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In apes, the mandibular third premolar (P3) is adapted for a role in honing the large upper canine. The role of honing was lost early in hominin evolution, releasing the tooth from this functional constraint and allowing it to respond to subsequent changes in masticatory demands. This led to substantial morphological changes, and as such the P3 has featured prominently in systematic analyses of the hominin clade. The application of microtomography has also demonstrated that examination of the enamel-dentine junction (EDJ) increases the taxonomic value of variations in crown morphology. Here we use geometric morphometric techniques to analyze the shape of the P3 EDJ in a broad sample of fossil hominins, modern humans, and extant apes (n = 111). We test the utility of P3 EDJ shape for distinguishing among hominoids, address the affinities of a number of hominin specimens of uncertain taxonomic attribution, and characterize the changes in P3 EDJ morphology across our sample, with particular reference to features relating to canine honing and premolar ‘molarization’. We find that the morphology of the P3 EDJ is useful in taxonomic identification of individual specimens, with a classification accuracy of up to 88%. The P3 EDJ of canine-honing apes displays a tall protoconid, little metaconid development, and an asymmetrical crown shape. Plio-Pleistocene hominin taxa display derived masticatory adaptations at the EDJ, such as the molarized premolars of Australopithecus africanus and Paranthropus, which have well-developed marginal ridges, an enlarged talonid, and a large metaconid. Modern humans and Neanderthals display a tall dentine body and reduced metaconid development, a morphology shared with premolars from Mauer and the Cave of Hearths. Homo naledi displays a P3 EDJ morphology that is unique among our sample; it is quite unlike Middle Pleistocene and recent Homo samples and most closely resembles Australopithecus, Paranthropus and early Homo specimens.

The taxonomy, phylogeny, and biology of theParanthropusspecies have been the center of debates since the earliest discoveryof the TM 1517 cranium (now attributed toParanthropus robustus)from the Plio-Pleistocene site of Kromdraai (South Africa) in 1938by R. Broom (Broom, 1938). In particular, whetherParanthropusboiseiandP. robustusrepresent two distinct taxa, and whether thetwo species emerged from a common ancestor (i.e., the hypothesisof a monophyletic group) remain largely unresolved (reviewed inConstantino and Wood, 2004,2007;Wood and Schroer, 2017).Besides the taxonomic and phylogenetic aspects, the presence oftwo contemporaneous‘robust’species in the eastern and southernAfrican hominin fossil records raises critical questions regardingpotential occurrences of homoplasies in the hominin clade, withsubstantial implications for our understanding of early hominin paleobiology (Wood and Schroer, 2017). Moreover, the absence ofassociated cranial and postcranial remains attributed toPara-nthropuscomplicate the reconstruction of the behavior of thisenigmatic‘megadont’hominin genus (reviewed in Constantino and Wood, 2007).

The Early Pleistocene site of Swartkrans in South Africa's Cradle of Humankind World Heritage Site has been significant for our understanding of the evolution of both early Homo and Paranthropus, as well as the earliest archaeology of southern Africa. Previous attempts to improve a faunal age estimate of the earliest deposit, Member 1, had produced results obtained with uranium-lead dating (U-Pb) on flowstones and cosmogenic burial dating of quartz, which placed the entire member in the range of1.7/1.8 Ma and2.3 Ma. In 2014, two simple burial dates for the Lower Bank, the earliest unit within Member 1, narrowed its age to between ca. 1.8 Ma and 2.2 Ma. A new dating program using the isochron method for burial dating has established an absolute age of 2.22 ± 0.09 Ma for a large portion of the Lower Bank, which can now be identified as containing the earliest Oldowan stone tools and fossils of Paranthropus robustus in South Africa. This date agrees within one sigma with the U-Pb age of 2.25 ± 0.08 Ma previously published for the flowstone underlying the Lower Bank and confirms a relatively rapid rate of accumulation for a large portion of the talus.

International audience; Reconstruction of the locomotor repertoire of the australopiths (Australopithecus and Paranthropus) has progressively integrated information from the mechanosensitive internal structure of the appendicular skeleton. Recent investigations showed that the arrangement of the trabecular network at the femoral head center is biomechanically compatible with the pattern of cortical bone distribution across the neck, both suggesting a full commitment to bipedalism in australopiths, but associated with a slightly altered gait kinematics compared to Homo involving more lateral deviation of the body center of mass over the stance limb. To provide a global picture in Paranthropus robustus of the trabecular architecture of the proximal femur across the head, neck and greater trochanter compartments, we applied techniques of virtual imaging to the variably preserved Early Pleistocene specimens SK 82, SK 97, SK 3121, SKW 19 and SWT1/LB-2 from the cave site of Swartkrans, South Africa. We also assessed the coherence between the structural signals from the center of the head and those from the trabecular network of the inferolateral portion of the head and the inferior margin of the neck, sampling the so-called vertical bundle, which in humans represents the principal compressive system of the joint. Our analyses show a functionally related trabecular organization in Pa. robustus that closely resembles the extant human condition, but which also includes some specificities in local textural arrangement. The network of the inferolateral portion of the head shows a humanlike degree of anisotropy and a bone volume fraction intermediate between the extant human and the African ape patterns. These results suggest slight differences in gait kinematics between Pa. robustus and extant humans. The neck portion of the vertical bundle revealed a less biomechanically sensitive signal. Future investigations on the australopith hip joint loading environment should more carefully investigate the trabecular structure of the trochanteric region and possible structural covariation between cortical bone distribution across the neck and site-specific trabecular properties of the arcuate bundle.

This study assessed variation in the supraorbital and orbital region of the Middle Pleistocene hominins (MPHs), sometimes called Homo heidelbergensis s.l., to test whether it matched the expectations of intraspecific variation. The morphological distinctiveness and relative variation of this region, which is relatively well represented in the hominin fossil record, was analyzed quantitatively in a comparative taxonomic framework. Coordinates of 230 3D landmarks (20) and sliding semilandmarks (210) were collected from 704 specimens from species of Homo, Australopithecus, Paranthropus, Gorilla, Pan, Papio, and Macaca. Results showed that the MPHs had expected levels of morphological distinctiveness and intragroup and intergroup variation in supraorbital and orbital morphology, relative to commonly recognized non-hominin catarrhine species. However, the Procrustes distances between this group and H. sapiens were significantly higher than expected for two closely related catarrhine species. Furthermore, this study showed that variation within the MPH could be similarly well contained within existing hypodigms of H. sapiens, H. neanderthalensis, and H. erectus s.l. Although quantitative assessment of supraorbital and orbital morphology did not allow differentiation between taxonomic hypotheses in later Homo, it could be used to test individual taxonomic affiliation and identify potentially anomalous individuals. This study confirmed a complicated pattern of supraorbital and orbital morphology in the MPH fossil record and raises further questions over our understanding of the speciation of H. sapiens and H. neanderthalensis and taxonomic diversity in later Homo.

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

International audience; Because of its exceptional degree of preservation and its geological age of ∼3.67 Ma, StW 573 makes an invaluable contribution to our understanding of early hominin evolution and paleobiology. The morphology of the bony labyrinth has the potential to provide information about extinct primate taxonomic diversity, phylogenetic relationships and locomotor behaviour. In this context, we virtually reconstruct and comparatively assess the bony labyrinth morphology in StW 573. As comparative material, we investigate 17 southern African hominin specimens from Sterkfontein, Swartkrans and Makapansgat (plus published data from two specimens from Kromdraai B), attributed to Australopithecus, early Homo or Paranthropus, as well as 10 extant human and 10 extant chimpanzee specimens. We apply a landmark-based geometric morphometric method for quantitatively assessing labyrinthine morphology. Morphology of the inner ear in StW 573 most closely resembles that of another Australopithecus individual from Sterkfontein, StW 578, recovered from the Jacovec Cavern. Within the limits of our sample, we observe a certain degree of morphological variation in the Australopithecus assemblage of Sterkfontein Member 4. Cochlear morphology in StW 573 is similar to that of other Australopithecus as well as to Paranthropus specimens included in this study, but it is substantially different from early Homo. Interestingly, the configuration of semicircular canals in Paranthropus specimens from Swartkrans differs from other fossil hominins, including StW 573. Given the role of the cochlea in the sensory-driven interactions with the surrounding environment, our results offer new perspectives for interpreting early hominin behaviour and ecology. Finally, our study provides additional evidence for discussing the phylogenetic polarity of labyrinthine traits in southern African hominins.

