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Objectives: Size-corrected tooth crown measurements were used to estimate phenetic affinities among Homo naledi (~335-236 ka) and 11 other Plio-Pleistocene and recent species. To assess further their efficacy, and identify dental evolutionary trends, the data were then quantitatively coded for phylogenetic analyses. Results from both methods contribute additional characterization of H. naledi relative to other hominins., Materials and Methods: After division by their geometric mean, scaled mesiodistal and buccolingual dimensions were used in tooth size apportionment analysis to compare H. naledi with Australopithecus africanus, A. afarensis, Paranthropus robustus, P. boisei, H. habilis, H. ergaster, H. erectus, H. heidelbergensis, H. neanderthalensis, H. sapiens, and Pan troglodytes. These data produce equivalently scaled samples unaffected by interspecific size differences. The data were then gap-weighted for Bayesian inference., Results: Congruence in interspecific relationships is evident between methods, and with many inferred from earlier systematic studies. However, the present results place H. naledi as a sister taxon to H. habilis, based on a symplesiomorphic pattern of relative tooth size. In the preferred Bayesian phylogram, H. naledi is nested within a clade comprising all Homo species, but it shares some characteristics with australopiths and, particularly, early Homo., Discussion: Phylogenetic analyses of relative tooth size yield information about evolutionary dental trends not previously reported in H. naledi and the other hominins. Moreover, with an appropriate model these data recovered plausible evolutionary relationships. Together, the findings support recent study suggesting H. naledi originated long before the geological date of the Dinaledi Chamber, from which the specimens under study were recovered., (© 2021 The Authors. American Journal of Physical Anthropology published by Wiley Periodicals LLC.)

Objectives: One of the most contentious issues in paleoanthropology is the nature of the last common ancestor of humans and our closest living relatives, chimpanzees and bonobos (panins). The numerical composition of the vertebral column has featured prominently, with multiple models predicting distinct patterns of evolution and contexts from which bipedalism evolved. Here, we study total numbers of vertebrae from a large sample of hominoids to quantify variation in and patterns of regional and total numbers of vertebrae in hominoids., Materials and Methods: We compile and study a large sample (N = 893) of hominoid vertebral formulae (numbers of cervical, thoracic, lumbar, sacral, caudal segments in each specimen) and analyze full vertebral formulae, total numbers of vertebrae, and super-regional numbers of vertebrae: presacral (cervical, thoracic, lumbar) vertebrae and sacrococcygeal vertebrae. We quantify within- and between-taxon variation using heterogeneity and similarity measures derived from population genetics., Results: We find that humans are most similar to African apes in total and super-regional numbers of vertebrae. Additionally, our analyses demonstrate that selection for bipedalism reduced variation in numbers of vertebrae relative to other hominoids., Discussion: The only proposed ancestral vertebral configuration for the last common ancestor of hominins and panins that is consistent with our results is the modal formula demonstrated by chimpanzees and bonobos (7 cervical-13 thoracic-4 lumbar-6 sacral-3 coccygeal). Hox gene expression boundaries suggest that a rostral shift in Hox10/Hox11-mediated complexes could produce the human modal formula from the proposal ancestral and panin modal formula., (© 2021 Wiley Periodicals LLC.)

Objectives: Increased use of three-dimensional (3D) imaging data has led to a need for methods capable of capturing rich shape descriptions. Semi-landmarks have been demonstrated to increase shape information but placement in 3D can be time consuming, computationally expensive, or may introduce artifacts. This study implements and compares three strategies to more densely sample a 3D image surface., Materials and Methods: Three dense sampling strategies: patch, patch-thin-plate spline (TPS), and pseudo-landmark sampling, are implemented to analyze skulls from three species of great apes. To evaluate the shape information added by each strategy, the semi or pseudo-landmarks are used to estimate a transform between an individual and the population average template. The average mean root squared error between the transformed mesh and the template is used to quantify the success of the transform., Results: The landmark sets generated by each method result in estimates of the template that on average were comparable or exceeded the accuracy of using manual landmarks alone. The patch method demonstrates the most sensitivity to noise and missing data, resulting in outliers with large deviations in the mean shape estimates. Patch-TPS and pseudo-landmarking provide more robust performance in the presence of noise and variability in the dataset., Conclusions: Each landmarking strategy was capable of producing shape estimations of the population average templates that were generally comparable to manual landmarks alone while greatly increasing the density of the shape information. This study highlights the potential trade-offs between correspondence of the semi-landmark points, consistent point spacing, sample coverage, repeatability, and computational time., (© 2021 Wiley Periodicals LLC.)

Objectives: One role of dental pulp is in the upkeep and maintenance of dentine. Under wear, odontoblasts in the pulp deposit tertiary dentine to ensure the sensitive internal dental tissues are not exposed and vulnerable to infection. It follows that there may be an adaptive advantage for increasing molar pulp volume in anthropoid primate taxa that are prone to high levels of wear. The relative volume of dental pulp is therefore predicted to covary with dietary abrasiveness (in the sense of including foods that cause high degrees of wear)., Materials and Methods: We examined relatively unworn lower second molars in pairs of species of extant hominoids, cebids, and pitheciids that vary in the abrasiveness of their diet (n = 36). Using micro-CT scans, we measured the percent of tooth that is pulp (PTP) as the ratio of pulp volume to that of the total volume of the tooth., Results: We found that in each pair of species, the taxa that consume a more abrasive diet had a significantly higher PTP than the closely related taxa that consume a softer diet., Conclusions: Our results point to an adaptive mechanism in the molars of taxa that consume abrasive diets and are thus subject to higher levels of wear. Our results provide additional understanding of the relationship between dental pulp and diet and may offer insight into the diet of extinct taxa such as Paranthropus boisei or into the adaptive context of the taurodont molars of Neanderthals., (© 2021 Wiley Periodicals LLC.)

Objectives: Many primates change their locomotor behavior as they mature from infancy to adulthood. Here we investigate how long bone cross-sectional geometry in Pan, Gorilla, Pongo, Hylobatidae, and Macaca varies in shape and form over ontogeny, including whether specific diaphyseal cross sections exhibit signals of periosteal adaptation or canalization., Materials and Methods: Diaphyseal cross sections were analyzed in an ontogenetic series across infant, juvenile, and adult subgroups. Three-dimensional laser-scanned long bone models were sectioned at midshaft (50% of biomechanical length) and distally (20%) along the humerus and femur. Traditional axis ratios acted as indices of cross-sectional circularity, while geometric morphometric techniques were used to study cross-sectional allometry and ontogenetic trajectory., Results: The humeral midshaft is a strong indicator of posture and locomotor profile in the sample across development, while the mid-femur appears more reflective of shifts in size. By comparison, the distal diaphyses of both limb elements are more ontogenetically constrained, where periosteal shape is largely static across development relative to size, irrespective of a given taxon's behavior or ecology., Discussion: Primate limb shape is not only highly variable between taxa over development, but at discrete humeral and femoral diaphyseal locations. Overall, periosteal shape of the humeral and femoral midshaft cross sections closely reflects ontogenetic transitions in behavior and size, respectively, while distal shape in both bones appears more genetically constrained across intraspecific development, regardless of posture or size. These findings support prior research on tradeoffs between function and safety along the limbs., (© 2020 Wiley Periodicals LLC.)

Objectives: Molar crenulation is defined as the accessory pattern of grooves that appears on the occlusal surface of many mammalian molars. Although frequently used in the characterization of species, this trait is often assessed qualitatively, which poses unavoidable subjective biases. The objective of this study is to quantitatively test the variability in the expression of molar crenulation in primates and its association with molar size and diet., Methods: The variability in the expression of molar crenulation in hominids (human, chimpanzee, gorilla, and orangutan) was assessed with fractal analysis using photographs of first, second and third upper and lower molars. After this, representative values for 29 primate species were used to evaluate the correlation between molar complexity, molar size, and diet using a phylogenetic generalized least squares regression., Results: Results show that there are statistically significant differences in fractal dimensions across hominid species in all molars, with orangutan molars presenting higher values of occlusal complexity. Our results indicate that there is no significant association between molar complexity and molar size or diet., Discussion: Our results show higher levels of occlusal complexity in orangutans, thus supporting previously published observations. Our analyses, however, do not indicate a clear association between molar complexity and molar size or diet, pointing to other factors as the major drivers of complexity. To our knowledge, our study is the first one to use fractal analysis to measure occlusal complexity in primates. Our results show that this approach is a rapid and cost-effective way to measure molar complexity., (© 2020 Wiley Periodicals LLC.)

