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.     

Improved age control on early Homo fossils from the upper Burgi Member at Koobi Fora,Kenya

To address questions regarding the evolutionary origin, radiation and dispersal of the genus Homo, it is crucial to be able to place the occurrence of hominin fossils in a high-resolution chronological framework. The period around 2 Ma (millions of years ago) in eastern Africa is of particular interest as it is at this time that a more substantial fossil record of the genus Homo is first found. Here we combine magnetostratigraphy and strontium (Sr) isotope stratigraphy to improve age control on hominin-bearing upper Burgi (UBU) deposits in Areas 105 and 131 on the Karari Ridge in the eastern Turkana Basin (Kenya). We identify the base of the Olduvai subchron (bC2n) plus a short isolated interval of consistently normal polarity that we interpret to be the Pre-Olduvai event. Combined with precession-forced (∼20 kyr [thousands of years]) wet–dry climate cycles resolved by Sr isotope ratios, the magnetostratigraphic data allow us to construct an age model for the UBU deposits. We provide detailed age constraints for 15 hominin fossils from Area 131, showing that key specimens such as cranium KNM-ER 1470, partial face KNM-ER 62000 and mandibles KNM-ER 1482, KNM-ER 1801, and KNM-ER 1802 can be constrained between 1.945 ± 0.004 and 2.058 ± 0.034 Ma, and thus older than previously estimated. The new ages are consistent with a temporal overlap of two species of early Homo that can be distinguished by their facial morphology. Further, our results show that in this time interval, hominins occurred throughout the wet–dry climate cycles, supporting the hypothesis that the lacustrine Turkana Basin was a refugium during regionally dry periods. By establishing the observed first appearance datum of a marine-derived stingray in UBU deposits at 2.058 ± 0.034 Ma, we show that at this time the Turkana Basin was hydrographically connected to the Indian Ocean, facilitating dispersal of fauna between these areas. From a biogeographical perspective, we propose that the Indian Ocean coastal strip should be considered as a possible source area for one or more of the multiple Homo species in the Turkana Basin from over 2 Ma onwards.     

New Australopithecus robustus fossils and associated U-Pb dates from Cooper's Cave (Gauteng, South Africa)

Australopithecus robustus is one of the best represented hominin taxa in Africa, with hundreds of specimens recovered from six fossil localities in the Bloubank Valley area of Gauteng Province, South Africa. However, precise geochronological ages are presently lacking for these fossil cave infills. In this paper, we provide a detailed geological background to a series of hominin fossils retrieved from the newly investigated deposit of Cooper's D (located partway between Sterkfontein and Kromdraai in the Bloubank Valley), including uranium-lead (U-Pb) ages for speleothem material associated with A. robustus. U-Pb dating of a basal speleothem underlying the entire deposit results in a maximum age of 1.526 (±0.088) Ma for Cooper's D. A second U-Pb date of ca. 1.4 Ma is produced from a flowstone layer above this basal speleothem; since this upper flowstone is not a capping flowstone, and fossiliferous sediments are preserved above this layer, some of the hominins might be slightly younger than the calculated age. As a result, we can broadly constrain the age of the hominins from Cooper's D to between 1.5 and approximately 1.4 Ma. Extinct fauna recorded in this comparatively young deposit raise the possibility that the Bloubank Valley region of South Africa represented a more stable environmental refugium for taxa relative to tectonically more active East Africa. The sediments of the deposit likely infilled rapidly during periods when arid conditions prevailed in the paleoenvironment, although it is unclear whether sediment deposition and bone deposition were necessarily contemporaneous occurrences. We reconstruct the paleoenvironment of Cooper's D as predominantly grassland, with nearby woodlands and a permanent water source. The hominin teeth recovered from Cooper's D are all from juveniles and can be confidently assigned to A. robustus. In addition, two juvenile mandibular fragments and an adult thoracic vertebra are tentatively attributed to A. robustus.     

Stratigraphic context and direct dating of the Neandertal mandible from Cova del Gegant (Sitges, Barcelona)

Stratigraphic study of the Cova del Gegant’s sedimentary fill revealed different cycles of accumulation of typical interior cave and delta facies. A precise chronology for these deposits, the faunal remains and stone tools contained therein was obtained by radiocarbon, U-Th and OSL. Our results indicate that the Upper Pleistocene archaeological sequence dates between 49.3 ± 1.8 ka BP, the U-Th age of the overlying flowstone, and 60.0 ± 3.9 ka BP, the OSL age of the basal deposits. We have also directly dated the site’s Neandertal mandible to 52.3 ± 2.3 ka by U-Th.     