A ∼1.52 Ma adult upper limb skeleton of Paranthropus boisei (KNM-ER 47000) recovered from the Koobi Fora Formation, Kenya (FwJj14E, Area 1A) includes most of the distal half of a right humerus (designated KNM-ER 47000B). Natural transverse fractures through the diaphysis of KNM-ER 470000B provide unobstructed views of cortical bone at two sections typically used for analyzing cross-sectional properties of hominids (i.e., 35% and 50% of humerus length from the distal end). Here we assess cross-sectional properties of KNM-ER 47000B and two other P. boisei humeri (OH 80-10, KNM-ER 739). Cross-sectional properties for P. boisei associated with bending/torsional strength (section moduli) and relative cortical thickness (%CA; percent cortical area) are compared to those reported for nonhuman hominids, AL 288-1 (Australopithecus afarensis), and multiple species of fossil and modern Homo. Polar section moduli (Zp) are assessed relative to a mechanically relevant measure of body size (i.e., the product of mass [M] and humerus length [HL]). At both diaphyseal sections, P. boisei exhibits %CA that is high among extant hominids (both human and nonhuman) and similar to that observed among specimens of Pleistocene Homo. High values for Zp relative to size (M × HL) indicate that P. boisei had humeral bending strength greater than that of modern humans and Neanderthals and similar to that of great apes, A. afarensis, and Homo habilis. Such high humeral strength is consistent with other skeletal features of P. boisei (reviewed here) that suggest routine use of powerful upper limbs for arboreal climbing.

Australopithecus anamensis is a pivotal species in human evolution. It is likely to be the direct ancestor of Australopithecus afarensis and the species that may have given rise to the Homo and Paranthropus lineages. It had a suite of adaptations for habitual bipedalism and a diet that differed from that of earlier hominin species. Under what environmental and ecological conditions did this suite of adaptations arise? The early Pliocene site of Kanapoi in the Lake Turkana Basin of Kenya has the largest sample of A. anamensis in eastern Africa and a rich record of fossil vertebrates. Most Kanapoi fossils are chronologically well constrained by radiometrically dated tephras between the ages of 4.2 and 4.1 million years ago. Sedimentological, isotopic, and faunal data indicate that the environments of Kanapoi during the early Pliocene had a complex range of vegetation types that included closed woodlands, shrubs, and grasslands near a river (for most of the sequence) or lake. These were dynamic landscapes that could shift rapidly from fluvial to lacustrine conditions, and then back. Australopithecus anamensis shared its environments with at least 10 species of very large herbivores, which undoubtedly played a major role in modifying the landscape by opening wooded areas and providing pathways for bipedal hominins. Hominins may have competed for terrestrial resources with abundant suids (Nyanzachoerus and Notochoerus) and for arboreal resources with monkeys (Parapapio being the most common cercopithecid). Kanapoi had a formidable group of predators that included a very abundant species of hyena (Parahyaena howelli), two sabre-tooth felids (Dinofelis and Homotherium), a giant otter (Enhydriodon cf. dikikae), and three species of crocodiles. Various measures of abundance indicate that A. anamensis was an important component of the Kanapoi early Pliocene ecosystems, and that its key adaptations allowed this species to thrive in complex and dynamic landscapes. (C) 2019 Elsevier Ltd. All rights reserved. Leakey Foundation Paleontological Scientific Trust (PAST) of South Africa National Science FoundationNational Science Foundation (NSF) University of Missouri Research Board info:eu-repo/semantics/publishedVersion

International audience; The thick-enameled, bunodont dentition shared by most early hominins has traditionally been interpreted as reflecting durophagy, especially in the robust genus Paranthropus. However, subsequent works on dental microwear textures (DMT) and biogeochemical compositions have challenged this hypothesis. Some authors argued that their robust morphology might have been driven by the consumption of mechanically challenging resources during periods of food scarcity. An experimental baseline using a model taxon with bunodont, thickenameled cheek teeth, could help better interpret DMT and test hypotheses regarding the consumption of mechanically challenging foods that could be fallback foods. Besides, earlier studies have shown that DMT can track subtle dietary variations in extant taxa. This study aims at testing the hypothesis that the consumption of various seeds has an impact on DMT of bunodont mammals despite similar overall diets. Trials were conducted on four groups of domestic pigs (Sus scrofa) all fed on mixed cereal and soy flours: the control group received only flours (n = 12), and the three others were supplemented with either 20% corn kernels (n = 6), 30% barley seeds (n =5), or 10 hazelnuts in shell per day (n = 6). We studied phases I and II facets on first molars and on fourth deciduous premolars, and applied a subsampling surface strategy to identify discriminative parameters among dietary groups. Principal Component Analyses show that DMT differ between pigs fed on different types of seeds. Our results also demonstrate that combining both crushing and shearing facets into analyses improves dietary discriminations. This study shows that variables that contribute most to dietary discriminations as selected from the subsampling strategy are mainly height parameters. These results thus support the idea that the consumption of seeds has an impact on the relief of surface textures.

The craniomandibular morphology of Homo naledi shows variable resemblances with species across Homo, which confounds an easy assessment of its phylogenetic position. In terms of skull shape, H. naledi has its closest affinities with Homo erectus, while mandibular shape places it closer to early Homo. From a tooth crown perspective, the smaller molars of H. naledi make it distinct from early Homo and H. erectus. Here, we compare the mandibular molar root morphology of six H. naledi individuals from the Dinaledi Chamber to those of African and Eurasian Plio-Pleistocene fossil hominins (totalling 183 mandibular first, second and third molars). The analysis of five root metric variables (cervical plane area, root length, root cervix volume, root branch volume, and root surface area) derived from microCT reconstructions reveals that the molar roots of H. naledi are smaller than those of Homo habilis, Homo rudolfensis, and H. erectus, but that they resemble those of three Homo sp. specimens from Swartkrans and Koobi Fora in size and overall appearance. Moreover, though H. naledi molar roots are similar in absolute size to Pleistocene Homo sapiens, they differ from H. sapiens in having a larger root volume for a given cervical plane area and less taurodont roots; the root cervix-to-branch proportions of H. naledi are comparable to those of Australopithecus africanus and species of Paranthropus. H. naledi also shares a metameric root volume pattern (M2 > M3 > M1) with Australopithecus and Paranthropus but not with any of the other Homo species (M2 > M1 > M3). Our findings therefore concur with previous studies that found that H. naledi shares plesiomorphic features with early Homo, Australopithecus, and Paranthropus. While absolute molar root size aligns H. naledi with Homo from North and South Africa, it is distinguishable from these in terms of root volumetric proportions.

The Drimolen Palaeocave System in the ‘Fossil Hominid Sites of South Africa’ UNESCO World Heritage Site is well known for numerous remains of early hominins such as Paranthropus robustus and early Homo. These hominin fossils, along with bone tools and notably diverse accumulation of non-hominin primates and fauna, have all been excavated from the 'Main Quarry' area of the site where extensive lime-mining took place. Here we report the first radiometric age of 1.712 ± 0.269 Ma for hominin bearing deposits associated with the DNH7 Paranthropus robustus cranium in the Main Quarry area of the site, which is consistent with recent biochronological estimates. This age is similar to recent estimates for Swartkrans Member 1 Hanging Remnant (somewhere between 2.3 and 1.8 Ma) which also contains Paranthropus and early Homo. Simultaneously, we integrate the newly radiometrically dated Main Quarry deposits with a new fossil deposit, the Drimolen Makondo, discovered in 2013, that is situated some 50 m up the hill to the west from the Main Quarry. It has experienced only limited disturbance from mining but much more extensive erosion. Preliminary excavations and analysis have revealed that the Makondo infill is older than the Main Quarry, dating to 2.706 ± 0.428 Ma. Its greater age is confirmed by biochronology. The Makondo thus overlap with the suggested end of deposition of Australopithecus bearing Sterkfontein deposits, although it is yet to yield any hominin remains. These new dates for the two Drimolen Palaeocave System deposits indicates that, contrary to prior age estimates, the Drimolen site as a whole records the critical hominin and faunal turnover in South African palaeocommunities that occurred around 2.3–1.7 Ma. Finally, as the Drimolen Makondo represents a rare example of a pre-2 Ma fossil bearing deposit in the Gauteng exposures of the Malmani dolomite, we also integrate our results into the greater South African record of palaeodeposit formation (most of which occur between ∼2.0 and 1.0 Ma). An analysis of the age of palaeocave infillings across the Malmani dolomite suggests that, as is classically the case with karst, the height within the dolomite is broadly correlated to their age, although with some notable exceptions that are likely related to localised geological features. Our analysis also indicates that most caves have undergone some form of secondary karstification related to a younger phase of cave formation, contrasting with models that suggest the cavities all formed at the same time and that infill is related to erosion and the opening up of cave passages. As such, the reason that few pre-2 Ma deposits have been identified in the Gauteng exposures of the Malmani dolomite is probably because these older caves have been eroded away. Identifying such early caves is critical in understanding whether earlier hominins may have once existed in South Africa or if erosion of older deposits (or an absence of speleogenesis at this time) has made such early periods absent from the geological record.

