Reappraisal of the taxonomic status of the cranium Stw 53 from the Plio/Pleistocene of Sterkfontein,in South Africa

A fossil skull, Stw 53, from the Plio/Pleistocene of Sterkfontein, in South Africa, has been referred toHomo habilis Leakey, Napier, andTobias, 1964. Reappraisal of its putative hominine affinity reveals a closer resemblance toAustralopithecus africanus Dart, 1925. The skull, as reconstructed, is too small forH. habilis; with no indication of brain expansion overA. africanus; has a facial angle outside the hominine range, but identical with that ofA. africanus; and whose teeth are not elongated but display buccolingual expansion. Although it was found in the same strata (Member 5) as stone tools, there is no causal connection. It has been dated faunistically at 2–1.5 my BP, but due to an unconformity it is suggested that it could be older. In spite of its late date, Stw 53 shows no intermediate characters which could support a trend towardsH. habilis orA. robustus Broom, 1938. It may, therefore, represent a relict population ofA. africanus.     

A complete second metatarsal (StW 89) from Sterkfontein Member 4, South Africa

The functional anatomy of the hominin foot has played a crucial role in studies of locomotor evolution in human ancestors and extinct relatives. However, foot fossils are rare, often isolated, and fragmentary. Here, we describe a complete hominin second metatarsal (StW 89) from the 2.0-2.6 million year old deposits of Member 4, Sterkfontein Cave, South Africa. Like many other fossil foot bones, it displays a mosaic of derived human-like features and primitive ape-like features. StW 89 possesses a domed metatarsal head with a prominent sulcus, indicating dorsiflexion at the metatarsophalangeal joint during bipedal walking. However, while the range of motion at the metatarsophalangeal joint is human-like in dorsiflexion, it is ape-like in plantarflexion. Furthermore, StW 89 possesses internal torsion of the head, an anatomy decidedly unlike that found in humans today. Unlike other hominin second metatarsals, StW 89 has a dorsoplantarly gracile base, perhaps suggesting more midfoot laxity. In these latter two anatomies, the StW 89 second metatarsal is quite similar to the recently described second metatarsal of the partial foot from Burtele, Ethiopia. We interpret this combination of anatomies as evidence for a low medial longitudinal arch in a foot engaged in both bipedal locomotion, but also some degree of pedal, and perhaps even hallucal, grasping. Additional fossil evidence will be required to determine if differences between this bone and other second metatarsals from Sterkfontein reflect normal variation in an evolving lineage, or taxonomic diversity.     

Continuous dental eruption identifies Sts 5 as the developmentally oldest fossil hominin and informs the taxonomy of Australopithecus africanus

The relatively small Australopithecus africanus specimen Sts 5 has figured prominently in taxonomic debates, and the determination of this specimen as a young male or an elderly female has the potential to offer a great deal of resolution on this question. Sts 5 has been argued to be either a small, immature male or a mature female based on a variety of characters.     

Stratigraphy, artefact industries and hominid associations for Sterkfontein, member 5

A revised stratigraphy for the early hominid site of Sterkfontein (Gauteng Province, South Africa) reveals a complex distribution of infills in the main excavation area between 2.8 and 1.4 m.y.a, as well as deposits dating to the mid to late Pleistocene. New research now shows that the Member 4 australopithecine breccia (2.8-2.6 Ma) extends further west than was previously thought, while a late phase of Member 4 is recognized in a southern area. The artefact-bearing breccias were defined sedimentologically as Member 5, but one supposed part of these younger breccias, the StW 53 infill, lacks in situ stone tools, although it does appear to post-date 2.6 Ma when artefacts first appear in the archaeological record. The StW 53 hominid, previously referred to Homo habilis, is here argued to be Australopithecus. The first artefact-bearing breccia of Member 5 is the Oldowan Infill, estimated at 2-1.7 Ma. It occupies a restricted distribution in Member 5 east and contains an expedient, flake-based tool industry associated with a few fossils of Paranthropos robustus. An enlarged cave opening subsequently admitted one or more Early Acheulean infills associated in Member 5 west with Homo ergaster. The artefacts attest to a larger site accumulation between ca. 1.7 and 1.4 Ma, with more intensive use of quartzite over quartz and a subtle but important shift to large flakes and heavier-duty tools. The available information on palaeoenvironments is summarized, showing an overall change from tropical to sub-tropical gallery forest, forest fringe and woodland conditions in Member 4 to more open woodland and grassland habitats in the later units, but with suggestions of a wet localized topography in the Paranthropus -bearing Oldowan Infill.     

