The DNH 7 skull of Australopithecus robustus from Drimolen (Main Quarry), South Africa.

Although the early hominin species Australopithecus robustus has been known for more than eight decades and is represented by hundreds of fossils from sites in South Africa, a complete, well-preserved skull has been elusive. DNH 7, an adult cranium and mandible from the Drimolen site, was identified, on the basis of its small size, as a presumptive female of A. robustus. Here, we provide a detailed comparative description of the specimen. In cranial, facial, and dental size, DNH 7 is confirmed to lie at the extreme small end of the A. robustus range of variation, along with a few fragmentary maxillofacial specimens from Swartkrans. In addition, relative to the classically derived craniofacial features of the Swartkrans+Kromdraai portions of the A. robustus hypodigm, primitive anatomy pervades the DNH 7 face, braincase, and cranial base. Taken together, these pieces of evidence place DNH 7 in a previously unfilled position on the robust Australopithecus morphocline, where the specimen highlights the morphological distinctions between southern and eastern African species of this group and epitomizes the anatomy expected of the group's last common ancestor.

The functional role of the Carabelli trait in early and late hominins.

The Carabelli trait is a dental feature that forms along the lingual margin of the protocone of deciduous and permanent maxillary molars. It is variably expressed, ranging from a small pit or furrow to a large cusp, and its development seems to be associated with crown size and molar cusp spatial configuration. The degree of expression of the Carabelli trait differs systematically between hominin taxa, and for this reason, it has been used extensively in the reconstruction of their phylogeny. However, the functional implications of having a large Carabelli trait remain unclear. In this study, we analyze the macrowear pattern of maxillary molars of early and late hominins using the occlusal fingerprint analysis method, an approach based on digital models of teeth that helps in reconstructing occlusal dynamics occurring during mastication. Tooth crowns with a small Carabelli cusp generally exhibit larger wear contact areas that extended cervically, while two additional new occlusal contact areas are common in teeth characterized by a large Carabelli cusp. These wear areas are created at the beginning of the chewing cycle, when occluding with the slopes of the lingual groove of the lower molars, between the metaconid and entoconid cusps. Advancing tooth wear leads to a slight enlargement of Carabelli occlusal contacts increasing their functional area. A steep inclination could be mechanically important in food reduction and in balancing the functional load distribution during mastication contacts. Steep wear areas are particularly developed in primates that process foods characterized by tough and fibrous textural properties. Future biomechanical and microwear texture analyses could provide additional information on the mechanical adaptation of this dental trait.

Contemporaneity of Australopithecus, Paranthropus, and early Homo erectus in South Africa.

Understanding the extinction of Australopithecus and origins of Paranthropus and Homo in South Africa has been hampered by the perceived complex geological context of hominin fossils, poor chronological resolution, and a lack of well-preserved early Homo specimens. We describe, date, and contextualize the discovery of two hominin crania from Drimolen Main Quarry in South Africa. At ~2.04 million to 1.95 million years old, DNH 152 represents the earliest definitive occurrence of Paranthropus robustus, and DNH 134 represents the earliest occurrence of a cranium with clear affinities to Homo erectus These crania also show that Homo, Paranthropus, and Australopithecus were contemporaneous at ~2 million years ago. This high taxonomic diversity is also reflected in non-hominin species and provides evidence of endemic evolution and dispersal during a period of climatic variability.

Root caries on a Paranthropus robustus third molar from Drimolen.

OBJECTIVES: Dental caries is often perceived as a modern human disease. However, their presence is documented in many early human groups, various nonhuman primates and, increasingly, our hominin ancestors and relatives. In this study, we describe an antemortem lesion on the root of a Paranthropus robustus third molar from Drimolen, South Africa, which likely represents another example of caries in fossil hominins. MATERIALS AND METHODS: The molar, DNH 40, is dated to 2.0-1.5 Ma and displays a lesion on the mesial root surface, extending from the cementoenamel junction 3 mm down toward the apex. The position and severity of the lesion was macroscopically recorded and micro-CT scanned to determine the extent of dentine involvement. RESULTS: A differential diagnosis indicates root caries, as the lesion is indistinguishable from clinical examples. Although necrotic in appearance, external tertiary dentine is evident on a micro CT scan. Gingival recession and/or continuous eruption of the tooth as a result of extensive occlusal wear would have occurred to facilitate caries formation. Therefore, the lesion is likely linked to relative old age of this individual. DISCUSSION: This new example increases the total number of carious lesions described in P. robustus teeth to 10, on occlusal, interproximal, and now, root surfaces. Beyond the consumption of caries-causing food, caries formation would have also required the presence of requisite intraoral cariogenic bacteria in this individual and the species. Of interest, the presence of tertiary dentine on the outward surface suggests the DNH 40 lesion may have been arrested, that is, no longer active, perhaps relating to a change in diet or oral microbiome just prior to the individual's death.

