Re-examination of the lower stratigraphy in the classic section, limeworks site, Makapansgat, South Africa

At the end of Member 2 times, remnant mammillary-form stalactites in the roof and wall contacts show that an irregular wedge-shaped space existed from the Classic Section out to the Cone area. It was into this space that hyenas and other denning animals were able to gain access from an as yet unfilled Cone area. During the mining operations of the 1920’s, Eitzman (1958) recollects entering an impressive bone breccia resembling a charnel house. We suggest that it is most probable that the Grey Breccia corresponds to his main breccia as only the area on top of Member 2, between the dolomite walls, roof and stalagmite boss has the required volume. It also lies at the end of an incline first dug by the miners, (still in existence), and at the end of which he was able to note some of the deposits. The bone breccias at the same level at the back of the Cone suggest that other dens were created at about this time. As the roof continued to retreat upslope and more material was washed in or fell in from the surface so more sediment was winnowed into the back of the Cone. At some stage this allowed prey animals to enter the Cone area and gain access between the speleothem barrier to the area of the Classic Section. As most Australopithecine fragments came from this breccia, it seems likely that, as with Swartkrans, the hominid fragments are the result of predation. Then as sediments continued to wash into the Cone area, the Classic Section and the back of the Cone became inaccessible for denning purposes. Large speleothem remnants embedded in sediment starting part-way up the Cone shows that a roof continued to exist in this area for some time. In fact, even today, although bits of speleothems can be found in the eroded hardened surface sediments there are few, if any, dolomite blocks. We presume that they were removed along with the upper part of the infilled, cavern along with the country rock as the surface continued to erode. Although it may not be possible to reconstruct a complete strati-graphic sequence involving the whole of the site, we have presented summary of evidence here that shows it is possible to trace stratigraphic relationships from the base of the Ancient Entrance to the top of the Cone and which includes the Grey Breccia and the two other bone breccias. Present work is focussed on completion of a new chronology by comparing our site magnetostratigraphy with the global polarity timescale. The speleothem deposition today in any of the caves of the Makapansgat area, with its annual rainfall of about 700mm, is extremely low. It is commonly recognised by karst geomorphologists (eg, Ford and Williams, 1989) that massive deposits of speleothem are characteristic of humid tropical to semi-tropical karsts. Hence, at the time of their formation some 3 Ma or more ago, the massive speleothem deposits are evidence for a warmer, and certainly a much wetter, climate than that of today.     

Sexual dimorphism in the face of Australopithecus africanus

Recently discovered crania of Australopithecus africanus from Sterkfontein Member 4 and Makapansgat enlarge the size range of the species and encourage a reappraisal of both the degree and pattern of sexual dimorphism. Resampling methodology (bootstrapping) is used here to establish that A. africanus has a greater craniofacial size range than chimpanzees or modern humans, a range which is best attributed to a moderately high degree of sexual dimorphism. Compared to other fossil hominins, this variation is similar to that of Homo habilis (sensu lato) but less than that of A. boisei. The finding of moderately high dimorphism is corroborated by a CV-based estimate and ratios between those specimens considered to be male and those considered to be female. Inferences about the pattern of craniofacial dimorphism in the A. africanus face currently rely on the relationship of morphology and size. Larger specimens, particularly Stw 505, show prominent superciliary eminences and glabellar regions, but in features related in part to canine size, such as the curvature of the infraorbital surface, large and small specimens of A. africanus are similar. In this respect, the pattern resembles that of modern humans more so than chimpanzees or lowland gorillas. A. africanus may also show novel patterns of sexual dimorphism when compared to extant hominines, such as in the form of the anterior pillar. However, males of the species do not exhibit characteristics of more derived hominins, such as A. robustus. Am J Phys Anthropol 108:97–127, 1999. © 1999 Wiley-Liss, Inc.     

Landscapes and their relation to hominin habitats: case studies from Australopithecus sites in eastern and southern Africa