Functional anatomy of the hand of Australopithecus africanus

The Sterkfontein hand bones, attributed to Australopithecus africanus, were analysed to determine potential hand function of the power grip type of this species. The metacarpus is as stable as that of modern humans, as indicated by the depth of the groove on the base of metacarpal 2, the styloid process of metacarpal 3, the base articular surface areas, and the ligament markings on the bases of the metacarpals. The flexion and rotation of metacarpal 5 might have been less than that of modern humans, due to a more marked ventral articular lip on the base. The metacarpus acts as a lever, acting in various planes. The extensor carpi ulnaris and extensor carpi radialis longus muscles were probably better developed than in modern humans. The extensor carpi radialis brevis and flexor carpi radialis muscles would probably have been as well developed as in modern humans. None of the long tendons have a mechanical disadvantage as compared to modern humans. The metacarpals have a high robusticity index. The proximal phalanges show some midshaft swelling, slightly greater curvature than in modern humans, and some side to side bowing: pongid features. The fibrous flexor sheath markings are well developed, but resemble those of modern humans rather than those of the pongids. A single middle phalanx resembles that of modern humans, and has well developed ridges for insertion of the flexor digitorum superficialis muscle. The distal phalanx of the thumb has a well developed region for insertion of the flexor pollicis longus muscle, and has a mechanical advantage over modern humans for action of this muscle at the interphalangeal joint. The features indicate that the hand of A. africanus was well adapted to powerful hand use, as in hammering, striking, chopping, scraping, and gouging actions, as well as for throwing and climbing activities.     

Adaptive and phylogenetic significance of ontogenetic sequences in Archaeolemur, subfossil lemur from Madagascar

Among the best known of recently extinct Malagasy lemurs is Archaeolemur, which is represented by many hundreds of specimens. The phylogenetic affinities of this taxon are unclear, especially in light of recent preliminary analysis of ancient DNA which does not support its previously accepted close relationship with the living Indridae. We examined the nearly complete skeletons of two adults and one juvenile and other less complete specimens to reconstruct aspects of the ontogeny of Archaeolemur. To compare the development of Archaeolemur to that of living strepsirrhines we collected data on Propithecus verreauxi, Eulemur fulvus, and Lemur catta. Additionally, because Archaeolemur exhibits some morphological convergences with distantly related papionins, we tested for convergence in the developmental patterns of Archaeolemur and Macaca fascicularis. Data include the status of tooth eruption, craniofacial sutural closure, and postcranial epiphyseal fusion, as well as linear measurements. We used discriminant function analysis and other tools to explore ontogenetic similarities and differences. The adaptive and phylogenetic significance of ontogenetic information is discussed. Our analysis shows that Archaeolemur displays a clear strepsirrhine pattern of development with only minor macaque convergences. Among the Strepsirrhini, Archaeolemur is slightly more similar developmentally to E. fulvus and L. catta than to P. verreauxi. Some of the distinctive features of the ontogeny of Archaeolemur may be related to diet, while others bear apparent testimony to a relatively rapid absolute pace of growth and development.     

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.     

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.     