The mandibular third premolar (P3) exhibits substantial differences in size and shape among hominoid taxa, and displays a number of discrete traits that have proven to be useful in studies of hominin taxonomy and phylogeny. Discrete traits at the enamel-dentine junction (EDJ) can be accurately assessed on moderately worn specimens, and often appear sharper than at the outer-enamel surface (OES). Here we use microtomography to image the P3 EDJ of a broad sample of extant apes, extinct hominins and modern humans (n = 100). We present typologies for three important premolar discrete traits at the EDJ (transverse crest, marginal ridge and buccal grooves), and score trait frequencies within our sample. We find that the transverse crest is variable in extant apes, while the majority of hominins display a transverse crest which runs directly between the two major premolar cusps. Some Neanderthals display a unique form in which the transverse crest fails to reach the protoconid. We find that mesial marginal ridge discontinuity is common in Australopithecus anamensis and Australopithecus afarensis while continuous marginal ridges largely characterize Australopithecus africanus and Paranthropus. Interrupted mesial and distal marginal ridges are again seen in Homo sapiens and Neanderthals. Premolar buccal grooves, previously identified at the OES as important for hominin systematics, are again found to show a number of taxon-specific patterns at the EDJ, including a clear difference between Australopithecus and Paranthropus specimens. However, their appearance may be dependent on the morphology of other parts of the crown such as the protoconid crest, and the presence of accessory dentine horns. Finally, we discuss rare variations in the form of dentine horns that underlie premolar cusps, and their potential homology to similar morphologies in other tooth positions.

The African savannas that many early hominins occupied likely experienced stark seasonality and contained mosaic habitats (i.e., combinations of woodlands, wetlands, grasslands, etc.). Most would agree that the bulk of dietary calories obtained by taxa such as Australopithecus and Paranthropus came from the consumption of vegetation growing across these landscapes. It is also likely that many early hominins were selective feeders that consumed particular plants/plant parts (e.g., leaves, fruit, storage organs) depending on the habitat and season within which they were foraging. Thus, improving our understanding of how the nutritional properties of potential hominin plant foods growing in modern African savanna ecosystems respond to season and vary by habitat will improve our ability to model early hominin dietary behavior. Here, we present nutritional analyses (crude protein and acid detergent fiber) of plants growing in eastern and southern African savanna habitats across both wet and dry seasons. We find that many assumptions about savanna vegetation are warranted. For instance, plants growing in our woodland habitats have higher average protein/fiber ratios than those growing in our wetland and grassland transects. However, we find that the effects of season and habitat are complex, an example being the unexpectedly higher protein levels we observe in the grasses and sedges growing in our Amboseli wetlands during the dry season. Also, we find significant differences between the vegetation growing in our eastern and southern African field sites, particularly among plants using the C4 photosynthetic pathway. This may have implications for the differences we see between the stable carbon isotope compositions and dental microwear patterns of eastern and southern African Paranthropus species, despite their shared, highly derived craniodental anatomy.

It has been suggested that a shift in diet is one of the key adaptations that distinguishes the genus Homo from earlier hominins, but recent stable isotopic analyses of fossils attributed to Homo in the Turkana Basin show an increase in the consumption of C4 resources circa 1.65 million years ago, significantly after the earliest evidence for Homo in the eastern African fossil record. These data are consistent with ingesting more C4 plants, more animal tissues of C4 herbivores, or both, but it is also possible that this change reflects factors unrelated to changes in the palaeobiology of the genus Homo. Here we use new and published carbon and oxygen isotopic data (n = 999) taken from large-bodied fossil mammals, and pedogenic carbonates in fossil soils, from East Turkana in northern Kenya to investigate the context of this change in the isotope signal within Homo. By targeting taxa and temporal intervals unrepresented or undersampled in previous analyses, we were able to conduct the first comprehensive analysis of the ecological context of hominin diet at East Turkana during a period crucial for detecting any dietary and related behavioural differences between early Homo (H. habilis and/or H. rudolfensis) and Homo erectus. Our analyses suggest that the genus Homo underwent a dietary shift (as indicated by δ13Cena and δ18Oena values) that is (1) unrelated to changes in the East Turkana vegetation community and (2) unlike patterns found in other East Turkana large mammals, including Paranthropus and Theropithecus. These data suggest that within the Turkana Basin a dietary shift occurred well after we see the first evidence of early Homo in the region. Carbon and oxygen isotope analysis of sediments and soils from hominin locales in Kenya coupled with results from hominin taxa suggest that a dietary shift from C3 to C4 resources occurred in the genus Homo circa 1.65 million years ago despite palaeoenvironmental continuity.

Changes in first metatarsal (MT1) morphology within the hominin clade are crucial for reconstructing the evolution of a forefoot adapted for human-like gait. Studies of the external morphology of the MT1 in humans, non-human apes, and fossil hominins have documented changes in its robusticity, epiphyseal shape and its articulation with the medial cuneiform. Here, we test whether trabecular structure in the MT1 reflects different loading patterns in the forefoot across extant large apes and humans, and within this comparative context, infer locomotor behavior in two fossil hominins from Swartkrans, South Africa. Microtomographic scans were collected from the MT1 of Pongo sp. (n = 6), Gorilla gorilla (n = 10), Pan troglodytes (n = 10), Homo sapiens (n = 11), as well as SKX 5017 (Paranthropus robustus), and SK 1813 (Hominin gen. sp. indet.). Trabecular structure was quantified within the head and base using a ‘whole-epiphysis’ approach with medtool 4.2. We found that modern humans displayed relatively higher bone volume fraction (BV/TV) in the dorsal region of each epiphysis and a higher overall degree of anisotropy (DA), whereas great apes showed higher BV/TV in the plantar regions, reflecting dorsiflexion at the metatarsophalangeal (MTP) joint in the former and plantarflexion in the latter. Both fossils displayed low DA, with SKX 5017 showing a hyper-dorsal concentration of trabecular bone in the head (similar to humans), while SK 1813 showed a more central trabecular distribution not seen in either humans or non-human apes. Additionally, we found differences between non-human apes, modern humans, and the fossil taxa in trabecular spacing (Tb.Sp.), number (Tb.N.), and thickness (Tb.th.). While low DA in both fossils suggests increased mobility of the MT1, differences in their trabecular distributions could indicate variable locomotion in these Pleistocene hominins (recognizing that the juvenile status of SK 1813 is a potential confounding factor). In particular, evidence for consistent loading in hyper-dorsiflexion in SKX 5017 would suggest locomotor behaviors beyond human-like toe off during terrestrial locomotion.

International audience; Studies of the australopith (Australopithecus and Paranthropus) proximal femur have increasingly integrated information from the local arrangement of the cortical and cancellous bone to allow functional-biomechanical inferences on the locomotor behavioral patterns. In Australopithecus africanus and Paranthropus robustus, the cancellous bone organization at the center of the femoral head shows principal strut orientation similar to that of fossil and recent humans, which indicates that australopiths were human-like in many aspects of their bipedalism. However, by combining outer morphology with superoinferior asymmetry in cortical bone thickness at the base of neck and mid-neck, it has been suggested that, while adapted for terrestrial bipedality, australopiths displayed a slightly altered gait kinematics compared to Homo. We used techniques of 2D and 3D virtual imaging applied to an X-ray microtomographic record to assess cortical bone distribution along the entire femoral neck compartment in four upper femora from Swartkrans, South Africa (SK 82, SK 97, SK 3121, and SWT1/LB-2) and compared the results to the extant human and chimpanzee conditions. Our results support and extend previous evidence for more symmetric superior and inferior femoral neck cortical thicknesses in P. robustus than in modern humans and show that the differences are even greater than previously reported. However, P. robustus and humans still share a trend of lateral-to-medial decrease in asymmetry of the superior/inferior cortical thickness ratio, while this pattern is reversed in chimpanzees. We also identified two features uniquely characterizing P. robustus: an accentuated contrast between the relatively thicker anterior and the thinner posterior walls, and a more marked lateral-to-medial thinning of both cortices compared to extant humans and chimpanzees, which indicate wider interspecific differences among hominids in structural organization of the proximal femur than previously reported. It remains to be ascertained if, and to what extent, these features also characterize the femoral neck of Australopithecus.

The KNM-ER 64060 dentition derives from a horizon that most likely dates to between 2.02 and 2.03 Ma. A proximate series of postcranial bones (designated KNM-ER 64061) derives from the same siltstone unit and may be associated with the dentition, but their separation on the surface of the site leaves some room for doubt. KNM-ER 64060 is one of fewer than ten hominin specimens from the Early Pleistocene of East Africa that comprises a full or nearly complete mandibular dentition. Its taxonomic attribution is potentially significant, especially if the postcranial elements are related. At least three, and probably four hominin species, including Paranthropus boisei and Homo erectus (= H. ergaster), are known at about this time in East Africa. Other penecontemporaneous fossils have been referred to a single, highly variable species, H. habilis, or two taxa, namely H. habilis and H. rudolfensis. Although the weight of evidence supports the attribution of these specimens to two species, there is notable lack of agreement over the assignation of individual fossils. We take a conservative approach and group all such specimens under the designation “early Homo sp.” for comparative purposes. KNM-ER 64060 is clearly attributable to Homo rather than Paranthropus. The preponderance of the evidence suggests that the affinities of KNM-ER 64060 are with fossils assigned to the early Homo sp. category rather than with H. erectus. This is indicated by the overall sizes of the KNM-ER 64060 canine, premolar and molar crowns, the size relationships of the P3 to P4, the relative narrowness of its premolar crowns, the cusp proportions of the M1 and especially those of the M2 and M3, and seemingly the possession of a two-rooted P4. Some of these comparisons suggest further that among the fossils comprising the early Homo sp. sample, the KNM-ER 64060 dentition exhibits greater overall similarity to specimens such as OH 7 and OH 16 that represent Homo habilis sensu stricto.