Ages-at-death distribution of the early Pleistocene hominin fossil assemblage from Drimolen (South Africa).

OBJECTIVES: A prevailing hypothesis in paleoanthropology is that early Pleistocene hominin bones were accumulated in South African caves by carnivores, which used those shelters, and the trees surrounding them, as refuge and feeding sites. We tested this hypothesis at the site of Drimolen, by comparing its hominin age-at-death distribution to that of the nearby and roughly contemporaneous site of Swartkrans. MATERIALS AND METHODS: We employed standard dental aging systems in order to categorize the Drimolen hominin teeth into age classes of 5 years each. We then compared the age-at-death distribution for Drimolen with the published data available for the Swartkrans hominins. RESULTS: Age-at-death distributions indicate that the age category "young adults" is the best represented age category at Swartkrans and the most poorly represented one at Drimolen. Moreover, Drimolen has a preponderance of infant specimens. Both sites have a low frequency of old adult specimens. CONCLUSIONS: Differences observed in frequencies of the age-at-death categories suggest different mechanisms of hominin skeletal accumulation at Drimolen and Swartkrans. Swartkrans' frequency curve reflects mortality in a population subjected to predation and is thus consistent with the carnivore-accumulating hypothesis. In contrast, the Drimolen curve is similar to that of wild populations of living apes. Living primates have been observed exploiting caves as sleeping shelters, for nutritional, security, drinking, and thermoregulatory purposes. We suggest that similar cave use by Pleistocene hominins can explain, in large part, the accumulation of hominin bones at Drimolen. Such a conclusion is another illustration of the growing awareness that a "one-size-fits-all" taphonomic model for South African early Pleistocene hominin sites is probably insufficient.

Hominin skeletal part abundances and claims of deliberate disposal of corpses in the Middle Pleistocene.

Humans are set apart from other organisms by the realization of their own mortality. Thus, determining the prehistoric emergence of this capacity is of significant interest to understanding the uniqueness of the human animal. Tracing that capacity chronologically is possible through archaeological investigations that focus on physical markers that reflect "mortality salience." Among these markers is the deliberate and culturally mediated disposal of corpses. Some Neandertal bone assemblages are among the earliest reasonable claims for the deliberate disposal of hominins, but even these are vigorously debated. More dramatic assertions center on the Middle Pleistocene sites of Sima de los Huesos (SH, Spain) and the Dinaledi Chamber (DC, South Africa), where the remains of multiple hominin individuals were found in deep caves, and under reported taphonomic circumstances that seem to discount the possibility that nonhominin actors and processes contributed to their formation. These claims, with significant implications for charting the evolution of the "human condition," deserve scrutiny. We test these assertions through machine-learning analyses of hominin skeletal part representation in the SH and DC assemblages. Our results indicate that nonanthropogenic agents and abiotic processes cannot yet be ruled out as significant contributors to the ultimate condition of both collections. This finding does not falsify hypotheses of deliberate disposal for the SH and DC corpses, but does indicate that the data also support partially or completely nonanthropogenic formational histories.

Macromammalian faunas, biochronology and palaeoecology of the early Pleistocene Main Quarry hominin-bearing deposits of the Drimolen Palaeocave System, South Africa.

The Drimolen Palaeocave System Main Quarry deposits (DMQ) are some of the most prolific hominin and primate-bearing deposits in the Fossil Hominids of South Africa UNESCO World Heritage Site. Discovered in the 1990s, excavations into the DMQ have yielded a demographically diverse sample of Paranthropus robustus (including DNH 7, the most complete cranium of the species recovered to date), early Homo, Papio hamadryas robinsoni and Cercopithecoides williamsi. Alongside the hominin and primate sample is a diverse macromammalian assemblage, but prior publications have only provided a provisional species list and an analysis of the carnivores recovered prior to 2008. Here we present the first description and analysis of the non-primate macromammalian faunas from the DMQ, including all 826 taxonomically identifiable specimens catalogued from over two decades of excavation. We also provide a biochronological interpretation of the DMQ deposits and an initial discussion of local palaeoecology based on taxon representation.The current DMQ assemblage consists of the remains of minimally 147 individuals from 9 Orders and 14 Families of mammals. The carnivore assemblage described here is even more diverse than established in prior publications, including the identification of Megantereon whitei, Lycyaenops silberbergi, and first evidence for the occurrence of Dinofelis cf. barlowi and Dinofelis aff. piveteaui within a single South African site deposit. The cetartiodactyl assemblage is dominated by bovids, with the specimen composition unique in the high recovery of horn cores and dominance of Antidorcas recki remains. Other cetartiodactyl and perissodactyl taxa are represented by few specimens, as are Hystrix and Procavia; the latter somewhat surprisingly so given their common occurrence at penecontemporaneous deposits in the region. Equally unusual (particularly given the size of the sample) is the identification of single specimens of giraffoid, elephantid and aardvark (Orycteropus cf. afer) that are rarely recovered from regional site deposits. Despite the diversity within the DMQ macromammalian faunas, there are few habitat- or biochronologically-sensitive species that provide specific ecologic or age boundaries for the deposits. Recovered species can only support the non-specific, mixed open-to-closed palaeohabitats around Drimolen that have been reconstructed for the other penecontemporaneous South African palaeokarst deposits. The identified Equus quagga ssp. specimens recovered from the floor of the current excavation (∾-4.5-5 m below datum) suggests that most, if not all the DMQ specimens, were deposited after 2.33 Ma. Simultaneously, the carnivore specimens (D. cf. barlowi, L. silberbergi) suggest earlier Pleistocene (pre- 2.0-1.8 Ma) to maximally 1.6 Ma deposition (D. aff. piveteaui) for most of the DMQ fossil assemblage.

