ArticlePDF Available

The bovids from Elandsfontein, South Africa, and their implications for the age, palaeoenvironment, and origins of the site


Abstract and Figures

The bovid fossils from Elandsfontein, south-western Cape Province, South Africa, comprise 7257 individually numbered specimens from 18 species. Taxonomic comparisons with Olduvai Gorge and other African sites and the high percentage of extinct forms imply that the bones accumulated in the earlier part of the Middle Quaternary, probably sometime between 700,000 and 400,000 years ago. By extension, this is also the most likely age for the skull cap of archaicHomo sapiens (‘Saldanha Man’) and for the occasional ‘late’ Acheulean stone artifacts that accompany the animal bones. In keeping with geomorphological observations and other aspects of the fauna, the bovids indicate a relatively grassy and moist environment, apparently during an interglaciation that differed significantly from the Holocene. Geomorphological context, the frequent occurrence of partial skeletons, bone damage, and skeletal part representation suggest that carnivore feeding on carcasses scattered across a Mid-Quaternary land surface was probably the main factor shaping the Elandsfontein bone assemblage. Porcupines may also have played a role, but there is little evidence for human activity. The Elandsfontein assemblage thus provides a useful ‘control’ for comparison with bone accumulations where context, associations, and bone damage indicate that people were heavily involved. For example, there are very few young animals in the otherwise attritional profile of ‘giant’ buffalo from Elandsfontein, probably because carnivores often rapidly and completely consumed young carcasses. This suggests that few young carcasses would be available for human scavenging and thus that archaeological attritional profiles in which young individuals are common probably reflect active human hunting, at least of young animals.
Content may be subject to copyright.
A preview of the PDF is not available
... This seasonal lack of forage renders the western coastal plains inhospitable to most ruminant grazers, except for the hartebeest (Radloff, 2008), which is remarkable in its ability to extract relatively high-quality grass leaf from senescent swards and persist through dry seasons and droughts (Kingdon and Hoffman, 2013). Clearly, the Pleistocene faunas are signaling other important environmental and ecological changes, perhaps including more humid environments and altered rainfall regimes (Klein and Cruz-Uribe, 1991;Cruz-Uribe et al., 2003;Klein et al., 2007), increased availability of spring-fed wetlands (Braun et al., 2013;Lehmann et al., 2016Lehmann et al., , 2018, or ungulate migrations without parallel in the CFR today (Marean, 2010;Faith and Thompson, 2013; but see Lehmann et al., 2018). ...
... Specifically, given previous suggestions that the Pleistocene faunas are signaling important changes in moisture availability (e.g. Klein and Cruz-Uribe, 1991;Grine and Klein, 1993;Cruz-Uribe et al., 2003;Klein et al., 2007;Stynder, 2009), we develop ecometric models for predicting aridity based on the distribution of ungulate dental traits across the present-day CFR. We show that these models are applicable across a broad range of southern African environments, and use them to reconstruct past aridity along the western coastal plains. ...
... The mid-Pleistocene fauna from the dunefield at Elandsfontein farm (~100 km NNW of Cape Town), referred to as Elandsfontein Main, consists primarily of specimens that were surface-collected in the 1950s and early 1960s (Klein and Cruz-Uribe, 1991;Klein et al., 2007). Comparison with welldated eastern African faunas suggests a likely age of between 1.0 and 0.6 Ma (Klein et al., 2007), although recent paleomagnetic evidence suggests a probable minimum age of 0.78 Ma (Braun et al., 2013). ...
The Pleistocene ungulate communities from the western coastal plains of South Africa's Cape Floristic Region (CFR) are diverse and dominated by grazers, in contrast to the region's Holocene and historical faunas, which are relatively species-poor and dominated by small-bodied browsers and mixed feeders. An expansion of grassy habitats is clearly implied by the Pleistocene faunas, but the presence of ruminant grazers that cannot survive the summer dry season typical of the region today suggests other important paleoecological changes. Here we use dental ecometrics to explore the paleoecological implications of the region's Pleistocene faunas. We show that the dental traits (hypsodonty and occlusal topography) of the ungulates that occurred historically in the CFR track annual and summer aridity, and we use these relationships to reconstruct past aridity. Our results indicate that the Pleistocene faunas signal paleoenvironments that were on average less arid than today, including during the summer, consistent with other lines of evidence that suggest a higher water table and expansion of well-watered habitats. Greater water availability can be explained by lower temperature and reduced evapotranspiration during cooler phases of the Pleistocene, probably coupled with enhanced groundwater recharge due to increased winter precipitation.