A well-preserved distal portion of a left humerus was discovered in 1976 during excavations directed by J. Chavaillon at the Gombore I site, in the Melka Kunture area (Ethiopia). The specimen, labelled Gombore IB-7594 (formally Melka Kunture 3, or MK3), was found in situ within unit 2 of level B, which is dated to >1.39 Ma and includes a rich Oldowan Paleolithic assemblage. Although MK3 has never been described in detail, it appeared in the literature several times and, from a taxonomic point of view, has been alternatively regarded as Homo , Australopithecus or Paranthropus . According to our analysis, MK3 exhibits a suite of features that fit the variability of the genus Homo and does not display any clear Australopithecus/Paranthropus affinity. Nevertheless, MK3 adds a great deal of variability to the genus Homo , at least as far as the Early Pleistocene fossil record is concerned. In particular, our quantitative approach, which combines traditional morphometric analyses and geometric morphometrics, highlights traits that are uncommon among the Plio-Pleistocene fossil record, while affinities with Mid-to-Late Pleistocene representatives of Homo are observed. In addition, the large size of MK3 suggests that this humerus belonged to an individual whose body weight approached 90 kg, far from the range of body size known for Homo representatives in the Early Pleistocene and as big as those of extant humans or even gorillas. We suggest that such peculiar features are of interest when regarded from an ecological perspective; thus, dimension and morphology of MK3 may be considered as an exaptation that became useful when early humans dispersed to high altitudes such as those of the upper Awash basin on the Ethiopian plateau, at heights above 2000 m.

The question of how many Australopithecus species lived at Sterkfontein Member 4 and Makapansgat continues to be controversial inasmuch as the fossils are poorly preserved, the stratigraphy is difficult to interpret, and the cranial, dental, and postcranial remains are mostly not associated. To proceed we applied the most intensive modern methods of 3D geometric morphometrics to dental form, specifically the shapes of the upper second molars (M2s) in a sample combining 13 Australopithecus, 11 Paranthropus, and 23 Homo. We analyzed outer and inner crown surfaces, as well as crown and cervical outlines both separately and together, using a total of 16 landmarks, 51 curve semilandmarks, and 48 pseudolandmarks over the four structures. Outer and inner enamel surfaces are highly correlated in this dataset, while crown outline is the least informative of the four structures. Homo was easily distinguished from both Australopithecus and Paranthropus by these methods, likewise Homo sapiens from Homo neanderthalensis. There were, however, no stable classes within the Australopithecus sample or between Australopithecus and Paranthropus. Instead, there was a gradient along which Australopithecus prometheus and Australopithecus africanus lie toward the extremes, with Paranthropus overlapping both. If there are indeed different species at this site, then either their M2 morphologies are uninformative or else the present sample is too small to make an accurate assessment. Our findings suggest that the variability of the Australopithecus specimens will be difficult to interpret authoritatively, independent of the method used.

Enamel thickness continues to be an important morphological character in hominin systematics and is\ud frequently invoked in dietary reconstructions of Plio-Pleistocene hominin taxa. However, to date, the\ud majority of published data on molar enamel thickness of Pliocene and early Pleistocene hominins derive\ud from naturally fractured random surfaces of a small number of specimens. In this study we systematically\ud analyze enamel thickness in a large sample of Plio-Pleistocene fossil hominins (n ¼ 99), extant hominoids\ud (n ¼ 57), and modern humans (n ¼ 30). Based on analysis of 2D mesial planes of section derived from\ud microtomography, we examine both average and relative enamel thickness, and the distribution of\ud enamel across buccal, occlusal, and lingual components of mandibular molars. Our results confirm the\ud trend of increasing enamel thickness during the Pliocene that culminates in the thick enamel of the\ud robust Australopithecus species, and then decreases from early Homo to recent modern humans. All\ud hominin taxa share a regional average enamel thickness pattern of thick occlusal enamel and greater\ud buccal than lingual enamel thickness. Pan is unique in exhibiting the thinnest average enamel thickness\ud in the occlusal basin. Statistical analysis indicates that among Pliocene hominins enamel thickness is a\ud weak taxonomic discriminator. The data underlying these results are included in a table in the Supplementary\ud Online Material.

Closely related species are likely to experience resource competition in areas where their ranges overlap. Fossil evidence suggests that hominins in East Africa c. 2-1.5 million years ago may have lived synchronically and sympatrically, and that competition may have contributed to the different tooth sizes observed in Homo and Paranthropus. To assess the likelihood that these taxa overlapped, we applied a character displacement model to the postcanine tooth size of fossil hominins and validated this model in populations of living primates. Mandibular fourth premolar (P4 ) crown size was measured from fossil taxa and from living primate species where dietary overlap is established. Dimensions of the P4 crown were fitted to a character matrix and described as the response variables of a generalized linear model that took taxon and location as input variables. The model recovered significant divergence in samples of closely related, living primates. When applied to fossil hominins the same model detected strong indications of character displacement between early Homo and Paranthropus (P = 0.002) on the basis of their P4 crown size. Our study is an example of how ecologically informed morphologies measured in appropriate extant referents can provide a comparative context for assessing community and ecological evolution in the fossil record.

The Sterkfontein Caves site is one of the richest early hominin fossil localities in Africa. More specifically, the fossiliferous deposits within the lower-lying Jacovec Cavern have yielded valuable hominin remains; prominent among them is the Australopithecus partial cranium StW 578. Due to the fragmentary nature of the braincase, the specimen has not yet been formally assigned to a species. In this context, we employ microtomography to quantify cranial thickness and composition of StW 578 in order to assess its taxonomic affinity. As comparative material, we investigate 10 South African hominin cranial specimens from Sterkfontein (StW 505, Sts 5, Sts 25, Sts 71), Swartkrans (SK 46, SK 48, SK 49) and Makapansgat (MLD 1, MLD 10, MLD 37/38), attributed to either Australopithecus or Paranthropus, as well as 10 extant human and 10 extant chimpanzee crania. Thickness variation in and structural arrangement of the inner and outer cortical tables and the diploe are automatically assessed at regular intervals along one parasagittal and one coronal section. Additionally, topographic cranial vault thickness distribution is visualized using color maps. Comparisons highlight an absolutely and relatively thickened condition of the StW 578 cranial vault versus those of other South African Plio-Pleistocene hominins. Moreover, in StW 578, as well as in the Australopithecus specimens Sts 5 and Sts 71 from Sterkfontein, the diploic layer contributes substantially to cumulative vault thickness (i.e., >60%). Within the comparative sample investigated here, StW 505 and Sts 71 from Sterkfontein Member 4, both attributed to Australopithecus, most closely resemble StW 578 in terms of cranial vault thickness values, tissue proportions, and two- and three-dimensional distributions. Including additional Plio-Pleistocene Australopithecus and Paranthropus crania from South and East Africa in future studies would further help establish morphological variability in these hominin taxa.

In 2013, 2014 new hominin remains were uncovered in the Dinaledi chamber of the Rising Star cave system in South Africa. In 2015 Berger and colleagues identified these remains as belonging to a new species Homo naledi (Berger et al., 2015). Subsequent comparative studies of the skull, postcrania and permanent dentition have supported this taxonomic affiliation (Harcourt-Smith et al., 2015; Kivell et al., 2015; Irish et al., 2018). The deciduous teeth can offer unique insights into hominin evolution. Due to their early onset and rapid development their morphology is thought to be under stronger genetic control and less influenced by environment than are the permanent teeth. In this study we compared the H. naledi deciduous teeth from the 2013–2014 excavations to samples representing much of the hominin clade including Australopithecus afarensis, Australopithecus africanus, Paranthropus boisei, Paranthropus robustus, early Homo, Homo antecessor, Homo erectus s.l., Homo floresiensis, Middle Pleistocene Homo, Homo neanderthalensis, early Homo sapiens and recent H. sapiens from Sub-Saharan Africa. By making such a broad morphological comparison, we aimed to contextualize the Dinaledi hominins and to further assess the validity of their taxonomic assignment. Our analysis of the deciduous teeth revealed a unique combination of features that mirror (but also expand) that found in the permanent teeth. This mosaic includes an asymmetrical lower canine with a distal tubercle, an upper first molar with a large hypocone and epicrista associated with a mesial cuspule, a molarized lower first molar resembling Paranthropus, and upper and lower second molars that resemble later Homo in their lack of accessory cusps. The unique combination of deciduous dental characters supports previous studies assigning H. naledi to a new species, although its phylogenetic position vis-a-vis other Homo species remains ambiguous.