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.     

Orbital forcing and the spread of C4 grasses in the late Neogene: stable isotope evidence from South African speleothems

Reconstructing Plio-Pleistocene African paleoenvironments is important for models of early hominin evolution, but is often hampered by low-resolution or discontinuous climatic data. Here, we present high-resolution stable oxygen and carbon isotope time series data from two flowstones (secondary cave deposits) from the South African hominin-bearing Makapansgat Valley. The age of the older of the two flowstones (Collapsed Cone) is constrained by magnetostratigraphy to approximately 4-5 Ma; the younger flowstone (Buffalo Cave) grew between 2.0-1.5 Ma, as determined by magnetostratigraphy and orbital tuning of the isotopic data. The carbon isotope data is used as a proxy for the proportion of C(4) grasses in the local environment and the oxygen isotope data reflects monsoon rainfall intensity. The carbon isotope evidence indicates that in the late Miocene/early Pliocene, the local environment was dominated by C(3) vegetation, whereas, in the Plio-Pleistocene, it was composed of a mixture of C(3) and C(4) vegetation. This suggests that C(4) grasses became a significant part of the Makapansgat Valley ecosystem at approximately 4-5 Ma, towards the end of the late Neogene global expansion of C(4) grasses. After this initial expansion, South Africa experienced further fluctuations in the proportion of C(3) and C(4) vegetation during the Plio-Pleistocene, in response to regional and global climatic changes. Most notably, the Buffalo Cave flowstone provides evidence for C(4) grass expansion at ca. 1.7 Ma that we suggest was a response to African aridity caused by the onset of the Walker Circulation in the Pacific Ocean at this time.     

Stratigraphy, U-Th chronology, and paleoenvironments at Gladysvale Cave: insights into the climatic control of South African hominin-bearing cave deposits

Gladysvale Cave is one of the few Plio-Pleistocene hominin-bearing cave sites in South Africa that contains a well-stratified cave fill with clastic sediments interspersed with flowstones. The clastic sediments can be divided into units based on the presence of intercalated flowstones, forming flowstone bounded units (FBU). Ten MC-ICP-MS uranium-series dates on several flowstone horizons in the Gladysvale Internal Deposit fan indicate deposition from the late mid-Pleistocene ( approximately 570 ka) to Holocene ( approximately 7 ka) during limited periods of higher effective moisture. Clastic sedimentation occurred during the interceding, presumably more arid, periods. This sequence is not consistent with earlier models for South African caves that simply assumed interglacial sedimentation and glacial erosion. (13)C/(12)C data suggest that flowstone tended to form during periods with higher proportions of C(3) plants in the local vegetation, while clastic sediments reflect higher proportions of C(4) grasses, although this is not always the case. We argue that flowstones are precipitated during periods of higher effective precipitation and restricted cave entrances, while clastic sediments accumulated during periods with more open vegetation. The sedimentary fill of the fossiliferous deposits are, therefore, highly episodic in nature, with large periods of time unlikely to be represented. This has serious implications for the other hominin-bearing caves close by, as these deposits are likely to be similarly episodic. This is especially pertinent when addressing extinction events and reconstructions of paleoenvironments, as large periods of time may be unrecorded. The Gladysvale Cave fill sediments may serve as a climatically forced chronostratigraphic model for these less well-stratified and well-dated Plio-Pleistocene sites.     

Combined ESR/U-series chronology of Acheulian hominid-bearing layers at Trinchera Galería site,Atapuerca, Spain

The Sierra de Atapuerca, northern Spain, is known from many prehistoric and palaeontological sites documenting human prehistory in Europe. Three major sites, Gran Dolina, Galería and Sima del Elefante, range in age from the oldest hominin of Western Europe dated to 1.1 to 1.3 Ma (millions of years ago) at Sima del Elefante to c.a. 0.2 Ma on the top of the Galería archaeological sequence. Recently, a chronology based on luminescence methods (Thermoluminescence [TL] and Infrared Stimulated Luminescence [IRSL]) applied to cave sediments was published for the Gran Dolina and Galería sites. The authors proposed for Galería an age of 450 ka (thousands of years ago) for the units lower GIII and GII, suggesting that the human occupation there is younger than the hominid remains of Sima de los Huesos (>530 ka) around 1 km away.     