Early hominin auditory capacities.

Studies of sensory capacities in past life forms have offered new insights into their adaptations and lifeways. Audition is particularly amenable to study in fossils because it is strongly related to physical properties that can be approached through their skeletal structures. We have studied the anatomy of the outer and middle ear in the early hominin taxa Australopithecus africanus and Paranthropus robustus and estimated their auditory capacities. Compared with chimpanzees, the early hominin taxa are derived toward modern humans in their slightly shorter and wider external auditory canal, smaller tympanic membrane, and lower malleus/incus lever ratio, but they remain primitive in the small size of their stapes footplate. Compared with chimpanzees, both early hominin taxa show a heightened sensitivity to frequencies between 1.5 and 3.5 kHz and an occupied band of maximum sensitivity that is shifted toward slightly higher frequencies. The results have implications for sensory ecology and communication, and suggest that the early hominin auditory pattern may have facilitated an increased emphasis on short-range vocal communication in open habitats.

Dental ontogeny in pliocene and early pleistocene hominins.

Until recently, our understanding of the evolution of human growth and development derived from studies of fossil juveniles that employed extant populations for both age determination and comparison. This circular approach has led to considerable debate about the human-like and ape-like affinities of fossil hominins. Teeth are invaluable for understanding maturation as age at death can be directly assessed from dental microstructure, and dental development has been shown to correlate with life history across primates broadly. We employ non-destructive synchrotron imaging to characterize incremental development, molar emergence, and age at death in more than 20 Australopithecus anamensis, Australopithecus africanus, Paranthropus robustus and South African early Homo juveniles. Long-period line periodicities range from at least 6-12 days (possibly 5-13 days), and do not support the hypothesis that australopiths have lower mean values than extant or fossil Homo. Crown formation times of australopith and early Homo postcanine teeth fall below or at the low end of extant human values; Paranthropus robustus dentitions have the shortest formation times. Pliocene and early Pleistocene hominins show remarkable variation, and previous reports of age at death that employ a narrow range of estimated long-period line periodicities, cuspal enamel thicknesses, or initiation ages are likely to be in error. New chronological ages for SK 62 and StW 151 are several months younger than previous histological estimates, while Sts 24 is more than one year older. Extant human standards overestimate age at death in hominins predating Homo sapiens, and should not be applied to other fossil taxa. We urge caution when inferring life history as aspects of dental development in Pliocene and early Pleistocene fossils are distinct from modern humans and African apes, and recent work has challenged the predictive power of primate-wide associations between hominoid first molar emergence and certain life history variables.

Brief communication: beyond the South African cave paradigm--Australopithecus africanus from Plio-Pleistocene paleosol deposits at Taung.

Following the discovery of the "Taung Child" (Australopithecus africanus) in 1924 in the Buxton-Norlim Limeworks near Taung, the fossil-bearing deposits associated with the Dart and Hrdlicka pinnacles have been interpreted as the mined remnants of cave sediments that formed within the Plio-Pleistocene Thabaseek Tufa: either as a younger cave-fill or as contemporaneous carapace caves. When combined with the Plio-Pleistocene dolomitic cave deposits from the "Cradle of Humankind," a rather restricted view emerges that South African early hominins derived from cave deposits, whereas those of east and central Africa are derived from fluvio-lacustrine and paleosol deposits. We undertook a sedimentological and paleomagnetic analysis of the pink-colored deposit (PCS) from which the "Taung Child" is purported to have derived and demonstrate that it is a calcrete, a carbonate-rich pedogenic sediment, which formed on the paleo-land surface. The deposit extends 100 s of meters laterally beyond the Dart and Hrdlicka Pinnacles where it is interbedded with the Thabaseek Tufa, indicating multiple episodes of calcrete development and tufa growth. The presence of in situ rhizoconcretions and insect trace fossils (Celliforma sp. and Coprinisphaera sp.) and the distinctive carbonate microfabric confirm that the pink deposit is a pedogenic calcrete, not a calcified cave sediment. Paleomagnetic and stratigraphic evidence indicates that a second, reversed polarity, fossil-bearing deposit (YRSS) is a younger fissure-fill formed within a solutional cavity of the normal polarity tufa and pink calcrete (PCS). These observations have implications for the dating, environment, and taphonomy of the site, and increase the likelihood of future fossil discoveries within the Buxton-Norlim Limeworks.

