Sex-biased sampling may influence Homo naledi tooth size variation.

A frequent source of debate in paleoanthropology concerns the taxonomic unity of fossil assemblages, with many hominin samples exhibiting elevated levels of variation that can be interpreted as indicating the presence of multiple species. By contrast, the large assemblage of hominin fossils from the Rising Star cave system, assigned to Homo naledi, exhibits a remarkably low degree of variation for most skeletal elements. Many factors can contribute to low sample variation, including genetic drift, strong natural selection, biased sex ratios, and sampling of closely related individuals. In this study, we tested for potential sex-biased sampling in the Rising Star dental sample. We compared coefficients of variation for the H. naledi teeth to those for eight extant hominoid samples. We used a resampling procedure that generated samples from the extant taxa that matched the sample size of the fossil sample for each possible Rising Star dental sex ratio. We found that variation at four H. naledi tooth positions-I2, M1, P4, M1-is so low that the possibility that one sex is represented by few or no individuals in the sample cannot be excluded. Additional evidence is needed to corroborate this inference, such as ancient DNA or enamel proteome data, and our study design does not address other potential factors that would account for low sample variation. Nevertheless, our results highlight the importance of considering the taphonomic history of a hominin assemblage and suggest that sex-biased sampling is a plausible explanation for the low level of phenotypic variation found in some aspects of the current H. naledi assemblage.

An examination of Homo naledi early juveniles recovered from the Rising Star cave system, South Africa.

BACKGROUND: Six Homo naledi early juveniles were recovered from U.W. 101 (Dinaledi Chamber), U.W. 102 (Lesedi Chamber), and U.W. 110 in the Rising Star cave system. AIM: This paper develops the information for the H. naledi early juvenile life stage, as defined by a combination of deciduous and permanent dentition, and the eruption of the first permanent molar. SUBJECTS AND METHODS: The growing number of young individuals recovered from the Rising Star cave system allows us to gain a better understanding of their variation, or lack thereof, and provides a basis to estimate broad ranges for age at death of the individuals. The individuals are identified and described through craniodental remains and spatial associations. RESULTS AND CONCLUSION: Our results show that the teeth are remarkably consistent across the localities in their metric and non-metric traits, and our analyses refine previous estimations on dental eruptions with the first permanent molar erupting first in the sequence among permanent teeth.

Descriptive catalog of Homo naledi dental remains from the 2013 to 2015 excavations of the Dinaledi Chamber, site U.W. 101, within the Rising Star cave system, South Africa.

More than 150 hominin teeth, dated to ∼330-241 thousand years ago, were recovered during the 2013-2015 excavations of the Dinaledi Chamber of the Rising Star cave system, South Africa. These fossils comprise the first large single-site sample of hominin teeth from the Middle Pleistocene of Africa. Though scattered remains attributable to Homo sapiens, or their possible lineal ancestors, are known from older and younger sites across the continent, the distinctive morphological feature set of the Dinaledi teeth supports the recognition of a novel hominin species, Homo naledi. This material provides evidence of African Homo lineage diversity that lasts until at least the Middle Pleistocene. Here, a catalog, anatomical descriptions, and details of preservation and taphonomic alteration are provided for the Dinaledi teeth. Where possible, provisional associations among teeth are also proposed. To facilitate future research, we also provide access to a catalog of surface files of the Rising Star jaws and teeth.

Ancient teeth, phenetic affinities, and African hominins: Another look at where Homo naledi fits in.

