Paranthropus Research Articles

Mobility and subsistence at the Willandra Lakes: A comparative analysis of femoral cross-sectional properties in the Lake Mungo 3 skeleton

Previous research suggests that some hominin postcranial features do not follow a linear path of increasing modernization through geological time. With respect to the distal humerus, in particular, the earliest known hominin specimens are reportedly among the most modern in morphology, while some later humeri appear further removed from the average modern human shape. Although Plio-Pleistocene humeri vary widely in size, previous studies have failed to account for size-related shape variation when making morphometric comparisons. This study reexamines hominin postcranial evolution in light of distal humeral allometry. Using two-dimensional landmark data, the relationship between specimen size and shape among modern humans is quantified using multivariate regression and principal components analysis of size-shape space. Fossils are compared with modern human shapes expected at a given size, as well as with the overall average human shape. The null hypothesis of humeral isometry in modern humans is rejected. Subsequently, if one takes allometry into account, the apparent pattern of hominin humeral evolution does not resemble the pattern described above. All 14 of the Plio-Pleistocene hominin fossils examined here share a similar pattern of shape differences from equivalently-sized modern humans, though they vary in the extent to which these differences are expressed. The oldest specimen in the sample (KNM-KP 271; Australopithecus anamensis) exhibits the least human-like elbow morphology. Similarly primitive morphology characterizes all younger species of Australopithecus as well as Paranthropus robustus. After 2 Ma, a subtly more human-like elbow morphology is apparent among specimens attributed to early Homo, as well as among isolated specimens that may represent either Homo or Paranthropus boisei. This study emphasizes the need to consider size-related shape variation when individual fossil specimens are compared with the average shape of a comparative group, particularly when specimens fall near an extreme of the comparative size distribution.

Mandibular development in Australopithecus robustus

Australopithecus robustus has a distinct mandibular anatomy, with a broad and deep corpus and a tall, relatively upright ramus. How this anatomy arose through development is unknown, as gross mandibular size and shape change have not been thoroughly examined quantitatively in this species. Herein, I investigate A. robustus mandibular growth by comparing its ontogenetic series with a sample of recent humans, examining age-related size variation in 28 linear measurements. Resampling is used to compare the amount of proportional size change occurring between tooth eruption stages in the small and fragmentary A. robustus sample, with that of a more complete human skeletal population. Ontogenetic allometry of corpus robusticity is also assessed with least squares regression. Results show that nearly all measurements experience greater average increase in A. robustus than in humans. Most notably, A. robustus corpus breadth undergoes a spurt of growth before eruption of M1 , likely due in part to delayed resorption of the ramus root on the lateral corpus. Between the occlusion of M1 and M2 , nearly all dimensions experience greater proportional size change in A. robustus. Nested resampling analysis affirms that this pattern of growth differences between species is biologically significant, and not a mere byproduct of the fossil sample size. Some species differences are likely a function of postcanine megadontia in A. robustus, although the causes of other differences are less clear. This study demonstrates an important role of the postnatal period for mandibular shape development in this species.

A 3-dimensional assessment of molar enamel thickness and distribution pattern in Gigantopithecus blacki

In order to investigate molar enamel thickness and distribution in Gigantopithecus blacki, two maxillary and three mandibular molars were micro-CT scanned, and the 3-dimensional (3D) shape of the enamel crown cap was digitally reconstructed. Results show that G. blacki molars have very thick enamel, over 6 mm in places. 3D average thickness of the entire crown relative to crown size was shown to be comparable to that of modern Homo sapiens, and thinner than in the very thick enameled robust Australopithecus condition. Durability of the tooth, calculated as the enamel volume relative to maximum horizontal area of the crown, was shown to be comparable to that of fossil hominids, including Australopithecus robustus. The thick occlusal enamel of G. blacki molars was found to be accompanied by a buccolingual gradient in which thicker enamel occurs on the cusps of the “functional” side. This is associated with higher crown height on the “functional” side of the crown, and implies an adaptation to differential buccolingual wear. These results indicate that G. blacki molars were adapted to heavy wear, but probably in a manner that was different from Pliocene and Pleistocene hominids.