Objectives: We aimed to investigate molar enamel development in fossil orangutans from Guangxi and shed light on the evolution of Asian great apes., Materials and Methods: We collected 32 fossil orangutan molars, most of which were from Guangxi apothecaries and the Guangxi Daxin Heidong cave, and prepared histological sections of each molar. We then characterized aspects of dental development, including long period line periodicity, number of Retzius lines and lateral enamel formation time, cuspal enamel thickness, and enamel formation time., Results: The long period line periodicity in fossil orangutans ranged from 9 to 10 days (mean, 9.09 days). The molar lateral enamel formation time ranged from 1.48 to 3.17 years (540-1,152 days). Cuspal enamel thickness in fossil orangutan molars ranged from 949 to 2,535 μm, and cuspal enamel formation time ranged from 0.64 to 1.87 years. Molar enamel formation time of fossil orangutans ranged from 2.47 to 4.67 years., Discussion: Long-period line periodicity of fossil orangutans from Guangxi was within the variation range of extant orangutans, and the average long period line periodicity (9.09 days) of fossil orangutans from Guangxi in this study was lower than the values for extant orangutans (9.5 days) and fossil orangutans (10.9 days) from Sumatra and Vietnam. Orangutan enamel thickness may have gradually decreased from the Middle Pleistocene to Holocene. Crown formation time of fossil orangutans was slightly longer than that of extant orangutans, and the M1 emergence age of fossil orangutans from Guangxi was about 4-6 years. These findings might indicate the regional difference or evolutionary changes in orangutans since Pleistocene. Dental development of the Guangxi fossil orangutans were more similar to that of Asian Miocene apes, suggesting the closer evolutionary relationship of orangutans to Miocene Asian fossil apes., (© 2020 Wiley Periodicals LLC.)

Objectives: The purpose of this study is to help elucidate the taxonomic relationship between Homo naledi and other hominins., Materials and Methods: Homo naledi deciduous maxillary and mandibular molars from the Dinaledi Chamber, South Africa were compared to those of Australopithecus africanus, Australopithecus afarensis, Paranthropus robustus, Paranthropus boisei, early Homo sp., Homo erectus, early Homo sapiens, Upper Paleolithic H. sapiens, recent southern African H. sapiens, and Neanderthals by means of morphometric analyses of crown outlines and relative cusp areas. The crown shapes were analyzed using elliptical Fourier analyses followed by principal component analyses (PCA). The absolute and relative cusp areas were obtained in ImageJ and compared using PCA and cluster analyses., Results: PCA suggests that the crown shapes and relative cusp areas of mandibular molars are more diagnostic than the maxillary molars. The H. naledi deciduous mandibular first and second molar (dm 1 and dm 2 ) do not have a strong affinity to any taxon in the comparative sample in all analyses. While the H. naledi dm 2 plots as an outlier in the relative cusp analysis, the H. naledi specimen fall closest to Australopithecus due to their relatively large metaconid, a primitive trait for the genus Homo. Although useful for differentiating Neanderthals from recent southern African H. sapiens and UP H. sapiens, the PCA of the relative cusp areas suggests that the deciduous maxillary second molars (dm 2 ) do not differentiate other groups. The three H. naledi dm 2 cuspal areas are variable and fall within the ranges of other Homo, as well as Australopithecus, and Paranthropus suggesting weak diagnostic utility., Discussion: This research provides another perspective on the morphology of, and variation within, H. naledi. The H. naledi deciduous molars do not consistently align with any genus or species in the comparative sample in either the crown shape or relative cusp analyses. This line of inquiry is consistent with other cranial and postcranial studies suggesting that H. naledi is unique., (© 2020 Wiley Periodicals LLC.)

Objectives: The paleontological description and comparative analysis using discrete morphology, morphometrics (linear and geometric) and cross-sectional geometry of three femoral diaphyseal sections from the Middle Pleistocene site of Hualongdong, China., Materials and Methods: The material consists of the original Hualongdong femoral fossils and available data on femoral diaphyses from Middle and Late Pleistocene archaic humans and Middle and earlier Upper Paleolithic modern humans. The methods include visual observation, diaphyseal diameters, cross-sectional parameters (transverse areas and second moments of area derived from micro-CT scans), and geometric morphometrics using semilandmark data., Results: The Hualong 11 midshaft section is similar to other Middle and Late Pleistocene archaic humans in being transversely broad and lacking a pilaster despite a prominent linea aspera. It clusters principally with archaic human femora in all measured parameters. The Hualong 15 and 16 subtrochanteric pieces are similar to many Middle Pleistocene and early modern human femora in being transversely broad. In particular, Hualong 15 exhibits a prominent lateral (gluteal) buttress, similar to many Upper Paleolithic femora but also the Lazaret and Krapina archaic ones. In addition, Hualong 15 has a small third trochanter, a common Upper Paleolithic but rare earlier feature., Discussion: The Hualong 11 femoral piece reinforces the general Middle Pleistocene pattern, especially for eastern Eurasia from which archaic human femora are rare. The subtrochanteric proportions of Hualong 15 and 16 reinforce the Early Pleistocene and (generally) Middle Pleistocene pattern of bone distributions, but their subperiosteal contours align them (along with those of the Lazaret and Krapina femora) with Upper Paleolithic ones. It is difficult to account for these proportions from the generally broad pelves of Pleistocene archaic humans., (© 2020 Wiley Periodicals LLC.)

Objectives: The primate foot has been extensively investigated because of its role in weight-bearing; however, the calcaneus has been relatively understudied. Here we examine entire gorilla calcaneal external shape to understand its relationship with locomotor behavior., Materials and Methods: Calcanei of Gorilla gorilla gorilla (n = 43), Gorilla beringei graueri (n = 20), and Gorilla beringei beringei (n = 15) were surface or micro-CT scanned. External shape was analyzed through a three-dimensional geometric morphometric sliding semilandmark analysis. Semilandmarks were slid relative to an updated Procrustes average in order to minimize the bending energy of the thin plate spline interpolation function. Shape variation was summarized using principal components analysis of shape coordinates. Procrustes distances between taxa averages were calculated and resampling statistics run to test pairwise differences. Linear measures were collected and regressed against estimated body mass., Results: All three taxa exhibit statistically different morphologies (p < .001 for pairwise comparisons). G. g. gorilla demonstrates an anteroposteriorly elongated calcaneus with a deeper cuboid pivot region and mediolaterally flatter posterior talar facet. G. b. beringei possesses the flattest cuboid and most medially-angled posterior talar facets. G. b. graueri demonstrates intermediate articular facet morphology, a medially-angled tuberosity, and an elongated peroneal trochlea., Discussion: Articular facet differences separate gorillas along a locomotor gradient. G. g. gorilla is adapted for arboreality with greater joint mobility, while G. b. beringei is adapted for more stereotypical loads associated with terrestriality. G. b. graueri's unique posterolateral morphology may be due to a secondary transition to greater arboreality from a more terrestrial ancestor., (© 2020 Wiley Periodicals LLC.)

Objectives: The Krapina rock shelter has yielded a large assemblage of early Neandertals. Although endocranial volume (ECV) has been estimated for four individuals from the site, several published values that appear in the literature warrant revisiting., Materials and Methods: We used virtual methods, including high-resolution surface models of fossils and 3D geometric morphometrics, to reconstruct endocasts and estimate ECV for five Krapina crania. We generated 10 reconstructions of each endocast to quantify missing data uncertainty. To assess the method and our ECV estimates, we applied these techniques to the Spy II Neandertal, and estimated ECV of a human reference endocast simulating the missing data of the Krapina fossils., Results: We obtained an average ECV estimate of 1,526 cm 3 for Spy II, consistent with previous research. Estimated ECV of juveniles Krapina 1 and 2 average 1,419 and 1,286 cm 3 , respectively. Estimates for the relatively complete adults Krapina 3 and 6 range from 1,247 to 1,310 cm 3 and 1,135 to 1,207 cm 3 , respectively, while the more fragmentary Krapina 5 averaged 1,397 cm 3 . The missing data simulation suggests more fragmentary crania yield more uncertain and possibly overestimated ECVs., Conclusions: We have provided new estimates of brain size of the Krapina Neandertals, including the first estimates for Krapina 2. Brain size at Krapina was similar to other pre-Würm Neandertals, within the range of but lower than the average of later Neandertals. Although the virtual approach overcomes many challenges of fossil preservation, our results are nevertheless subject to future revision., (© 2020 Wiley Periodicals LLC.)

Objectives: Enamel thickness features prominently in hominoid evolutionary studies. To date, however, studies of enamel thickness in humans, great apes, and their fossil relatives have focused on the permanent molar row. Comparatively little research effort has been devoted to tissue proportions within deciduous teeth. Here we attempt to fill this gap by documenting enamel thickness variation in the deciduous dentition of extant large-bodied hominoids., Materials and Methods: We used microcomputed tomography to image dental tissues in 80 maxillary and 78 mandibular deciduous premolars of Homo sapiens, Pan troglodytes, Gorilla, and Pongo. Two-dimensional virtual sections were created from the image volumes to quantify average (AET) and relative (RET) enamel thickness, as well as its distribution across the crown., Results: Our results reveal no significant differences in enamel thickness among the great apes. Unlike the pattern present in permanent molars, Pongo does not stand out as having relatively thicker-enameled deciduous premolars than P. troglodytes and Gorilla. Humans, on the other hand, possess significantly thicker deciduous premolar enamel in comparison to great apes. Following expectations from masticatory biomechanics, we also find that the "functional" side (protocone, protoconid) of deciduous premolars generally possesses thicker enamel than the "nonfunctional" side., Discussion: Our study lends empirical support to anecdotal observations that patterns of AET and RET observed for permanent molars of large-bodied apes do not apply to deciduous premolars. By documenting enamel thickness variation in hominoid deciduous teeth, this study provides the comparative context to interpret rates and patterns of wear of deciduous teeth and their utility in life history reconstructions., (© 2020 Wiley Periodicals LLC.)

