Re-appraisal of the stratigraphy and determination of new U-Pb dates for the Sterkfontein hominin site, South Africa

Sterkfontein Caves is the single richest early hominin site in the world with deposits yielding one or more species of Australopithecus and possible early Homo, as well as an extensive faunal collection. The inability to date the southern African cave sites accurately or precisely has hindered attempts to integrate the hominin fossil evidence into pan-African scenarios about human evolutionary history, and especially hominin biogeography. We have used U-Pb and U-Th techniques to date sheets of calcium carbonate flowstone inter-bedded between the fossiliferous sediments. For the first time, absolute age ranges can be assigned to the fossil-bearing deposits: Member 2 is between 2.8 ± 0.28 and 2.6 ± 0.30 Ma and Member 4 between 2.65 ± 0.30 and 2.01 ± 0.05 Ma. The age of 2.01 ± 0.05 Ma for the top of Member 4 constrains the last appearance of Australopithecus africanus to 2 Ma. In the Silberberg Grotto we have reproduced the U-Pb age of ∼2.2 Ma of for the flowstones associated with StW573. We believe that these deposits, including the fossil and the flowstones, accumulated rapidly around 2.2 Ma. The stratigraphy of the site is complex as sediments are exposed both in the underground chambers and at surface. We present a new interpretation of the stratigraphy based on surface mapping, boreholes logs and U-Pb ages. Every effort was made to retain the Member system, however, only Members 2 and 4 are recognized in the boreholes. We propose that the deposits formally known as Member 3 are in fact the distal equivalents of Member 4. The sediments of Members 2 and 4 consisted of cone-like deposits and probably never filled up the cave. The U-Th ages show that there are substantial deposits younger than 400 ka in the underground cave, underlying the older deposits, highlighting again that these cave fills are not simple layer-cakes.     

Dietary adaptations of South African australopiths: inference from enamel prism attitude

The angle at which enamel prisms approach the wear surface holds information with regard to the stiffness of the tissue, as well as its wear resistance. Hence, analyses of prism orientation may shed light on questions of whether the thick enamel in hominins has evolved to confer stiffness or wear resistance to the teeth and may thus inform about the diet and behavioural ecology of these species. This was explored for Paranthropus robustus and Australopithecus africanus, whereby a distinction was made between prisms at the Phase I and Phase II facets. The results were compared with those obtained for Theropithecus, Macaca, and Potamochoerus for whom behavioural and/or experimental data are available, and were interpreted against simple mechanical principles. The South African hominins differ significantly in their relationships between wear facets and prism angulations. Teeth of P. robustus are better adapted to more vertical loads during mastication (Phase I), whereas those of A. africanus are better adapted to cope with more laterally-directed loads (Phase II) commonly associated with roll-crush and mastication. Overall, teeth of P. robustus appear stiffer, while those of A. africanus seem more wear resistant.     

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.     

Carnivora from the Plio-Pleistocene hominin site of Drimolen, Gauteng, South Africa

Drimolen is one of the newest and most productive hominin sites in South Africa, and is dated on faunal grounds between 2.0 Ma to 1.5 Ma. This paper provides the first overview of the Carnivora from Drimolen, updating the previously published preliminary faunal list, and describing all currently prepared craniodental and postcranial material. The Drimolen specimens are described in comparison with other modern and fossil South African carnivore material. The carnivores cover a range of taxa including hyaenids, felids, canids and herpestids. Most notable amongst these are the sabretooth Dinofelis aff. piveteaui craniodental and postcranial remains, which are described in detail, and a Chasmaporthetes nitidula cranium. The genus Chasmaporthetes is found at three other sites in the area - Sterkfontein, Swartkrans and Coopers D. There are two models for the geographic origin of Dinofelis piveteaui, in that it may have arisen in either eastern or southern Africa. These possibilities are discussed in the light of the new South African Dinofelis material, as the Drimolen material appears to represent a more primitive form with affinities with D. piveteaui. Fossil leopard material from Kromdraai B and Drimolen is also discussed, as the metapodia assigned to P. pardus from these two sites are very small, but lie within the variation of modern leopards. Such size differences in fossil postcrania may have implications for the niches that these animals may have occupied in the past.     