Degradation and Utilization of Xylans by the Rumen AnaerobePrevotella bryantii(formerlyP. ruminicolasubsp.brevis) B14

Freshly harvested whole cells from cultures ofP. bryantiiB₁4 grown with oat spelt xylan (OSX) as an energy source showed less than 25% of the enzyme activity against OSX, and less than 15% of the activity against birchwood xylan (BWX) and carboxymethylcellulose, that was detectable in sonicated cell preparations. This indicates that much of this hydrolytic activity is either periplasmic, membrane-associated or intracellular and may be concerned with the processing of transported oligosaccharides.P. bryantiiB₁4 cultures were able to utilise up to 45% and 51% of the total pentose present in OSX and BWX, respectively, after 24 h, but could utilize 84% of a water-soluble fraction of BWX. Analysis of the xylan left undegraded after incubation withP. bryantiishowed that while xylose and arabinose were removed to a similar extent, uronic acids were utilized to a greater extent than xylose. Predigestion of xylans with two cloned xylanases from the cellulolytic rumen anaerobeRuminococcus flavefaciensgave little increase in overall pentose utilization suggesting that externalP. bryantiixylanases are as effective as the clonedR. flavefaciensenzymes in releasing products that can be utilised byP. bryantiicells. The xylanase system ofP. bryantiiis able to efficiently utilise not only xylo-oligosaccharides but also larger water-soluble xylan fragments.     

Purification and properties of Pinus taeda arginase from germinated seedlings

In germinated loblolly pine (Pinus taeda L.) seeds arginine accumulates in the seedling during its growth immediately following germination. The enzyme arginase (L-arginine amidinohydrolase, EC 3.5.3.1) is responsible for hydrolyzing this arginine into ornithine and urea. Loblolly pine arginase was purified to homogeneity from seedling cotyledons by chromatographic separation on DE-52 cellulose, Matrex Green and arginine-linked Sepharose 4B. The enzyme was purified 148-fold and a single polypeptide band was identified as arginase. The molecular mass was determined to be 140 kDa by FPLC, while the subunit size was shown to be 37 kDa by SDS-PAGE, predicting a homotetramer holoprotein. Removal of manganese from the enzyme abolishes catalytic activity, which can be restored by incubating the protein with Mn²⁺. Antibodies, raised against the arginase subunit, are able to immunotitrate arginase activity and are monospecific for arginase on immunoblots.     

Newly discovered fossil- and artifact-bearing deposits, uranium-series ages, and Plio-Pleistocene hominids at Swartkrans Cave, South Africa

We report on new research at Swartkrans Cave, South Africa, that provides evidence of two previously unrealized artifact- and fossil-bearing deposits. These deposits underlie a speleothem dated by the uranium-thorium disequilibrium technique to 110,000 ± 1,980 years old, the first tightly constrained, geochronological date available for the site. Recovered fauna from the two underlying deposits—including, prominently, the dental remains of Paranthropus (Australopithecus) robustus from the uppermost layer (Talus Cone Deposit)—indicate a significantly older, late Pliocene or early Pleistocene age for these units. The lowest unit (LB East Extension) is inferred to be an eastward extension of the well-known Lower Bank of Member 1, the earliest surviving infill represented at the site. The date acquired from the speleothem also sets the maximum age of a rich Middle Stone Age lithic assemblage.     

Two new parrotfishes of the genus Scarus from the Central and South Pacific, with further examples of sexual dichromatism