Objectives: The primary aim of this study was to describe the insertion sites of the ligaments holding the flexor digitorum profundus and superficialis muscles (flexor ridges) in proximal phalanges 2-5 of African apes and modern humans. To interpret differences in flexor ridge size based on general behavioral differences among taxa., Materials and Methods: We analyzed 3D models of manual proximal phalanges 2-5 from 29 gorillas (Gorilla beringei and Gorilla gorilla), 30 chimpanzees (Pan troglodytes) and 36 recent modern humans. Flexor ridges (mm 2 ) were compared within and across genera., Results: Gorillas and chimpanzees had larger flexor ridges for phalanges 2-4 than humans and this difference subsists when controlling for body size. Each genus had a unique insertion size pattern across the digits, with the most heterogeneous pattern found in chimpanzees, followed by humans, and lastly gorillas. These patterns corresponded strongly to the differences in the size of the phalanges within each genus, except for phalanx 5 in humans, which had a larger flexor ridge than expected., Discussion: When comparing these genera, the flexor ridges signal differences between taxa that use their hands for manipulation and locomotion (gorillas and chimpanzees) and taxa that use them exclusively for manipulation (humans). This functional signal was also apparent in the PP5 of humans, whose larger FR may be indicating the high recruitment of this digit during forceful precision grip characteristic of humans., (© 2020 Wiley Periodicals LLC.)

Objectives: While many attempts have been made to estimate body mass in hominins from lower limb bone dimensions, the upper limb has received far less attention in this regard. Here we develop new body mass estimation equations based on humeral articular breadths in a large modern human sample and apply them to 95 Plio-Pleistocene specimens., Materials and Methods: Humeral head superoinferior and total distal articular mediolateral breadths were measured in a morphologically diverse sample of 611 modern human skeletons whose body masses were estimated from bi-iliac breadth and reconstructed stature. Reduced major axis regressions were used to compute body mass estimation equations. Consistency of the resulting estimates with those derived previously using lower limb bone equations was assessed in matched Plio-Pleistocene individuals or samples., Results: In the modern reference sample, the new humeral body mass estimation equations exhibit only slightly lower precision compared to the previously derived lower limb bone equations. They give generally similar estimates for Pleistocene Homo, after accounting for the different shape of the humeral head articular surface in archaic Middle and Late Pleistocene Homo, except for distal humeral estimates for Late Pleistocene specimens, which average somewhat below lower limb estimates. Humeral equations give body mass estimates for australopiths that appear much too high, except for Australopithecus sediba. A chimpanzee-based distal humeral articular formula appears to work well for larger australopith specimens., Discussion: The new formulae provide a more secure foundation for estimating hominin body mass from humeri than previously available equations., (© 2020 Wiley Periodicals, Inc.)

Objectives: Because trabecular bone volume fraction (BV/TV) is influenced by variations in physical activity recent declines in BV/TV in humans are often attributed to modern sedentary lifestyles. This study tests the hypothesis that presumed variations in mechanical loading between groups can predict the observed BV/TV patterns in humans, chimpanzees and gorillas in two bones: the calcaneus which experiences high and well characterized impact forces, and the C2 vertebrae which experiences reduced locomotor forces., Materials and Methods: BV/TV and other structural variables were quantified from high-resolution microCT scans in gorillas, chimpanzees, and four Homo sapiens populations: Pleistocene, semi-sedentary Natufians; Holocene hunter-gatherers from Point Hope, Alaska; Holocene nomadic pastoralists from medieval Europe; and modern, sedentary Americans., Results: In the calcaneal tuberosity, Natufian BV/TV was 36, 46, and 46% greater than Alaskans (p = .02), Europeans (p = .005) and modern Americans (p = .002), respectively, but not significantly different from apes. BV/TV was not significantly different between modern Americans and Alaskans or Europeans. In the C2, Natufian BV/TV was 53 and 25% greater than in the Alaskan (p = .0001) and European (p = .048) populations., Discussion: These results suggest that phenomena other than or in addition to variations in physical activity are needed to explain BV/TV patterns observed in H. sapiens, and point to a systemic decline in H. sapiens BV/TV after the Pleistocene., (© 2020 Wiley Periodicals LLC.)

Objectives: The Pleistocene taxon Paranthropus robustus was established in 1938 following the discovery at Kromdraai B, South Africa, of the partial cranium TM 1517a and associated mandible TM 1517b. Shortly thereafter, a distal humerus (TM 1517g), a proximal ulna (TM 1517e), and a distal hallucial phalanx (TM 1517k) were collected nearby at the site, and were considered to be associated with the holotype. TM 1517a-b represents an immature individual; however, no analysis of the potentially associated postcranial elements has investigated the presence of any endostructural remnant of recent epiphyseal closure. This study aims at tentatively detecting such traces in the three postcranial specimens from Kromdraai B., Materials and Methods: By using μXCT techniques, we assessed the developmental stage of the TM 1517b's C-M3 roots and investigated the inner structure of TM 1517g, TM 1517e, and TM 1517k., Results: The M2 shows incompletely closed root apices and the M3 a half-completed root formation stage. The distal humerus was likely completely fused, while the proximal ulna and the distal hallucial phalanx preserve endosteal traces of the diaphyseo-epiphyseal fusion process., Discussion: In the hominin fossil record, there are few unambiguously associated craniodental and postcranial remains sampling immature individuals, an essential condition for assessing the taxon-specific maturational patterns. Our findings corroborate the original association of the craniodental and postcranial remains representing the P. robustus type specimen. As with other Plio-Pleistocene hominins, the odonto-postcranial maturational pattern of TM 1517 more closely fits an African great ape rather than the extant human pattern., (© 2020 The Authors. American Journal of Physical Anthropology published by Wiley Periodicals, Inc.)

Objectives: The dentition of Olduvai Hominid 1 (OH1) exhibits an anomalous pattern of dental wear that was originally attributed to either intentional cultural modification (filing) or plant processing behaviors. A differential diagnosis of the wear and assessment of the biological affinity of OH1 is presented., Materials and Methods: Macroscopic and microscopic observations of all labial and buccal tooth surfaces were undertaken to assess wear patterns. A multivariate analysis of mandibular morphology of OH1 compared to other Late Pleistocene, Holocene, and recent modern humans was used to ascertain biological affinity., Results: The morphological variation of the OH1 mandible is closely aligned with variation in penecontemporaneous fossils from Africa and outside that of recent humans. The concave wear facets exposing dentin on the labial surfaces of all three preserved mandibular incisors is confirmed. Substantial loss of labial/buccal surfaces was documented on the surfaces of all in situ maxillary and mandibular canines, premolars, and molars ranging from distinct facets with well-defined edges, to blunting or "polishing" around areas of maximum buccal curvature. The wear on both the anterior and postcanine teeth closely resemble that caused by adornments ("labrets") worn in lower-lip and buccal facial piercings known from bioarchaeological and ethnographic contexts. The wear pattern suggests that the OH1 wore three facial piercings-two buccal/lateral and a medial one in the lower lip., Discussion: Our findings suggest that the expression of social identities through intentional body modification is more diverse than previously documented elsewhere in Africa during the Late Pleistocene (i.e., ablation) and Early Holocene (i.e., ablation, chipping, and filing)., (© 2020 Wiley Periodicals, Inc.)

Objective: Distal femoral metaphyseal surface morphology is highly variable in extant mammals. This variation has previously been linked to differences in locomotor behavior. We perform the first systematic survey and description of the development of this morphology in extant hominoids., Materials and Methods: We collected 3D surface laser scans of the femora of 179 human and great ape individuals throughout all subadult stages of development. We qualitatively and quantitatively describe metaphyseal surface morphology., Results: We find that the metaphysis is topographically simple in all hominoids during the fetal and infant periods relative to later developmental periods, and in apes it develops significant complexity throughout development. Humans, by contrast, retain relatively flat metaphyseal surfaces throughout ontogeny., Discussion: Major shifts in morphology appear to coincide with major shifts in locomotor behavior, suggesting that metaphyseal morphology is developmentally plastic and highly dependent on the biomechanical loadings at the knee joint. This is consistent with a large body of biomedical research, which demonstrates the primacy of mechanical forces in determining growth plate ossification patterns. Additionally, specific metaphyseal morphology appears highly correlated with specific locomotor modes, suggesting that metaphyseal surface morphology will be useful for reconstructing the locomotor behavior of fossil primate taxa., (© 2020 Wiley Periodicals, Inc.)

Objectives: The dexterity of fossil hominins is often inferred by assessing the comparative manual anatomy and behaviors of extant hominids, with a focus on the thumb. The aim of this study is to test whether trabecular structure is consistent with what is currently known about habitually loaded thumb postures across extant hominids., Materials and Methods: We analyze first metacarpal (Mc1) subarticular trabecular architecture in humans (Homo sapiens, n = 10), bonobos (Pan paniscus, n = 10), chimpanzees (Pan troglodytes, n = 11), as well as for the first time, gorillas (Gorilla gorilla gorilla, n = 10) and orangutans (Pongo sp., n = 1, Pongo abelii, n = 3 and Pongo pygmaeus, n = 5). Using a combination of subarticular and whole-epiphysis approaches, we test for significant differences in relative trabecular bone volume (RBV/TV) and degree of anisotropy (DA) between species., Results: Humans have significantly greater RBV/TV on the radiopalmar aspects of both the proximal and distal Mc1 subarticular surfaces and greater DA throughout the Mc1 head than other hominids. Nonhuman great apes have greatest RBV/TV on the ulnar aspect of the Mc1 head and the palmar aspect of the Mc1 base. Gorillas possessed significantly lower DA in the Mc1 head than any other taxon in our sample., Discussion: These results are consistent with abduction of the thumb during forceful "pad-to-pad" precision grips in humans and, in nonhuman great apes, a habitually adducted thumb that is typically used in precision and power grips. This comparative context will help infer habitual manipulative and locomotor grips in fossil hominins., (© 2019 Wiley Periodicals, Inc.)

