Carpal kinematics and morphological correlates of wrist ulnar deviation mobility in nonhuman anthropoid primates.

OBJECTIVES: Primates employ wrist ulnar deviation during a variety of locomotor and manipulative behaviors. Extant hominoids share a derived condition in which the ulnar styloid process has limited articulation or is completely separated from the proximal carpals, which is often hypothesized to increase ulnar deviation range of motion. Acute angulation of the hamate's triquetral facet is also hypothesized to facilitate ulnar deviation mobility and mechanics. In this study, we test these longstanding ideas. METHODS: Three-dimensional (3D) carpal kinematics were examined using a cadaveric sample of Pan troglodytes, Pongo sp., and five monkey species. Ulnar styloid projection and orientation of the hamate's triquetral facet were quantified using 3D models. RESULTS: Although carpal rotation patterns in Pan and Pongo were uniquely similar in some respects, P. troglodytes exhibited overall kinematic similarity with large terrestrial cercopithecoids (Papio and Mandrillus). Pongo, Macaca, and Ateles had high wrist ulnar deviation ranges of motion, but Pongo did this via a unique mechanism. In Pongo, the triquetrum functions as a distal carpal rather than part of the proximal row. Ulnar styloid projection and wrist ulnar deviation range of motion were not correlated but ulnar deviation range of motion and the triquetrohamate facet orientation were correlated. CONCLUSIONS: Increased ulnar deviation mobility is not the function of ulnar styloid withdrawal in hominoids. Instead, this feature probably reduces stress on the ulnar side wrist or is a byproduct of adaptations that increase supination. Orientation of the hamate's triquetral facet offers some potential to reconstruct ulnar deviation mobility in extinct primates.

Morphological and evolutionary insights into the keystone element of the human foot's medial longitudinal arch.

The evolution of the medial longitudinal arch (MLA) is one of the most impactful adaptations in the hominin foot that emerged with bipedalism. When and how it evolved in the human lineage is still unresolved. Complicating the issue, clinical definitions of flatfoot in living Homo sapiens have not reached a consensus. Here we digitally investigate the navicular morphology of H. sapiens (living, archaeological, and fossil), great apes, and fossil hominins and its correlation with the MLA. A distinctive navicular shape characterises living H. sapiens with adult acquired flexible flatfoot, while the congenital flexible flatfoot exhibits a 'normal' navicular shape. All H. sapiens groups differentiate from great apes independently from variations in the MLA, likely because of bipedalism. Most australopith, H. naledi, and H. floresiensis navicular shapes are closer to those of great apes, which is inconsistent with a human-like MLA and instead might suggest a certain degree of arboreality. Navicular shape of OH 8 and fossil H. sapiens falls within the normal living H. sapiens spectrum of variation of the MLA (including congenital flexible flatfoot and individuals with a well-developed MLA). At the same time, H. neanderthalensis seem to be characterised by a different expression of the MLA.

Human and African ape myosin heavy chain content and the evolution of hominin skeletal muscle.

Humans are unique among terrestrial mammals in our manner of walking and running, reflecting 7 to 8 Ma of musculoskeletal evolution since diverging with the genus Pan. One component of this is a shift in our skeletal muscle biology towards a predominance of myosin heavy chain (MyHC) I isoforms (i.e. slow fibers) across our pelvis and lower limbs, which distinguishes us from chimpanzees. Here, new MyHC data from 35 pelvis and hind limb muscles of a Western gorilla (Gorilla gorilla) are presented. These data are combined with a similar chimpanzee dataset to assess the MyHC I content of humans in comparison to African apes (chimpanzees and gorillas) and other terrestrial mammals. The responsiveness of human skeletal muscle to behavioral interventions is also compared to the human-African ape differential. Humans are distinct from African apes and among a small group of terrestrial mammals whose pelvis and lower limb muscle is slow fiber dominant, on average. Behavioral interventions, including immobilization, bed rest, spaceflight and exercise, can induce modest decreases and increases in human MyHC I content (i.e. -9.3% to 2.3%, n = 2033 subjects), but these shifts are much smaller than the mean human-African ape differential (i.e. 31%). Taken together, these results indicate muscle fiber content is likely an evolvable trait under selection in the hominin lineage. As such, we highlight potential targets of selection in the genome (e.g. regions that regulate MyHC content) that may play an important role in hominin skeletal muscle evolution.