Termites in the hominin diet: A meta-analysis of termite genera, species and castes as a dietary supplement for South African robust australopithecines

Termite foraging by chimpanzees and present-day modern humans is a well-documented phenomenon, making it a plausible hypothesis that early hominins were also utilizing this resource. Hominin termite foraging has been credited by some to be the explanation for the unexpected carbon isotope signatures present in South African hominin teeth, which suggest the diet was different from that of extant non-human great apes, consisting of a significant amount of resources that are not from woody-plants. Grass-eating termites are one potential resource that could contribute to the carbon signature. However, not all termites eat grasses, and in fact, the termites that are most widely consumed by chimpanzees and by many present-day human populations at best have a mixed diet that includes small amounts of grasses.Here I review the ecology of termites and how it affects their desirability as a food resource for hominins, and conduct a meta-analysis of nutritional values for various genera, species and castes from the literature. Termites are very diverse, even within species, and this variability affects both their carbon signatures and nutritional value, hindering generalizations regarding the contribution of termites to the hominin diet. It is concluded here that a combination of soldiers and alates of the genus Macrotermes be used to model the insectivory component of the Plio-Pleistocene hominin diet due to their significant amounts of energy-yielding nutrients and potential role as a critical resource for supporting larger-brained hominins.

Geometric properties and comparative biomechanics of Homo floresiensis mandibles

The hypodigm of Homo floresiensis from the cave of Liang Bua on Flores Island in the archipelago of Indonesia includes two mandibles (LB1/2 and LB6/1). The morphology of their symphyses and corpora has been described as sharing similarities with both australopiths and early Homo despite their Late Pleistocene age. Although detailed morphological comparisons of these mandibles with those of modern and fossil hominin taxa have been made, a functional analysis in the context of masticatory biomechanics has yet to be performed.Utilizing data on cortical bone geometry from computed tomography scans, we compare the mechanical attributes of the LB1 and LB6 mandibles with samples of modern Homo, Pan, Pongo, and Gorilla, as well as fossil samples of Paranthropus robustus, Australopithecus africanus and South African early Homo. Structural stiffness measures were derived from the geometric data to provide relative measures of mandibular corpus strength under hypothesized masticatory loading regimes. These mechanical variables were evaluated relative to bone area, mandibular length and estimates of body size to assess their functional affinities and to test the hypothesis that the Liang Bua mandibles can be described as scaled-down variants of either early hominins or modern humans.Relative to modern hominoids, the H. floresiensis material appears to be relatively strong in terms of rigidity in torsion and transverse bending, but is relatively weak under parasagittal bending. Thus, they are ‘robust’ relative to modern humans (and comparable with australopiths) under some loads but not others. Neither LB1 nor LB6 can be described simply as ‘miniaturized’ versions of modern human jaws since mandible length is more or less equivalent in Homo sapiens and H. floresiensis. The mechanical attributes of the Liang Bua mandibles are consistent with previous inferences that masticatory loads were reduced relative to australopiths but remained elevated relative to modern Homo.

News and views: Response to ‘Non-metric dental traits and hominin phylogeny’ by Carter et al., with additional information on the Arizona State University Dental Anthropology System and phylogenetic ‘place’ of Australopithecus sediba

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.

Regional diversity patterns in African bovids, hyaenids, and felids during the past 3 million years: the role of taphonomic bias and implications for the evolution of Paranthropus