Reconstructing the geomorphic history of Liang Bua, Flores, Indonesia: a stratigraphic interpretation of the occupational environment

Liang Bua, in Flores, Indonesia, was formed as a subterranean chamber over 600 ka. From this time to the present, a series of geomorphic events influenced the structure of the cave and cave deposits, creating a complex stratigraphy. Within these deposits, nine main sedimentary units have been identified. The stratigraphic relationships between these units provide the evidence needed to reconstruct the geomorphic history of the cave. This history was dominated by water action, including slope wash processes, channel formation, pooling of water, and flowstone precipitation, which created waterfalls, cut-and-fill stratigraphy, large pools of water, and extensive flowstone cappings. The reconstructed sequence of events over the last 190 k.yr. has been summarized by a series of time slices that demonstrate the nature of the occupational environment in Liang Bua. The earliest artifacts at the site, dated to ∼190 ka, testify to hominin presence in the area, but the reconstructions suggest that occupation of the cave itself may not have been possible until after ∼100 ka. At ∼95 ka, channel erosion of a basal unit, which displays evidence of deposition in a pond environment, created a greater relief on the cave floor, and formed remanent areas of higher ground that later became a focus for hominin occupation from 74-61 ka by the west wall and in the center of the cave, and from ∼18-17 ka by the east wall. These zones have been identified according to the sloping nature of the stratigraphy and the distribution of artifacts, and their locations have implications for the archaeological interpretation of the site.     

An ecological reassessment of the southern African carnivore guild: a case study from Member 4, Sterkfontein, South Africa

The southern African late Pliocene to early Pleistocene carnivore guild was much larger than that of the present day. Understanding how this guild may have functioned is important for the reconstruction of carnivore-hominin interactions and to assess the potential for hominin scavenging in southern Africa. In modern ecosystems, the coexistence of larger carnivore species is constrained by several factors, which include high levels of interspecific competition. Here, the composition of the fossil carnivore guild is examined using Sterkfontein Member 4 (Cradle of Humankind, South Africa) as a case study. Sterkfontein Member 4 contains 10 larger carnivore taxa (body mass > 21.5 kg) and may also contain two Australopithecus species. Two possible causes of higher numbers of carnivore species in the South African fossil record are initially considered. First, that there is a bias introduced through comparing assemblages of differing sizes; second, carnivore biodiversity may have been artificially inflated due to previous taxonomic splitting of carnivore species, such as Crocuta. These possibilities are rejected and modern ecological data are used to construct a simple spatial model to determine how many carnivores could have co-existed. Although the resulting model indicates that the carnivore taxa present in Member 4 could have co-occurred, modern ecological studies indicate that it is highly unlikely that they would have co-existed simultaneously. Considering the complex depositional processes that operate in the southern African cave sites, it is proposed that the larger carnivore guild observed in the Sterkfontein Member 4 fossil assemblage is a palimpsest created by time-averaging. In light of this, we suggest that sites which have a large number of carnivore taxa should be examined for time-averaging, while those sites which have relatively few species may be a better reflection of carnivore communities.     

Les derniers Homo heidelbergensis et leurs descendants les néandertaliens : datation des sites d’Orgnac 3, du Lazaret et de Zafarraya

This article presents the dating results recently obtained on three archaeological sites in Europe. At Orgnac 3 (Ardèche, France) from where the last Homo heidelbergensis fossils are associated with the first evidence of levallois technique, two speleothem formations from the 5b–6–7^th layers were U-Th dated with MC-ICPMS, giving an age range of 319–255 ka (2σ) (MIS 8–9), while the volcanic ash-bearing second layer was dated by ⁴⁰Ar/³⁹Ar, obtaining a preliminary date of 308.2 ± 6.8 ka (2σ). The combined ESR/U-Th dating of red deer enamel teeth from Lazaret cave (Alpes-Maritimes, France) attributed ages of 120–190 ka to the Acheulean and pre-Mousterian layers (MIS 6), which is in agreement with previous TIMS U-Th dates between 108 and 44 ka on calcite samples from the overlying TRA trench (MIS 5, 4, 3). At Zafarraya (Andalousie, Espagne), a number of ¹⁴C measurements on charcoal samples as well as combined ESR/U-Th dates on Capra and Equus dental enamels assigned the Mousterian artefacts and neandertalian fossils-bearing deposits an age interval between 42 and 34 ka (MIS 3).     