A new species of Homo, Homo naledi, was described in 2015 based on the hominin skeletal remains from the Dinaledi Chamber of the Rising Star cave system, South Africa. Subsequent craniodental comparative analyses, both phenetic and cladistic, served to support its taxonomic distinctiveness. Here we provide a new quantitative analysis, where up to 78 nonmetric crown and root traits of the permanent dentition were compared among samples of H. naledi (including remains from the recently discovered Lesedi Chamber) and eight other species from Africa: Australopithecus afarensis, Australopithecus africanus, Paranthropus boisei, Paranthropus robustus, Homo habilis, Homo erectus, Middle Pleistocene Homo sp., and Pleistocene and Holocene Homo sapiens. By using the mean measure of divergence distance statistic, phenetic affinities were calculated among samples to evaluate interspecific relatedness. The objective was to compare the results with those previously obtained, to assess further the taxonomic validity of the Rising Star hominin species. In accordance with earlier findings, H. naledi appears most similar dentally to the other African Homo samples. However, the former species is characterized by its retention and full expression of features relating to the main cusps, as well as the root numbers, with a near absence of accessory traits-including many that, based on various cladistic studies, are plesiomorphic in both extinct and extant African hominins. As such, the present findings provide additional support for the taxonomic validity of H. naledi as a distinct species of Homo.

Homo naledi pelvic remains from the Dinaledi Chamber, South Africa.

In the hominin fossil record, pelvic remains are sparse and are difficult to attribute taxonomically when they are not directly associated with craniodental material. Here we describe the pelvic remains from the Dinaledi Chamber in the Rising Star cave system, Cradle of Humankind, South Africa, which has produced hominin fossils of a new species, Homo naledi. Though this species has been attributed to Homo based on cranial and lower limb morphology, the morphology of some of the fragmentary pelvic remains recovered align more closely with specimens attributed to the species Australopithecus afarensis and Australopithecus africanus than they do with those of most (but not all) known species of the genus Homo. As with A. afarensis and A. africanus, H. naledi appears to have had marked lateral iliac flare and either a weakly developed or non-existent acetabulocristal buttress or a distinct, albeit weakly developed, acetabulospinous buttress. At the same time, H. naledi has robust superior pubic and ischiopubic rami and a short ischium with a narrow tuberoacetabular sulcus, similar to those found in modern humans. The fragmentary nature of the Dinaledi pelvic assemblage makes the attribution of sex and developmental age to individual specimens difficult, which in turn diminishes our ability to identify the number of individuals represented in the assemblage. At present, we can only confidently say that the pelvic fossils from Rising Star represent at least four individuals based on the presence of four overlapping right ischial fossils (whereas a minimum of 15 individuals can be identified from the Dinaledi dental assemblage). A primitive, early Australopithecus-like false pelvis combined with a derived Homo-like true pelvis is morphologically consistent with evidence from the lower ribcage and proximal femur of H. naledi. The overall similarity of H. naledi ilia to those of australopiths supports the inference, drawn from the observation of primitive pelvic morphology in the extinct species Homo floresiensis, that there is substantial variation in pelvic form within the genus Homo. In the light of these findings, we urge caution in making taxonomic attributions-even at the genus level-of isolated fossil ossa coxae.

Patterns of lateral enamel growth in Homo naledi as assessed through perikymata distribution and number.

Perikymata, incremental growth lines visible on tooth enamel surfaces, differ in their distribution and number among hominin species, although with overlapping patterns. This study asks: (1) How does the distribution of perikymata along the lateral enamel surface of Homo naledi anterior teeth compare to that of other hominins? (2) When both perikymata distribution and number are analyzed together, how distinct is H. naledi from other hominins? A total of 19 permanent anterior teeth (incisors and canines) of H. naledi were compared, by tooth type, to permanent anterior teeth of other hominins: Australopithecus afarensis, Australopithecus africanus, Paranthropus robustus, Paranthropus boisei, Homo ergaster/Homo erectus, other early Homo, Neandertals, and modern humans, with varying sample sizes. Repeated measures analyses of the percentage of perikymata per decile of reconstructed crown height yielded several statistically significant differences between H. naledi and other hominins. Canonical variates analysis of percentage of perikymata in the cervical half of the crown together with perikymata number revealed that, in 8 of 19 cases, H. naledi teeth were significantly unlikely to be classified as other hominins, while exhibiting least difference from modern humans (especially southern Africans). In a cross-validated analysis, 68% of the H. naledi teeth were classified as such, while 32% were classified as modern human (most often southern African). Of 313 comparative teeth use for this analysis, only 1.9% were classified as H. naledi. What tends to differentiate H. naledi anterior tooth crowns from those of most other hominins, including some modern humans, is strongly skewed perikymata distributions combined with perikymata numbers that fall in the middle to lower ranges of hominin values. H. naledi therefore tends toward a particular combination of these features that is less often seen in other hominins. Implications of these data for the growth and development of H. naledi anterior teeth are considered.

