New craniodental fossils of Paranthropus robustus from Kromdraai, South Africa (2014-2017 excavations).

Since the initial discovery of Paranthropus robustus at the site of Kromdraai in 1938, the hypodigm of this species has been expanded by subsequent work at the localities of Swartkrans and Drimolen, with a few fossils also known from Cooper's D, Gondolin and Sterkfontein Member 5. Beginning in 2014, systematic excavations at Kromdraai uncovered a large and previously unknown fossiliferous area, shedding light on Units O and P in the earliest part of the site's stratigraphic sequence. The aim of this paper is to provide detailed descriptions and illustrations of 30 P. robustus craniodental specimens recovered between 2014 and 2017 within the Unit P deposits at Kromdraai. This new sample predates all prior conspecific specimens found at this site (including the holotype of P. robustus from Kromdraai, TM 1517). Its basic dental morphology dimensions and cranial features are compared in a preliminary analysis with other P. robustus samples. The P. robustus sample from Kromdraai Unit P documents previously unknown portions of the P. robustus juvenile cranium. The new dental and cranial remains aid in the exploration of potential morphological distinctions between site-specific P. robustus samples and are compared favorably in size and morphology with the small P. robustus specimens from Drimolen (e.g., DNH 7). These findings do not support the hypothesis that the specimens from Drimolen belong to a different taxonomic group. Instead, they reinforce the presence of a significant degree of sexual dimorphism within P. robustus. The Kromdraai Unit P specimens also contribute to the biodemographic profile of P. robustus. The notable prevalence of infants (i.e., juvenile individuals before the emergence of their first permanent molars) mirrors the natural mortality profiles observed in wild chimpanzees. This suggests a closer resemblance in the processes of accumulation in Kromdraai Unit P and Drimolen than at Swartkrans.

Variation, sexual dimorphism, and enlargement of the frontal sinus with age in adult South Africans.

OBJECTIVES: To document frontal sinus volume (FSV) in a sample of sub-Saharan Africans with a view to evaluating claims that such populations exhibit comparatively small sinuses. This study also addresses questions related to sexual dimorphism, incidence of sinus aplasia, and the possibility that FSV continues to increase through adulthood. MATERIALS AND METHODS: FSV was measured from CT scans of adult crania from the Dart Collection. Sex and age were known for each individual. Linear cranial dimensions were used to compute a geometric mean from which a scaled FSV was computed for each cranium. RESULTS: FSV does not differ significantly between sexes, but females exhibit a higher incidence of aplasia. There is considerable variation in FSV in this sample, with the average ranking among the higher means reported for other population samples. The incidence of FS aplasia falls within the range of values recorded for other population samples. Although our study is cross-sectional rather than longitudinal, there is strong evidence that FSV continues to increase with age throughout adulthood. DISCUSSION: The FSV mean of our sample contradicts the notion that sub-Saharan Africans possess small sinuses. In a global context, geography (climate and altitude) does not appear to be related to FSV. The absence of sexual dimorphism in our sample is unexpected, as significant dimorphism has been reported for most other population samples. Our results support other indications that the frontal sinus continues to expand throughout adulthood, especially in females, and that it is likely due to bone resorption.

Metopism in adult South Africans and its relationship to frontal sinus size.

This study documents the incidences of complete and partial metopism and their possible relationship to frontal sinus volume (FSV) in a sample of modern adult black South Africans with a view to evaluating the hypothesis that metopism affects frontal sinus hypoplasia. FSV was measured from CT scans and the incidence of metopism was recorded from direct observations of dried cadaveric crania. The sex of each individual was known. Four linear cranial dimensions were used to compute a geometric mean by which to scale FSV. The incidence of partial metopism (38%) is comparable to that reported for other population samples, although there is considerable variation among these global sample frequencies. It is significantly more common in male than female South Africans. FSV in individuals with complete metopism is smaller than average but not inordinately so. On the other hand, FSV is significantly larger in individuals with partial metopism than in those that do not present with this sutural remnant. The data on FSV in individuals with and without partial metopism contradict the hypothesis that there is a relationship between partial metopism and frontal sinus hypoplasia. As such, the metopic remnant evinced by the Late Pleistocene cranium from Hofmeyr, South Africa is unlikely to be related to its very small FSV.