Objective: This study aims to develop a comparative basis for assessing the developmental stage of KNM-ER 42700 based on the ontogenetic pattern of the ectocranial surface of the basicranium in modern humans and chimpanzees., Materials and Methods: A total of 33 landmarks were collected from an ontogenetic sample of modern humans (80), chimpanzees (51), and 12 individuals classified as Homo erectus s.l. Ontogenetic trajectories were analyzed, and common aspects were extracted for the purpose of discriminating age groups. A regression of size on the extracted shape variables was used to investigate common ontogenetic allometry., Results: The basicranial development of chimpanzees and humans follows different trajectories; however, similarities are also present. The common shape component of development extracted can be used to define age groups in both chimpanzees and modern humans. The extracted shape component presents a similar ontogenetic and static-allometric pattern in these two species. The developmental stages of H. erectus s.l. specimens were attributed following these common traits. Our analysis correctly assigned developmental stages to those specimens of H. erectus for which developmental ages are known., Discussion: The component used for assessing the developmental stage has an ontogenetic allometric component. However, this shape component can discriminate age group irrespective of size and is no longer related to size when static allometry is considered. Adult H. erectus s.l. specimens were attributed to the adult category. KNM-WT 15000 fell with the late juvenile age group, whereas D2700 plotted in the region of overlap between the juvenile and adult age groups and Mojokerto with the younger age groups, as predicted by their known developmental ages. KNM-ER 42700 fell within the adult variability despite its incompletely fused spheno-occipital synchondrosis., (© 2019 The Authors. American Journal of Physical Anthropology published by Wiley Periodicals, Inc.)

Objectives: The taxonomic status of isolated hominoid teeth from the Asian Pleistocene has long been controversial due to difficulties distinguishing between pongine and hominin molars given their high degree of morphometrical variation and overlap. Here, we combine nonmetric and geometric morphometric data to document a dental pattern that appears to be taxonomically diagnostic among Pongo. We focus on the protoconule, a cuspule of well-documented evolutionary history, as well as on shape differences of the mesial fovea of the upper molars., Materials and Methods: We examined 469 upper molars of eight hominid genera (Australopithecus, Paranthropus, Homo, Meganthropus, Sivapithecus, Pan, Gorilla, and Pongo), including representatives of Homo erectus and extinct and recent Pongo. Analyses were conducted at the enamel-dentine junction to overcome the limitations introduced by dental wear., Results: We found that a moderate or large protoconule is present in ~80% of Pleistocene and extant Pongo. Conversely, a moderate to pronounced protoconule in hominins, Meganthropus, and African great apes occurs in low frequencies (~0-20%). Canonical variate analyses for the mesial fovea show that Pleistocene and extant Pongo cluster together and are clearly differentiated from all other groups, except for Sivapithecus., Discussion: This study suggests that the protoconule and the shape of the mesial fovea in upper molars are useful features for the taxonomic identification of isolated hominid teeth. By identifying these new features, our results can contribute to the better understanding of hominoid evolutionary history and biogeography during the Asian Pleistocene. However, we emphasize that the reported features should be used in combination with other diagnostic variables for the most accurate taxonomic assessments., (© 2019 Wiley Periodicals, Inc.)

Objectives: Morphological variation within the southern African hypodigm of Paranthropus has been the focus of major interest since the earliest discoveries in the "Cradle of Humankind." Given the relevance of the bony labyrinth for investigating fossil primate paleobiodiversity, this article aims to provide additional evidence for assessing the degree of regional variation within Paranthropus through the comparative analysis of the inner ear of DNH 22., Materials and Methods: As comparative material, 18 southern African hominin specimens from Sterkfontein, Swartkrans, and Makapansgat (plus published data from Kromdraai B), attributed to Australopithecus, early Homo or Paranthropus, as well as 10 extant human and 10 extant common chimpanzee specimens are investigated. A landmark-based geometric morphometric method is applied for quantitatively assessing labyrinthine morphology. Additionally, cochlear parameters and oval window area are measured., Results: In terms of semicircular canal and cochlear shape, DNH 22 most resembles the Paranthropus specimen SKW 18 from Swartkrans. Both specimens differ from the other Paranthropus specimens investigated in this study by an anteroposteriorly large posterior semicircular canal and a cochlea with loose turns in the apical portion. Conversely, the oval window area in DNH 22 closely fits the range observed in Paranthropus from Swartkrans and Kromdraai B., Discussion: The inner ear of the DNH 22 specimen represents a unique opportunity to provide further insight into the early hominin labyrinthine variation pattern. In particular, the description of DNH 22 raises critical questions on the diversity of the vestibular system and evolutionary pattern of the auditory apparatus in Paranthropus., (© 2019 Wiley Periodicals, Inc.)

Objectives: Topographic estimates of dental relief are now commonly used to make dietary inferences from the teeth of extant and extinct primates. We thoroughly compared commonly used relief estimates in an effort to help researchers decide which variable best suits their objectives., Materials and Methods: We combined a total of three datasets: five theoretical models built to compare the effect of tooth complexity and basin depth on relief estimates, a dataset of 110 lower molars of prosimians, and a dataset of 25 upper molars of apes. We investigated intra-mesh variation and tooth average relief, estimated from slope and three different relief indices, according to four criteria: (1) the ability to map relief on topographic maps, (2) the correlation with other relief estimates, (3) the ability to separate high-relief molars of folivores from deep-relief molars of insectivores in prosimians, and (4) the influence of surface complexity on relief estimates in apes., Results: We found that polygon slope and relief index are linked by a mathematical relation. Tooth average slope and all relief indices are strongly correlated. In contrast, relief estimates are moderately correlated to cusp elevation. One relief index of four relief estimates had an excellent ability to separate high-relief from deep-relief molars in prosimians, whereas slope could not separate them. No significant effect of tooth complexity on dental relief could be detected in apes., Conclusions: Because slope and relief indices are highly correlated, it is strongly recommended not to combine them in multivariate analysis. Still, slope and relief indices show interesting differences in scaling, graphical representation, computation method, and ability to separate high-relief and deep-relief molars. Our results also suggest that slope and relief indices can vary independently of tooth complexity and are moderately affected by mean cusp elevation in apes., (© 2019 Wiley Periodicals, Inc.)

Objectives: The femoral remains recovered from the Lesedi Chamber are among the most complete South African fossil hominin femora discovered to date and offer new and valuable insights into the anatomy and variation of the bone in Homo naledi. While the femur is one of the best represented postcranial elements in the H. naledi assemblage from the Dinaledi Chamber, the fragmentary and commingled nature of the Dinaledi femoral remains has impeded the assessment of this element in its complete state., Materials and Methods: Here we analyze and provide descriptions of three new relatively well-preserved femoral specimens of H. naledi from the Lesedi Chamber: U.W. 102a-001, U.W. 102a-003, and U.W. 102a-004. These femora are quantitatively and qualitatively compared to multiple extinct hominin femoral specimens, extant hominid taxa, and, where possible, each other., Results: The Lesedi femora are morphologically similar to the Dinaledi femora for all overlapping regions, with differences limited to few traits of presently unknown significance. The Lesedi distal femur and mid-diaphysis preserve anatomy previously unidentified or unconfirmed in the species, including an anteroposteriorly expanded midshaft and anteriorly expanded patellar surface. The hypothesis that the Lesedi femoral sample may represent two individuals is supported., Discussion: The Lesedi femora increase the range of variation of femoral morphology in H. naledi. Newly described features of the diaphysis and distal femur are either taxonomically uninformative or Homo-like. Overall, these three new femora are consistent with previous functional interpretations of the H. naledi lower limb as belonging to a species adapted for long distance walking and, possibly, running., (© 2019 Wiley Periodicals, Inc.)

Objectives: The basicranium and face are two integrated bony structures displaying great morphological diversity across primates. Previous studies in hominids determined that the basicranium is composed of two independent modules: the midline basicranium, mostly influenced by brain size, and the lateral basicranium, predominantly associated with facial shape. To better assess how morphological integration impacts the evolution of primate cranial shape diversity, we test to determine whether the relationships found in hominids are retained across the order., Materials and Methods: Three-dimensional landmarks (29) were placed on 143 computed tomography scans of six major clades of extant primate crania. We assessed the covariation between midline basicranium, lateral basicranium, face, and endocranial volume using phylogenetically informed partial least squares analyses and phylogenetic generalized least squares models., Results: We found significant integration between lateral basicranium and face and between midline basicranium and face. We also described a significant correlation between midline basicranium and endocranial volume but not between lateral basicranium and endocranial volume., Discussion: Our findings demonstrate a significant and pervasive integration in the craniofacial structures across primates, differing from previous results in hominids. The uniqueness of module organization in hominids may explain this distinction. We found that endocranial volume is significantly integrated to the midline basicranium but not to the lateral basicranium. This finding underlines the significant effect of brain size on the shape of the midline structures of the cranial base in primates. With the covariations linking the studied features defined here, we suggest that future studies should focus on determining the causal links between them., (© 2019 Wiley Periodicals, Inc.)