The enigmatic molar from Gondolin, South Africa: Implications for Paranthropus paleobiology

The specific attribution of the large hominin M₂ (GDA-2) from Gondolin has significant implications for the paleobiology of Paranthropus. If it is a specimen of Paranthropus robustus it impacts that species' size range, and if it belongs to Paranthropus boisei it has important biogeographic implications. We evaluate crown size, cusp proportions and the likelihood of encountering a large-bodied mammal species in both East and South Africa in the Early Pleistocene. The tooth falls well outside the P. robustus sample range, and comfortably within that for penecontemporaneous P. boisei. Analyses of sample range, distribution and variability suggest that it is possible, albeit unlikely to find a M₂ of this size in the current P. robustus sample. However, taphonomic agents - carnivore (particularly leopard) feeding behaviors - have likely skewed the size distribution of the Swartkrans and Drimolen P. robustus assemblage. In particular, assemblages of large-bodied mammals accumulated by leopards typically display high proportions of juveniles and smaller adults. The skew in the P. robustus sample is consistent with this type of assemblage. Morphological evidence in the form of cusp proportions is congruent with GDA-2 representing P. robustus rather than P. boisei. The comparatively small number of large-bodied mammal species common to both South and East Africa in the Early Pleistocene suggests a low probability of encountering an herbivorous australopith in both. Our results are most consistent with the interpretation of the Gondolin molar as a very large specimen of P. robustus. This, in turn, suggests that large, presumptive male, specimens are rare, and that the levels of size variation (sexual dimorphism) previously ascribed to this species are likely to be gross underestimates.     

Plio-Pleistocene evolution of the genus Castor (Rodentia, Mammalia) in Europe: C. fiber plicidens of Pietrafitta (Perugia, Central Italy)

In the lignite sediments of Pietrafitta (Tiberino Basin, Umbria, Central Italy), a rich fossil assemblage of vertebrate, invertebrate and plant remains belonging to the Farneta Faunal Unit (Late Villafranchian, Early Pleistocene) was collected. Among them fossil beaver cranial remains are relatively abundant. Western-Central Europe Villafranchian beaver remains were assigned to C. plicidens by some authors because molar occlusal surface patterns show complex enamel crenulations. Several recent authors have classified them as C. fiber while analysing other morphological patterns. Our samples have been compared to Plio-Pleistocene fossil remains and to living European populations of the genus Castor. New morphometric parameters on molar occlusal surfaces have been defined and statistical analyses (One-Way MANOVA, Principal Component Analysis and Canonical Discriminant Analysis) have been performed on them. The results point out a subspecific separation between the Late Villafranchian beaver of Pietrafitta, Quarata and San Giovanni in Valdarno (Umbria and Tuscany) and C. fiber populations. St. Vallier (France) Late Pliocene and Mosbach 2 (Germany) Middle Pleistocene beavers classified respectively by Viret and Friant as C. plicidens, show a C. f. fiber molar teeth pattern. Therefore, C. fiber plicidens did not occur in Central-Western Europe and this subspecific name may be used only for the local populations of Valdarno and Tiberino Basin (San Giovanni in Valdarno, Quarata, Pietrafitta and a few localities of the same area), at that time peripheral populations, probably semi-isolated during the Late Villafranchian, and at the southern limit of the geographic range of C. fiber.     

A comparative study of frontal bone morphology among Pleistocene hominin fossil groups

Features of the frontal bone that are conventionally used to distinguish among fossil hominin groups were quantitatively examined. Fifty-five fossil crania dating from the early to the late Pleistocene were analyzed. Using a modified pantograph, outlines of the frontal bone were collected along the midsagittal and two parasagittal planes. The profile from nasion to bregma, as well as two profiles above the medial and lateral sections of the orbit, respectively, extending from the orbital margin to the coronal suture were traced. The outlines were measured using Elliptical Fourier Function Analysis (EFFA), which enabled a quantification of aspects of the frontal bone that have historically been described primarily in nonmetric or linear terms. Four measurements were obtained: 1) overall morphology as expressed in the Fourier harmonic amplitudes; 2) maximum projection of the supraorbital torus at three points along the browridge (glabella and the medial and lateral aspects of the torus above the orbit); 3) maximum distance of the frontal squama from the frontal chord, capturing forehead curvature; and 4) nasion-bregma chord length. The results indicate that the midsagittal profile is significantly different among all Pleistocene groups in analyses that include both size and shape, as well as size-adjusted data. Homo erectus is significantly different from the late Pleistocene groups (Neandertals and early modern H. sapiens) in glabellar projection. Anatomically modern humans are significantly different from all other groups in both raw and size-standardized analyses of all three outlines that captured overall morphology, as well as forehead curvature and lateral supraorbital torus prominence, and middle Pleistocene Homo are significantly different in both medial and lateral overall parasagittal form. However, for the majority of analyses there were no significant differences among the Pleistocene archaic groups in supraorbital torus projection, frontal squama curvature, nasion-bregma chord length, or overall frontal bone morphology.     