Though late Middle Pleistocene in age, Homo naledi is characterized by a mosaic of Australopithecus-like (e.g., curved fingers, small brains) and Homo-like (e.g., elongated lower limbs) traits, which may suggest it occupied a unique ecological niche. Ecological reconstructions inform on niche occupation, and are particularly successful when using dental material. Tooth shape (via dental topography) and size were quantified for four groups of South African Plio-Pleistocene hominins (specimens of Australopithecus africanus, Paranthropus robustus, H. naledi, and Homo sp.) on relatively unworn M2s to investigate possible ecological differentiation in H. naledi relative to taxa with similar known geographical ranges. H. naledi has smaller, but higher-crowned and more wear resistant teeth than Australopithecus and Paranthropus. These results are found in both lightly and moderately worn teeth. There are no differences in tooth sharpness or complexity. Combined with the high level of dental chipping in H. naledi, this suggests that, relative to Australopithecus and Paranthropus, H. naledi consumed foods with similar fracture mechanics properties but more abrasive particles (e.g., dust, grit), which could be due to a dietary and/or environmental shift(s). The same factors that differentiate H. naledi from Australopithecus and Paranthropus may also differentiate it from Homo sp., which geologically predates it, in the same way. Compared to the great apes, all hominins have sharper teeth, indicating they consumed foods requiring higher shear forces during mastication. Despite some anatomical similarities, H. naledi likely occupied a distinct ecological niche from the South African hominins that predate it.

Paranthropus boisei was first described in 1959 based on fossils from the Olduvai Gorge and now includes many fossils from Ethiopia to Malawi. Knowledge about its postcranial anatomy has remained elusive because, until recently, no postcranial remains could be reliably attributed to this taxon. Here, we report the first associated hand and upper limb skeleton (KNM-ER 47000) of P. boisei from 1.51 to 1.53 Ma sediments at Ileret, Kenya. While the fossils show a combination of primitive and derived traits, the overall anatomy is characterized by primitive traits that resemble those found in Australopithecus, including an oblique scapular spine, relatively long and curved ulna, lack of third metacarpal styloid process, gracile thumb metacarpal, and curved manual phalanges. Very thick cortical bone throughout the upper limb shows that P. boisei had great upper limb strength, supporting hypotheses that this species spent time climbing trees, although probably to a lesser extent than earlier australopiths. Hand anatomy shows that P. boisei, like earlier australopiths, was capable of the manual dexterity needed to create and use stone tools, but lacked the robust thumb of Homo erectus, which arguably reflects adaptations to the intensification of precision grips and tool use. KNM-ER 47000 provides conclusive evidence that early Pleistocene hominins diverged in postcranial and craniodental anatomy, supporting hypotheses of competitive displacement among these contemporaneous hominins.

The African Plio-Pleistocene hominins known as australopiths evolved derived craniodental features frequently interpreted as adaptations for feeding on either hard, or compliant/tough foods. Among australopiths, Paranthropus boisei is the most robust form, exhibiting traits traditionally hypothesized to produce high bite forces efficiently and strengthen the face against feeding stresses. However, recent mechanical analyses imply that P. boisei may not have been an efficient producer of bite force and that robust morphology in primates is not necessarily strong. Here we use an engineering method, finite element analysis, to show that the facial skeleton of P. boisei is structurally strong, exhibits a strain pattern different from that in chimpanzees (Pan troglodytes) and Australopithecus africanus, and efficiently produces high bite force. It has been suggested that P. boisei consumed a diet of compliant/tough foods like grass blades and sedge pith. However, the blunt occlusal topography of this and other species suggests that australopiths are adapted to consume hard foods, perhaps including grass and sedge seeds. A consideration of evolutionary trends in morphology relating to feeding mechanics suggests that food processing behaviors in gracile australopiths evidently were disrupted by environmental change, perhaps contributing to the eventual evolution of Homo and Paranthropus.

Phylogenetic analyses require evolutionarily independent characters, but there is no consensus, nor has there been a clear methodology presented on how to define character independence in a phylogenetic context, particularly within a complex morphological structure such as the skull. Following from studies of craniofacial development, we hypothesize that the premaxilla is an independent evolutionary module with two integrated characters that have traditionally been treated as independent. We test this hypothesis on a large sample of primate skulls and find evidence supporting the premaxilla as an independent module within the larger module of the palate. Additionally, our data indicate that the convexity of the nasoalveolar clivus and the contour of the alveolus are integrated within the premaxilla. We show that the palate itself is composed of two distinct modules: the FNP-derived premaxillae and the mxBA1-derived maxillae and palatines. Application of our data to early African hominin facial morphology suggests that at least three separate transitions contributed to robust facial morphology: 1) an increase in the size of the post-canine dentition housed within the maxillae and palatines, 2) modification of the premaxilla generating a concave clivus and reduced incisor alveolus, and 3) modification of the zygomatic, shifting the zygomatic root and lateral face anteriorly. These data lend support to the monophyly of Paranthropus boisei and Paranthropus robustus, and provide mounting evidence in favor of a Paranthropus clade. This study also highlights the utility of applying developmental evidence to studies of morphological evolution.

The inhibitory cascade is a mathematical model for interpreting the relative size of the occlusal surfaces of mammalian molars in terms of developmental mechanisms. The cascade is derived from experimental studies of mouse molars developed in culture, and has been tested and applied to the dentitions of rodents, ungulates, carnivores, and platyrrhines. Results from such applications have provided new information regarding the origins of plesiomorphic traits in mammalian clade and how derived morphologies may arise. In this study we apply the inhibitory cascade model to the postcanine dentition of a sample of Old World primates that includes fossil hominins. The results of this study suggest that the inhibitory cascade (i.e. M1 < M2 < M3 ) describes the relative sizes of the molar occlusal areas of Old World primates and is likely the plesiomorphic condition for this clade. Within that clade, whereas most Old World monkeys have a M1 < M2 < M3 pattern, most apes have a M1 < M2 ≈ M3 pattern. This modified cascade suggests that greater levels of inhibition (or less activation) are acting on the posterior molars of apes, thus facilitating the reduction of M3 s within the apes. With the exception of the baboon genus Papio, extant congeners typically share the same molar inhibitory cascade. The differences in the relative size relationships observed in the molar and premolar-molar cascades of the species included in the fossil hominin genus Paranthropus suggest that although large postcanine teeth are a shared derived trait within this genus, the developmental basis for postcanine megadontia may not be the same in these two Paranthropus taxa. Our results show that phenotypic characters such as postcanine megadontia may not reflect common development.

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.

Australian Archaeomagnetism Laboratory, Department of Archaeology, Environment and Community Planning, Faculty of Humanities and Social Sciences, La Trobe University, Melbourne Campus, Bundoora, 3086 VIC, Australia Department of Anatomy and Developmental Biology, School of Biomedical Sciences, Faculty of Medicine, Nursing & Health Sciences, Monash University, Clayton, Melbourne, 3800 VIC, Australia

Foraging is constrained by the energy within resources and the mechanics of acquisition and assimilation. Thick molar enamel, a character trait differentiating hominins from African apes, is predicted to mitigate the mechanical costs of chewing obdurate foods. The classic expression of hyperthick enamel together with relatively massive molars, termed megadontia, is most evident in Paranthropus, a lineage of hominins that lived about 2.7–1.2 million years ago. Among contemporary primates, thicker molar enamel corresponds with the consumption of stiffer, deformation-resistant foods, possibly because thicker enamel can better resist cracking under high compressive loads. Accordingly, plant underground storage organs (USOs) are thought to be a central food resource for hominins such as Paranthropus due to their abundance, isotopic composition, and mechanical properties. Here, we present a process-based model to investigate foraging constraints as a function of energetic demands and enamel wear among human ancestors. Our framework allows us to determine the fitness benefits of megadontia, and to explore under what conditions stiff foods such as USOs are predicted to be chosen as fallback, rather than preferred, resources. Our model predictions bring consilience to the noted disparity between functional interpretations of megadontia and microwear evidence, particularly with respect to Paranthropus boisei.