Objectives: In this study, we reexamined the body mass estimate for the Homo erectus specimen KNM-ER 5428 based on talus dimensions. Previous estimates of >90 kg for this fossil are large in comparison to body mass estimates for other H. erectus specimens., Materials and Methods: The study sample consisted of tali and femora of 132 modern cadaver males from a documented body mass skeletal collection. We recorded the talus trochlear mediolateral (TTML) breadth and femoral head diameter (FHD) for each modern human specimen, and obtained KNM-ER 5428's TTML values from the literature. We developed regression formulae based on TTML using the body mass estimated from FHD for the entire human sample and for known body masses from a normal-BMI subsample, and then used these formulae to calculate body mass for KNM-ER 5428. In addition, we examined the range of body masses for individuals with TTML measurements comparable to KNM-ER 5428., Results: The body masses of normal-BMI individuals with a TTML ≥32.3 mm (the smaller of the two fossil measurements from the literature) ranged between 60.3 and 86.2 kg and averaged 72.3 kg. The body masses of normal-BMI individuals with a TTML ≥33.7 mm (the larger measurement) ranged between 63.5 and 86.2 kg with a mean of 73.6 kg. The correlations between TTML and body mass are moderate. Revised body mass point estimates for KNM-ER 5428 ranged between 69.2 and 81.6 kg based on TTML, and average 70.5 and 76.0 kg., Discussion: Results suggest previously published body mass estimates of KNM-ER 5428's are too large. Its body mass was likely between 70 and 76 kg rather than >90 kg., (© 2019 Wiley Periodicals, Inc.)

Objectives: Purported evolutionary shifts in shoulder structure have been linked to changes in hominin behavior and adaptation. Researchers use clavicle morphology to infer these shifts. However, there is a lack of empirical data underlying such predictive relationships. This study investigates how clavicle morphology affects articulated shoulder girdle and upper thorax configuration in humans., Materials and Methods: Landmarks and scalar measurements on the clavicle, scapula, and ribs 1-3 were collected from three-dimensional computed tomographic scans of living humans. Covariation between disarticulated and articulated morphology was assessed using partial least squares and regression analyses., Results: We found support for hypotheses linking combined dimensions of the clavicle, ribs, and scapula to resting protraction. Individuals with relatively short clavicles tend to exhibit protracted and elevated resting positions of the scapula. It is more difficult to predict superoinferior configuration, which is only minimally affected by clavicle curvature. Instead, the superoinferior position of the scapula on the thorax is governed equally by clavicle orientation and rib declination. Shoulder breadth is determined primarily by clavicle length, but orientation has a comparable effect. Therefore, reliable reconstructions of shoulder breadth can be established using clavicle length, together with consideration of orientation. Relationships between clavicle and thorax morphology are weaker., Discussion: Understanding the determinants of variation in human shoulder structure informs interpretation of skeletal remains. Our investigations describe how important aspects of shoulder structure can be inferred from disarticulated clavicles and we provide the attendant predictive equations. Future work on interspecific variation will improve skeletal reconstruction for more ancient hominins., (© 2019 Wiley Periodicals, Inc.)

Objectives: There is continuing controversy over the number of taxa documented by the Dmanisi hominins. Variation may reflect age and sex differences within a single population. Alternatively, two (or more) distinct species may be present. Our null hypothesis states that just one population is represented at the site., Materials and Methods: We assess the likely sources of variation in endocranial capacity, craniofacial and mandibular morphology, and the expression of characters related to aging and sex dimorphism. We use the coefficient of variation and a modified version of Levene's test for equal variances to compare trait variation at Dmanisi with that in fossil hominins and modern Homo sapiens from Africa., Results: Skull 5 presents a low, massive vault, and a muzzle-like lower face. Other individuals have larger brains and more globular vaults. Despite such variation, the five crania share numerous features. All of the mandibles possess marginal tubercles, mandibular tori, and a distinctive patterning of mental foramina. Relative variation at Dmanisi is comparable to that in selected reference groups. Further growth anticipated in Skull 3, age-related remodeling affecting the D2600 mandible, pathology, and sex dimorphism can account for much of the interindividual variation observed. The preponderance of evidence supports our null hypothesis., Discussion: Sources of the variation within ancient Homo assemblages remain poorly understood. Skull 5 is a very robust male, with a brain smaller than that of both a juvenile (Skull 3) and a probable female (Skull 2). Skull 1 has the largest brain, but cranial superstructures do not clearly mark this individual as male or female. It is likely that the Dmanisi hominins represent a single paleospecies of Homo displaying a pattern of sex dimorphism not seen in living hominids., (© 2018 Wiley Periodicals, Inc.)

Objectives: A prevailing hypothesis in paleoanthropology is that early Pleistocene hominin bones were accumulated in South African caves by carnivores, which used those shelters, and the trees surrounding them, as refuge and feeding sites. We tested this hypothesis at the site of Drimolen, by comparing its hominin age-at-death distribution to that of the nearby and roughly contemporaneous site of Swartkrans., Materials and Methods: We employed standard dental aging systems in order to categorize the Drimolen hominin teeth into age classes of 5 years each. We then compared the age-at-death distribution for Drimolen with the published data available for the Swartkrans hominins., Results: Age-at-death distributions indicate that the age category "young adults" is the best represented age category at Swartkrans and the most poorly represented one at Drimolen. Moreover, Drimolen has a preponderance of infant specimens. Both sites have a low frequency of old adult specimens., Conclusions: Differences observed in frequencies of the age-at-death categories suggest different mechanisms of hominin skeletal accumulation at Drimolen and Swartkrans. Swartkrans' frequency curve reflects mortality in a population subjected to predation and is thus consistent with the carnivore-accumulating hypothesis. In contrast, the Drimolen curve is similar to that of wild populations of living apes. Living primates have been observed exploiting caves as sleeping shelters, for nutritional, security, drinking, and thermoregulatory purposes. We suggest that similar cave use by Pleistocene hominins can explain, in large part, the accumulation of hominin bones at Drimolen. Such a conclusion is another illustration of the growing awareness that a "one-size-fits-all" taphonomic model for South African early Pleistocene hominin sites is probably insufficient., (© 2019 Wiley Periodicals, Inc.)

Objectives: We provide descriptions and functional interpretations of 11 >2.0 Ma hominin vertebral and upper limb fossils from Sterkfontein., Materials and Methods: We employed taphonomic methods to describe postmortem damage observed on the fossils. We used osteometric tools and measurements to generate quantitative descriptions, which were added to qualitative descriptions of the fossils. These observations were then interpreted using published data on the same skeletal elements from extant and extinct hominoid taxa., Results: Six of the fossils carry carnivore tooth marks. Two vertebrae show morphologies that are consistent with fully developed lordosis of the lumbar spine, but which are not completely consistent with bipedal loading of the same intensity and/or frequency as reflected in the lumbars of modern humans. A clavicle shows a combination of humanlike and apelike features, the latter of which would have endowed its hominin with good climbing abilities. When combined, analyses of fragmentary radius and ulna fossils yield more ambiguous results., Discussion: The new fossil collection presents a mix of bipedal and climbing features. It is unclear whether this mix indicates that all Sterkfontein hominins of >2.0 Ma were terrestrial bipeds who retained adaptations for climbing or whether the collection samples two differently adapted, coeval hominins, Australopithecus africanus and Australopithecus prometheus, both of which are represented at Sterkfontein by skull remains. Regardless, the significant frequency of tooth-marked fossils in the sample might indicate that predation was a selection pressure that maintained climbing adaptations in at least some Sterkfontein hominins of this period., (© 2018 Wiley Periodicals, Inc.)

Objectives: The aim of this study is to explore the root and root canal morphology of Homo fossil occupying China during the Middle Pleistocene period. Human occupation and evolutionary dynamics in East Asia during the Middle Pleistocene period is one of the most intriguing issues in paleoanthropology, with the coexistence of multiple lineages and regional morphs suggesting a complex population interaction scenario. Although premolar root and canal morphology has certain phylogenetic, taxonomic, and functional implications, its morphological diversity, possible evolutionary trend and characteristics regarding Middle Pleistocene hominins inhabiting East Asia are still insufficiently understood; where these populations fits within the Homo lineage (with respect to root and pulp canal structure) needs to be explored., Materials and Methods: Using microtomography, we directly observed and assessed the nonmetric variability of root and canal forms in maxillary and mandibular premolars of Chinese Middle Pleistocene Homo (N = 19), and compared our observed variations with Eurasian Early Pleistocene specimens from the Asia continent (N = 1) and Java (N = 2), as well as with Neanderthals (N = 28) and recent modern humans (N = 67)., Results: A total number of nine types of root-canal forms were recorded. As a whole, the Chinese Middle Pleistocene record shows an evolutionary trend toward a modern human-like condition (a reduction of root/canal number and a simplification of root surface structure). We documented primitive signals like high percentage of Tomes' root in lower premolars. A considerable occurrence of incompletely separated root branches and bifid root and canal apices, representing evolutionary transformation from multi-root to single-root condition was also noticed. The results were compared with previous publications on Early and Middle Pleistocene Homo in East Africa, North Africa, and Eurasia., Conclusion: This work provides new original data, incorporates the latest human fossil discoveries and suggests that analyzing the variation of premolar root structural organization, notably integrating together root/canal form and number, could possibly contribute to taxonomic and phylogenetic assessments. The mid-Middle Pleistocene populations, or "classic" Homo erectus, in our study show closer affinity to Early and Middle Pleistocene hominins in Eurasia, than to East African early Homo, which supports the suggestion that at least some of the Early Pleistocene hominin groups in Eurasia contribute to the later population; on the other hand, it is still difficult to clearly trace the evolutionary fate of those late Middle Pleistocene populations (roughly assigned as archaic Homo sapiens through a craniodental perspective). More comparable materials from the Early to Middle Pleistocene period as well as precise chronological framework is needed to further explore the evolutionary trends of archaic hominins in the Asian continent before the arrival of modern humans., (© 2019 Wiley Periodicals, Inc.)