A new hominin from the Basal Member of the Hadar Formation, Dikika, Ethiopia, and its geological context

In this paper we report for the first time hominin remains from the Basal Member of the Hadar Formation at Dikika, in the Awash Valley of Ethiopia, dating to greater than 3.4 Ma. The new fossil, DIK-2-1, is a fragment of a left mandible and associated dentition. The mandible is attributed to Australopithecus afarensis. However, the new fossil exhibits some metric and morphological features that have not previously been seen in the A. afarensis hypodigm, increasing the already impressive degree of variation in the mandibular sample of the species.     

First occurrence of early Homo in the Nachukui Formation (West Turkana, Kenya) at 2.3-2.4 Myr

Cognitive abilities and techno-economic behaviours of hominids in the time period between 2.6-2.3 Myr have become increasingly well-documented. This time period corresponds to the oldest evidence for stone tools at Gona (Kada Gona, West Gona, EG 10-12, OGS 6-7), Hadar (AL 666), lower Omo valley (Ftji1, 2 & 5, Omo 57, Omo 123) in Ethiopia, and West Turkana (Lokalalei sites -LA1 & LA2C-) in Kenya. In 2002 a new palaeoanthropological site (LA1alpha), 100 meters south of the LA1 archaeological site, produced a first right lower molar of a juvenile hominid (KNM-WT 42718). The relative small size of the crown, its marked MD elongation and BL reduction, the relative position of the cusps, the lack of a C6 and the mild expression of a protostylid, reinforced by metrical analyses, demonstrate the distinctiveness of this tooth compared with Australopithecus afarensis, A. anamensis, A. africanus and Paranthropus boisei, and its similarity to early Homo. The LA1alpha site lies 2.2 m above the Ekalalei Tuff which is slightly younger than Tuff F dated to 2.34+/-0.04 Myr. This juvenile specimen represents the oldest occurrence of the genus Homo in West Turkana.     

Three new species and two new combinations in the Aizoaceae from the Western and Northern Cape of South Africa

Three new species of Aizoaceae from the Western Cape are described. Octopoma tanquanum Klak and Vlokia montana Klak are dwarf shrubby or mat-forming succulents which belong to the Ruschieae in the Ruschioideae. Mesembryanthemum knolfonteinense Klak of the Mesembryanthemoideae is a geophyte. Octopoma tanquanum inhabits relatively low lying arid areas within the Tanqua Karoo and the Little Karoo and is thought to be closely allied to the two species of Octopoma found in the Little Karoo. Both V. montana and M. knolfonteinense grow at relatively high altitudes within the fynbos biome. In addition, Ruschia littlewoodii L.Bolus is transferred to Phiambolia, and two new combinations are made in Antimima for Ruschia hexamera L.Bolus and Ruschia radicans L.Bolus.     

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.     

New species of Cercopithecoides from Haasgat, North West Province, South Africa

Analyses of new cercopithecid fossil specimens from the South African site of Haasgat point to craniofacial affinities with the genus Cercopithecoides. Detailed metric and non-metric comparisons with South African Cercopithecoides williamsi, and other East African Cercopithecoides species, Cercopithecoides kimeui, Cercopithecoides meaveae, Cercopithecoides kerioensis, and Cercopithecoides alemyehui demonstrate that the Haasgat fossils have distinct craniofacial morphology and dental metrics. Specifically, material from Haasgat probably represents one of the smaller Cercopithecoides, differing from the others in its particular suite of features that vary within the genus. It is unique in its more vertical ramus, associated with a relatively lengthened mandibular body. Haasgat Cercopithecoides has a particularly narrow interorbital region between relatively larger ovoid orbits, with articulation of the maxillary bones at a suture above the triangular nasal bones. Furthermore, the maxillary arcade is more rounded than other Cercopithecoides, converging at the M² and M³. The conclusion drawn from this analysis is that the Pleistocene Haasgat fossils are colobines representing a distinct taxon of Cercopithecoides, Cercopithecoides haasgati, thus adding a second species of the genus to southern Africa.     