Homo floresiensis: a cladistic analysis

The announcement of a new species, Homo floresiensis, a primitive hominin that survived until relatively recent times is an enormous challenge to paradigms of human evolution. Until this announcement, the dominant paradigm stipulated that: 1) only more derived hominins had emerged from Africa, and 2) H. sapiens was the only hominin since the demise of Homo erectus and Homo neanderthalensis. Resistance to H. floresiensis has been intense, and debate centers on two sets of competing hypotheses: 1) that it is a primitive hominin, and 2) that it is a modern human, either a pygmoid form or a pathological individual. Despite a range of analytical techniques having been applied to the question, no resolution has been reached. Here, we use cladistic analysis, a tool that has not, until now, been applied to the problem, to establish the phylogenetic position of the species. Our results produce two equally parsimonious phylogenetic trees. The first suggests that H. floresiensis is an early hominin that emerged after Homo rudolfensis (1.86 Ma) but before H. habilis (1.66 Ma, or after 1.9 Ma if the earlier chronology for H. habilis is retained). The second tree indicates H. floresiensis branched after Homo habilis.     

Geological summary of the Busidima Formation (Plio-Pleistocene) at the Hadar paleoanthropological site, Afar Depression, Ethiopia

The Hadar paleoanthropological site in Ethiopia preserves a record of hominin evolution spanning from approximately 3.45 Ma to 0.8 Ma. An angular unconformity just above the ca. 2.95 Ma BKT-2 complex divides the sediments into the Hadar Formation (ca. 3.8-2.9 Ma) and the Busidima Formation (ca. 2.7-0.15 Ma). The unconformity is likely a response to a major tectonic reorganization in the Afar Depression, and activation of the As Duma fault near the Ethiopian Escarpment (west of Hadar) created a half-graben in which the Busidima Formation was deposited. The pattern and character of sedimentation in the region changed dramatically above the unconformity, as cut-and-fill channel conglomerates and silt-dominated paleosols that comprise the Busidima Formation stand in sharp contrast to the underlying deposits of the Hadar Formation. Conglomerate deposition has been related to both the perennial, axial paleo-Awash and ephemeral, escarpment-draining tributaries. Overbank silts have yielded fossils attributed to early Homo and Oldowan stone tools. Numerous tuffaceous deposits exist within the Busidima Formation, but they are often spatially limited, fine-grained, and reworked. Recent work on the tephrostratigraphic framework of the Busidima Formation at Hadar has identified at least 12 distinct vitric tephras and established the first geochemical-based correlations between Hadar and the neighboring project areas of Gona and Dikika. Compared to Gona and Dikika, where Busidima Formation sediments are exposed over large areas, the highly discontinuous sediments at Hadar comprise less than 40 m in composite section and are exposed over an area of <20 km², providing only snapshots into the 2.7-0.15 Ma window. The stratigraphic record at Hadar confirms the complex depositional history of the Busidima Formation, and also provides important details on regional stratigraphic correlations and the pattern of deposition and erosion in the lower Awash Valley reflective of its tectonic history.     

Stretching the time span of hominin evolution at Kromdraai (Gauteng,South Africa): Recent discoveries

The Plio-Pleistocene locality of Kromdraai B has yielded the type specimen of Paranthropus robustus, as well as 27 additional fossil hominin specimens. In a number of both cranial and dental features, the states shown by the Kromdraai Paranthropus are more conservative when compared to the more derived conditions displayed by both South African conspecifics and the post-2.3 Ma eastern African Paranthropus boisei. Since 2014, we excavated the earliest known infilling of the Kromdraai cave system in a previously unexplored area. This new locality provided as yet 2200 identifiable macrovertebrate fossils, including 22 hominins, all tied in the earliest part of the stratigraphic sequence, representing three distinct depositional periods. Since we report here, for the first time, the occurrence of fossil hominins in Members 1 and 2, our discoveries stretch the time span of hominin evolution at Kromdraai and contribute to a better understanding of the origin of Paranthropus in southern Africa.     