Objectives: Mandibular corpus robusticity (corpus breadth/corpus height) is the most commonly utilized descriptor of the mandibular corpus in the great ape and hominin fossil records. As a consequence of its contoured shape, linear metrics used to characterize mandibular robusticity are inadequate to quantify the shape of the mandibular corpus. Here, we present an alternative to the traditional assessment of mandibular shape by analyzing the outline of the mandibular corpus in cross-section using landmarks and semilandmarks., Materials and Methods: Outlines of the mandibular corpus in cross-section between M 1 and M 2 were quantified in a sample of hominoids and analyzed using generalized Procrustes analysis, Procrustes ANOVA, CVA, and cluster analysis. Corpus breadth and width were also collected from the same sample and analyzed using regression, ANOVA, and cluster analysis., Results: Analysis of corpus outline shape revealed significant differences in mandibular corpus shape that are independent of size and sex at the genus level across hominoids. Cluster analysis based on the analysis of corpus outline shape results in almost all specimens grouping based on taxonomic affinity (99.28% correct classification). Comparison of these results to results using traditional measures of mandibular robusticity shows that analysis of the outline of the corpus in cross-section discriminate extant great apes more reliably., Conclusion: The strong taxonomic signal revealed by this analysis indicates that quantification of the outline of the mandibular corpus more fully captures mandibular corpus shape and offers the potential for greater power in discriminating among taxa in the hominoid fossil record., (© 2018 Wiley Periodicals, Inc.)

Objectives: Middle Pleistocene fossil hominins, often summarized as Homo heidelbergensis sensu lato, are difficult to interpret due to a fragmentary fossil record and ambiguous combinations of primitive and derived characters. Here, we focus on one aspect of facial shape and analyze shape variation of the dental arcades of these fossils together with other Homo individuals., Materials and Methods: Three-dimensional landmark data were collected on computed tomographic scans and surface scans of Middle Pleistocene fossil hominins (n = 8), Homo erectus s.l. (n = 4), Homo antecessor (n = 1), Homo neanderthalensis (n = 13), recent (n = 52) and fossil (n = 19) Homo sapiens. To increase sample size, we used multiple multivariate regression to reconstruct complementary arches for isolated mandibles, and explored size and shape differences among maxillary arcades., Results: The shape of the dental arcade in H. erectus s.l. and H. antecessor differs markedly from both Neanderthals and H. sapiens. The latter two show subtle but consistent differences in arcade length and width. Shape variation among Middle Pleistocene fossil hominins does not exceed the amount of variation of other species, but includes individuals with more primitive and more derived morphology, all more similar to Neanderthals and H. sapiens than to H. erectus s.l., Discussion: Although our results cannot reject the hypothesis that the Middle Pleistocene fossil hominins belong to a single species, their shape variation comprises a more primitive morph that represents a likely candidate for the shape of the last common ancestor of Neanderthals and H. sapiens, and a more derived morph resembling Neanderthals. The arcade shape difference between Neanderthals and H. sapiens might be related to different ways to withstand mechanical stress., (© 2018 Wiley Periodicals, Inc.)

Objectives: As a complex, polygenic trait, brain size has likely been influenced by a range of direct and indirect selection pressures for both cognitive and non-cognitive functions and capabilities. It has been hypothesized that hominin brain expansion was, in part, a correlated response to selection acting on aerobic capacity (Raichlen & Polk, 2013). According to this hypothesis, selection for aerobic capacity increased the activity of various signaling molecules, including those involved in brain growth. One key molecule is brain-derived neurotrophic factor (BDNF), a protein that regulates neuronal development, survival, and plasticity in mammals. This review updates, partially tests, and expands Raichlen and Polk's (2013) hypothesis by evaluating evidence for BDNF as a mediator of brain size., Discussion: We contend that selection for endurance capabilities in a hot climate favored changes to muscle composition, mitochondrial dynamics and increased energy budget through pathways involving regulation of PGC-1α and MEF2 genes, both of which promote BDNF activity. In addition, the evolution of hairlessness and the skin's thermoregulatory response provide other molecular pathways that promote both BDNF activity and neurotransmitter synthesis. We discuss how these pathways contributed to the evolution of brain size and function in human evolution and propose avenues for future research. Our results support Raichlen and Polk's contention that selection for non-cognitive functions has direct mechanistic linkages to the evolution of brain size in hominins., (© 2018 American Association of Physical Anthropologists.)

Objectives: Several studies have investigated potential functional signals in the trabecular structure of the primate proximal humerus but with varied success. Here, we apply for the first time a "whole-epiphyses" approach to analysing trabecular bone in the humeral head with the aim of providing a more holistic interpretation of trabecular variation in relation to habitual locomotor or manipulative behaviors in several extant primates and Australopithecus africanus., Materials and Methods: We use a "whole-epiphysis" methodology in comparison to the traditional volume of interest (VOI) approach to investigate variation in trabecular structure and joint loading in the proximal humerus of extant hominoids, Ateles and A. africanus (StW 328)., Results: There are important differences in the quantification of trabecular parameters using a "whole-epiphysis" versus a VOI-based approach. Variation in trabecular structure across knuckle-walking African apes, suspensory taxa, and modern humans was generally consistent with predictions of load magnitude and inferred joint posture during habitual behaviors. Higher relative trabecular bone volume and more isotropic trabeculae in StW 328 suggest A. africanus may have still used its forelimbs for arboreal locomotion., Discussion: A whole-epiphysis approach to analysing trabecular structure of the proximal humerus can help distinguish functional signals of joint loading across extant primates and can provide novel insight into habitual behaviors of fossil hominins., (© 2018 Wiley Periodicals, Inc.)

Objectives: Prior examination of the ontogeny of Hominoid talo-crural joint morphology using Singular Warp analysis suggested both a genetic and epigenetic signal during development. This question is examined using trajectory analysis and its implications for the Hominin fossil record explored., Materials and Methods: Trajectory analysis was used to examine talo-crural joint shape at different molar eruption stages during development among a cohort of 221 specimens of Homo sapiens, Gorilla gorilla, Pan troglodytes, and Pan paniscus. Trajectory length, angle, and shape were compared among species. Trajectories that showed a consistent pattern of shape change were predicted to show a genetic signal, while change in that pattern an epigenetic signal., Results: The trajectories of all four hominoid tali were consistent during M1-M2 development, but there was a change in Pan paniscus alone at M2-M3. The tibial trajectories were again consistent in M1-M2 development, however, only Pan paniscus persisted in this trajectory at M2-M3 while the other three changed., Discussion: Trajectory analysis demonstrated an epigenetic signal in Pan paniscus in the talus and among the other three hominoids in the tibia. Behavioral (epigenetic) changes may impact hominoid ankle shape in extant species. These changes may reflect differences in locomotor repertoire altering shape, as has been suggested in the Hominin fossil record. Further examination of closely related hominoid species may provide insights into potential behavioral influences in the rapid change and diversity of Hominin talo-crural shape in the Plio-pleistocene., (© 2018 Wiley Periodicals, Inc.)

Objectives: Uncinate processes are protuberances on the cranial surface of subaxial cervical vertebrae that assist in stabilizing and guiding spinal motion. Shallow uncinate processes reduce cervical stability but confer an increased range of motion in clinical studies. Here we assess uncinate processes among extant primates and model cervical kinematics in early fossil hominins., Materials and Methods: We compare six fossil hominin vertebrae with 48 Homo sapiens and 99 nonhuman primates across 20 genera. We quantify uncinate morphology via geometric morphometric methods to understand how uncinate process shape relates to allometry, taxonomy, and mode of locomotion., Results: Across primates, allometry explains roughly 50% of shape variation, as small, narrow vertebrae feature the relatively tallest, most pronounced uncinate processes, whereas larger, wider vertebrae typically feature reduced uncinates. Taxonomy only weakly explains the residual variation, however, the association between Uncinate Shape and mode of locomotion is robust, as bipeds and suspensory primates occupy opposite extremes of the morphological continuum and are distinguished from arboreal generalists. Like humans, Australopithecus afarensis and Homo erectus exhibit shallow uncinate processes, whereas A. sediba resembles more arboreal taxa, but not fully suspensory primates., Discussion: Suspensory primates exhibit the most pronounced uncinates, likely to maintain visual field stabilization. East African hominins exhibit reduced uncinate processes compared with African apes and A. sediba, likely signaling different degrees of neck motility and modes of locomotion. Although soft tissues constrain neck flexibility beyond limits suggested by osteology alone, this study may assist in modeling cervical kinematics and positional behaviors in extinct taxa., (© 2018 Wiley Periodicals, Inc.)