... The ecosystems of the mid-Pleistocene of the Cape Floristic Region are recorded in another nearby dunefield locality known as Elandsfontein. Over 100 years of investigation into the prehistory of Elandsfontein have produced one of the largest faunal assemblages available from the African mid-Pleistocene.39,79,80 Excavations at Elandsfontein have recorded the presence of hominins that produced Acheulean tools32,79,81 and indicate that hominins had some behavioral association with the fauna found at the locality.82 ...
... Over 100 years of investigation into the prehistory of Elandsfontein have produced one of the largest faunal assemblages available from the African mid-Pleistocene.39,79,80 Excavations at Elandsfontein have recorded the presence of hominins that produced Acheulean tools32,79,81 and indicate that hominins had some behavioral association with the fauna found at the locality.82 Explorations of the isotopic ecology of the region suggest that hominins occupied this environment at a time when the region hosted an abundant large mammalian fauna in association with an ecosystem that differs dramatically from the contemporary low-nutrient strandveld that exists in the region today.32,[83][84][85] ...
Despite advances in our understanding of the geographic and temporal scope of the Paleolithic record, we know remarkably little about the evolutionary and ecological consequences of changes in human behavior. Recent inquiries suggest that human evolution reflects a long history of interconnections between the behavior of humans and their surrounding ecosystems (e.g., niche construction). Developing expectations to identify such phenomena is remarkably difficult because it requires understanding the multi-generational impacts of changes in behavior. These long-term dynamics require insights into the emergent phenomena that alter selective pressures over longer time periods which are not possible to observe, and are also not intuitive based on observations derived from ethnographic time scales. Generative models show promise for probing these potentially unexpected consequences of human-environment interaction. Changes in the uses of landscapes may have long term implications for the environments that hominins occupied. We explore other potential proxies of behavior and examine how modeling may provide expectations for a variety of phenomena.
... Archaeological records point toward a dominance of large bodied ruminant and non-ruminant grazing species during the LGM on the PAP(Copeland et al., 2016;Hodgkins et al., 2020;Klein, 1972bKlein, , 1983Klein and Cruz-Uribe, 1991;, providing a stark contrast to what we see in the low nutrient fynbos ecosystem found along the coastline today.Although it is possible to identify the areas where the highest number of large herbivores were likely to occur, it remains difficult to know their exact densities. The size and management of protected areas in current landscapes strongly constrains the distribution, populations and diversity of large herbivores. ...
... 42.8 mm) fromNelson Bay Cave(Faith Personal communication)) bovid. Body mass predictions from Nelson Bay Cave represent an intermediary between extremely high predictions (841 kg calculated from the length of the third molar from Elandsfontein, South Africa(Klein and Cruz-Uribe, 1991)) and extremely low predictions (200 kg,Smith et al., 2003). M. priscus was a bovid that became extinct just prior to the Pleistocene-Holocene transition(Table 4.1). ...