Hovk 1 and the Middle and Upper Paleolithic of Armenia: a preliminary framework

The territory of present day Armenia is a geographic contact zone between the Near East and the northern Caucasus. Armenian Middle and Upper Paleolithic records are both few and patchy as a result of the historical paucity of systematic archaeological research in the country. Consequently, it is currently difficult to correlate the Armenian Middle and Upper Paleolithic records with those from other neighboring regions. We present new archaeological and chronometric data (luminescence, U-Th, and ¹⁴C) from our ongoing research at Hovk 1 Cave in northeast Armenia. We discuss in particular two activity phases in Hovk 1 Cave for which we have outline chronometric data: (1) an early Middle Paleolithic occupational phase, dated by optically stimulated luminescence (OSL) to 104 ± 9.8 ka BP_OSL; and (2) a Paleolithic occupational phase characterized by microlithic flakes dated by AMS ¹⁴C to 39,109 ± 1,324 calibrated years BP_Hulu. The two phases are separated by a hiatus in hominin occupation corresponding to MIS 4 and an episode in early MIS 3. These chronometric data, taken together with the preliminary paleoenvironmental reconstruction of the Hovk 1 Cave and environment, suggest that these activity phases represent short-lived and seasonal use of the cave presumably by small groups of hunters during episodes of mild climate. Neither tool manufacture nor butchery appears to have taken place within the cave, and consequently, the archaeological record included, for the most part, finished tools and blanks. We address the chronology and techno-typological aspects of Hovk 1 lithics in relation to: (1) the Paleolithic records of Armenia, and (2) the broader interregional context of early Middle Paleolithic hominin occupation and the Middle-Upper Paleolithic transition in the Caucasus.     

New fossils of Australopithecus anamensis from Kanapoi,West Turkana,Kenya (2003–2008)

Renewed fieldwork from 2003 through 2008 at the Australopithecus anamensis type-site of Kanapoi, Kenya, yielded nine new fossils attributable to this species. These fossils all date to between 4.195 and 4.108 million years ago. Most were recovered from the lower fluvial sequence at the site, with one from the lacustrine sequence deltaic sands that overlie the lower fluvial deposits but are still below the Kanapoi Tuff. The new specimens include a partial edentulous mandible, partial maxillary dentition, two partial mandibular dentitions, and five isolated teeth. The new Kanapoi hominin fossils increase the sample known from the earliest Australopithecus, and provide new insights into morphology within this taxon. They support the distinctiveness of the early A. anamensis fossils relative to earlier hominins and to the later Australopithecus afarensis. The new fossils do not appreciably extend the range of observed variation in A. anamensis from Kanapoi, with the exception of some slightly larger molars, and a canine tooth root that is the largest in the hominin fossil record. All of the Kanapoi hominins share a distinctive morphology of the canine–premolar complex, typical early hominin low canine crowns but with mesiodistally longer honing teeth than seen in A. afarensis, and large, probably dimorphic, canine tooth roots. The new Kanapoi specimens support the observation that canine crown height, morphology, root size and dimorphism were not altered from a primitive ape-like condition as part of a single event in human evolution, and that there may have been an adaptive difference in canine function between A. anamensis and A. afarensis.     

Correlation of the KHS Tuff of the Kibish Formation to volcanic ash layers at other sites, and the age of early Homo sapiens (Omo I and Omo II)

Hominin specimens Omo I and Omo II from Member I of the Kibish Formation, Ethiopia are attributed to early Homo sapiens, and an age near 196 ka has been suggested for them. The KHS Tuff, within Member II of the Kibish Formation has not been directly dated at the site, but it is believed to have been deposited at or near the time of formation of sapropel S6 in the Mediterranean Sea. Electron microprobe analyses suggest that the KHS Tuff correlates with the WAVT (Waidedo Vitric Tuff) at Herto, Gona, and Konso (sample TA-55), and with Unit D at Kulkuletti in the Ethiopian Rift Valley. Konso sample TA-55 is older than 154 ka, and Unit D at Kulkuletti is dated at 183 ka. These correlations and ages provide strong support for the age originally suggested for the hominin remains Omo I and Omo II, and for correlation of times of deposition in the Kibish region with formation of sapropels in the Mediterranean Sea. The Aliyo Tuff in Member III of the Kibish Formation is dated at 104 ka, and correlates with Gademotta Unit 15 in the Ethiopian Rift Valley.     

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.