Scarus lauia (Jordan & Evermann) and Scarus formosus Valenciennes are junior synonyms oi Scarus dubius Bennett, the former based on the terminal phase and the latter on the initial phase. Scarus dubius is endemic to the Hawaiian Islands, one of five species of the genus in these islands. The initial phase of Scarus fiavipectoralis Schultz is grey to brown with whitish streaks on the abdomen, a dark caudal fin and yellow at the pectoral base. The range of this species is extended from the Philippines to the Marshall, Solomon, and Caroline Islands and the Great Barrier Reef. Scarusfasciatus Valenciennes is a junior svnonym of of the western Pacific 5. rivulatus Valenciennes. The initial phase is grey to brown with three pale stripes on the abdomen. Scarus lepidus Jenyns is a junior synonym of the wide-ranging Indo-Pacific S. globiceps Valenciennes. The initial phase is coloured like that of S. rivulatus. Scarus janthochir Bleeker, S. chlorodon Jenyns, and S. singaporensis Bleeker are junior synonyms of 5. prasiognathos Valenciennes; it occurs from the western Pacific to the Maldive Islands. Scarus elerae Jordan & Seale and S. urbanus (Smith) are junior synonyms of the wide-ranging Indo-Pacific S. tricolor Bleeker based on the terminal phase. Scarus dimidiatus Bleeker occurs in the western Pacific east to Samoa. Its initial phase, very similar to that of the western Indian Ocean S. scaber, is yellowish, shading to whitish ventrally, with five slightly diagonal dark bars on the upper half of the body. Callyodon mutabilis Gray is a junior synonym of S. quoyi Valenciennes. Scarus spinus (Kner) is the valid name for the distinctive bullet-headed parrotfish with a greenish yellow snout which was misidentified as S.formosus by Schultz. The initial phase is dark brown with pale bars. The terminal phase of the Pacific parrotfish Scarus schlegeli Bleeker is distinctive in possessing a pale bar on the side (yellow dorsally, light green ventrally). The drab initial phase with alternating dark and light bars was misidentified as S. venosus Valenciennes by Schultz, a junior synonym of S. psittacus Forsskǎl. The terminal phase of the western Pacific Scarus bleekeri de Beaufort has a characteristic large whitish patch bordered with blue-green on the cheek. The initial phase is dark brown, the edges of the scales narrowly orangish, shading to orange-red ventrally, with faint yellowish bars, caudal peduncle and caudal fin. A closely related species from Indonesia and the Andaman Sea is tentatively identified as 5. troschelii Bleeker. The terminal phase has a band of blue-green curving from the corner of the mouth to below the eye and back down to the pectoral base. Scarus japanensis (Bloch), identified in most recent papers as S. capistratoides Bleeker, is the terminal phase and senior synonym of the dark brown, red-tailed S. pyrrhurus (Jordan & Seale); it is restricted to the western Pacific (easternmost record, Samoa). Scarus gibbus Ruppell is differentiated into three populations based on colour. Scarus koputea sp. nov., unique in having four rows of scales on the cheek, is described from the Marquesas Islands. Scarus longipinnis sp. nov. is described from Pitcairn, Rapa, and the Capricorn Group of the Great Barrier Reef; its pelvic lins are longer and the dorsal fin more elevated than other Scarus.     

The cation status of some indigenous plants from a Cango valley limestone-sandstone transition,South Africa

Leaf tissue of four representative species from a limestone-sandstone transition, together with their respective soils, were analysed for cations. The species studied comprisedLeucospermum cuneiforme (Proteaceae) from the acid sandstone,Indigofera sp. nov. (Fabaceae) from the base-rich limestone,Elytropappus rhinocerotis (Asteraceae) from the transition zone andDodonaea angustifolia (Sapindaceae) which occured on all three substrates. The sandstone substrate is low in cations, particularly K and it is suggested that the Leucospermum may supplement its K nutrition with Na. Despite growing on soils of differing cation content, Dodonaea maintained a constant leaf cation content over the three substrates.     

Dental variation ofProconsulfrom the Tinderet region, Kenya

The morphology of mandibular molars of the twoProconsulspecies,Proconsul majorandProconsul africanus, from the Tinderet region, Kenya was analyzed. While the molar size variability within the TinderetP. majorwas slightly greater than those of local African ape subspecies, the shape variability was comparable. Because the twoProconsulspecies show some differences in cusp areal proportions,P. majoris not just a largerP. africanusin molar morphology. Napak specimens are generally similar to the TinderetP. major.