Understanding the variations in structure and abundance of animals and what leads to their distribution within the landscape has captured the attention of ecologists for centuries. Importantly, knowledge of current behaviour of large mammals can be used to inform historic population dynamics and is essential to understanding how early humans used large mammals as a foraging resource. Central to this thesis and improving our understanding of large herbivores is the Palaeo-Agulhas Plain (PAP) where large mammalian herbivores formed a key food resource for early humans. The PAP, now submerged off the southern Cape of South Africa, formed a novel ecosystem during lower sea levels. Characterised by large expanses of nutrient rich grasslands and large grazing herbivores, the PAP stands in stark contrast to the nutrient poor fynbos ecosystems that is in the southern Cape today. In this thesis I focus on the Last Glacial Maximum (LGM; ~20 ka) when the PAP was last fully exposed to answer questions relating to the habitat use and range distribution of large herbivores. Importantly, through the Paleoscape Project, modelled climate, soil and vegetation have made these recreations of large mammals possible. Using modelled climate and vegetation this thesis aims to model the large herbivore communities and understand the influence of early humans on the PAP during the LGM for successful integration into the PaleoscapeABM (the PAP agent-based model). To improve our understanding of large mammals on the PAP I identified five large herbivores that became extinct on the PAP since the LGM and modelled their behavioural and physical traits using k-Nearest Neighbour imputation. I predicted the biomass of large herbivores across the PAP using actual biomass of large herbivores from 39 protected areas across South Africa (spanning five functional groups to include the extinct species) across a rainfall gradient and different biomes. The distribution of large herbivores would likely have been driven by similar top-down and bottom-up drivers we see in large herbivore ecology today. Knowing this, I created a predictive model for large mammals by predicting the probability of occurrence of functional groups of large herbivores in relation to environmental drivers and humans. Results showed that all species (except Antidorcas australis) were adapted to the grassy environment of the PAP and these specialisations likely contributed to their extinction along with changing climates and intensified hunting from humans. When predicting herbivore viii biomass, biome was the most important factor influencing the relationship between herbivores and rainfall. In general, large herbivore biomass increased with rainfall across biomes, except for grassland. Finally, I showed the probability of occurrence of large herbivores was influenced by early humans, water availability and a landscape of fear on the PAP. Through this thesis I have successfully provided detailed accounts of the biomass and probability of occurrence of large herbivores on the PAP. Importantly, this information can be seamlessly integrated into the PaleoscapeABM. Finally, I highlight the importance of this knowledge in understanding early humans, the potential shortcomings of this study and resulting areas where research needs to be focused.
... Ultimately, this revealed that all faunal specimens were, at best, only minimally identifiable to taxonomic size class due to cultural fragmentation and/or diagenesis. Thus, minimally identifiable specimens were compared to general taxonomic size to assess the approximate size of fauna processed at the site (Klein and Cruz-Uribe 1991). ...
The middle Tanana Valley near Fairbanks, Alaska has been the subject of nearly a century of archaeological research focused on the earliest inhabitants of the region. Recent research at Niidhaayh Na’ (XBD-110) provides new information about human behavior and technological organization at the Pleistocene-Holocene transition. This multicomponent site is located on the Delta moraine and overlooks Delta Creek (Niidhaayh Na’). The results of the first seasons of full-scale excavation research at the site, begun in 2017, reveal two lithic workshops dating to ca. 11,800 and 9500 calendar years ago, associated with core fragments, tools, debitage, and intact faunal remains. Future research at the site will advance archaeological understandings of human adaptive decision-making during the late glacial period in central Alaska, with implications for our understanding of the first Americans and human behavior more generally.
... The unselective nature of non-ruminants (most of which consume varying proportions of browse and grass; Hempson et al., 2015b) enable them to consume larger quantities of lower quality vegetation than ruminant herbivores (Duncan et al., 1990;Cromsigt et al., 2009), whereas ruminant herbivores can digest high quality forage more effectively (Duncan et al., 1990;Illius et al., 2002;Cromsigt et al., 2009). Archaeological records point toward a dominance of large-bodied ruminant and non-ruminant grazing species during the LGM on the PAP (Klein, 1972(Klein, , 1983Klein and Cruz-Uribe, 1991;Copeland et al., 2016;Hodgkins et al., 2020;Venter et al., 2020), providing a stark contrast to what we see in the low nutrient fynbos ecosystem found along the coastline today . ...