A pilot study of local anesthesia training using a mixed-reality haptic fidelity model.

PURPOSE/OBJECTIVES: One of the most difficult local anesthetic blocks to master in dentistry is the inferior alveolar nerve block (IANB). Historically, dental students have practiced local anesthesia on one another. At the University of Colorado, these practice sessions have been limited to one required laboratory session. The predictability and confidence of student IANB success have not been high in the past. Therefore, the objective of this study was to investigate the impact of a novel IANB simulator, built on a three dimensional (3D)-printed mixed-reality haptic model, for second-year dental students to practice on prior to their laboratory session. METHODS: Thirty-nine student participants volunteered to practice with the IANB simulator. Participants were divided into two groups, Group A and Group B. Self-reported confidence and injection-specific accuracy were measured during IANB simulator practice and the laboratory session. During lab, partner numbness was assessed as a measure of IANB success. Groups A (n = 20) and B (n = 19) practiced with the simulator before and after laboratory, respectively. Injection domains were not assessed during Group B's practice with the IANB simulator. RESULTS: Self-reported confidence increased for both groups (p < 0.001). However, for anesthetic success, Group A exhibited significantly greater success (52.6%) than Group B (17.6%) (p = 0.029). CONCLUSION: Self-reported confidence in performing an IANB improved and higher anesthetic success was achieved for Group A. Further investigation is necessary to determine the long-term impact of using the IANB simulator in dental education.

A geometric morphometric approach to investigate primate proximal phalanx diaphysis shape.

Current approaches to quantify phalangeal curvature assume that the long axis of the bone's diaphysis approximates the shape of a portion of a circle (included angle method) or a parabola (second-degree polynomial method). Here we developed, tested, and employed an alternative geometric morphometrics-based approach to quantify diaphysis shape of proximal phalanges in humans, apes and monkeys with diverse locomotor behaviors. 100 landmarks of the central longitudinal axis were extracted from 3D surface models and analyzed using 2DGM methods, including Generalized Procrustes Analyses. Principal components analyses were performed and PC1 scores (>80% of variation) represented the dorsopalmar shape of the bone's central longitudinal axis and separated taxa consistently and in accord with known locomotor behavioral profiles. The most suspensory taxa, including orangutans, hylobatids and spider monkeys, had significantly lower PC1 scores reflecting the greatest amounts of phalangeal curvature. In contrast, bipedal humans and the quadrupedal cercopithecoid monkeys sampled (baboons, proboscis monkeys) exhibited significantly higher PC1 scores reflecting flatter phalanges. African ape (gorillas, chimpanzees and bonobos) phalanges fell between these two extremes and were not significantly different from each other. PC1 scores were significantly correlated with both included angle and the a coefficient of a second-degree polynomial calculated from the same landmark dataset, but had a significantly higher correlation with included angles. Our alternative approach for quantifying diaphysis shape of proximal phalanges to investigate dorsopalmar curvature is replicable and does not assume a priori either a circle or parabola model of shape, making it an attractive alternative compared with existing methodologies.

An infant burial from Arma Veirana in northwestern Italy provides insights into funerary practices and female personhood in early Mesolithic Europe.

The evolution and development of human mortuary behaviors is of enormous cultural significance. Here we report a richly-decorated young infant burial (AVH-1) from Arma Veirana (Liguria, northwestern Italy) that is directly dated to 10,211-9910 cal BP (95.4% probability), placing it within the early Holocene and therefore attributable to the early Mesolithic, a cultural period from which well-documented burials are exceedingly rare. Virtual dental histology, proteomics, and aDNA indicate that the infant was a 40-50 days old female. Associated artifacts indicate significant material and emotional investment in the child's interment. The detailed biological profile of AVH-1 establishes the child as the earliest European near-neonate documented to be female. The Arma Veirana burial thus provides insight into sex/gender-based social status, funerary treatment, and the attribution of personhood to the youngest individuals among prehistoric hunter-gatherer groups and adds substantially to the scant data on mortuary practices from an important period in prehistory shortly following the end of the last Ice Age.