Reconstructing patterns of Plio-Pleistocene mammalian faunal exchange between eastern and southern Africa may help us to better understand patterns of evolution within the hominin clade. However, differences in geological context, taphonomic history and collection methods, as well as a lack of a precise chronology in one of the regions, complicate attempts to compare the faunas of the two regions, but access to new comprehensive datasets encouraged us to re-examine this critical time period in the African paleontological record. In this study, we examine the biogeographic histories of three terrestrial African mammalian families whose fossil records span the past 3 million years to test hypotheses related to the evolutionary history of the hominin genus Paranthropus. We used presence/absence data for 117 species from 38 genera within the family Bovidae and 34 species from 15 genera within the families Hyaenidae and Felidae from 52 eastern African and 40 southern African fossil localities. These assemblages were placed into 500 ka time slices and compared at both the genus and species level using the Jaccard index of faunal similarity. Our findings indicate that sampling biases have more effect on the patterns of interchange between eastern and southern African Bovidae than they do the patterns of interchange seen in the Hyaenidae and Felidae. However, even when these biases are taken into account there are persistent differences in the degree of interchange within and between these families. These findings suggest that mammalian groups (including hominins) can have very different histories of exchange between eastern and southern Africa over the past 3 million years.There is no a priori reason why any of the three families we examined is a suitable proxy for the eastern and southern African hyper-megadont and megadont hominin taxa presently included in Paranthropus, but of the three we suggest that the Bovidae is likely to come closest to being an appropriate proxy. The results of this comparative study are consistent with relatively independent evolutionary trajectories in the two regions for the hominins that are presently included in the genus Paranthropus.

A new partial temporal bone of a juvenile hominin from the site of Kromdraai B (South Africa)

The site of Kromdraai B (KB) (Gauteng, South Africa) has yielded a minimum number of nine hominins including the type specimen of Paranthropus robustus (TM 1517), the only partial skeleton of this species known to date. Four of these individuals are juveniles, one is a subadult and four are young adults. They all occur with a macrofaunal assemblage spread across the succession of at least two time periods that occurred in South Africa approximately two million years ago. Here we report on an additional, newly discovered petrous temporal bone of a juvenile hominin, KB 6067. Following the description of KB 6067, we assess its affinities with Australopithecus africanus, P. robustus and early Homo. We discuss its developmental age and consider its association with other juvenile hominin specimens found at Kromdraai B. KB 6067 probably did not reach five years of age and in bony labyrinth morphology it is close to P. robustus, but also to StW 53, a specimen with uncertain affinities. However, its cochlear and oval window size are closer to some hominin specimens from Sterkfontein Member 4 and if KB 6067 is indeed P. robustus this may represent a condition that is evolutionarily less derived than that shown by TM 1517 and other conspecifics sampled so far. The ongoing fieldwork at KB, as well as the petrography and geochemistry of its deposits, will help to determine when the various KB breccias accumulated, and how time may be an important factor underlying the variation seen among KB 6067 and the rest of the fossil hominin sample from this site.

Cercopithecoid primate postcranial fossils from Cooper's D, South Africa

Among several highly fossiliferous localities in the Bloubank Valley (Gauteng, South Africa), the Cooper's Cave System has been known since 1938 and has produced a rich fossil assemblage, including some remains of the early hominin Paranthropus robustus. In 2001, excavations began at a new locality, Cooper's D, which dates to ∼1.4-1.5Ma. Although hominins are relatively rare in the assemblage, remains of cercopithecoid primates are much more common. Craniodental fossils currently indicate the possible presence of at least three large-bodied cercopithecoid primate genera at Cooper's D: Gorgopithecus, Papio, and Theropithecus. In this study, we identify and describe >100 cercopithecoid primate postcranial fossils representing all regions of the appendicular skeleton. The specimens come from several age classes and size morphs; more than one third of the fossils described are from sub-adult and juvenile individuals. The adult postcranial fossils vary substantially in size, with body masses estimated between 30 and 60kg (from 16 of the better preserved specimens). The functional morphology of the postcranial remains indicate that these elements come from animals that likely utilized terrestrial substrates, but they remain difficult to definitively attribute to Gorgopithecus, Theropithecus, or Papio given the absence of associated skeletons. The smaller specimens likely belong to Papio while the larger ones can be attributed to the other two genera. Because Cooper's D has also yielded fossils of the early hominin Paranthropus robustus, this raises the question of how these four large-bodied, mostly terrestrial primates sympatrically utilized the landscape.