Brief communication: Dental microwear and diet of Homo naledi.

OBJECTIVES: A recent study of dental chipping suggested that Homo naledi teeth were exposed to "acute trauma" on a regular basis during life, presumably from the consumption of grit-laden foods. This follows debate concerning the etiology of dental chips in South African hominin teeth that dates back more than half a century. Some have argued that antemortem chips result from consumption of hard foods, such as nuts and seeds or bone, whereas others have claimed that exogenous grit on roots and tubers are responsible. Here we examine the dental microwear textures of H. naledi, both to reconstruct aspects of diet of these hominins and to assess the possibility that hard foods (gritty or otherwise) are the culprits for the unusually high antemortem chip incidence reported. METHODS: We made high-resolution replicas of original molars and found that ten individuals preserve antemortem wear. These were scanned by white-light scanning confocal profilometry and analyzed using scale-sensitive fractal analysis. Resulting data were compared with those published for other fossil hominins and extant non-human primates. RESULTS: Our results indicate that H. naledi had complex microwear textures dominated by large, deep pits. The only known fossil hominin with higher average texture complexity is Paranthropus robustus, and the closest extant primates in a comparative baseline series appear to be the hard-object feeder, Cercocebus atys, and the eurytopic generalist, Papio ursinus. CONCLUSIONS: This study suggests that H. naledi likely consumed hard and abrasive foods, such as nuts or tubers, at least on occasion, and that these might well be responsible for the pattern of chipping observed on their teeth.

The upper limb of Homo naledi.

The evolutionary transition from an ape-like to human-like upper extremity occurred in the context of a behavioral shift from an upper limb predominantly involved in locomotion to one adapted for manipulation. Selection for overarm throwing and endurance running is thought to have further shaped modern human shoulder girdle morphology and its position about the thorax. Homo naledi (Dinaledi Chamber, Rising Star Cave, Cradle of Humankind, South Africa) combines an australopith-like cranial capacity with dental characteristics akin to early Homo. Although the hand, foot, and lower limb display many derived morphologies, the upper limb retains many primitive traits. Here, we describe the H. naledi upper extremity (excluding the hand) in detail and in a comparative context to evaluate the diversity of clavicular, scapular, humeral, radial, and ulnar morphology among early hominins and later Homo. Homo naledi had a scapula with a markedly cranially-oriented glenoid, a humerus with extremely low torsion, and an australopith-like clavicle. These traits indicate that the H. naledi scapula was situated superiorly and laterally on the thorax. This shoulder girdle configuration is more similar to that of Australopithecus and distinct from that of modern humans, whose scapulae are positioned low and dorsally about the thorax. Although early Homo erectus maintains many primitive clavicular and humeral features, its derived scapular morphology suggests a loss of climbing adaptations. In contrast, the H. naledi upper limb is markedly primitive, retaining morphology conducive to climbing while lacking many of the derived features related to effective throwing or running purported to characterize other members of early Homo.

Skull diversity in the Homo lineage and the relative position of Homo naledi.

The discovery of Homo naledi has expanded the range of phenotypic variation in Homo, leading to new questions surrounding the mosaic nature of morphological evolution. Though currently undated, its unique morphological pattern and possible phylogenetic relationships to other hominin taxa suggest a complex evolutionary scenario. Here, we perform geometric morphometric analyses on H. naledi cranial and mandibular remains to investigate its morphological relationship with species of Homo and Australopithecus. We use Generalized Procrustes analysis to place H. naledi within the pattern of known hominin skull diversity, distributions of Procrustes distances among individuals to compare H. naledi and Homo erectus, and neighbor joining trees to investigate the potential phenetic relationships between groups. Our goal is to address a set of hypotheses relating to the uniqueness of H. naledi, its affinity with H. erectus, and the age of the fossils based on skull morphology. Our results indicate that, cranially, H. naledi aligns with members of the genus Homo, with closest affiliations to H. erectus. The mandibular results are less clear; H. naledi closely associates with a number of taxa, including some australopiths. However, results also show that although H. naledi shares similarities with H. erectus, some distances from this taxon - especially small-brained members of this taxon - are extreme. The neighbor joining trees place H. naledi firmly within Homo. The trees based on cranial morphology again indicate a close relationship between H. naledi and H. erectus, whereas the mandibular tree places H. naledi closer to basal Homo, suggesting a deeper antiquity. Altogether, these results emphasize the unique combination of features (H. erectus-like cranium, less derived mandible) defining H. naledi. Our results also highlight the variability within Homo, calling for a greater focus on the cause of this variability, and emphasizing the importance of using the total morphological package for species diagnoses.