Evolutionary trends of the lateral foot in catarrhine primates: Contextualizing the fourth metatarsal of Australopithecus afarensis.

In 2000, a complete fourth metatarsal (Mt4) of the ∼3- to 4-Million-year-old hominin Australopithecus afarensis was recovered in Hadar, Ethiopia. This metatarsal presented a mostly human-like morphology, suggesting that a rigid lateral foot may have evolved as early as ∼3.2 Ma. The lateral foot is integral in providing stability during the push off phase of gait and is key in understanding the transition to upright, striding bipedalism. Previous comparisons of this fossil were limited to Pan troglodytes, Gorilla gorilla, and modern humans. This study builds on previous studies by contextualizing the Mt4 morphology of A. afarensis (A.L. 333-160) within a diverse comparative sample of nonhuman hominoids (n = 144) and cercopithecids (n = 138) and incorporates other early hominins (n = 3) and fossil hominoids that precede the Pan-Homo split (n = 4) to better assess the polarity of changes in lateral foot morphology surrounding this divergence. We investigate seven morphological features argued to be functionally linked to human-like bipedalism. Our results show that some human-like characters used to assess midfoot and lateral foot stiffness in the hominin fossil record are present in our Miocene ape sample as well as in living cercopithecids. Furthermore, modern nonhuman hominoids can be generally distinguished from other species in most metrics. These results suggest that the possession of a rigid foot in hominins could represent a conserved trait, whereas the specialized pedal grasping mechanics of extant apes may be more derived, in which case some traits often used to infer bipedal locomotion in early hominins may, instead, reflect a lower reliance on pedal grasping. Another possibility is that early hominins reverted from modern ape Mt4 morphology into a more plesiomorphic condition when terrestrial bipedality became a dominant behavior. More fossils dating around the Pan-Homo divergence time are necessary to test these competing hypotheses.

Homo naledi pollical metacarpal shaft morphology is distinctive and intermediate between that of australopiths and other members of the genus Homo.

Homo naledi fossils from the Rising Star cave system provide important insights into the diversity of hand morphology within the genus Homo. Notably, the pollical (thumb) metacarpal (Mc1) displays an unusual suite of characteristics including a median longitudinal crest, a narrow proximal base, and broad flaring intrinsic muscle flanges. The present study evaluates the affinities of H. naledi Mc1 morphology via 3D geometric morphometric analysis of shaft shape using a broader comparative sample (n = 337) of fossil hominins, recent humans, apes, and cercopithecoid monkeys than in prior work. Results confirm that the H. naledi Mc1 is distinctive from most other hominins in being narrow at the proximal end but surmounted by flaring muscle flanges distally. Only StW 418 (Australopithecus cf. africanus) is similar in these aspects of shape. The gracile proximal shaft is most similar to cercopithecoids, Pan, Pongo, Australopithecus afarensis, and Australopithecus sediba, suggesting that H. naledi retains the condition primitive for the genus Homo. In contrast, Neandertal Mc1s are characterized by wide proximal bases and shafts, pinched midshafts, and broad distal flanges, while those of recent humans generally have straight shafts, less robust muscle flanges, and wide proximal shafts/bases. Although uncertainties remain regarding character polarity, the morphology of the H. naledi thumb might be interpreted as a retained intermediate state in a transformation series between the overall gracility of the shaft and the robust shafts of later hominins. Such a model suggests that the addition of broad medial and lateral muscle flanges to a primitively slender shaft was the first modification in transforming the Mc1 into the overall more robust structure exhibited by other Homo taxa including Neandertals and recent Homo sapiens in whose shared lineage the bases and proximal shafts became expanded, possibly as an adaptation to the repeated recruitment of powerful intrinsic pollical muscles.

Strength properties of extant hominoid hallucal and pollical metapodials.