Herbivore distribution throughout Africa is strongly linked to mean annual precipitation. We use that relationship to predict functional group composition of herbivore communities during the last glacial maximum (ca. 21 ka) on the now submerged Palaeo-Agulhas Plain (PAP), South Africa. We used metabolic large herbivore biomass (MLHB) from 39 South African protected areas, in five functional groups (characterized by behavior and physiology). We examined how modern factors influenced MLHB and considered the effects of biome, annual rainfall, percentage winter rainfall, and protected area size. Overall, biome was the most important factor influencing the relationship between MLHB and rainfall. In general, MLHB increased with rainfall, but not for the grassland biome. Outside grasslands, most functional groups' metabolic biomass increased with increasing rainfall, irrespective of biome, except for medium-sized social mixed feeder species in savanna and thicket. Protected area size was influential for medium-sized social mixed feeders and large browsers and rainfall influenced medium-sized social mixed feeders, offering some perspectives on spatial constraints on past large herbivore biomass densities. These results improve our understanding of the likely herbivore community composition and relative biomass structure on the PAP, an essential driver of how early humans utilized large mammals as a food resource.
... The infundibula of moderately worn maxillary molars are often complex, with pronounced mesiodistal constriction. This complexity differs from more primitive specimens of Connochaetes (Klein and Cruz-Uribe, 1991;Bibi et al., 2017), as well as Rusingoryx , which tend to have simpler infundibula. The lingual lobes of the maxillary molars and the buccal lobes of the mandibular molars are fairly evenly rounded, with the mandibular specimens tending to have a distinct circular outline. ...
We report on the Late Pleistocene (36-12 ka) mammals from Kibogo in the Nyanza Rift of western Kenya, providing (1) a systematic description of the mammal remains, (2) an assessment of their paleoenvironmental implications, and (3) an analysis of the biogeographic implications of non-analog species associations. Kibogo has yielded one of the largest paleontological assemblages from the Late Pleistocene of eastern Africa, and it is dominated by grassland ungulates (e.g., equids and alcelaphin antelopes), including an assortment of extralimital (e.g., Equus grevyi, Ceratotherium simum, Redunca arundinum) and extinct species (Syncerus antiquus, Damaliscus hypsodon, Megalotragus sp.). The composition of the fauna, in conjunction with the soils and topography of the region, indicate the local presence of edaphic grassland situated within a broader environment that was substantially grassier and likely drier than at present. In contrast to non-analog faunas from higher latitudes (e.g., North America and western Eurasia), the climatic niches of non-analog species associations strongly overlap, indicating that non-analog climate regimes during the Late Pleistocene of eastern Africa are not necessary to account for the former association of presently allopatric species. The Kibogo faunas add to a growing body of evidence implying that the composition of present-day African herbivore communities is distinct from those of the geologically recent past.
... The Bovidae have been used in many studies to indicate habitat type (Gentry 1970;Scott 1979Scott , 1985Vrba 1980b;Greenacre & Vrba 1984;Soulounias & Dawson-Saunders 1988;Klein & Cruz-Uribe 1991;Plummer & Bishop 1994;Spencer 1995Spencer , 1997Kappelman et al. 1997). These studies include the fimctional morphology of cranial and postcranial elements, the extension of modem habitat associations to infer those of fossil taxa. ...
Plio-Pleistocene mammalian faunal evolution has been causally linked to global climate change. This study investigates the extent to which climate affected the faunal representation of large mammals in the fossil assemblages from the Turkana Basin, in East Africa during the Plio-Pleistocene. The research project involved the collection and study of both modem and fossil specimens Relative abundances of fossil mammalian families and the Bovidae are explored. To interpret climatically driven faunal change, comparisons are made with fossil faunas from the Nachukui, Shungura and Koobi Fora Formations. The combined relative abundances of bovid tribes through time are used to infer changes in habitat representation. The reduncines, wet habitats indicators, are abundant around 2.5 ma in contrast to the Shungura Formation represented by closed-dry habitats. This suggests that local rather than global scale climatic influences are affecting habitat representation. At 1.7 ma the inferred habitat in the Nachukui Formation shows similarity with more distant localities, namely the Olduvai Basin in Tanzania, where more arid and open environments become increasingly dominant. In a second part of this study, data were collected from modem antelope remains to develop new body weight estimation equations, essential for the estimation of body weights of fossil antelopes from the Nachukui Formation assemblage. Dental remains are less accurate estimators of body weight than are postcranial remains. The body weight estimation equations developed in this study are more accurate than published estimation equations when compared. The body weights of Nachukui Formation fossil bovids were estimated using the equations developed in this study. Variation in body weight representation of bovid tribes through time, in the context of changing habitats, and the use of bovid body weights in taxonomic identification of isolated teeth are explored for the Nachukui Formation.