Viewpoints: Feeding mechanics, diet, and dietary adaptations in early hominins

Inference of feeding adaptation in extinct species is challenging, and reconstructions of the paleobiology of our ancestors have utilized an array of analytical approaches. Comparative anatomy and finite element analysis assist in bracketing the range of capabilities in taxa, while microwear and isotopic analyses give glimpses of individual behavior in the past. These myriad approaches have limitations, but each contributes incrementally toward the recognition of adaptation in the hominin fossil record. Microwear and stable isotope analysis together suggest that australopiths are not united by a single, increasingly specialized dietary adaptation. Their traditional (i.e., morphological) characterization as "nutcrackers" may only apply to a single taxon, Paranthropus robustus. These inferences can be rejected if interpretation of microwear and isotopic data can be shown to be misguided or altogether erroneous. Alternatively, if these sources of inference are valid, it merely indicates that there are phylogenetic and developmental constraints on morphology. Inherently, finite element analysis is limited in its ability to identify adaptation in paleobiological contexts. Its application to the hominin fossil record to date demonstrates only that under similar loading conditions, the form of the stress field in the australopith facial skeleton differs from that in living primates. This observation, by itself, does not reveal feeding adaptation. Ontogenetic studies indicate that functional and evolutionary adaptation need not be conceptually isolated phenomena. Such a perspective helps to inject consideration of mechanobiological principles of bone formation into paleontological inferences. Finite element analysis must employ such principles to become an effective research tool in this context.

Local ecological profile for Paranthropus robustus in South Africa using stable carbon isotopes from associated bovid teeth

The biomechanical characterization of lower limb long bones in the chrono-ecogeographically diverse species Homo erectus is a fundamental step for assessing evolutionary changes in locomotor mode and body shape that occurred within the genus Homo. However, the samples available for the Early and earlier Middle Pleistocene are small and widely scattered in time and space, thus limiting our understanding of the nature and polarity of morphological trends. Compared to the African fossil record, loading histories based on detailed biomechanical assessment of diaphyseal strength in Indonesian H. erectus lower limb long bones have not been assessed. By using a microtomographic record (μCT), we performed a quantitative analysis of the biomechanical properties and structural organization of Kresna 11, a late Early Pleistocene adult H. erectus femoral shaft from the Sangiran Dome, Central Java. Relative to the modern human condition, Kresna 11 shows the predominant mediolateral cortical thickening (hypertrophy) and the distal displacement of the minimum diaphyseal breadth characteristic of early Homo femora, associated nonetheless with relatively modest cortical thickness within the mid-proximal portion. Synthetic functional imaging of the shaft through the planar representation of its inner structure has revealed distal thickening of the medial cortex, a feature previously unreported in H. erectus. The increase in relative mediolateral bending strength observed in Kresna 11 supports the hypothesis that, rather than simply reflecting differences in patterns of locomotor loading, biomechanical properties of the femoral shaft in archaic Homo are strongly influenced by body shape, i.e., variations in pelvic breadth and femoral neck length.

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

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

Evidence for dietary change but not landscape use in South African early hominins

The dichotomy between early Homo and Paranthropus is justified partly on morphology. In terms of diet, it has been suggested that early Homo was a generalist but that Paranthropus was a specialist. However, this model is challenged and the issue of the resources used by Australopithecus, the presumed common ancestor, is still unclear. Laser ablation profiles of strontium/calcium, barium/calcium and strontium isotope ratios in tooth enamel are a means to decipher intra-individual diet and habitat changes. Here we show that the home range area was of similar size for species of the three hominin genera but that the dietary breadth was much higher in Australopithecus africanus than in Paranthropus robustus and early Homo. We also confirm that P. robustus relied more on plant-based foodstuffs than early Homo. A South African scenario is emerging in which the broad ecological niche of Australopithecus became split, and was then occupied by Paranthropus and early Homo, both consuming a lower diversity of foods than Australopithecus.