Functional comparisons of cortical bone strength properties between hominoid hallucal and pollical metapodials (Mt1 and Mc1, respectively) are lacking. Determining which of these two elements is stronger, and by how much, could be informative because the hallux and pollex are used differently both within and among extant hominoids during locomotion and manipulation (i.e., functional differentiation between autopod pairs). Here, we compare Mt1 and Mc1 midshaft cortical area, polar section modulus, and polar second moment of area, calculated from high-resolution computed tomography images in humans (n = 21), chimpanzees (n = 47), gorillas (n = 24), orangutans (n = 20), siamangs (n = 8), and gibbons (n = 21). Intraindividual comparisons between bones within species were made using paired t-tests. Log10-transformed Mt1:Mc1 ratios were created to assess relative strength asymmetry between bones, and interspecific comparisons of these proportions were made using analyses of variance. Absolute strength differences between the Mt1 and Mc1 for all variables were significantly larger in the Mt1 for all species (p < 0.05). Significant differences across species in Mt1:Mc1 proportions were also found, thereby demonstrating that strength asymmetry between bones differs among taxa (p < 0.05); asymmetry was lowest in orangutans, intermediate in gorillas, and greatest in humans, chimpanzees, siamangs, and gibbons. These findings support the hypothesis that the Mt1 is better adapted structurally than the Mc1 for bearing mechanical loads during weight support of locomotion in all extant hominoids and that pedal hallucal grasping likely engenders higher loads than manual pollical grasping in nonhuman hominoids. Thus, functional differentiation in autopod use within and among hominoids is reflected in hallucal and pollical metapodial strength properties.

Evaluation of Articular Surface Similarity of Hemi-Hamate Grafts and Proximal Middle Phalanx Morphology: A 3D Geometric Morphometric Approach.

PURPOSE: Hemi-hamate arthroplasty has been described as a viable treatment option for unstable proximal interphalangeal joint fracture-dislocations. The procedure uses a dorsal distal hamate osteochondral graft to recreate the injured volar middle phalanx (MP) proximal base. The purpose of this study was to evaluate the similarity in shape of these articular surfaces using quantitative 3-dimensional methods. METHODS: Three-dimensional virtual renderings were created from laser scans of the articular surfaces of the dorsal distal hamate and the volar MP bases of the index, middle, ring, and little fingers from cadaveric hands of 25 individuals. Three-dimensional landmarks were obtained from the articular surfaces of each bone and subjected to established geometric morphometric analytical approaches to quantify shape. For each individual, bone shapes were evaluated for covariation using 2-block partial least-squares and principal component analyses. RESULTS: No statistically significant covariation was found between the dorsal distal hamate and volar MP bases of the middle, ring, or little digits. Whereas the volar MP bases demonstrated relative morphologic uniformity among the 4 digits both within and between individuals, the dorsal distal hamates exhibited notable variation in articular surface morphology. CONCLUSIONS: Despite the early to midterm clinical success of hemi-hamate arthroplasty, there is no statistically significant, uniform similarity in shape between the articular surfaces of the dorsal distal hamate and the volar MP base. In addition, there is wide variation in the articular morphology of the hamate among individuals. CLINICAL RELEVANCE: The lack of uniform similarity in shape between the dorsal distal hamate and the volar MP base may result in unpredictable outcomes in HHA. It is recommended that the variation in hamate morphology be considered while reconstructing the injured volar MP base in the procedure.

Evolution and function of the hominin forefoot.

The primate foot functions as a grasping organ. As such, its bones, soft tissues, and joints evolved to maximize power and stability in a variety of grasping configurations. Humans are the obvious exception to this primate pattern, with feet that evolved to support the unique biomechanical demands of bipedal locomotion. Of key functional importance to bipedalism is the morphology of the joints at the forefoot, known as the metatarsophalangeal joints (MTPJs), but a comprehensive analysis of hominin MTPJ morphology is currently lacking. Here we present the results of a multivariate shape and Bayesian phylogenetic comparative analyses of metatarsals (MTs) from a broad selection of anthropoid primates (including fossil apes and stem catarrhines) and most of the early hominin pedal fossil record, including the oldest hominin for which good pedal remains exist, Ardipithecus ramidus Results corroborate the importance of specific bony morphologies such as dorsal MT head expansion and "doming" to the evolution of terrestrial bipedalism in hominins. Further, our evolutionary models reveal that the MT1 of Ar. ramidus shifts away from the reconstructed optimum of our last common ancestor with apes, but not necessarily in the direction of modern humans. However, the lateral rays of Ar. ramidus are transformed in a more human-like direction, suggesting that they were the digits first recruited by hominins into the primary role of terrestrial propulsion. This pattern of evolutionary change is seen consistently throughout the evolution of the foot, highlighting the mosaic nature of pedal evolution and the emergence of a derived, modern hallux relatively late in human evolution.