Variation in enamel thickness within the genus Homo.

Recent humans and their fossil relatives are classified as having thick molar enamel, one of very few dental traits that distinguish hominins from living African apes. However, little is known about enamel thickness in the earliest members of the genus Homo, and recent studies of later Homo report considerable intra- and inter-specific variation. In order to assess taxonomic, geographic, and temporal trends in enamel thickness, we applied micro-computed tomographic imaging to 150 fossil Homo teeth spanning two million years. Early Homo postcanine teeth from Africa and Asia show highly variable average and relative enamel thickness (AET and RET) values. Three molars from South Africa exceed Homo AET and RET ranges, resembling the hyper thick Paranthropus condition. Most later Homo groups (archaic European and north African Homo, and fossil and recent Homo sapiens) possess absolutely and relatively thick enamel across the entire dentition. In contrast, Neanderthals show relatively thin enamel in their incisors, canines, premolars, and molars, although incisor AET values are similar to H. sapiens. Comparisons of recent and fossil H. sapiens reveal that dental size reduction has led to a disproportionate decrease in coronal dentine compared with enamel (although both are reduced), leading to relatively thicker enamel in recent humans. General characterizations of hominins as having 'thick enamel' thus oversimplify a surprisingly variable craniodental trait with limited taxonomic utility within a genus. Moreover, estimates of dental attrition rates employed in paleodemographic reconstruction may be biased when this variation is not considered. Additional research is necessary to reconstruct hominin dietary ecology since thick enamel is not a prerequisite for hard-object feeding, and it is present in most later Homo species despite advances in technology and food processing.

Early hominin dental remains from the Plio-Pleistocene site of Drimolen, South Africa.

The Plio-Pleistocene fossil hominin site of Drimolen is located approximately 5.5km north of the other well-known South African Plio-Pleistocene sites (Sterkfontein, Swartkrans, Kromdraai, Coopers). It was discovered by A.W. Keyser in 1992. Systematic excavations led to the recovery of a remarkable number of fossil vertebrate taxa, including hominins. Most of the specimens collected consist of isolated teeth or those in jaws. The aim of this paper is to provide a morphological description of the dental specimens. The taxonomic allocation of each specimen is also reported, either confirming or revising previous provisional attributions. The analysis confirms the occurrence of two hominin species, Paranthropus robustus and Homo sp. With over 80 fossil hominin specimens recovered so far, Drimolen is the second largest sample of P. robustus, after Swartkrans. At Drimolen, P. robustus is represented mostly by craniodental specimens (63) among which are 47 isolated teeth and the remainder are maxillary and mandibular fragments with teeth. The assemblage markedly increases the dental sample of P. robustus. Furthermore, the Drimolen sample includes tooth classes not present in the Swartkrans or Kromdraai samples. The new tooth classes include both deciduous upper lateral incisors (DNH 31) and canines (DNH 23). In the dental sample described here there are nine specimens probably attributable to Homo, although a specific attribution is not yet possible. These specimens expand the small sample of early Homo from South African sites. Basic dimensions (MD and BL) of the Drimolen dental remains are compared in a preliminary analysis with other hominin samples. This analysis delineates the Drimolen P. robustus dental sample as characterized by smaller teeth overall than the Swartkrans sample (and in some cases also smaller than the Kromdraai sample), as well as a greater size range.

Extended male growth in a fossil hominin species.

In primates that are highly sexually dimorphic, males often reach maturity later than females, and young adult males do not show the size, morphology, and coloration of mature males. Here we describe extended male development in a hominin species, Paranthropus robustus. Ranking a large sample of facial remains on the basis of dental wear stages reveals a difference in size and robusticity between young adult and old adult males. Combined with estimates of sexual dimorphism, this pattern suggests that male reproductive strategy focused on monopolizing groups of females, in a manner similar to that of silverback gorillas. However, males appear to have borne a substantial cost in the form of high rates of predation.