The mid-face of lower Pleistocene hominins and its bearing on the attribution of SK 847 and StW 53

SK 847 and StW 53 have often been cited as evidence for early Homo in South Africa. To examine whether midfacial morphology is in agreement with these attributions, we analyze Euclidean distances calculated from 3-D coordinates on the maxillae of SK 847 and StW 53, as well as Australopithecus africanus (Sts 5, Sts 71), Paranthropus robustus (SK 46, SK 48, SK 52, SK 83), early Homo (KNM-ER 1813, KNM-ER 1805, KNM-ER 3733, KNM-WT 15000), P. boisei (KNM-ER 406, KNM-WT 17000, KNM-WT 17400), Gorilla gorilla (n=116), Homo sapiens (n=342), Pan paniscus (n=21) and P. troglodytes (n=65). Multivariate analyses separate extant hominoids suggesting we have captured taxonomic affinity. With the exception of SK 847 and SK 52, South African fossils tend to cluster together. P. robustus differs substantially from East African robust megadonts. SK 847 and StW 53 resemble the East African Homo specimens that are the most australopith-like, such as KNM-WT 15000 and KNM-ER 1813. The resemblance between StW 53 and Homo is driven partly by similarities in maxillary size. When distances are scaled, StW 53 aligns with A. africanus, while SK 847 clusters primarily with early Homo.

Dental microwear and stable isotopes inform the paleoecology of extinct hominins

Determining the diet of an extinct species is paramount in any attempt to reconstruct its paleoecology. Because the distribution and mechanical properties of food items may impact postcranial, cranial, mandibular, and dental morphologies related to their procurement, ingestion, and mastication, these anatomical attributes have been studied intensively. However, while mechanical environments influence skeletal and dental features, it is not clear to what extent they dictate particular morphologies. Although biomechanical explanations have been widely applied to extinct hominins in attempts to retrodict dietary proclivities, morphology may say as much about what they were capable of eating, and perhaps more about phylogenetic history, than about the nature of the diet. Anatomical attributes may establish boundary limits, but direct evidence left by the foods that were actually (rather than hypothetically) consumed is required to reconstruct diet. Dental microwear and the stable light isotope chemistry of tooth enamel provide such evidence, and are especially powerful when used in tandem. We review the foundations for microwear and biogeochemistry in diet reconstruction, and discuss this evidence for six early hominin species (Ardipithecus ramidus, Australopithecus anamensis, Au. afarensis, Au. africanus, Paranthropus robustus, and P. boisei). The dietary signals derived from microwear and isotope chemistry are sometimes at odds with inferences from biomechanical approaches, a potentially disquieting conundrum that is particularly evident for several species.

New hominid fossils from Member 1 of the Swartkrans formation, South Africa

Member 1 of the Swartkrans Formation is comprised of two sedimentary infills, the Lower Bank (LB) and the Hanging Remnant (HR). Together, the LB and HR preserve fossils of early Homo and Paranthropus robustus, Earlier Stone Age lithic artifacts, purported bone digging tools and butchered animal bones. Collectively, this evidence was the first to establish the co-existence of two early Pleistocene hominid species and also led to inferences of plant root harvesting and meat-eating by one or both of those species. P. robustus is the more abundant of the two hominids at Swartrkrans, represented in Member 1 by hundreds of fossils that derive from at least 99 individuals. Thus, Swartkrans Member 1 stands as the world's single largest repository of that extinct species. Here we add to the Member 1 sample of hominid fossils with descriptions of 14 newly discovered specimens.

Three‐dimensional evaluation of root canal morphology in lower second premolars of early and middle pleistocene human populations from atapuerca (Burgos, Spain)