We have analyzed the phytolith content of 37 samples from the paleosol capped by Tuff IC dated 1.84 Ma within Middle Bed I at Olduvai Gorge, Tanzania. Samples were obtained from different trenches sampling an area of about 25 ha to document the paleovegetation associated with the abundant faunal remains and hominin artifacts in FLK, AMK, PTK and DS sites. The analysis reveals the abundance of globular granulate and forest indicator phytoliths (from 3% to 86% relative to the total number of phytoliths in the assemblages) attributed to woody plants, and of short cell grass phytoliths (up to 22%). The presence of palm phytoliths (up to 57%) attests for meso-haline soils in the area in agreement with the presence of the saline/alkaline Olduvai paleolake and one (or more) groundwater discharge areas (freshwater springs and/or wetland). Phytoliths provide botanical evidence for a mixed paleovegetation dominated by forest and woodland with patches of open-spaces occupied by grasses, prior to the deposition of Tuff IC. The presence of fern phytoliths in some assemblages suggest shaded and humid habitat, which supports the presence of freshwater (watercourses or springs) that might have attracted hominins and other animals. Yet, we have found that our phytolith extractions (residues) also include micrometric zeolites (chabazite, clinoptilolite, phillipsite), which attest that the phytolith signal in the Zinj paleosols is certainly affected by dissolution. This study completes the spatial analysis of the area and reveals that the wooded area extends north, south, and west of FLK Zinj, with palm trees preferentially located in the periphery, like the Zinj, AMK, and DS sites. The phytolith-inferred distribution of plants does not support the hypothesis that topography was influential in the location of these sites, particularly given that trees and/or shrubs were dominant all over. Rather, it is the presence of freshwater ponds that seem to have been the attractive factor for hominins at this particular place and time.

KNM-ER 47000 is a fossil hominin upper limb skeleton from the Koobi Fora Formation, Kenya (FwJj14E, Area 1A) that includes portions of the scapula, humerus, ulna, and hand. Dated to ∼1.52 Ma, the skeleton could potentially belong to one of multiple hominin species that have been documented in the Turkana Basin during this time, including Homo habilis, Homo erectus, and Paranthropus boisei. Although the skeleton lacks associated craniodental material, the partial humerus (described here) preserves anatomical regions (i.e., distal diaphysis, elbow joint) that are informative for taxonomic identification among early Pleistocene hominins. In this study, we analyze distal diaphyseal morphology and the shape of the elbow region to determine whether KNM-ER 47000 can be confidently attributed to a particular species. The morphology of the KNM-ER 47000 humerus (designated KNM-ER 47000B) is compared to that of other early Pleistocene hominin fossil humeri via the application of multivariate ordination techniques to both two-dimensional landmark data (diaphysis) and scale-free linear shape data (elbow). Distance metrics reflecting shape dissimilarity between KNM-ER 47000B and other fossils (and species average shapes) are assessed in the context of intraspecific variation within modern hominid species (Homo sapiens, Pan troglodytes, Gorilla gorilla, Pongo pygmaeus). Our comparative analyses strongly support attribution of KNM-ER 47000 to P. boisei. Compared to four other partial skeletons that have (justifiably or not) been attributed to P. boisei, KNM-ER 47000 provides the most complete picture of upper limb anatomy in a single individual. The taxonomic identification of KNM-ER 47000 makes the skeleton an important resource for testing future hypotheses related to P. boisei upper limb function and the taxonomy of isolated early Pleistocene hominin remains.

Placing the biological adaptations of Pleistocene hominins within a well-resolved ecological framework has been a longstanding goal of paleoanthropology. This effort, however, has been challenging due to the discontinuous nature of paleoecological data spanning many important periods in hominin evolution. Sediments from the Upper Burgi (1.98–1.87 Ma), KBS (1.87–1.56 Ma) and Okote (1.56–1.38 Ma) members of the Koobi Fora Formation at East Turkana in northern Kenya document an important time interval in the evolutionary history of the hominin genera Homo and Paranthropus. Although much attention has been paid to Upper Burgi and KBS member deposits, far less is known regarding the East Turkana paleoecosystem during Okote Member times. This study pairs spatially-resolved faunal abundance data with stable isotope geochemistry from mammalian enamel to investigate landscape-scale ecosystem variability during Okote Member times. We find that during this period 1) taxa within the East Turkana large mammal community were distributed heterogeneously across space, 2) the abundance of C3 and C4 vegetation varied between East Turkana subregions, and 3) the Karari subregion, an area with abundant evidence of hominin stone tool manufacture, had significantly more C3 vegetation than regions closer to the central axis of the Turkana Basin (i.e., Ileret and Koobi Fora). These findings indicate that the East Turkana paleoecosystem during the Okote Member was highly variable across space and provided a complex adaptive landscape for Pleistocene hominins.

KNM-ER 47000A is a new 1.52 Ma hominin scapular fossil belonging to an associated partial skeleton from the Koobi Fora Formation, Kenya (FwJj14E, Area 1A). This fossil effectively doubles the record of Early Pleistocene scapulae from East Africa, with KNM-WT 15000 (early African Homo erectus) preserving the only other known scapula to date. KNM-ER 47000A consists of a complete glenoid cavity preserving a portion of the scapular spine and neck, the proximal half of the acromion, and a majority of the axillary border. A sufficient amount of anatomy is preserved to compare KNM-ER 47000A with scapulae of several Australopithecus species, extinct Homo, and living hominoids. The glenohumeral joint of KNM-ER 47000A is more laterally oriented than those of great apes and Australopithecus, aligning it closely with KNM-WT 15000 and modern humans. While this morphology does not imply a strong commitment to arboreality, its scapular spine is obliquely oriented—as in gorillas and some Australopithecus fossils—particularly when compared to the more horizontal orientation seen in KNM-WT 15000 and modern humans. Such a spine orientation suggests a narrow yet long infraspinous region, a feature that has been attributed to suspensory taxa. Accordingly, the morphology of KNM-ER 47000A presents conflicting behavioral implications. Nonetheless, a multivariate consideration of the available scapular traits aligns KNM-ER 47000A and Australopithecus with great apes, whereas KNM-WT 15000 resembles modern humans. The scapular morphology of KNM-ER 47000A is unique among fossil and extant hominoids and its morphological differences from KNM-WT 15000 strengthen the attribution of KNM-ER 47000 to Paranthropus boisei as opposed to early Homo. As the first evidence of scapular morphology in P. boisei, KNM-ER 47000A provides important new information on variation in hominin shoulder and upper limb anatomy from this critical period of hominin evolutionary history.

The life histories of early hominins are commonly characterized as being like those of great apes. However, the life histories of the extant great apes differ considerably from one another. Moreover, the extent to which their life histories correlate with the two aspects of morphology used to infer the life histories of fossil species, brain size and dental development, has remained subject to debate. Increased knowledge of great ape life histories and, more recently, dental development —in particular ages at first molar emergence— now make it clearer that the latter is strongly associated with important life-history attributes, whereas brain size, as reflected by cranial capacity, is less informative. Here we estimate ages at M1 emergence in several infant/juvenile individuals of Australopithecus and Paranthropus based on previous estimates of ages at death, determined through dental histology. These are uniformly earlier than would be predicted either by adult cranial capacity or by comparison to ages at M1 emergence in free-living extant great apes. This suggests that either, 1) the life histories of the early hominins were faster than those of all extant great apes; 2) there was selection for rapid initial dental development and presumably early weaning, but that early hominin life histories were otherwise more prolonged and consistent with adult cranial capacities; or 3) the ages at death have been systematically underestimated, resulting in underestimates of the ages at M1 emergence. We investigate the implications of each of these alternatives and, where possible, explore evidence that might support one over the others.

Hominin fossils of gracile and robust australopith groups were found both in East and in South Africa. It is unclear, however, whether all robusts belong to a monophyletic Paranthropus clade, as the craniofacial resemblance among robust australopiths might only be a superficial correlate of similar masticatory adaptations and not evidence of shared ancestry. It has been suggested that the East African Australopithecus/Paranthropus boisei and the South African A./P. robustus might be convergent allometric variants of their gracile geographical neighbors A. afarensis and A. africanus. Here we approach the phylogenetic questions about robust and gracile australopiths from an “evo-devo” perspective, examining how simple alterations of development could contribute to the shape differences among hominin species. Using geometric morphometrics we compare gracile and robust australopith crania in the context of the allometric scaling patterns of Pan troglodytes, P. paniscus, and Gorilla gorilla. We examine support for two alternative evolutionary scenarios based on predictions derived from quantitative genetics models: either (1) A./P. robustus evolved in South Africa from the gracile A. africanus, or (2) A./P. robustus is a local variant of the eastern African A./P. boisei. We use developmental simulations to demonstrate that some robust characteristics (wide faces, anteriorly placed zygomatics, and facial dishing) can be predicted by allometric scaling along the ontogenetic trajectory of the gracile A. africanus. We find, however, that the facial differences between A. africanus specimens (Taung, Sts 5, Sts 71, and Stw 505) and A./P. robustus specimen SK 48 cannot be explained by allometric scaling alone. Facial shape differences between A./P. robustus SK 48 and A./P. boisei (KNM-ER 732, KNM-ER 406, OH 5) and the A./P. aethiopicus specimen KNM-WT 17000, on the other hand, can largely be explained by allometric scaling. This is consistent with a close evolutionary relationship of these robust taxa.