Objectives: Great apes provide a point of reference for understanding the evolution of locomotion in hominoids and early hominins. We assessed (1) the extent to which great apes use diagonal sequence, diagonal couplet gaits, like other primates, (2) the extent to which gait and posture vary across great apes, and (3) the role of body mass and limb proportions on ape quadrupedal kinematics., Methods: High-speed digital video of zoo-housed bonobos (Pan paniscus, N = 8), chimpanzees (Pan troglodytes, N = 13), lowland gorillas (Gorilla gorilla, N = 13), and orangutans (Pongo spp. N = 6) walking over-ground at self-selected speeds were used to determine the timing of limb touch-down, take-off, and to measure joint and segment angles at touch-down, midstance, and take-off., Results: The great apes in our study showed broad kinematic and spatiotemporal similarity in quadrupedal walking. Size-adjusted walking speed was the strongest predictor of gait variables. Body mass had a negligible effect on variation in joint and segment angles, but stride frequency did trend higher among larger apes in analyses including size-adjusted speed. In contrast to most other primates, great apes did not favor diagonal sequence footfall patterns, but exhibited variable gait patterns that frequently shifted between diagonal and lateral sequences., Conclusion: Similarities in the terrestrial walking kinematics of extant great apes likely reflect their similar post-cranial anatomy and proportions. Our results suggest that the walking kinematics of orthograde, suspensory Miocene ape species were likely similar to living great apes, and highlight the utility of videographic and behavioral data in interpreting primate skeletal morphology., (© 2018 Wiley Periodicals, Inc.)

Objectives: The locomotor and manipulative abilities of australopithecines are highly debated in the paleoanthropological context. Australopithecus afarensis and Australopithecus sediba likely engaged in arboreal locomotion and, especially the latter, in certain activities implying manipulation. Nevertheless, their degree of arboreality and the relevance of their manipulative skills remain unclear. Here we calculate the pronation efficiency of the forearm (E rot ) in these taxa to explore their arboreal and manipulative capabilities using a biomechanical approach., Materials and Methods: Three-dimensional humeral images and upper limb measurements of A.L. 288-1 (Au. afarensis) and MH2 (Au. sediba) were used to calculate E rot using a previously described biomechanical model., Results: Maximal E rot in elbow flexion occurs in a rather supinated position of the forearm in Au. afarensis, similarly to Pan troglodytes. In elbow extension, maximal E rot in this fossil taxon occurs in the same forearm position as in Pongo spp. In Au. sediba the forearm positions where E rot is maximal are largely coincident with those for Hylobatidae., Conclusions: The pattern in Au. afarensis suggests relevant arboreal capabilities, which would include vertical climbing, although it is suggestive of poorer manipulative skills than in modern humans. The similarity between Au. sediba and Hylobatidae is difficult to interpret, but the differences between Au. sediba and Au. afarensis suggest that the capacity to rotate the forearm followed different evolutionary processes in these australopithecine species. Although functional inferences from the upper limb are complex, the observed differences between both taxa point to the existence of two distinct anatomical models., (© 2017 Wiley Periodicals, Inc.)

Objectives: A nearly complete hominin fossil cranium from Dali in Shaanxi Province, China was excavated in 1978. We update and expand on previous research by providing a multivariate analysis of the specimen relative to a large sample of Middle and Late Pleistocene hominins., Materials and Methods: We apply principal components analysis, discriminant function analysis, and a method of assessing group membership based on a soft independent model of class analogy (SIMCA) to the study of Dali's cranial morphology. We evaluate Dali's affinities within the context of Middle and Late Pleistocene Homo patterns of craniofacial morphology., Results: When just the facial skeleton is considered, Dali aligns with Middle Paleolithic H. sapiens and is clearly more derived than African or Eurasian Middle Pleistocene Homo. When just the neurocranium is considered, Dali is most similar to African and Eastern Eurasian but not Western European Middle Pleistocene Homo. When both sets of variables are considered together, Dali exhibits a unique morphology that is most closely aligned with the earliest H. sapiens from North Africa and the Levant., Discussion: These results add perspective to our previous view of as Dali a "transitional" form between Chinese H. erectus and H. sapiens. Athough no taxonomic allocation is appropriate at this time for Dali, it appears to represent a population that played a more central role in the origin of Chinese H. sapiens. Dali's affinities can be understood in the context of Wu's Continuity with Hybridization scenario and a braided-stream network model of gene flow. Specifically, we propose that Pleistocene populations in China were shaped by periods of isolated evolutionary change within local lineages at certain times, and gene flow between local lineages or between Eastern and Western Eurasia, and Africa at other times, resulting in contributions being made in different capacities to different regions at different times., (© 2017 Wiley Periodicals, Inc.)

Objective: Herein we introduce a newly recovered partial calvaria, OH 83, from the upper Ndutu Beds of Olduvai Gorge, Tanzania. We present the geological context of its discovery and a comparative analysis of its morphology, placing OH 83 within the context of our current understanding of the origins and evolution of Homo sapiens., Materials and Methods: We comparatively assessed the morphology of OH 83 using quantitative and qualitative data from penecontemporaneous fossils and the W.W. Howells modern human craniometric dataset., Results: OH 83 is geologically dated to ca. 60-32 ka. Its morphology is indicative of an early modern human, falling at the low end of the range of variation for post-orbital cranial breadth, the high end of the range for bifrontal breadth, and near average in frontal length., Discussion: There have been numerous attempts to use cranial anatomy to define the species Homo sapiens and identify it in the fossil record. These efforts have not met wide agreement by the scientific community due, in part, to the mosaic patterns of cranial variation represented by the fossils. The variable, mosaic pattern of trait expression in the crania of Middle and Late Pleistocene fossils implies that morphological modernity did not occur at once. However, OH 83 demonstrates that by ca. 60-32 ka modern humans in Africa included individuals that are at the fairly small and gracile range of modern human cranial variation., (© 2017 Wiley Periodicals, Inc.)

Objectives: The aim of this study is to compare the degree and patterning of inter- and intra-individual metameric variation in South African australopiths, early Homo and modern humans. Metameric variation likely reflects developmental and taxonomical issues, and could also be used to infer ecological and functional adaptations. However, its patterning along the early hominin postcanine dentition, particularly among South African fossil hominins, remains unexplored., Materials and Methods: Using microfocus X-ray computed tomography (µXCT) and geometric morphometric tools, we studied the enamel-dentine junction (EDJ) morphology and we investigated the intra- and inter-individual EDJ metameric variation among eight australopiths and two early Homo specimens from South Africa, as well as 32 modern humans., Results: Along post-canine dentition, shape changes between metameres represented by relative positions and height of dentine horns, outlines of the EDJ occlusal table are reported in modern and fossil taxa. Comparisons of EDJ mean shapes and multivariate analyses reveal substantial variation in the direction and magnitude of metameric shape changes among taxa, but some common trends can be found. In modern humans, both the direction and magnitude of metameric shape change show increased variability in M 2 -M 3 compared to M 1 -M 2 . Fossil specimens are clustered together showing similar magnitudes of shape change. Along M 2 -M 3 , the lengths of their metameric vectors are not as variable as those of modern humans, but they display considerable variability in the direction of shape change., Conclusion: The distalward increase of metameric variation along the modern human molar row is consistent with the odontogenetic models of molar row structure (inhibitory cascade model). Though much remains to be tested, the variable trends and magnitudes in metamerism in fossil hominins reported here, together with differences in the scale of shape change between modern humans and fossil hominins may provide valuable information regarding functional morphology and developmental processes in fossil species., (© 2017 Wiley Periodicals, Inc.)

Objectives: This study takes a new approach to interpreting dental development in Pleistocene Homo in comparison with recent modern humans. As rates of dental development and skeletal growth are correlated given age in modern humans, using age and skeletal growth in tandem yields more accurate dental development estimates. Here, I apply these models to fossil Homo to obtain more individualized predictions and interpretations of their dental development relative to recent modern humans., Materials and Methods: Proportional odds logistic regression models based on three recent modern human samples (N = 181) were used to predict permanent mandibular tooth development scores in five Pleistocene subadults: Homo erectus/ergaster, Neanderthals, and anatomically modern humans (AMHs). Explanatory variables include a skeletal growth indicator (i.e., diaphyseal femoral length), and chronological age., Results: AMHs Lagar Velho 1 and Qafzeh 10 share delayed incisor development, but exhibit considerable idiosyncratic variation within and across tooth types, relative to each other and to the reference samples. Neanderthals Dederiyeh 1 and Le Moustier 1 exhibit delayed incisor coupled with advanced molar development, but differences are reduced when femoral diaphysis length is considered. Dental development in KNM-WT 15,000 Homo erectus/ergaster, while advanced for his age, almost exactly matches the predictions once femoral length is included in the models., Discussion: This study provides a new interpretation of dental development in KNM-WT 15000 as primarily reflecting his faster rates of skeletal growth. While the two AMH specimens exhibit considerable individual variation, the Neanderthals exhibit delayed incisor development early and advanced molar development later in ontogeny., (© 2017 Wiley Periodicals, Inc.)