Inter-ray variation in metatarsal strength properties in humans and African apes: Implications for inferring bipedal biomechanics in the Olduvai Hominid 8 foot.

When measured as a ratio of mean midshaft diameter to bone length, the OH 8 fossil hominin foot exhibits a metatarsal (Mt) robusticity pattern of 1 > 5 > 3 > 4 > 2, which differs from the widely perceived "common" modern human pattern (1 > 5 > 4 > 3 > 2); African apes generally exhibit a third pattern (1 > 2 > 3 > 4 > 5). Largely because of the relative ranking of Mt2 and Mt5, OH 8 metatarsals structurally resemble the pattern exhibited by bipedal humans more than the pattern of quadrupedal and climbing African apes. Considering only these three phenotypes, however, discounts the potentially important functional implications of variation in modern human (and African ape) metatarsal robusticity patterns, suggesting that they are not useful for interpreting the specific biomechanics of a bipedal gait in fossils (i.e., whether it was modern human-like or not). Using computed tomography scans to quantify metatarsal midshaft cross-sectional geometry in a large sample of Homo (n=130), Gorilla (n=44) and Pan (n=80), we documented greater variation in metatarsal robusticity patterns than previously recognized in all three groups. While apes consistently show a 1 > 2 > 3 > 4 > 5 pattern in our larger sample, there does not appear to be a similarly precise single "common" human pattern. Rather, human metatarsals converge towards a 1 > 4/5 > 2/3 pattern, where metatarsals 4 and 5, and metatarsals 2 and 3, often "flip" positions relative to each other depending on the variable examined. After reassessing what a "common" human pattern could be based on a larger sample, the previously described OH 8 pattern of 1 > 5 > 3 > 4 > 2 is only observed in some humans (<6%) and almost never in apes (<0.5%). Although this suggests an overall greater similarity to (some) humans than to any ape in loading of the foot, the relatively rare frequency of these humans in our sample underscores potential differences in loading experienced by the medial and lateral columns of the OH 8 foot compared to modern humans.

Kinematics of the anthropoid os centrale and the functional consequences of scaphoid-centrale fusion in African apes and hominins.

In most primates, the os centrale is interposed between the scaphoid, trapezoid, trapezium, and head of the capitate, thus constituting a component of the wrist's midcarpal complex. Scaphoid-centrale fusion is among the clearest morphological synapomorphies of African apes and hominins. Although it might facilitate knuckle-walking by increasing the rigidity and stability of the radial side of the wrist, the exact functional significance of scaphoid-centrale fusion is unclear. If fusion acts to produce a more rigid radial wrist that stabilizes the hand and limits shearing stresses, then in taxa with a free centrale, it should anchor ligaments that check extension and radial deviation, but exhibit motion independent of the scaphoid. Moreover, because the centrale sits between the scaphoid and capitate (a major stabilizing articulation), scaphoid-centrale mobility should correlate with scaphocapitate mobility in extension and radial deviation. To test these hypotheses, the centrale's ligamentous binding was investigated via dissection in Pongo and Papio, and the kinematics of the centrale were quantified in a cadaveric sample of anthropoids (Pongo sp., Ateles geoffroyi, Colobus guereza, Macaca mulatta, and Papio anubis) using a computed-tomography-based method to track wrist-bone motion. Results indicate that the centrale rotates freely relative to the scaphoid in all taxa. However, centrale mobility is only correlated with scaphocapitate mobility during extension in Pongo-possibly due to differences in overall wrist configuration between apes and monkeys. If an extant ape-like wrist characterized early ancestors of African apes and hominins, then scaphoid-centrale fusion would have increased midcarpal rigidity in extension relative to the primitive condition. Although biomechanically consistent with a knuckle-walking hominin ancestor, this assumes that the trait evolved specifically for that biological role, which must be squared with contradictory interpretations of extant and fossil hominoid morphology. Regardless of its original adaptive significance, scaphoid-centrale fusion likely presented a constraint on early hominin midcarpal mobility.