Taxonomic attribution of the KNM-ER 1500 partial skeleton from the Burgi Member of the Koobi Fora Formation, Kenya.

Paranthropus boisei is well represented in the eastern African fossil record by craniodental remains, but very few postcranial fossils can be securely attributed to this taxon. For this reason, KNM-ER 1500 from East Turkana, Kenya, is especially important. KNM-ER 1500 is a badly weathered and fragmented postcranial skeleton associated with a small piece of mandibular corpus. It derives from the Burgi Member, which has yielded diagnostic craniodental fossils attributable to P. boisei, Homo habilis, Homo rudolfensis and Homo erectus. Although it has been proposed that KNM-ER 1500 may be attributable to P. boisei based on the small mandibular fragment, this hypothesis remained challenging to test. Here we re-examine the preserved portions of KNM-ER 1500 and reassess support for its taxonomic attribution. There are compelling features of the mandible, proximal femur, and especially the proximal radius that support attribution of KNM-ER 1500 to P. boisei. These features include the absolute width of the mandible and its lack of a lateral intertoral sulcus, an anteroposteriorly compressed femoral neck with a distinctive posteroinferior marginal ridge, the rim of the radial head that is proximodistally uniform in thickness around its circumference, and a long radial neck that is elliptical in cross section. No feature serves to align KNM-ER 1500 with Homo to the exclusion of Paranthropus. KNM-ER 1500 was a small-bodied individual and attributing this specimen to P. boisei confirms that significant postcranial-size dimorphism was present in this species.

Hominin fossils from Kromdraai and Drimolen inform Paranthropus robustus craniofacial ontogeny.

Ontogeny provides critical information about the evolutionary history of early hominin adult morphology. We describe fossils from the southern African sites of Kromdraai and Drimolen that provide insights into early craniofacial development in the Pleistocene robust australopith Paranthropus robustus. We show that while most distinctive robust craniofacial features appear relatively late in ontogeny, a few do not. We also find unexpected evidence of independence in the growth of the premaxillary and maxillary regions. Differential growth results in a proportionately larger and more postero-inferiorly rotated cerebral fossa in P. robustus infants than in the developmentally older Australopithecus africanus juvenile from Taung. The accumulated evidence from these fossils suggests that the iconic SK 54 juvenile calvaria is more likely early Homo than Paranthropus. It is also consistent with the hypothesis that P. robustus is more closely related to Homo than to A. africanus.

Hypercementosis in Late Pleistocene Homo sapiens fossils from Klasies River Main Site, South Africa.

OBJECTIVE: To examine early Homo sapiens fossils from the Late Pleistocene site of Klasies River Main Site, South Africa for evidence of hypercementosis. The specimens represent seven adult individuals dated to between 119,000 and 58,000 years ago. These observations are contextualized in relation to the incidences of hypercementosis among recent human populations and fossil human samples and the potential etiologies of hypercementosis. DESIGN: The fossils were investigated utilizing micro-CT and nano-CT scanning to visualize and measure cementum apposition on permanent incisor, premolar and molar roots. Cementum thickness was measured at mid-root level, and the volume of the cementum sleeve was calculated for the two fossil specimens that display marked hypercementosis. RESULTS: Two of the fossils display no evidence of cementum hypertrophy. Three exhibit moderate cementum thickening, barely attaining the quantitative threshold for hypercementosis. Two evince marked hypercementosis. One of the Klasies specimens with marked hypercementosis is judged to be an older individual with periapical abscessing. The second specimen is a younger adult, and seemingly similar in age to other Klasies fossils that exhibit only minimal cementum apposition. However, this second specimen exhibits dento-alveolar ankylosis of the premolar and molars. CONCLUSIONS: These two fossils from Klasies River Main Site provide the earliest manifestation of hypercementosis in Homo sapiens.

Changing perspectives on early hominin diets.