p>The core argument demonstrates that the archaeological notion of transitions is untenable. They structure the past into blocks of time, thereby amalgamating behaviour patterns and establishing universal interpretations that are situated outside of hominid action. Within the current framework a transition is a historical junction point in chronological time, organised according to change and variation in archaeological assemblages. Several models have been proffered to explain change, but the underlying framework through which transitions are established has rarely been questioned, because of their key role in the interpretation of hominid evolution. This traditional framework is critiques and two themes are addressed to re-contextualise Middle Pleistocene archaeological interpretation. Firstly, in an exploration of the concepts of temporality and the taskscape, it is argued that time and space are mutually produced through hominid action. This alters the interpretation of change and variation, which is my second theme. I conclude that they exist in unison, as change is a constant although inconsistent process of transformation. Undermining the notion of fixed points of transition renders research focusing on origin points, and therefore modern humans origins, implausible. Current discourse on hominid identity draws on the structural opposition of 'modern' versus 'archaic' humans for interpretation. In contrast, I locate hominid identities through the exploration of social praxis , offering a way of linking recent social theory with the practice of lithic analysis to interpret changing hominid identities. The transformation from the Acheulean to the Middle Stone Age and Middle Palaeolithic is characterised in five case studies that analyse Middle Pleistocene lithic assemblages from the UK, France and South Africa. I demonstrate that there is no single identity for Acheulean, Middle Stone Age or Middle Palaeolithic hominids, and show how non-linear transformations in the detailed analysis of lithic artefacts and the surrounding taskscape can portray changing relations in hominid social life.</p
Full-text available
The Cooper’s D deposit has been dated by U-Pb at a maximum age of 1.37 Ma ± 0.113 and has yielded seven Hominini remains, six of them are attributed to the Southern African endemic species Paranthropus robustus Broom, 1938. However, the taxonomic composition of the faunal assemblage recovered at this site, especially ruminants, remains poorly understood. This paper updates the previous palaeontological and biochronological works and provides the first taxonomic description and paleoecological analysis of the whole bovid material from Cooper’s D. We propose a minimum age of 1.0 Ma for the Cooper’s D bone accumulation. Thus, we conclude that the assemblage accumulated between 1.375 ± 0.113 and 1.0 Ma based on the complementary results obtained from radiometric dates and biochronology. The paleoecological analysis suggests that Cooper’s D is likewise dominated by grassland with sparse covered habitat and probable water sources in the vicinity. Finally, we show a strong statistically significant negative correlation between the presence of grassland-adapted bovid species and P. robustus. This result indicates that P. robustus was more likely to be an eurytopic species and dietary generalist than an exclusive grazer.
Analyses of faunal remains are a key means of inferring palaeoenvironmental change. In this paper, the use of faunal remains as a proxy for environmental conditions from Marine Isotope Stage 6 to the Holocene in southern Africa is reviewed. The focus of this review is on large herbivore abundance and how these fluctuate temporally and regionally in accordance with palaeo-climatic shifts. Here, southern Africa is divided into four eco-regions loosely based on climatic, biotic and zoogeographic traits: the Cape Floristic Region, the arid and semi-arid region, the savanna and grassland region, and the wetter eastern region. The relative abundance of large herbivores within these regions are noted, and temporal trends are inferred. On the whole, most eco-regions maintain similar herbivore compositions over time showing the regional ecological resilience of these taxa to local-scale environmental change. Yet some changes in faunal frequencies are apparent. The Cape Floristic Region shows evidence of significant faunal turnover from the Late Pleistocene to the Holocene. Here, grazers are significantly more abundant during glacial periods, probably linked to the terrestrial expansion of the palaeo-Agulhas coastal plain. Shifts in ungulate abundance in the currently xeric central interior, also indicate wetter periods in the Pleistocene. Holocene faunas are generally similar to historic distributions but shifts between xeric and mesic periods are also evident.