The aim of this study is to describe the morphology of the roots and root canals of permanent lower second premolars (LP4s) with fully developed roots of five hominin groups: Homo sp. (ATE9-1 specimen) from Atapuerca-Sima del Elefante locality, H. antecessor (ATD6-4 and ATD6-125) from Atapuerca-Gran Dolina TD6 locality, H. heidelbergensis from Atapuerca-Sima de los Huesos locality, H. neanderthalensis from Krapina, Regourdou, and Abri Bourgeois-Delaunay localities, and two contemporary H. sapiens groups. The teeth were scanned by means of microtomography. The roots were divided into three virtual segments by three planes: cemento-enamel junction (CEJ), mid-root (MR), and mid-apex (MA). Volumetric and planar direct measurements of the whole teeth and each segment were taken. Descriptive statistical analyses and nonparametric Mann-Whiney test were performed to test for significant differences (P < 0.025) between groups. ATE9-1 and Gran Dolina-TD6 fossils present intricate radicular complexes that might be transitional between the morphologies of Australopithecus robustus and African early Homo and the derived conditions typically found in later Homo. In H. neanderthalensis and H. heidelbergensis, the root canals are wide, with small apical convergence. This trait is particularly pronounced in the Sima de los Huesos sample which may reflect a particularity of this population. Our study demonstrates the potential of hominin roots and root canals as untapped sources of taxonomic information when the tooth crown is fragmented. Future studies, including more fossil specimens and species will shed light in the polarity of the morphologies observed.

New fossil evidence and diet analysis of Gigantopithecus blacki and its distribution and extinction in South China

The present paper reports the recently discovered fossil teeth of Early Pleistocene Gigantopithecus blacki and associated mammalian fauna from Baeryan Cave, Bijie County in Guizhou Province, and also reviews briefly the known fossil sites of Gigantopithecus in south China of Pleistocene. In Early Pleistocene, Gigantopithecus had a wider distribution, but withdrew southward in Middle Pleistocene to a limited area mainly in South China, and it disappeared in the Late Pleistocene according to the present fossil records. Diet and habitat analysis from carbon isotope evidence is used to investigate the reasons for the extinction of Gigantopithecus, which fed on a pure C3 diet and lived in a forest habitat. It was clearly different from early hominins in South and East Africa, such as Australopithecus africanus, Paranthropus robustus and Paranthropus boisei, which had C4 diets. The extinction of Gigantopithecus was also related to the great changes of climate and environment of the Pleistocene, especially the last one million years, during which Homo became more and more prosperous and exerted great pressure on G. blacki.

Hard-Object Feeding in Sooty Mangabeys (Cercocebus atys) and Interpretation of Early Hominin Feeding Ecology

Morphology of the dentofacial complex of early hominins has figured prominently in the inference of their dietary adaptations. Recent theoretical analysis of craniofacial morphology of Australopithecus africanus proposes that skull form in this taxon represents adaptation to feeding on large, hard objects. A modern analog for this specific dietary specialization is provided by the West African sooty mangabey, Cercocebus atys. This species habitually feeds on the large, exceptionally hard nuts of Sacoglottis gabonensis, stereotypically crushing the seed casings using their premolars and molars. This type of behavior has been inferred for A. africanus based on mathematical stress analysis and aspects of dental wear and morphology. While postcanine megadontia, premolar enlargement and thick molar enamel characterize both A. africanus and C. atys, these features are not universally associated with durophagy among living anthropoids. Occlusal microwear analysis reveals complex microwear textures in C. atys unlike those observed in A. africanus, but more closely resembling textures observed in Paranthropus robustus. Since sooty mangabeys process hard objects in a manner similar to that proposed for A. africanus, yet do so without the craniofacial buttressing characteristic of this hominin, it follows that derived features of the australopith skull are sufficient but not necessary for the consumption of large, hard objects. The adaptive significance of australopith craniofacial morphology may instead be related to the toughness, rather than the hardness, of ingested foods.

No brain expansion in Australopithecus boisei

The endocranial volumes of robust australopithecine fossils appear to have increased in size over time. Most evidence with temporal resolution is concentrated in East African Australopithecus boisei. Including the KNM-WT 17000 cranium, this sample comprises 11 endocranial volume estimates ranging in date from 2.5 million to 1.4 million years ago. But the sample presents several difficulties to a test of trend, including substantial estimation error for some specimens and an unusually low variance. This study reevaluates the evidence, using randomization methods and a related test using an explicit model of variability. None of these tests applied to the A. boisei endocranial volume sample produces significant evidence for a trend in that species, whether or not the early KNM-WT 17000 specimen is included.