Locomotor Hand Postures, Carpal Kinematics During Wrist Extension, and Associated Morphology in Anthropoid Primates.

The biomechanics of wrist extension (or dorsiflexion) are important for understanding functional adaptation of the primate hand because extension mobility varies with habitual locomotor hand posture and facilitates certain manipulative tasks. Here, intercarpal kinematics are employed to investigate mechanisms underlying wrist extension in a sample of anthropoids representing various arboreal and terrestrial locomotor modes. Carpal kinematics are studied using computed-tomography of cadaveric forelimbs, and these data are combined with a morphometric analysis of biomechanically-informative anatomical features. The results indicate that stiff-wristed knuckle-walking chimpanzees and digitigrade baboons are characterized by low ranges of motion (ROMs) at the radiocarpal and midcarpal complexes. Palmigrade-capable monkeys have high extension ROMs at both the radiocarpus and midcarpus, while palmigrade-capable orangutans achieve wrist extension through moderate radiocarpal ROMs and high midcarpal ROMs. Morphometrics demonstrate that a more projecting dorsal ridge of the distal radius corresponds with low-to-moderate radioscaphoid mobility in the apes, but that baboons resemble palmigrade-capable monkeys in this metric. Thus, the dorsal ridge of the radius alone is not a good indicator of wrist mobility and hand posture. However, the extent of the lunate's articular arc on the dorsum of the capitate head is correlated with midcarpal mobility across taxa. These findings suggest that although a precise relationship between wrist extension ROM and morphology is difficult to define, the presence of a pronounced dorsal ridge combined with an abbreviated dorsal capitate articular arc reflects limited overall dorsiflexion with attendant constraints on the adoption of palmigrade hand postures. Anat Rec, 300:382-401, 2017. © 2016 Wiley Periodicals, Inc.

The evolutionary origin and population history of the grauer gorilla.

Gorillas living in western central Africa (Gorilla gorilla) are morphologically and genetically distinguishable from those living in eastern central Africa (Gorilla beringei). Genomic analyses show eastern gorillas experienced a significant reduction in population size during the Pleistocene subsequent to geographical isolation from their western counterparts. However, how these results relate more specifically to the recent biogeographical and evolutionary history of eastern gorillas remains poorly understood. Here we show that two rare morphological traits are present in the hands and feet of both eastern gorilla subspecies at strikingly high frequencies (>60% in G. b. graueri; ∼28% in G. b. beringei) in comparison with western gorillas (<1%). The intrageneric distribution of these rare traits suggests that they became common among eastern gorillas after diverging from their western relatives during the early to middle Pleistocene. The extremely high frequencies observed among grauer gorillas-which currently occupy a geographic range more than ten times the size of that of mountain gorillas-imply that grauers originated relatively recently from a small founding population of eastern gorillas. Current paleoenvironmental, geological, and biogeographical evidence supports the hypothesis that a small group of eastern gorillas likely dispersed westward from the Virungas into present-day grauer range in the highlands just north of Lake Kivu, either immediately before or directly after the Younger Dryas interval. We propose that as the lowland forests of central Africa expanded rapidly during the early Holocene, they became connected with the expanding highland forests along the Albertine Rift and enabled the descendants of this small group to widely disperse. The descendant populations significantly expanded their geographic range and population numbers relative to the gorillas of the Virunga Mountains and the Bwindi-Impenetrable Forest, ultimately resulting in the grauer gorilla subspecies recognized today. This founder-effect hypothesis offers some optimism for modern conservation efforts to save critically endangered eastern gorillas from extinction.

The hand of Homo naledi.

A nearly complete right hand of an adult hominin was recovered from the Rising Star cave system, South Africa. Based on associated hominin material, the bones of this hand are attributed to Homo naledi. This hand reveals a long, robust thumb and derived wrist morphology that is shared with Neandertals and modern humans, and considered adaptive for intensified manual manipulation. However, the finger bones are longer and more curved than in most australopiths, indicating frequent use of the hand during life for strong grasping during locomotor climbing and suspension. These markedly curved digits in combination with an otherwise human-like wrist and palm indicate a significant degree of climbing, despite the derived nature of many aspects of the hand and other regions of the postcranial skeleton in H. naledi.