Body size, brain size, and sexual dimorphism in Homo naledi from the Dinaledi Chamber.

Homo erectus and later humans have enlarged body sizes, reduced sexual dimorphism, elongated lower limbs, and increased encephalization compared to Australopithecus, together suggesting a distinct ecological pattern. The mosaic expression of such features in early Homo, including Homo habilis, Homo rudolfensis, and some early H. erectus, suggests that these traits do not constitute an integrated package. We examined the evidence for body mass, stature, limb proportions, body size and dental size dimorphism, and absolute and relative brain size in Homo naledi as represented in the Dinaledi Chamber sample. H. naledi stature and body mass are low compared to reported values for H. erectus, with the exception of some of the smaller bodied Dmanisi H. erectus specimens, and overlap with larger Australopithecus and early Homo estimates. H. naledi endocranial volumes (465-560 cc) and estimates of encephalization quotient are also similar to Australopithecus and low compared to all Homo specimens, with the exception of Homo floresiensis (LB1) and the smallest Dmanisi H. erectus specimen (D4500). Unlike Australopithecus, but similar to derived members of genus Homo, the Dinaledi assemblage of H. naledi exhibits both low levels of body mass and dental size variation, with an estimated body mass index of sexual dimorphism less than 20%, and appears to have an elongated lower limb. Thus, the H. naledi bauplan combines features not typically seen in Homo species (e.g., small brains and bodies) with those characteristic of H. erectus and more recent Homo species (e.g., reduced mass dimorphism, elongated lower limb).

The skull of Homo naledi.

The species Homo naledi was recently named from specimens recovered from the Dinaledi Chamber of the Rising Star cave system in South Africa. This large skeletal sample lacks associated faunal material and currently does not have a known chronological context. In this paper, we present comprehensive descriptions and metric comparisons of the recovered cranial and mandibular material. We describe 41 elements attributed to Dinaledi Hominin (DH1-DH5) individuals and paratype U.W. 101-377, and 32 additional cranial fragments. The H. naledi material was compared to Plio-Pleistocene fossil hominins using qualitative and quantitative analyses including over 100 linear measurements and ratios. We find that the Dinaledi cranial sample represents an anatomically homogeneous population that expands the range of morphological variation attributable to the genus Homo. Despite a relatively small cranial capacity that is within the range of australopiths and a few specimens of early Homo, H. naledi shares cranial characters with species across the genus Homo, including Homo habilis, Homo rudolfensis, Homo erectus, and Middle Pleistocene Homo. These include aspects of cranial form, facial morphology, and mandibular anatomy. However, the skull of H. naledi is readily distinguishable from existing species of Homo in both qualitative and quantitative assessments. Since H. naledi is currently undated, we discuss the evolutionary implications of its cranial morphology in a range of chronological frameworks. Finally, we designate a sixth Dinaledi Hominin (DH6) individual based on a juvenile mandible.

The postcranial skeletal maturation of Australopithecus sediba.