The evolution of hominin growth and life history has long been a subject of intensive research, but it is only recently that paleoanthropologists have considered the ontogenetic basis of human morphological evolution. To date, most human EvoDevo studies have focused on developmental patterns in extant African apes and humans. However, the Old World monkey tribe Papionini, a diverse clade whose members resemble hominins in their ecology and population structure, has been proposed as an alternative model for human craniofacial evolution. This paper reviews prior studies of papionin development and socioecology and presents new analyses of juvenile shape variation and ontogeny to address fundamental questions concerning primate cranial development, including: (1) When are cranial shape differences between species established? (2) How do epigenetic influences modulate early-arising pattern differences? (3) How much do postnatal developmental trajectories vary? (4) What is the impact of developmental variation on adult cranial shape? and, (5) What role do environmental factors play in establishing adult cranial form? Results of this inquiry suggest that species differences in cranial morphology arise during prenatal or earliest postnatal development. This is true even for late-arising features that develop under the influence of epigenetic factors such as mechanical loading. Papionins largely retain a shared, ancestral pattern of ontogenetic shape change, but large size and sexual dimorphism are associated with divergent developmental trajectories, suggesting differences in cranial integration. Developmental simulation studies indicate that postnatal ontogenetic variation has a limited influence on adult cranial morphology, leaving early morphogenesis as the primary determinant of cranial shape. The ability of social factors to influence craniofacial development in Mandrillus suggests a possible role for phentotypic plasticity in the diversification of primate cranial form. The implications of these findings for taxonomic attribution of juvenile fossils, the developmental basis of early hominin characters, and hominin cranial diversity are discussed.

The large, bunodont postcanine teeth in living sea otters (Enhydra lutris) have been likened to those of certain fossil hominins, particularly the 'robust' australopiths (genus Paranthropus). We examine this evolutionary convergence by conducting fracture experiments on extracted molar teeth of sea otters and modern humans (Homo sapiens) to determine how load-bearing capacity relates to tooth morphology and enamel material properties. In situ optical microscopy and x-ray imaging during simulated occlusal loading reveal the nature of the fracture patterns. Explicit fracture relations are used to analyze the data and to extrapolate the results from humans to earlier hominins. It is shown that the molar teeth of sea otters have considerably thinner enamel than those of humans, making sea otter molars more susceptible to certain kinds of fractures. At the same time, the base diameter of sea otter first molars is larger, diminishing the fracture susceptibility in a compensatory manner. We also conduct nanoindentation tests to map out elastic modulus and hardness of sea otter and human molars through a section thickness, and microindentation tests to measure toughness. We find that while sea otter enamel is just as stiff elastically as human enamel, it is a little softer and tougher. The role of these material factors in the capacity of dentition to resist fracture and deformation is considered. From such comparisons, we argue that early hominin species like Paranthropus most likely consumed hard food objects with substantially higher biting forces than those exerted by modern humans.

Palaeomagnetic analysis was conducted on speleothems from Members 1–5 at Sterkfontein Cave, South Africa. Palaeomagnetic analysis of siltstone and speleothem from the bulk of Member 4 indicate a reversed magnetic polarity that dates the deposits and its Australopithecus africanus fossils to between 2.58 and ∼2.16 Ma. Further confirmation of this age comes in the form of two short normal polarity events correlated to the Reunion (∼2.16 Ma) and Huckleberry Ridge (∼2.05 Ma) events in speleothem capping the bulk of Member 4 and coeval with deposition of the final phase of Member 4, including A. africanus fossil Sts 5. At ∼2.16–2.05 Ma, Sts 5 is the youngest representative of A. africanus yet discovered. Palaeomagnetic analysis of the Silberberg Grotto deposits identifies a single short geomagnetic field event in flowstone overlying the StW 573 Australopithecus fossil, which is suggested to represent the Reunion event at ∼2.16 Ma. This further supports the uranium lead age estimates of 2.3–2.2 Ma for the StW 573 fossil. Based on a reversed polarity for the deposits below the skeleton it cannot be older than 2.58 Ma. If StW 573 is considered to be a second species of Australopithecus then this indicates that two species of Australopithecus are present at Sterkfontein between 2.6 and 2.0 Ma. All of the Member 5 deposits date to less than 1.8 Ma based on a comparison of palaeomagnetic, faunal, and electron spin resonance age estimates. The StW 53 fossil bearing infill (M5A) is intermediate in age between Member 4 and the rest of Member 5 (B-C) at around 1.78–1.49 Ma. The rest of Member 5 (B-C) containing Oldowan and Acheulian stone tools and Homo and Paranthropus fossils was deposited gradually between 1.40 and 1.07 Ma, much younger than previously suggested.

Morphological variation related to differential loading is well known for many craniomandibular elements. Yet, the function of the hard palate, and in particular the manner in which cortical and trabecular bone of the palate respond to masticatory loads, remains more ambiguous. Here, experimental data are presented that address the naturalistic influence of biomechanical loading on the postweaning development and structure of the hard palate. A rabbit model was used to test the hypothesis that variation in the morphology of the hard palate is linked to variation in masticatory stresses. Rabbit siblings were divided as weanlings into soft and hard/tough dietary treatment groups of 10 subjects each and were raised for 15 weeks until subadulthood. MicroCT analyses indicate that rabbits subjected to elevated masticatory loading developed hard palates with significantly greater bone area, greater cortical bone thickness along the oral lamina, and thicker anterior palates. Such diet-induced levels of palatal plasticity are comparable to those for other masticatory elements, which likely reflect osteogenic responses for maintaining the functional integrity of the palate vis-à-vis elevated stresses during unilateral mastication. These data support a role for mechanical loading in the determination of palatal morphology, especially its internal structure, in living and fossil mammals such as the hominin Paranthropus. Furthermore, these findings have potential implications for the evolution of the mammalian secondary hard palate as well as for clinical considerations of human oral pathologies.

The diet of early human ancestors has received renewed theoretical interest since the discovery of elevated δ13C values in the enamel of Australopithecus africanus and Paranthropus robustus. As a result, the hominin diet is hypothesized to have included C4 grass or the tissues of animals which themselves consumed C4 grass. On mechanical grounds, such a diet is incompatible with the dental morphology and dental microwear of early hominins. Most inferences, particularly for Paranthropus, favor a diet of hard or mechanically resistant foods. This discrepancy has invigorated the longstanding hypothesis that hominins consumed plant underground storage organs (USOs). Plant USOs are attractive candidate foods because many bulbous grasses and cormous sedges use C4 photosynthesis. Yet mechanical data for USOs—or any putative hominin food—are scarcely known. To fill this empirical void we measured the mechanical properties of USOs from 98 plant species from across sub-Saharan Africa. We found that rhizomes were the most resistant to deformation and fracture, followed by tubers, corms, and bulbs. An important result of this study is that corms exhibited low toughness values (mean = 265.0 J m−2) and relatively high Young’s modulus values (mean = 4.9 MPa). This combination of properties fits many descriptions of the hominin diet as consisting of hard-brittle objects. When compared to corms, bulbs are tougher (mean = 325.0 J m−2) and less stiff (mean = 2.5 MPa). Again, this combination of traits resembles dietary inferences, especially for Australopithecus, which is predicted to have consumed soft-tough foods. Lastly, we observed the roasting behavior of Hadza hunter-gatherers and measured the effects of roasting on the toughness on undomesticated tubers. Our results support assumptions that roasting lessens the work of mastication, and, by inference, the cost of digestion. Together these findings provide the first mechanical basis for discussing the adaptive advantages of roasting tubers and the plausibility of USOs in the diet of early hominins.

In this review we attempt to reconstruct the evolutionary history of hominin life history from extant and fossil evidence. We utilize demographic life history theory and distinguish life history variables, traits such as weaning, age at sexual maturity, and life span, from life history-related variables such as body mass, brain growth, and dental development. The latter are either linked with, or can be used to make inferences about, life history, thus providing an opportunity for estimating life history parameters in fossil taxa. We compare the life history variables of modern great apes and identify traits that are likely to be shared by the last common ancestor of Pan-Homo and those likely to be derived in hominins. All great apes exhibit slow life histories and we infer this to be true of the last common ancestor of Pan-Homo and the stem hominin. Modern human life histories are even slower, exhibiting distinctively long post-menopausal life spans and later ages at maturity, pointing to a reduction in adult mortality since the Pan-Homo split. We suggest that lower adult mortality, distinctively short interbirth intervals, and early weaning characteristic of modern humans are derived features resulting from cooperative breeding. We evaluate the fidelity of three life history-related variables, body mass, brain growth and dental development, with the life history parameters of living great apes. We found that body mass is the best predictor of great ape life history events. Brain growth trajectories and dental development and eruption are weakly related proxies and inferences from them should be made with caution. We evaluate the evidence of life history-related variables available for extinct species and find that prior to the transitional hominins there is no evidence of any hominin taxon possessing a body size, brain size or aspects of dental development much different from what we assume to be the primitive life history pattern for the Pan-Homo clade. Data for life history-related variables among the transitional hominin grade are consistent and none agrees with a modern human pattern. Aside from mean body mass, adult brain size, crown and root formation times, and the timing and sequence of dental eruption of Homo erectus are inconsistent with that of modern humans. Homo antecessor fossil material suggests a brain size similar to that of Homo erectus s. s., and crown formation times that are not yet modern, though there is some evidence of modern human-like timing of tooth formation and eruption. The body sizes, brain sizes, and dental development of Homo heidelbergensis and Homo neanderthalensis are consistent with a modern human life history but samples are too small to be certain that they have life histories within the modern human range. As more life history-related variable information for hominin species accumulates we are discovering that they can also have distinctive life histories that do not conform to any living model. At least one extinct hominin subclade, Paranthropus, has a pattern of dental life history-related variables that most likely set it apart from the life histories of both modern humans and chimpanzees.

