is considered to be a second species of Australopithecus then this indicates that two species of Australopithecus are present at Sterkfontein between 2.6 and 2.0 Ma. All of the Member 5 deposits date to less than 1.8 Ma based on a comparison of palaeomagnetic, faunal, and electron spin resonance age estimates. The StW 53 fossil bearing infill (M5A) is intermediate in age between Member 4 and the rest of Member 5 (B-C) at around 1.78-1.49 Ma. The rest of Member 5 (B-C) containing Oldowan and Acheulian stone tools and Homo and Paranthropus fossils was deposited gradually between 1.40 and 1.07 Ma, much younger than previously suggested.     

Principal components analysis of distal humeral shape in Pliocene to recent African hominids: the contribution of geometric morphometrics

The shape of the distal humerus in Homo, Pan (P. paniscus and P. troglodytes), Gorilla, and six australopithecines is compared using a geometric approach (Procrustes superimposition of landmarks). Fourteen landmarks are defined on the humerus in a two-dimensional space. Principal components analysis (PCA) is performed on all superimposed coordinates. I have chosen to discuss the precise place of KNM-KP 271 variously assigned to Australopithecus anamensis, Homo sp., or Praeanthropus africanus, in comparison with a sample of australopithecines. AL 288-1, AL 137-48 (Hadar), STW 431 (Sterkfontein), and TM 1517 (Kromdraai) are commonly attributed to Australopithecus afarensis (the two former), Australopithecus africanus, and Paranthropus robustus, respectively, while the taxonomic place of KNM-ER 739 (Homo or Paranthropus?) is not yet clearly defined. The analysis does not emphasize a particular affinity between KNM-KP 271 and modern Homo, nor with A. afarensis, as previously demonstrated (Lague and Jungers [1996]     

From apes to humans: locomotion as a key feature for phylogeny

If bipedalism has often been considered to be of a crucial interest for understanding hominid evolution, the acceptance of locomotor features to build phylogenies is still far from being a reality in the field. Especially for hominid evolution, it still seems to be difficult to accept that traits, other than craniodental ones, can be useful for defining the major dichotomies. The recent discovery of Australopithecus anamensis suggests a challenging view of the major dichotomy between apes and humans. Whilst it is widely accepted that Ardipithecus ramidus is ancestral to Australopithecus anamensis, which in its turn is ancestral to Australopithecus afarensis and then to later hominids, the postcranial adaptations, which should be taken into account, suggest another branching pattern. Based on the fact that by 4.0 million years two different locomotor patterns can be identified in hominids, two lineages would appear to be present: the "Australopithecine" lineage (with Australopithecus afarensis or Ardipithecus ramidus if the latter is really a hominid sensu stricto) and the "Hominine" lineage (with Australopithecus anamensis = Praeanthropus africanus).     

New craniodental fossils of papionin monkeys from Cooper's D,South Africa

Papionin monkey fossils are common in the Plio‐Pleistocene aged karst cave deposits northwest of Johannesburg in South Africa. These deposits have yielded important primate and other vertebrate fauna since their discovery in the early part of the 20th century. In this article, we describe new primate cranial and dental specimens from excavations at the site of Cooper's D in the Sterkfontein Valley that date to around 1.5 million years ago. Unlike other localities in southern Africa, most of the new fossils are referred to Theropithecus oswaldi oswaldi, an extinct gramnivorous monkey related to the living gelada. Diagnostic features of T. o. oswaldi crania and teeth include large, thickly enameled molars with tall, columnar cusps, and high molar relief, an upright mandibular ramus, postorbital constriction, and anterior fusion of temporal lines. Also present in the new sample are teeth referred to Papio sp., which show low crowned bunodont molars, and a number of indeterminate papionin teeth and skull fragments. The presence of T. o. oswaldi at Cooper's D extends the list of known localities where the taxon is found, and may indicate the presence of an open, grassland environment in the area during the early Pleistocene. The abundance of theropith fossils at Cooper's suggests that Papio was not consistently the most common papionin in southern Africa over the past three million years. Am J Phys Anthropol 151:613–629, 2013. © 2013 Wiley Periodicals, Inc.     