We examine the links between geomorphological processes, specific landscape features, surface water drainage, and the creation of suitable habitats for hominins. The existence of mosaic (i.e., heterogeneous) habitats within hominin site landscape reconstructions is typically explained using models of the riverine and gallery forest settings, or the pan or lake setting. We propose a different model: the Tectonic Landscape Model (TLM), where tectonic faulting and volcanism disrupts existing pan or river settings at small-scales (∼10-25 km). Our model encompasses the interpretation of the landscape features, the role of tectonics in creating these landscapes, and the implications for hominins. In particular, the model explains the underlying mechanism for the creation and maintenance of heterogeneous habitats in regions of active tectonics. We illustrate how areas with faulting and disturbed drainage patterns would have been attractive habitats for hominins, such as Australopithecus, and other fauna. Wetland areas are an important characteristic of surface water disturbance by fault activity; therefore we examine the tectonically-controlled Okavango Delta (Botswana) and the Nylsvley wetland (South Africa) as modern examples of how tectonics in a riverine setting significantly enhance the faunal and floral biodiversity. While tectonic landscapes may not have been the only type of attractive habitats to hominins, we propose a suite of landscape, faunal, and floral indicators, which when recovered together suggest that site environments may have been influenced by tectonic and/or volcanic activity while hominins were present. For the fossil sites, we interpret the faulting and landscapes around australopithecine-bearing sites of the Middle Awash (Ethiopia) and Makapansgat, Taung, and Sterkfontein (South Africa) to illustrate these relationships between landscape features and surface water bodies. Exploitation of tectonically active landscapes may explain why the paleoenvironmental signals, anatomy, diets, as well as the fauna associated with Australopithecus appear largely heterogeneous through time and space. This hypothesis is discussed in light of potential preservation and time-averaging effects which may affect patterns visible in the fossil record. The model, however, offers insight into the landscape processes of how such habitats are formed. The landscape features and range of habitat conditions, specifically the wetter, down-dropped plains and drier, uplifted flanks persist in close proximity for as long as the fault motion continues. The Tectonic Landscape Model provides an alternative explanation of why mixed habitats may be represented at certain sites over longer timescales.     

Biogeochemical and craniometric investigation of dietary ecology, niche separation, and taxonomy of Plio-Pleistocene cercopithecoids from the Makapansgat Limeworks

Three sympatric fossil cercopithecoid genera (Cercopithecoides, Parapapio, and Theropithecus) occur in Members 3 and 4 at the Makapansgat Limeworks hominin locality, South Africa, and their presence in a single ecosystem suggest a certain degree of ecological and/or dietary differentiation between taxa. Here, we explore the extent of dietary niche separation amongst these taxa using stable isotope (13C/12C, 18O/16O) and trace-element (Sr, Ba, Ca) analyses of fossil tooth enamel. In particular we searched for evidence of subtle niche separation between the more closely related, morphologically similar taxa of the genus Parapapio, as uncertainties exist around their taxonomy and taxonomic identification. Given these uncertainties, craniometric analyses were also performed to ground the dietary interpretations in a morphological context. The results found no clear taxonomic signal in the craniometric data for the Parapapio sample, and further indicate that this sample was no more variable morphologically than a single, geographically circumscribed, extant chacma baboon sample. In contrast, two overlapping dietary ecologies were found within this same Makapansgat Parapapio sample. Additionally, two widely differing dietary ecologies were found within the Cercopithecoides williamsi sample, while results for Theropithecus darti indicate a predominantly C4 diet. Hence, although biogeochemical dietary indicators point towards distinct dietary ecologies within and between fossil genera at Makapansgat, within the genus Parapapio disjunctions exist between the dietary categories and the taxonomic assignment of specimens.     

Bone breccias, bone dumps, and sedimentary sequences of the western Limeworks, Makapansgat, South Africa

The sediments in the western side of the Makapansgat Limeworks were either precipitated as speleothems, represented in the earlier massive deposits, or were deposited as coarsening clastic sediments, mainly representing later deposits. Between the earlier deposits and the main sedimentary phase, the stratigraphic sequence was inverted twice to a considerable height by the unusual deposition of subaqueous speleothem. Bone-bearing deposits, including the Main Quarry Bone Breccia and the well-known Grey Breccia belong, in time, to the lower part of the clastic deposits called the Red Silts. Australopith fossils have been found in the Grey Breccia dumped material and, in situ, from the dolomite clast breccia on the Main Quarry entrance buttress. Whatever the problems may be in provenancing some of the material from the Limeworks dumps, there is no doubt that the three rows of blocks on the southern side of the dumps belong to the Grey Breccia, and other rows contain red sediment sufficient for them to be safely associated with the Red Silts. There is no reason why this material should not be prepared with confidence as to its stratigraphic provenance. In any case, stratigraphic evidence, presented here, shows that there is little difference in time between the deposition of the bone-bearing breccias.     