OBJECTIVES: In 2008, an immature hominin defined as the holotype of the new species Australopithecus sediba was discovered at the 1.9 million year old Malapa site in South Africa. The specimen (MH1) includes substantial post-cranial skeletal material, and provides a unique opportunity to assess its skeletal maturation. METHODS: Skeletal maturity indicators observed on the proximal and distal humerus, proximal ulna, distal radius, third metacarpal, ilium and ischium, proximal femur and calcaneus were used to assess the maturity of each bone in comparison to references for modern humans and for wild chimpanzees (Pan troglodytes). RESULTS: In comparison to humans the skeletal maturational ages for Au. sediba correspond to between 12.0 years and 15.0 years with a mean (SD) age of 13.1 (1.1) years. In comparison to the maturational pattern of chimpanzees the Au. sediba indicators suggest a skeletal maturational age of 9-11 years. Based on either of these skeletal maturity estimates and the body length at death of MH1, an adult height of 150-156 cm is predicted. DISCUSSION: We conclude that the skeletal remains of MH1 are consistent with an ape-like pattern of maturity when dental age estimates are also taken into consideration. This maturity schedule in australopiths is consistent with ape-like estimates of age at death for the Nariokotome Homo erectus remains (KMN-WT 15000), which are of similar postcranial immaturity to MH1. The findings suggest that humans may have distinctive and delayed post-cranial schedules from australopiths and H. erectus, implicating a recent evolution of somatic and possibly life history strategies in human evolution.

Comparative biomechanics of Australopithecus sediba mandibles.

Fossils attributed to Australopithecus sediba are described as having phylogenetic affinities with early Homo to the exclusion of other South African australopiths. With respect to functional anatomy of mastication, one implication of this hypothesis is that A. sediba mandibles should exhibit absolutely and relatively reduced stiffness and strength in comparison to Australopithecus africanus and Paranthropus robustus jaws. Examination of cortical bone distribution in the MH 1 and MH 2 mandibles of A. sediba (evaluated against samples of Pan, early and modern Homo as well as A. africanus and P. robustus) indicate that the A. sediba mandibular corpus was geometrically similar to other South African australopiths. In particular, enhanced torsional rigidity is characteristic of all South African australopiths including A. sediba. These findings are consistent with a hypothesis that masticatory mechanics may have been similar to other australopiths (and distinct from exemplars of early Homo), and as such suggest that A. sediba's mandibles were functionally suited to consume hard and tough objects. Recent mechanical modeling of the A. sediba cranium, however, has been interpreted as indicating that this species was relatively poorly adapted to produce large bite forces and likely experienced relatively modest strains in its facial skeleton. This paradox - that the cranium signals a departure from the australopith morphotype whereas the mandibles conform to a hypodigm of australopith grade - can be resolved, in part, if it is acknowledged that mechanical performance variables offer imperfect insight into what constitutes feeding adaptations.

The apportionment of tooth size and its implications in Australopithecus sediba versus other Plio-pleistocene and recent African hominins.

OBJECTIVES: Australopithecus sediba is characterized further by providing formerly unpublished and refined mesiodistal and buccolingual crown measurements in the MH1 and MH2 specimens. After size correction, these data were compared with those in other fossil and recent samples to facilitate additional insight into diachronic hominin affinities. MATERIALS AND METHODS: Six comparative samples consist of fossil species: A. africanus, A. afarensis, Homo habilis, Paranthropus robustus, P. boisei, and H. erectus. Others comprise H. sapiens and Pan troglodytes. Re-estimates of "actual" dimensions in damaged A. sediba teeth were effected through repeated measurements by independent observers. X-ray synchrotron microtomography allowed measurement of crowns obscured by matrix and noneruption. Tooth size apportionment analysis, an established technique for intraspecific comparisons, was then applied at this interspecific level to assess phenetic affinities using both within- and among-group data. RESULTS: Comparison of these highly heritable dimensions identified a general trend for smaller posterior relative to larger anterior teeth (not including canines), contra Paranthropus, that allies A. sediba with other australopiths and Homo; however, specific reductions and/or shape variation in the species' canines, third premolars, and anterior molars relative to the other teeth mirror the patterning characteristic of Homo. DISCUSSSION: Of all samples, including east African australopiths, A. sediba appears most like H. habilis, H. erectus and H. sapiens regarding how crown size is apportioned along the tooth rows. These findings parallel those in prior studies of dental and other skeletal data, including several that suggest A. sediba is a close relative of, if not ancestral to, Homo.

