A quantitative assessment of variation in Holocene Khoesan crania from South Africa's western, south-western, southern and south-eastern coasts and coastal forelands

Includes bibliographical references (leaves 274-325)

Stable isotopic composition of fossil mammal teeth and environmental change in southwestern South Africa during the Pliocene and Pleistocene

The past 5 million years mark a global change from the warmer, more stable climate of the Pliocene to the initiation of glacial-interglacial cycles during the Pleistocene. Marine core sediment records located off the coast of southwestern Africa indicate aridification and intensified upwelling in the Benguela Current over the Pliocene and Pleistocene. However, few terrestrial records document environmental change in southwestern Africa over this time interval. Here we synthesize new and published carbon and oxygen isotope data of the teeth from large mammals (>6 kg) at Langebaanweg (~5 million years ago, Ma), Elandsfontein (1.0 – 0.6 Ma), and Hoedjiespunt (0.35 – 0.20 Ma), to evaluate environmental change in southwestern Africa between the Pliocene and Pleistocene. The majority of browsing and grazing herbivores from these sites yield enamel 13 C values within the range expected for animals with a pure C3 diet, however some taxa have enamel 13C values that suggest the presence of small amounts C4 grasses at times during the Pleistocene. Considering that significant amounts of C4 grasses require a warm growing season, these results indicate that the winter rainfall zone, characteristic of the region today, could have been in place for the past 5 million years. The average 18O value of the herbivore teeth increases ~4.4‰ between Langebaanweg and Elandsfontein for all taxa except suids. This increase may solely be a function of a change in hydrology between the fluvial system at Langebaanweg and the spring-fed environments at Elandsfontein, or a combination of factors that include depositional context, regional circulation and global climate. However, an increase in regional aridity or global cooling between the early Pliocene and mid-Pleistocene cannot explain the entire increase in enamel 18O values. Spring-fed environments like those at Elandsfontein may have 75 provided critical resources for mammalian fauna in the mid-Pleistocene within an increasingly arid southwestern Africa ecosystem

Stable isotope ecology of Cape dune mole-rats (Bathyergus suillus) from Elandsfontein, South Africa: implications for C4 vegetation and hominin paleobiology in the Cape Floral Region

The archaeological and paleontological records from the west coast of South Africa have potential to provide insights into ecosystem dynamics in the region during the mid Pleistocene. Although the fossil record suggests an ecosystem quite different than that of the region today, we understand little about the ecological factors that contributed to this disparity. The site of Elandsfontein (EFT) dates to between 1.0 and 0.6 million years ago (Ma), preserves in situ lithic and faunal materials found in direct association with each other, and provides the rare opportunity to examine the relationship between hominin behavioural variability and landscape heterogeneity in a winter rainfall ecosystem. In this study, we examine the stable carbon isotopic composition of a large sample (n = 81) of Cape dune mole-rats (Bathyergus suillus) and contemporaneous large mammals (> 6 kg; n = 194) from EFT. We find that δ13C values of B. suillus are significantly different to those of contemporaneous large mammals from EFT indicating a significant presence of plants utilizing the C4 photosynthetic pathway during the mid-Pleistocene, in contrast to present C3 dominated ecosystems along the west coast of South Africa. Additionally, we find that artifact density at EFT localities is positively correlated with δ13C values in B. suillus enamel suggesting that evidence of more intense hominin occupation may be associated with the presence of more C4 vegetation. Lastly, we hypothesize that this unique distribution of vegetation 1) provided abundant resources for both hominin and non-hominin taxa and 2) may have concentrated hominin and animal behavior in certain places on the ancient landscape

The use of faunal evidence to reconstruct site history and Hoedjiespunt 1 (HDP1), Western Cape