Artifact occurrences that can be integrated within the Acheulian Industrial Complex were recognized and described from Southern Africa towards the end of the last century. It was appreciated on geomorphological grounds even at that time that these Acheulian occurrences are of high antiquity despite the fact that they are frequently exposed at or near the present day surface. The earliest relatively dated artifact occurrences are known from the Australopithecine breccia sites of Sterkfontein and Swartkrans although there is the strong possibility that artifact occurrences of like or greater age exist elsewhere, as for example in the Vaal River basin. The Sterkfontein and Swartkrans occurrences have been variously described as Developed Oldowan and as Acheulian, but indicate an age in excess of 1 million years for the appearance of a stone tool manufacturing tradition in Southern Africa. At the other end of the time scale, the Acheulian is defined in terms of the consistent manufacture of bifaces. It persisted until some 100,000 years ago when replaced by a complex of the flake industries, some occurrences of which can be related to marine deposits of last interglacial age. The primary focus of initial Acheulian studies was on discerning evolutionary time trends in biface manufacture. Many of the sites studied were occurrences in geological context which perforce have proved limited in allowing discussion of content and inter-assemblage variability. Recently, more occurrences have been excavated in a primary or semiprimary context and make it possible to begin to discuss some aspects of the demography, subsistence, and behavior of the Acheulian populations through the Middle Pleistocene in Southern Africa. A limiting factor, however, is still the paucity of occurrences with preserved faunas or floras in association with artifactual materials. The Pleistocene faunas of southern Africa are known in general terms only and primarily indicate long-term environmental stability. Indeed as a stable, large geographical area presenting a mosaic of suitable habitats for Acheulian populations, southern Africa may have been an important population center in the Middle Pleistocene.
A revision of the Hippotragini from the Makapansgat Limeworks is proposed: a new species of Hippotragus is described, Hippotragus cookei, and fossils previously referred to cf. Oryx gazella and Hippotragus gigas are assigned to this species. H. gigas is present in the Pleistocene Member 5, but not in the Pliocene Grey Breccia, Member 3, as had been formerly supposed. A new genus and species from Member 3, Wellsiana torticornuta, tentatively referred to ?Hippotragini, is described based on a frontlet that had been assigned to Damaliscus sp. (aff. albifrons) by Wells and Cooke. The horncore piece previously referred to Aepyceros cf. melampus may belong to the same species as ?Hippotragini sp. nov. (Gentry, 1986) from the Laetolil Beds, Tanzania. The hippotragine fossils here discussed again emphasize that Makapansgat Member 3 contains a Pliocene assemblage that is more ancient than was originally thought, with particular affinities with the Laetolil Beds and also with the Pinjor Formation of the Siwaliks in India and Pakistan. - Author
Bones of recent mammals in the Amboseli Basin, southern Kenya, exhibit distinctive weathering characteristics that can be related to the time since death and to the local conditions of temperature, humidity and soil chemistry. A categorization of weathering characteristics into six stages, recognizable on descriptive criteria, provides a basis for investigation of weathering rates and processes. The time necessary to achieve each successive weathering stage has been calibrated using known-age carcasses. Most bones decompose beyond recognition in 10 to 15 yr. Bones of animals under 100 kg and juveniles appear to weather more rapidly than bones of large animals or adults. Small-scale rather than widespread environmental factors seem to have greatest influence on weathering characteristics and rates. Bone weathering is potentially valuable as evidence for the period of time represented in recent or fossil bone assemblages, including those on archeological sites, and may also be an important tool in censusing populations of animals in modern ecosystems.
Large cats, canids, bears, and hyenas create distinctive types of damage when they gnaw bones. This paper describes the diagnostic characteristics of damage done by each taxon to femora and tibiae of herbivores with body weights = or >300kg. Pleistocene and Recent fossil collections that include gnawed bones might provide data on the presence of carnivores whose own remains are not found in the collections. Information might also be gained about predator and scavenger utilization of prey carcasses, often a reflection of prey vulnerability or availability in past communities.-Author