Homo naledi, a new species of the genus Homo from the Dinaledi Chamber, South Africa.

Homo naledi is a previously-unknown species of extinct hominin discovered within the Dinaledi Chamber of the Rising Star cave system, Cradle of Humankind, South Africa. This species is characterized by body mass and stature similar to small-bodied human populations but a small endocranial volume similar to australopiths. Cranial morphology of H. naledi is unique, but most similar to early Homo species including Homo erectus, Homo habilis or Homo rudolfensis. While primitive, the dentition is generally small and simple in occlusal morphology. H. naledi has humanlike manipulatory adaptations of the hand and wrist. It also exhibits a humanlike foot and lower limb. These humanlike aspects are contrasted in the postcrania with a more primitive or australopith-like trunk, shoulder, pelvis and proximal femur. Representing at least 15 individuals with most skeletal elements repeated multiple times, this is the largest assemblage of a single species of hominins yet discovered in Africa.

Functional aspects of metatarsal head shape in humans, apes, and Old World monkeys.

Modern human metatarsal heads are typically described as "dorsally domed," mediolaterally wide, and dorsally flat. Despite the apparent functional importance of these features in forefoot stability during bipedalism, the distinctiveness of this morphology has not been quantitatively evaluated within a broad comparative framework. In order to use these features to reconstruct fossil hominin locomotor behaviors with any confidence, their connection to human bipedalism should be validated through a comparative analysis of other primates with different locomotor behaviors and foot postures, including species with biomechanical demands potentially similar to those of bipedalism (e.g., terrestrial digitigrady). This study explores shape variation in the distal metatarsus among humans and other extant catarrhines using three-dimensional geometric morphometrics (3 DGM). Shape differences among species in metatarsal head morphology are well captured by the first two principal components of Procrustes shape coordinates, and these two components summarize most of the variance related to "dorsal doming" and "dorsal expansion." Multivariate statistical tests reveal significant differences among clades in overall shape, and humans are reliably distinguishable from other species by aspects of shape related to a greater degree of dorsal doming. Within quadrupeds, terrestrial species also trend toward more domed metatarsal heads, but not to the extent seen in humans. Certain aspects of distal metatarsus shape are likely related to habitual dorsiflexion of the metatarsophalangeal joints, but the total morphological pattern seen in humans is distinct. These comparative results indicate that this geometric morphometric approach is useful to characterize the complexity of metatarsal head morphology and will help clarify its relationship with function in fossil primates, including early hominins.

Functional morphology of the hallucal metatarsal with implications for inferring grasping ability in extinct primates.

Primate evolutionary morphologists have argued that selection for life in a fine branch niche resulted in grasping specializations that are reflected in the hallucal metatarsal (Mt1) morphology of extant "prosimians", while a transition to use of relatively larger, horizontal substrates explains the apparent loss of such characters in anthropoids. Accordingly, these morphological characters-Mt1 torsion, peroneal process length and thickness, and physiological abduction angle-have been used to reconstruct grasping ability and locomotor mode in the earliest fossil primates. Although these characters are prominently featured in debates on the origin and subsequent radiation of Primates, questions remain about their functional significance. This study examines the relationship between these morphological characters of the Mt1 and a novel metric of pedal grasping ability for a large number of extant taxa in a phylogenetic framework. Results indicate greater Mt1 torsion in taxa that engage in hallucal grasping and in those that utilize relatively small substrates more frequently. This study provides evidence that Carpolestes simpsoni has a torsion value more similar to grasping primates than to any scandentian. The results also show that taxa that habitually grasp vertical substrates are distinguished from other taxa in having relatively longer peroneal processes. Furthermore, a longer peroneal process is also correlated with calcaneal elongation, a metric previously found to reflect leaping proclivity. A more refined understanding of the functional associations between Mt1 morphology and behavior in extant primates enhances the potential for using these morphological characters to comprehend primate (locomotor) evolution.