Hoedjiespunt 1 (HDP 1 ), is one of few later Middle Pleistocene to earlier Late Pleistocene African sites to yield well provenanced MSA hominid fossils, lending special significance to this site. The vertebrate fauna from this location, which consists of a palaeontological and an archaeological site, is described and analysed using both the taphonomic and controlled comparison approaches. The information obtained via this study allows for a better understanding of the context in which and the conditions under which these two sites were formed. Stratigraphic evidence and spatial information, suggest that the bones in the palaeontological site were in all likelihood accumulated in a cavity, thus postdating the sediments in which they occur. Circumstantial evidence, in addition to Klein and Cruz-Uribe's (1984) criteria for distinguishing assemblages accumulated by hyaenas from those accumulated by people, points towards the brown hyaena as the most likely accumulator of this assemblage. It is also suspected that the bone in the archaeological assemblage, may postdate the sediments in which they occur. This is suggested by the presence at the site, of tools manufactured out of calcrete, similar to the calcrete carapace which caps the stratigraphic sequence. Although density mediated destruction seems to have been the major cause of discrepancies in skeletal part abundance in the palaeontological site, it was found not to have been severe. The composition of species represented in the two assemblages differ. It was found that, apart from containing a small percentage of marine animals, the palaeontological site is dominated by grazing ungulates and carnivores. This assemblage was accumulated during a period of lowered sea level, or "glacial". On the other hand, the sample from the archaeological site contains proportionally fewer ungulates and carnivores, more small animals and more marine animals, reflecting a period of marine transgression, or "interglacial"

Hypercarnivory, durophagy or generalised carnivory in the Mio-Pliocene hyaenids of South Africa?

Carnivorans, the members of the order Carnivora, exhibit wide dietary diversity – from overwhelmingly herbivorous species (like the giant and red pandas) to species that specialise in the consumption of flesh (like the hypercarnivorous felids). Throughout the evolution of this order, many craniodental forms have emerged and gone extinct – notably the sabretooth felids that existed until the late Pleistocene. However, one carnivoran lineage, remarkable for its extreme masticatory adaptations, persists – the bone-cracking hyaenids. Three of the four extant members of this family (Crocuta crocuta, Hyaena hyaena and Parahyaena brunnea) are among the most durophagous mammals to have ever lived. The fourth extant hyaenid – the aardwolf (Proteles cristatus) – also exhibits impressive, although wholly different, masticatory adaptations as one of the most derived mammalian insectivores. How and when did the level of durophagy evident in extant bone-cracking hyenas evolve, and how do Mio-Pliocene hyenas compare to the extant members of the order in terms of their own dietary specialisations? An examination of the premolars of the Mio-Pliocene hyaenids from Langebaanweg, South Africa suggests that modern levels of durophagy appeared relatively recently. Results from an analysis of dental radii-of-curvature and premolar intercuspid notches suggest that these hyenas were neither bone crackers nor flesh specialists, but were dietary generalists

Early to mid-Holocene South African Later Stone Age human crania exhibit a distinctly Khoesan morphological pattern

The sample of South African early to mid-Holocene Later Stone Age (LSA) human crania is small and quite fragmentary, limiting our knowledge of human craniofacial morphology for this period. Previous limited analyses have described the morphology displayed by these early crania as a combination of Khoesan and non-Khoesan traits. Although essentially Khoesan-like in terms of facial morphology, their overall large size and robust neurocranial structure were regarded as atypical of Khoesan craniofacial morphology, leading to questions about the role of these early populations in the ancestry of recent Khoesan populations. Here we provide a quantitative analysis in which we compare five well-preserved pre-5000 BP LSA crania with (i) a large sample of post-5000 BP LSA Khoesan crania; and (ii) a sample of crania from recent South African Bantu-speakers. We show that these pre-5000 BP crania fall comfortably within the range of variation observed for the post-5000 BP Khoesan sample, in terms of both size and shape, suggesting that distinctive Khoesan craniofacial morphology was already present in South African LSA populations by the first half of the Holocene

A rare example of a human burial with a stone-packed grave shaft in the Western Cape Province, South Africa

In June 2003, rescue work near the town of Wolseley in the Tulbagh District, Western Cape Province, recovered evidence of a grave containing human skeletal remains placed in a reclined, seated position, with the knees pulled up under the head (vertically-flexed). A layer of 16 stones sealed the grave shaft just above the head of the skeleton, but well below the surface. Red ochre powder lay around the feet and the skull. Graves with stone-packed shafts, where the individual is buried in a vertically-flexed position, are extremely rare in the archaeological record of the Western Cape Province, and are more commonly associated with probable Khoekhoe burials in the Northern Cape (Schapera 1930; Dreyer & Meiring 1937; Inskeep 1986; Morris 1992). Here we provide an account of this unique occurrence and speculate on the cultural and biological identity of the individual