Molecular evidence indicates that the last common ancestor of the genus Pan and the hominin clade existed between 8 and 4 million years ago (Ma). The current fossil record indicates the Pan-Homo last common ancestor existed at least 5 Ma and most likely between 6 and 7 Ma. Together, the molecular and fossil evidence has important consequences for interpreting the evolutionary history of the hand within the tribe Hominini (hominins). Firstly, parsimony supports the hypothesis that the hand of the last common ancestor most likely resembled that of an extant great ape overall (Pan, Gorilla, and Pongo), and that of an African ape in particular. Second, it provides a context for interpreting the derived changes to the hand that have evolved in various hominins. For example, the Australopithecus afarensis hand is likely derived in comparison with that of the Pan–Homo last common ancestor in having shorter fingers relative to thumb length and more proximo-distally oriented joints between its capitate, second metacarpal, and trapezium. This evidence suggests that these derived features evolved prior to the intensification of stone tool-related hominin behaviors beginning around 2.5 Ma. However, a majority of primitive features most likely present in the Pan-Homo last common ancestor are retained in the hands of Australopithecus, Paranthropus/early Homo, and Homo floresiensis. This evidence suggests that further derived changes to the hands of other hominins such as modern humans and Neandertals did not evolve until after 2.5 Ma and possibly even later than 1.5 Ma, which is currently the earliest evidence of Acheulian technology. The derived hands of modern humans and Neandertals may indicate a morphological commitment to tool-related manipulative behaviors beyond that observed in other hominins, including those (e.g. H. floresiensis) which may be descended from earlier tool-making species.

Tali of several hominin taxa are preserved in the fossil record and studies of the external morphology of these often show a mosaic of human-like and ape-like features. This has contributed to a growing recognition of variability characterizing locomotor kinematics of Australopithecus. In contrast, locomotor kinematics of another Plio-Pleistocene hominin, Paranthropus, are substantially less well-documented, in part, because of the paucity of postcranial fossils securely attributed to the genus. Since the talus transmits locomotor-based loads through the ankle and its internal structure is hypothesized to reflect accommodation to such loads, it is a cornerstone structure for reconstructing locomotor kinematics. Here we quantify and characterize trabecular bone morphology within tali attributed to Australopithecus africanus (StW 102, StW 363, StW 486) and Paranthropus robustus (TM 1517), making quantitative comparisons to modern humans, extant non-human apes, baboons, and a hominin talus attributed to Paranthropus boisei (KNM-ER 1464). Using high-resolution images of fossil tali (25 μm voxels), nine trabecular bone subregions of interest beneath the articular surface of the talar trochlea were segmented to quantify localized patterns in distribution and primary strut orientation. It was found that trabecular strut orientation and shape, in some cases, can discriminate amongst species characterized by different locomotor foot kinematics. Discriminant function analyses using standard trabecular bone structural properties align TM 1517 with Pan and Gorilla, while other hominin tali structurally most resemble those of baboons. In primary strut orientation, Paranthropus tali (KNM-ER 1464 and TM 1517) resemble the human condition in the anterior-medial subregion, where strut orientation appears positioned to distribute compressive loads medially and distally toward the talar head. In A. africanus tali (particularly StW 486), primary strut orientation in this region resembles that of apes. These results suggest that Paranthropus may have had a human-like medial weight shift during the last half of stance phase but Australopithecus did not.

We propose that a key change in the evolution of hominids from the last common ancestor shared with chimpanzees was the substitution of plant underground storage organs (USOs) for herbaceous vegetation as fallback foods. Four kinds of evidence support this hypothesis: (1) dental and masticatory adaptations of hominids in comparison with the African apes; (2) changes in australopith dentition in the fossil record; (3) paleoecological evidence for the expansion of USO-rich habitats in the late Miocene; and (4) the co-occurrence of hominid fossils with root-eating rodents. We suggest that some of the patterning in the early hominid fossil record, such as the existence of gracile and robust australopiths, may be understood in reference to this adaptive shift in the use of fallback foods. Our hypothesis implicates fallback foods as a critical limiting factor with far-reaching evolutionary effects. This complements the more common focus on adaptations to preferred foods, such as fruit and meat, in hominid evolution.

A previous study of strontium/calcium (Sr/Ca) ratios in Paranthropus suggested that it consumed more animal foods than was previously believed. However, that study looked at Sr/Ca in fossil bone, which is known to be highly susceptible to diagenesis. Enamel, in contrast, is resistant to post-mortem alteration making it a more appropriate material for Sr/Ca analysis of Plio-Pleistocene fossils. Yet, we know virtually nothing about Sr/Ca in the enamel of modern African mammals, much less fossil taxa. To address this gap, we studied Sr/Ca in tooth enamel from modern mammals in the greater Kruger National Park, South Africa, as well as fossil fauna from the Sterkfontein Valley. Grazing herbivores have the highest Sr/Ca, followed by browsers and carnivores in both modern and fossil fauna. This similarity in ecological Sr/Ca patterning between modern and fossil fauna shows that diagenesis has not obscured the primary dietary signals. Australopithecus has significantly higher Sr/Ca than Paranthropus, and higher Sr/Ca than fossil papionins, browsers, and carnivores. Paranthropus has lower Sr/Ca than grazers, but its Sr/Ca is higher or equal to that of fossil papionins, browsers, and carnivores. Thus, Sr/Ca for both hominins is relatively high, and provides no direct evidence for omnivory in either taxon. The consumption of underground resources or insects are among the possible explanations for the highly elevated Sr/Ca in Australopithecus.

The relationship between climatic change and human evolution can be framed in terms of three major hypotheses. A modern version of the long-held savanna hypothesis posits that the expansion of grassland ecosystems in Africa was driven by global climatic change and led to the divergence of hominins from the apes and to the origin of the Homo clade. A related idea suggests that hominins originated in the late Miocene, and Homo in the late Pliocene, as constituents of broader pulses of faunal turnover synchronized by episodes of global climatic change. A more recent concept, the variability selection hypothesis, emphasizes the importance of fluctuating climates and environments, rather than any single trend, in shaping human adaptation and evolution. Here we evaluate these ideas for the Plio-Pleistocene in light of new analyses of fossil mammals from the Turkana Basin of Kenya and Ethiopia. Our results show that between 4 and 1 Ma (million years ago), there were profound faunal changes in the Turkana Basin. The most important of these changes include significant shifts in the abundance of the common families of mammals, episodes of high faunal turnover, and an increase in the number and abundance of species that show adaptations to grassland ecosystems. Episodes of relatively high faunal turnover occurred in the intervals 3.4–3.2, 2.8– 2.6, 2.4–2.2, and 2.0–1.8 Ma. Paranthropus and Homo appear in the Turkana Basin during successive intervals of high turnover at 2.8–2.6 and at 2.4–2.2 Ma, while the appearance of Homo erectus is coupled to a major episode of turnover and grassland expansion after 2 Ma. Thus, there was not a single turnover pulse of relevance to late Pliocene hominins, but multiple events that successively led to the appearance of Paranthropus, early Homo, and H. erectus. Our results also show evidence of large-scale, 100 ky-periodicity shifts in the fauna beginning at 2.5 Ma, during the time that Homo and lithic artifacts first appear in the Turkana Basin, lending support to the variability selection hypothesis [Science 273 (1996) 922; Potts R., 1996b. Humanity’s Descent: The Consequences of Ecological Instability. Avon Books, New York.] during the latest Pliocene. The savanna hypothesis may not explain the divergence of hominins from other apes, but it could be correct in stressing the importance of grasslands to the early evolution of Homo. The fundamental importance of grasslands may lie in the complexity and heterogeneity they added to the range of habitats available to the early species of the genus Homo. The turnover pulse hypothesis [Vrba, E.S., 1988. Late Pliocene climatic events and hominid evolution. In: Grine, F.E (Ed.). Evolutionary History of the ‘‘Robust’’ Australopithecines. Aldine, New York, pp. 405–426; Vrba, E.S., 1995. The fossil record of African antelopes (Mammalia, Bovidae) in relation to human evolution and paleoclimate. In: Vrba, E.S., Denton, G.H., Partridge, T.C., Burckle, L.H. (Eds.). Paleoclimate and Evolution, with Emphasis on Human Origins. Yale Univ. Press, New Haven, pp. 385–424.] may be correct in linking critical events in human evolution to broader pulses of faunal change ultimately driven by climate, but our