It is axiomatic that knowledge of the diets of extinct hominin species is central to any understanding of their ecology and our evolution. The importance of diet in the paleontological realm has led to the employment of multiple approaches in its elucidation. Some of these have deep historical roots, while others are dependent upon more recent technical and methodological advances. Historically, studies of tooth size, shape, and structure have been the gold standard for reconstructing diet. They focus on species-level adaptations, and as such, they can set theoretical brackets for dietary capabilities within the context of specific evolutionary moments. Other methods (e.g., analyses of dental calculus, biogeochemistry, and dental microwear) have only been developed within the past few decades, but are now beginning to yield evidence of the actual foods consumed by individuals represented by fossil remains. Here we begin by looking at these more "direct" forms of evidence of diet before showing that, when used in conjunction with other techniques, these "multi-proxy" approaches can raise questions about traditional interpretations of early hominin diets and change the nature of paleobiological interpretations.

An updated analysis of hominin phylogeny with an emphasis on re-evaluating the phylogenetic relationships of Australopithecus sediba.

The discovery and description of Australopithecus sediba has reignited the debate over the evolutionary history of the australopiths and the genus Homo. It has been suggested that A. sediba may be an ancestor of Homo because it possesses a mosaic of derived Homo-like and primitive australopith-like traits. However, an alternative hypothesis proposes that the majority of the purported Homo-like craniodental characters can be attributed to the juvenile status of the type specimen, MH1. We conducted an independent character assessment of the craniodental morphology of A. sediba, with particular emphasis on evaluating whether the ontogenetic status of MH1 may have affected its purported Homo-like characteristics. In doing so, we have also expanded fossil hypodigms to incorporate the new Australopithecus anamensis cranium from Woranso-Mille (MRD-VP-1/1), as well as recently described Paranthropus robustus cranial remains from Drimolen (DNH 7, DNH 155). Morphological character data were analyzed using both standard parsimony and Bayesian techniques. In addition, we conducted a series of Bayesian analyses constrained to evaluate the hypothesis that Australopithecus africanus and A. sediba are sister taxa. Based on the results of the parsimony and Bayesian analyses, we could not reject the hypothesis that A. sediba shares its closest phylogenetic affinities with the genus Homo. Therefore, based on currently available craniodental evidence, we conclude that A. sediba is plausibly the terminal end of a lineage that shared a common ancestor with the earliest representatives of Homo. We caution, however, that the discovery of new A. sediba fossils preserving adult cranial morphology or the inclusion of postcranial characters may ultimately necessitate a re-evaluation of this hypothesis.

The taxonomic attribution of African hominin postcrania from the Miocene through the Pleistocene: Associations and assumptions.

Postcranial bones may provide valuable information about fossil taxa relating to their locomotor habits, manipulative abilities and body sizes. Distinctive features of the postcranial skeleton are sometimes noted in species diagnoses. Although numerous isolated postcranial fossils have become accepted by many workers as belonging to a particular species, it is worthwhile revisiting the evidence for each attribution before including them in comparative samples in relation to the descriptions of new fossils, functional analyses in relation to particular taxa, or in evolutionary contexts. Although some workers eschew the taxonomic attribution of postcranial fossils as being less important (or interesting) than interpreting their functional morphology, it is impossible to consider the evolution of functional anatomy in a taxonomic and phylogenetic vacuum. There are 21 widely recognized hominin taxa that have been described from sites in Africa dated from the Late Miocene to the Middle Pleistocene; postcranial elements have been attributed to 17 of these. The bones that have been thus assigned range from many parts of a skeleton to isolated elements. However, the extent to which postcranial material can be reliably attributed to a specific taxon varies considerably from site to site and species to species, and is often the subject of considerable debate. Here, we review the postcranial remains attributed to African hominin taxa from the Late Miocene to the Middle and Late Pleistocene and place these assignations into categories of reliability. The catalog of attributions presented here may serve as a guide for making taxonomic decisions in the future.

Frontal sinuses and human evolution.

The frontal sinuses are cavities inside the frontal bone located at the junction between the face and the cranial vault and close to the brain. Despite a long history of study, understanding of their origin and variation through evolution is limited. This work compares most hominin species' holotypes and other key individuals with extant hominids. It provides a unique and valuable perspective of the variation in sinuses position, shape, and dimensions based on a simple and reproducible methodology. We also observed a covariation between the size and shape of the sinuses and the underlying frontal lobes in hominin species from at least the appearance of Homo erectus. Our results additionally undermine hypotheses stating that hominin frontal sinuses were directly affected by biomechanical constraints resulting from either chewing or adaptation to climate. Last, we demonstrate their substantial potential for discussions of the evolutionary relationships between hominin species.