Revisiting the age of the Florisbad hominin material.

In 1996, Grün and coworkers provided an ESR age determination for a hominin molar from Florisbad, South Africa, at 259 ± 35 ka. The most anatomically informative hominin specimen from Florisbad is a fragmentary craniofacial specimen, attributed by many to early Homo sapiens, which has been assumed to derive from the same individual as this tooth. Since 1996, evidence about the evolutionary context for the later Middle Pleistocene and its associated hominins in Africa had markedly changed. If the Florisbad date is accurate and if the molar is associated with the craniofacial specimen, this evidence may suggest that Homo naledi coexisted in time and geography with H. sapiens in southern Africa. Thus, the accuracy for all dates needs critical investigation. This study examines the published record of excavation and subsequent analyses for the Florisbad material. We find that the contemporary records raise doubt about the contemporaneity of the tooth and calvaria. The site's complex stratigraphy and the details about the tooth's discovery also pose challenges for the previous ESR age estimate. Because the fragmentary cranial specimen has value for morphological comparisons, developing additional means of understanding its geological age is necessary. The current data do not demonstrate the presence of H. sapiens in southern Africa at ~260 ka.

Immature remains and the first partial skeleton of a juvenile Homo naledi, a late Middle Pleistocene hominin from South Africa.

Immature remains are critical for understanding maturational processes in hominin species as well as for interpreting changes in ontogenetic development in hominin evolution. The study of these subjects is hindered by the fact that associated juvenile remains are extremely rare in the hominin fossil record. Here we describe an assemblage of immature remains of Homo naledi recovered from the 2013-2014 excavation season. From this assemblage, we attribute 16 postcranial elements and a partial mandible with some dentition to a single juvenile Homo naledi individual. The find includes postcranial elements never before discovered as immature elements in the sub-equatorial early hominin fossil record, and contributes new data to the field of hominin ontogeny.

Distinct mandibular premolar crown morphology in Homo naledi and its implications for the evolution of Homo species in southern Africa.

Homo naledi displays a combination of features across the skeleton not found in any other hominin taxon, which has hindered attempts to determine its placement within the hominin clade. Using geometric morphometrics, we assess the morphology of the mandibular premolars of the species at the enamel-dentine junction (EDJ). Comparing with specimens of Paranthropus, Australopithecus and Homo (n = 97), we find that the H. naledi premolars from the Dinaledi chamber consistently display a suite of traits (e.g., tall crown, well-developed P3 and P4 metaconid, strongly developed P3 mesial marginal ridge, and a P3 > P4 size relationship) that distinguish them from known hominin groups. Premolars from a second locality, the Lesedi Chamber, are consistent with this morphology. We also find that two specimens from South Africa, SK 96 (usually attributed to Paranthropus) and Stw 80 (Homo sp.), show similarities to the species, and we discuss a potential evolutionary link between H. naledi and hominins from Sterkfontein and Swartkrans.

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.

A case of benign osteogenic tumour in Homo naledi: Evidence for peripheral osteoma in the U.W. 101-1142 mandible.

The reported incidence of neoplasia in the extinct hominin record is rare. We describe here the first palaeopathological analysis of an osteogenic lesion in the extinct hominin Homo naledi from Dinaledi Cave (Rising Star), South Africa. The lesion presented as an irregular bony growth, found on the right lingual surface of the body of the adult mandible U.W. 101-1142. The growth was macroscopically evaluated and internally imaged using micro-focus x-ray computed tomography (µCT). A detailed description and differential diagnosis were undertaken using gross and micromorphology, and we conclude that the most probable diagnosis is peripheral osteoma - a benign osteogenic neoplasia. These tumours are cryptic in clinical expression, though they may present localised discomfort and swelling. It has been suggested that muscle traction may play a role in the development and expression of these tumours. The impact of this lesion on the individual affected is unknown. This study adds to the growing corpus of palaeopathological data from the South African fossil record, which suggests that the incidence of neoplastic disease in deep prehistory was more prevalent than traditionally accepted. The study also highlights the utility of micro-computed tomography in assisting accurate diagnoses of ancient pathologies.