Size and shape variation in the proximal femur of Australopithecus africanus

Aside from use as estimates of body mass dimorphism and fore to hind limb joint size comparisons, postcranial elements have not often contributed to assessments of variation in Australopithecus africanus. Meanwhile, cranial, facial, and dental size variation is interpreted to be high or moderately high. Further, the cranial base and face express patterns of structural (shape) variation, which are interpreted by some as evidence for the presence of multiple species. Here, the proximal femur is used to consider postcranial size and shape variation in A. africanus. Original fossils from Makapansgat and Sterkfontein, and samples from Homo, Pan, Gorilla, and Pongo were measured. Size variation was assessed by comparing the A. africanus coefficient of variation to bootstrapped distributions of coefficient of variation samples for each taxon. Shape variation was assessed from isometrically adjusted shape variables. First, the A. africanus standard deviation of log transformed shape variables was compared to bootstrapped distributions of logged standard deviations in each taxon. Second, shape variable based Euclidean distances between fossil pairs were compared to pairwise Euclidean distance distributions in each reference taxon. The degree of size variation in the A. africanus proximal femur is consistent with that of a single species, and is most comparable to Homo and Pan, lower than A. afarensis, and lower than some estimates of cranial and dental variation. Some, but not all, shape variables show more variation in A. africanus than in extant taxa. The degree of shape difference between some fossils exceeds the majority of pairwise differences in the reference taxa. Proximal femoral shape, but not size, variation is consistent with high estimates of A. africanus cranial variation.     

Microwear evidence for Plio-Pleistocene bovid diets from Makapansgat Limeworks Cave, South Africa

Makapansgat Limeworks Cave is a well-known Australopithecus africanus bearing locality that has spawned a considerable amount of paleoecological research because of its hominin component. Most recently, the paleoecology of this Plio-Pleistocene site has been studied by determining the diet and habitat of other extinct taxa, particularly the bovids. The diets of seven bovids (Aepyceros sp., Gazella vanhoepeni, Makapania broomi, Parmularius braini, Redunca darti, Tragelaphus sp. aff. T. angasii, and Tragelaphus pricei) have now been classified using taxonomic uniformitarianism, ecomorphology, stable carbon isotopes, and mesowear analysis. Here, dental microwear is applied to the same bovids for additional comparison and to further elucidate the strengths and weaknesses of each method. The different dietary proxy methods noted provide a temporal continuum, with genetic signals such as ecomorphology and taxonomic uniformitarianism indicating behavioral adaptations over geologic time, while nongenetic data such as stable carbon isotopes and mesowear reflect different aspects of average diet over extended portions of an animal's life, and dental microwear provides dietary snapshots.Microwear separated an extant baseline of ten bovid species into expected dietary categories and the Makapansgat bovids clearly fell into two groups with the same degree of separation as between extant grazers and browsers. The results indicate that a multidisciplinary approach produces a more accurate and robust reconstruction of past diets. In sum, the microwear analysis is in-line with the isotope and mesowear results, which suggest a stronger browsing component than either taxonomic uniformitarianism or ecomorphology imply.     

Combining isotopic and ecomorphological data to refine bovid paleodietary reconstruction: a case study from the Makapansgat Limeworks hominin locality

The relationship between environmental change and hominin evolution remains obscure. For the most part, this stems from the difficulty of reconstructing ancient hominin habitats. Bovids are among the most frequently utilized paleoenvironmental indicators, but little is known about the habitat preferences of extinct taxa. It is generally assumed that fossil bovids both ate the same things and occupied the same habitats as their closest extant relatives. We test the first part of this assumption by reconstructing the diets of seven bovids from Makapansgat Limeworks, South Africa. Since diet and habitat are linked, these reconstructions have implications for our understanding of fossil bovid habitat tolerances. Ecomorphological and stable carbon isotope analyses are employed, allowing us to take advantage of the strengths and overcome the weaknesses of both. In most cases, fossil bovids did have similar diets to their extant relatives, and probably occupied similar habitats. Gazella vanhoepeni and Aepyceros sp., however, were almost exclusive browsers, and not mixed feeders like their living counterparts.     

Morphometric variation in the papionin muzzle and the biochronology of the South African Plio‐Pleistocene karst cave deposits

Papionin monkeys are widespread, relatively common members of Plio‐Pleistocene faunal assemblages across Africa. For these reasons, papionin taxa have been used as biochronological indicators by which to infer the ages of the South African karst cave deposits. A recent morphometric study of South African fossil papionin muzzle shape concluded that its variation attests to a substantial and greater time depth for these sites than is generally estimated. This inference is significant, because accurate dating of the South African cave sites is critical to our knowledge of hominin evolution and mammalian biogeographic history. We here report the results of a comparative analysis of extant papionin monkeys by which variability of the South African fossil papionins may be assessed. The muzzles of 106 specimens representing six extant papionin genera were digitized and interlandmark distances were calculated. Results demonstrate that the overall amount of morphological variation present within the fossil assemblage fits comfortably within the range exhibited by the extant sample. We also performed a statistical experiment to assess the limitations imposed by small sample sizes, such as typically encountered in the fossil record. Results suggest that 15 specimens are sufficient to accurately represent the population mean for a given phenotype, but small sample sizes are insufficient to permit the accurate estimation of the population standard deviation, variance, and range. The suggestion that the muzzle morphology of fossil papionins attests to a considerable and previously unrecognized temporal depth of the South African karst cave sites is unwarranted. Am J Phys Anthropol 2010. © 2009 Wiley‐Liss, Inc.