A common mechanism drives the alignment between the micro- and macroevolution of primate molars.

A central challenge for biology is to reveal how different levels of biological variation interact and shape diversity. However, recent experimental studies have indicated that prevailing models of evolution cannot readily explain the link between micro- and macroevolution at deep timescales. Here, we suggest that this paradox could be the result of a common mechanism driving a correlated pattern of evolution. We examine the proportionality between genetic variance and patterns of trait evolution in a system whose developmental processes are well understood to gain insight into how such alignment between morphological divergence and genetic variation might be maintained over macroevolutionary time. Primate molars present a model system by which to link developmental processes to evolutionary dynamics because of the biased pattern of variation that results from the developmental architecture regulating their formation. We consider how this biased variation is expressed at the population level, and how it manifests through evolution across primates. There is a strong correspondence between the macroevolutionary rates of primate molar divergence and their genetic variation. This suggests a model of evolution in which selection is closely aligned with the direction of genetic variance, phenotypic variance, and the underlying developmental architecture of anatomical traits.

Hunter-Schreger Band configuration in human molars reveals more decussation in the lateral enamel of 'functional' cusps than 'guiding' cusps.

OBJECTIVES: Enamel prism decussation, which manifests as Hunter-Schreger Bands (HSB), is considered a mechanism to mitigate crack propagation. During the chewing cycle, the 'functional' cusps that are involved in Phase II crushing and grinding experience more complex patterns of stress than do those that 'guide' the molars into occlusion (Phase I). This study examines HSB configuration in the lateral enamel of human molars to identify potential differences between these cusps as predicted from their functional distinctions. DESIGN: Measurements were recorded from scanning electron micrographs of sections through the mesial cusps of unworn permanent molars. For each section, HSB packing density and the relative thickness of decussated enamel were quantified in the cuspal and middle segments of lateral enamel over the guiding and functional cusps. RESULTS: No clear trend from first to third molars in HSB configuration was found in either jaw. In maxillary molars, the functional cusp displays higher HSB packing density in the cuspal and middle segments, and relatively thicker decussated enamel in the cuspal segment than does the guiding cusp. In mandibular molars, the functional cusp displays higher HSB packing density in the middle segment than does the guiding cusp, but no difference in relative thickness was found between them. Enamel of mandibular molars shows weaker decussation than maxillary molars. CONCLUSIONS: The results suggest that guiding cusps are intrinsically more susceptible to crack propagation than functional cusps in human permanent molars. Structural factors such as enamel decussation should be considered when interpreting enamel chipping patterns in dietary contexts.

Variation in Middle Stone Age mandibular molar enamel-dentine junction topography at Klasies River Main Site assessed by diffeomorphic surface matching.

The morphology and variability of the Middle Stone Age (MSA) hominin fossils from Klasies River Main Site have been the focus of investigation for more than four decades. The mandibular remains have figured prominently in discussions relating to robusticity, size dimorphism, and symphyseal morphology. Variation in corpus size between the robust SAM-AP 6223 and the diminutive SAM-AP 6225 mandibles is particularly impressive, and the difference between the buccolingual diameters of their M2s significantly exceeds recent human sample variation. SAM-AP 6223 and SAM-AP 6225 are the only Klasies specimens with homologous teeth (M2 and M3) that permit comparisons of crown morphology. While the differences in dental trait expression at the outer enamel surfaces of these molars are slight, diffeomorphic surface analyses of their underlying enamel-dentine junction (EDJ) topographies reveal differences that are well beyond the means of pairwise differences among comparative samples of Later Stone Age (LSA) Khoesan and recent African homologues. The EDJs of both SAM-AP 6225 molars and the SAM-AP 6223 M3 fall outside the envelopes that define the morphospace of these two samples. Although the radiocarbon dated LSA individuals examined here differ by a maximum of some 7000 years, and the two Klasies jaws may differ by perhaps as much as 18,000 years, it is difficult to ascribe their differences to time alone. With reference to the morphoscopic traits by which the SAM-AP 6223 and SAM-AP 6225 EDJs differ, the most striking is the expression of the protoconid cingulum. This is very weakly developed on the SAM-AP 6223 molars and distinct in SAM-AP 6225. As such, this diminutive fossil exhibits a more pronounced manifestation of what is likely a plesiomorphic feature, thus adding to the morphological mosaicism that is evident in the Klasies hominin assemblage. Several possible explanations for the variation and mosaicism in this MSA sample are discussed.