Sex at Sterkfontein: 'Mrs. Ples' is still an adult female

The important question of whether the Australopithecus africanus hypodigm is taxonomically heterogeneous revolves largely around the interpretation of the morphological variation exhibited by the fossils from Sterkfontein. The sex assignment of these specimens is a critical component in the evaluation of their morphological variability. The Sts 5 cranium is pivotal in this regard because it is the most complete and undistorted specimen attributed to A. africanus. Although it has traditionally been regarded as an adult female, this view has been challenged. In particular, it has been argued recently that Sts 5 is a juvenile and that this, together with alveolar bone loss that has supposedly reduced the size of the canine socket, has led to its misinterpretation as a female. Virtual reconstruction of the M³ roots (and/or alveoli) contradicts arguments that these teeth were erupting at the time of death. Regardless, canine emergence and root completion are well ahead of M³ development in juvenile australopiths from Sterkfontein. Thus, even if the M³ root of Sts 5 was incomplete, its canine root would have been fully formed. Measurements of palate depth indicate that the alveolar margins of Sts 5 have not suffered from much (if any) bone loss in the region of the C/P³; any additional bone would result in a palate of truly exceptional depth. Therefore, the dimensions of the canine alveolus of Sts 5 can be regarded as proxies for those of the canine root. The canine root of Sts 5 is among the smallest recorded for any Sterkfontein australopith, which provides strong support for Robert Broom's initial attribution of sex to this specimen. There is no evidence to contradict the assertion that ‘Mrs. Ples’ is an adult female.     

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.     

Brief communication: Possible third molar impactions in the hominid fossil record

Impacted third molars affect 15%–20% of modern Americans and Western Europeans. In contrast, third molar impactions have not been reported in the early hominid fossil record. It is uncertain whether the lack of reports reflects an absence of impactions or a failure to recognize them. This communication is intended to raise awareness of the possibility of impactions by describing the appearance of impacted teeth and by noting two possible instances of impaction in early hominids. Specifically, the mandibular third molars of the Sterkfontein specimen, STS52b (Australopithecus africanus), and the left maxillary third molar of the Lake Turkana specimen, KNM-WT17400 (Australopithecus boisei), are positioned in a manner which suggests that they would not have erupted normally. Both specimens also exhibit strong crowding of the anterior dentition, providing further support for the view that these individuals lacked sufficient space for normal eruption of the third molars. Other published reports of dental crowding in the hominid fossil record are noted, and it is suggested that more attention be paid to dental impaction and dental crowding in hominid evolution. © 1993 Wiley-Liss, Inc.     

Role of carnivores in the accumulation of the Sterkfontein Member 4 hominid assemblage: a taphonomic reassessment of the complete hominid fossil sample (1936-1999)

New taphonomic data on the Sterkfontein Member 4 (South Africa) fossil hominid assemblage are presented. The previous estimate of hominid individuals represented in the deposit (45) is increased to 87. New minimum numbers of hominid skeletal elements are provided, and incidences of bone surface damage inflicted by prehistoric biological agents are summarized. The hominid sample from Member 4 is composed predominately of gnathic remains and has a paucity of postcrania. This dearth of postcrania limits, to some extent, inferences about the formation of the Sterkfontein assemblage. However, carnivore tooth marks on some fossil specimens and an overall broad similarity in patterns of skeletal part representation between Sterkfontein and primate bone assemblages created by extant carnivores suggest that carnivores did have some involvement in the accumulation of the fossil hominid assemblage. Thus, this study provides support for the “carnivore‐collecting hypothesis” of Brain (Brain [ 1981 ] The Hunters or the Hunted? Chicago: University of Chicago Press), which implicates large carnivores as prominent collecting agents of hominid body parts in Sterkfontein Member 4. Evidence of bone surface damage is, however, too scant to make confident inferences about specific carnivore taxon/taxa involved in hominid bone collection at the site. Am J Phys Anthropol, 2004. © 2004 Wiley‐Liss, Inc.