Two new species of Asteraceae (tribe Anthemideae,subtribe Pentziinae) from the Cape Floristic Region of South Africa

Two new species of Asteraceae (tribe Anthemideae, subtribe Pentziinae) from the Cape Floristic Region of South Africa are described. Marasmodes schlechteri Magee & J.C.Manning, a local endemic from the lowlands between Piketberg and the Olifants River Mountains, is distinguished by its spreading leaves with axillary fascicles and relatively large, solitary capitula with obconical involucre borne on very short lateral shoots. Pentzia trifida Schltr. ex. Magee & J.C.Manning is a well-collected limestone fynbos endemic that is readily distinguished by the regularly trifid leaves, broadly cyathiform involucres with lanceolate to oblanceolate bracts, and the well developed pappus extending the entire length of the relatively short corolla tube.     

A closer look at the “Protopithecus” fossil assemblages: new genus and species from Bahia,Brazil

The recently extinct large-bodied New World monkey Protopithecus brasiliensis Lund 1836 was named based on a distal humerus and proximal femur found in the Lagoa Santa cave system in the southeastern Brazilian state of Minas Gerais. These bones are from an animal about twice the size of the largest extant platyrrhines. One hundred and seventy-five years later, a nearly complete skeleton was discovered in the Toca da Boa Vista caves in the neighboring state of Bahia and was allocated to the same taxon as it was the first platyrrhine fossil of comparable size found since the originals. Our detailed study of the equivalent elements, however, reveals important morphological differences that do not correspond to intraspecific variation as we know it in related platyrrhine taxa. The presence of both an expanded brachioradialis flange on the humerus and gluteal tuberosity on the femur of the Bahian skeleton distinguishes it from the Lagoa Santa fossil as well as from all other platyrrhines. Further cranial and postcranial evidence suggests a closer relationship of the former with the alouattine Alouatta, while the limited Lund material fits more comfortably with the ateline clade. Therefore, we propose to limit P. brasiliensis Lund to the distal humerus and proximal femur from Lagoa Santa and erect a new genus and species for the skeleton from Toca da Boa Vista. Cartelles coimbrafilhoi was a large-bodied frugivore with a relatively small brain and diverse locomotor repertoire including both suspension and climbing that expands the range of platyrrhine biodiversity beyond the dimensions of the living neotropical primates.     

Pelargonium elandsmontanum (Geraniaceae: section Hoarea), a new species from the Western Cape Province, South Africa

The new species Pelargonium elandsmontanum is a local endemic from Elandsberg Nature Reserve near Malmesbury in the Western Cape Province, South Africa. One of six species of sect. Hoarea with just the posterior two petals developed, it resembles P. ternifolium in its trifoliolate leaves and pink petals but is distinguished from that species by the short, stout petioles, rhombic, acute leaflets with the upper surface glabrous or with spreading hairs (vs cuneate, apically incised leaflets with both surfaces adpressed-hairy), and five (vs four) fertile stamens.     

Juvenile ornithopod (Dinosauria: Rhabdodontidae) remains from the Upper Cretaceous (Lower Campanian, Gosau Group) of Muthmannsdorf (Lower Austria)

The fragmentary remains of a juvenile rhabdodontid ornithopod from the Coal-bearing Complex of the Gosau Group (Lower Campanian, Grünbach syncline) at Muthmannsdorf near Wiener Neustadt, Lower Austria are revised. The material, probably belonging to a single individual, includes a right dentary (lectotype of Iguanodon suessi Bunzel, 1871, designated herein), teeth, a fragmentary parietal, fragments of scapula, ?radius, femur, tibia, two vertebrae (lost) and a manual ungual.The lectotype dentary does not provide clear autapomorphies or sufficient diagnostic features to determine its position within the Rhabdodontidae at generic level. By this “Iguanodon suessi” Bunzel, 1871 and the genus “Mochlodon” Seeley, 1881, to which it was latter referred as type species, cannot be characterized sufficiently by differential diagnosis and these are best considered nomina dubia. Based upon combined character comparisons (mainly postcranial features) the Muthmannsdorf ornithopod is referred herein to Zalmoxes Weishampel, Jianu, Csiki and Norman, 2003, a genus so far known from the late Maastrichtian of Romania. It probably but not evidently represents a yet unnamed species, most closely related to Zalmoxes shqiperorum Weishampel, Jianu, Csiki and Norman, 2003. At the present state of knowledge the Austrian material is not further diagnostic at the species level and kept in open nomenclature as Zalmoxes sp.