Isotopic evidence for the timing of the dietary shift toward C4 foods in eastern African Paranthropus.

New approaches to the study of early hominin diets have refreshed interest in how and when our diets diverged from those of other African apes. A trend toward significant consumption of C4 foods in hominins after this divergence has emerged as a landmark event in human evolution, with direct evidence provided by stable carbon isotope studies. In this study, we report on detailed carbon isotopic evidence from the hominin fossil record of the Shungura and Usno Formations, Lower Omo Valley, Ethiopia, which elucidates the patterns of C4 dietary utilization in the robust hominin Paranthropus The results show that the most important shift toward C4 foods occurred at ∼2.37 Ma, within the temporal range of the earliest known member of the genus, Paranthropus aethiopicus, and that this shift was not unique to Paranthropus but occurred in all hominins from this fossil sequence. This uptake of C4 foods by hominins occurred during a period marked by an overall trend toward increased C4 grazing by cooccurring mammalian taxa from the same sequence. However, the timing and geographic patterns of hominin diets in this region differ from those observed elsewhere in the same basin, where environmental controls on the underlying availability of various food sources were likely quite different. These results highlight the complexities of dietary responses by hominins to changes in the availability of food resources.

Human manual distal phalanges from the Middle Stone Age deposits of Klasies River Main Site, Western Cape Province, South Africa.

Two new distal manual phalanges from the Middle Stone Age deposits of Klasies River Main Site are described. One (SAM-AP 6387) likely derives from ray II or ray III, whereas the other (SAM-AP 6388) is from the thumb. Both derive from a late adolescent or fully adult individual. They were recovered by H. Deacon from the same stratigraphic unit (submember W or possibly submember R) of the Shell and Sand Member of Cave 1, which places them between 100 and 90 ka. Both are comparatively small elements, and the possibility that they came from the same hand cannot be discounted at this time. These bones add to the meager and all too fragmentary postcranial human fossil sample from the Late Pleistocene of South Africa. These two specimens provide some additional evidence pertaining to the morphological attributes of the distal phalanges of the Middle Stone Age inhabitants of South Africa. Together with the distal pollical phalanx from Die Kelders (SAM-AP 6402), they are relatively small in comparison with homologs from recent human samples as well as Late Pleistocene specimens from Eurasia. Given their small sizes, the distal pollical phalanges from Klasies and Die Kelders are not dissimilar to Holocene Khoesan homologs. As expected, the Klasies elements differ noticeably from Neandertal homologs, especially in the narrowness of their shafts and distal tuberosities.

Clarifying relationships between cranial form and function in tapirs, with implications for the dietary ecology of early hominins.

Paleontologists and paleoanthropologists have long debated relationships between cranial morphology and diet in a broad diversity of organisms. While the presence of larger temporalis muscle attachment area (via the presence of sagittal crests) in carnivorans is correlated with durophagy (i.e. hard-object feeding), many primates with similar morphologies consume an array of tough and hard foods-complicating dietary inferences of early hominins. We posit that tapirs, large herbivorous mammals showing variable sagittal crest development across species, are ideal models for examining correlations between textural properties of food and sagittal crest morphology. Here, we integrate dietary data, dental microwear texture analysis, and finite element analysis to clarify the functional significance of the sagittal crest in tapirs. Most notably, pronounced sagittal crests are negatively correlated with hard-object feeding in extant, and several extinct, tapirs and can actually increase stress and strain energy. Collectively, these data suggest that musculature associated with pronounced sagittal crests-and accompanied increases in muscle volume-assists with the processing of tough food items in tapirs and may yield similar benefits in other mammals including early hominins.