Objectives: Trabecular microstructure of limb bone epiphyses has been used to elucidate the relationship between skeletal form and behavior among mammals. Such studies have often relied on the analysis of a single volume of interest (VOI). Here we present a method for evaluating variation in bone microstructure across articular surfaces by leveraging sliding semilandmarks., Methods: Two samples were used to demonstrate the proposed methodology and test the hypothesis that microstructural variables are homogeneously distributed: tali from two ape genera (Pan and Pongo, n = 9) and modern human distal femora (n = 10). Sliding semilandmarks were distributed across articular surfaces and used to locate the position of multiple VOIs immediately deep to the cortical shell. Trabecular bone properties were quantified using the BoneJ plugin for ImageJ. Nonparametric MANOVA tests were used to make group comparisons and differences were explored using principal components analysis and visualized using color maps., Results: Tests reveal that trabecular parameters are not distributed homogeneously and identify differences between chimpanzee and orangutan tali with regards to trabecular spacing and degree of anisotropy, with chimpanzee tali being more anisotropic and having more uniformly spaced trabeculae. Human males and females differed in the pattern of trabecular spacing with males having more uniform trabecular spacing across the joint surface., Conclusions: The proposed procedure quantifies variation in trabecular bone parameters across joint surfaces and allows for meaningful statistical comparisons between groups of interest. Consequently it holds promise to help elucidate links between trabecular bone structure and animal behavior., (© 2017 Wiley Periodicals, Inc.)

Objectives: In 2008, an immature hominin defined as the holotype of the new species Australopithecus sediba was discovered at the 1.9 million year old Malapa site in South Africa. The specimen (MH1) includes substantial post-cranial skeletal material, and provides a unique opportunity to assess its skeletal maturation., Methods: Skeletal maturity indicators observed on the proximal and distal humerus, proximal ulna, distal radius, third metacarpal, ilium and ischium, proximal femur and calcaneus were used to assess the maturity of each bone in comparison to references for modern humans and for wild chimpanzees (Pan troglodytes)., Results: In comparison to humans the skeletal maturational ages for Au. sediba correspond to between 12.0 years and 15.0 years with a mean (SD) age of 13.1 (1.1) years. In comparison to the maturational pattern of chimpanzees the Au. sediba indicators suggest a skeletal maturational age of 9-11 years. Based on either of these skeletal maturity estimates and the body length at death of MH1, an adult height of 150-156 cm is predicted., Discussion: We conclude that the skeletal remains of MH1 are consistent with an ape-like pattern of maturity when dental age estimates are also taken into consideration. This maturity schedule in australopiths is consistent with ape-like estimates of age at death for the Nariokotome Homo erectus remains (KMN-WT 15000), which are of similar postcranial immaturity to MH1. The findings suggest that humans may have distinctive and delayed post-cranial schedules from australopiths and H. erectus, implicating a recent evolution of somatic and possibly life history strategies in human evolution., (© 2017 Wiley Periodicals, Inc.)

Objectives: During the last 13 years, the late Early Pleistocene Gran Dolina-TD6-2 level (Sierra de Atapuerca, northern Spain) has yielded an additional sample of 26 dental specimens attributed to Homo antecessor. In this report, we present a descriptive and comparative study of the six deciduous teeth., Methods: We provide external and internal morphological descriptions following classical terminology, as well as the mesiodistal and buccolingual measurements of the teeth. The internal morphology was described by means of micro-CT technique., Results: The TD6 deciduous teeth preserve primitive features regarding the Homo clade, such as the presence of styles in lower and upper canines and developed anterior and posterior foveae in the dm 2 . However, other features related to the complexity of the crown morphology (e.g., cingulum) are not present in this sample. Furthermore, the great reduction of the talonid of the dm 1 s is also noteworthy. Despite the limited comparative evidence, the presence of a remarkably well-developed tuberculum molare in the dm 1 and dm 1 s from TD6 can be also considered a derived feature in the genus Homo. The TD6 hominins exhibit dental dimensions similar to those of other Pleistocene hominins. The dm 1 s are buccolingually elongated and the buccolingual diameter of ATD6-93 is the largest recorded so far in the Homo fossil record., Conclusions: This study expands the list of plesiomorphic features of H. antecessor, and provides some information on the evolutionary status of this species. However, the identification of some advanced traits evinces a step towards the derived morphology of European Pleistocene teeth. The study of the deciduous dentition confirms the mosaic pattern of H. antecessor morphology revealed in previous studies of this hominin sample., (© 2017 Wiley Periodicals, Inc.)

Objectives: The higher primate scapula has been subject to many explanations of the putative "adaptive value" of its individual traits. However, the shift from the bone's position in above branch quadrupeds to its more posterolateral position in recent hominoids obviously required fundamental changes to its general form. We hypothesize that most features argued to be individually adaptive are more likely secondary consequences of changes in its fundamental bauplan, a view more consistent with modern developmental biology., Materials and Methods: We tested this hypothesis with scapular metrics and angles from a broad anthropoid sample., Results: Our results support our hypothesis. Contrary to earlier predictions, vertebral border length differs little relative to body size in anthropoids, inferior angle position primarily reflects mediolateral scapular breadth, and supraspinous and infraspinous fossa sizes largely reflect scapular spine orientation. Suspensory taxa have cranially oriented glenoids, whereas slow clamberers and humans do not. Australopithecus most closely resembles the latter., Discussion: Most scapular features can be explained by only two primary changes: (1) reduction in mediolateral breadth and (2) change in the glenoid position relative to the vertebral border with increased reliance on suspension, which led to a more cranially angled scapular spine. Virtually all other scapular traits appear to be byproducts of these two changes. Based on fossil morphology, hominids 1 were derived from a last common ancestor primarily adapted for clambering and not for suspension. Scapular form in early hominids such as Australopithecus is therefore primitive and largely reflects the genus's general clambering heritage., (© 2017 Wiley Periodicals, Inc.)

Objectives: To update histological data on crown initiation and completion ages in southern Africans. To evaluate implications of these data for studies that: (a) rely on these data to time linear enamel hypoplasias (LEHs), or, (b) use these data for comparison to fossil hominins., Materials and Methods: Initiation ages were calculated on 67 histological sections from southern Africans, with sample sizes ranging from one to 11 per tooth type. Crown completion ages for southern Africans were calculated in two ways. First, actual derived initiation ages were added to crown formation times for each histological section to obtain direct information on the crown completion ages of individuals. Second, average initiation ages from this study were added to average crown formation times of southern Africans from the Reid and coworkers previous studies that were based on larger samples., Results: For earlier-initiating tooth types (all anterior teeth and first molars), there is little difference in ages of initiation and crown completion between this and previous studies. Differences increase as a function of initiation age, such that the greatest differences between this and previous studies for both initiation and crown completion ages are for the second and third molars., Discussion: This study documents variation in initiation ages, particularly for later-initiating tooth types. It upholds the use of previously published histological aging charts for LEHs on anterior teeth. However, this study finds that ages of crown initiation and completion in second and third molars for this southern African sample are earlier than previously estimated. These earlier ages reduce differences between modern humans and fossil hominins for these developmental events in second and third molars., (© 2017 Wiley Periodicals, Inc.)

Objectives: This study presents the first detailed morphological description and comparison of a Middle Pleistocene hominin mandibular fragment (PA 831) and associated teeth from the Hexian site in Eastern China. We aim to investigate where the Hexian mandible fits within the genus Homo variability in the light of an increased and better characterized Asian fossils record., Methods: Comparative samples include Pleistocene Homo mandibles and teeth from Africa, Asia, and Europe, as well as earlier African hominins (Australopithecus and early Homo) and Holocene recent humans. Both conventional morphological description and metric analysis were used. In addition, virtual reconstructions of the enamel dentine junction (EDJ) surface, pulp cavity, and roots with micro-CT were used to the mandible and teeth., Results: The Hexian mandible is characterized by a plesiomorphic structural pattern for the Homo clade, with strong corpus robustness and a subparallel and low-positioned mylohyoid line that differentiates the swollen subalveolar planum from the shallow subalveolar fossa. Features that are derived compared to early Homo include a moderately curved dental arcade, a well-developed lateral prominence placed at the M 2 -M 3 level, and multiple mental foramina. The Hexian mandible's complex enamel surface and strong, stout root structure are primitive traits for the Homo clade. Finally, the highly crenulated "dendrite-like" EDJ found in the molars may represent a dental feature specific to the continental Asian Homo erectus, but more data is needed to confirm this., Conclusions: Mandibular and dental features indicate that the Hexian mandible and teeth differ from northern Chinese H. erectus and European Middle Pleistocene hominins, but show some affinities with the Early Pleistocene specimens from Africa (Homo ergaster) and Java (H. erectus), as well as the Middle-Late Pleistocene mandible from Penghu, Taiwan. Compared to contemporaneous continental Asian hominin populations, the Hexian fossils may represent the survival of a primitive hominin, with more primitive morphologies than other contemporaneous or some chronologically older Asian hominin specimens., (© 2017 Wiley Periodicals, Inc.)