Secular trends in cranial size and shape among black South Africans over the late 19th and 20th centuries.

BACKGROUND: Previous studies of secular change in cranial size among black South Africans have produced conflicting results. AIM: We re-examined cranial size change in this population during the 19th and 20th century by evaluating its relationship with individual year-of-birth, and the significance of trends among eight decennial cohorts. SUBJECTS AND METHODS: This study is based on 102 male and 89 female adults born between 1865 and 1959. Linear regressions were employed to evaluate possible relationships between year-of-birth and cranial dimensions; ANOVAs were used to evaluate the significance of long-term trends among decennial cohorts. RESULTS: No analysis revealed a secular change in cranial length in either sex; however, the ANOVA for cranial length in the combined sex sample was significant. There is no secular trend in female cranial breadth, but males display a negative trend in this dimension. This results in a secular trend for increased male dolichocephaly. CONCLUSIONS: The factors that underlie the negative secular trend in male cranial breadth and the absence of a secular trend in overall cranial size in this population are unclear. Nevertheless, these observations accord with findings related to stature and long bone strength in this population and are consistent with observations for other sub-Saharan African populations.

Secular decline in limb bone strength among South African Africans during the 19th and 20th centuries.

OBJECTIVES: South African Africans have been reported to have experienced negative or null secular trends in stature and other measures of skeletal structure across the 19th and 20th centuries, presumably due to poor living conditions during a time of intensifying racial discrimination. Here, we investigate whether any secular trend is apparent in limb bone strength during the same period. MATERIALS AND METHODS: Cadaver-derived skeletons (n = 221) were analyzed from female and male South African Africans who were born between 1839 and 1970, lived in and around Johannesburg, and died between 1925 and 1991 when they were 17-90 years of age. For each skeleton, a humerus and femur were scanned using computed tomography, and mid-diaphyseal cross-sectional geometric properties were calculated and scaled according to body size. RESULTS: In general linear mixed models accounting for sex, age at death, and skeletal element, year of birth was a significant (p < .05) negative predictor of size-standardized mid-diaphyseal cortical area (a proxy for resistance to axial loading) and polar moment of area (a proxy for resistance to bending and torsion), indicating a temporal trend toward diminishing limb bone strength. No significant interactions were detected between year of birth and age at death, suggesting that the decline in limb bone strength was mainly due to changes in skeletal maturation rather than severity of age-related bone loss. DISCUSSION: Limb bone strength is thus potentially another feature of the skeletal biology of South African Africans that was compromised by poor living conditions during the 19th and 20th centuries.

An examination of the cross-sectional geometrical properties of the long bone diaphyses of Holocene foragers from Roonka, South Australia.

OBJECTIVES: This study examines long bone diaphyseal rigidity and shape of hunter-gatherers at Roonka to make inferences about subsistence strategies and mobility of inhabitants of semi-arid southeastern Australia. Roonka is a cemetery site adjacent to the Lower Murray River, which contains over 200 individuals buried throughout the Holocene. Archaeological evidence indicates that populations living near this river corridor employed mobile, risk averse foraging strategies. METHODS: This prediction of lifestyle was tested by comparing the cross-sectional geometric properties of the humerus, radius, ulna, femur, tibia, and fibula of individuals from Roonka to samples of varying subsistence strategies. Bilateral asymmetry of the upper limb bones was also examined. RESULTS: Roonka males and females have moderately high lower limb diaphyseal rigidity and shape. In the upper limb, females have low rigidity and bilateral asymmetry while males have moderately high rigidity and bilateral asymmetry. This pattern is similar to other foraging groups from Australia and southern Africa that have behaviorally adapted to arid and semi-arid environments. DISCUSSION: Lower limb results suggest that populations in the Lower Murray River Valley had relatively elevated foraging mobility. Upper limb rigidity and bilateral asymmetry indicate a sexual division of labor at Roonka. Females resemble other samples that had mixed subsistence strategies that involved hunting, gathering, and processing tasks. Males display a pattern similar to groups that preferentially hunted large game, but that supplemented this source with smaller game and riverine resources.