Comparative analysis of femoral biomechanical neck length in primates.

The unique abductor capability of the human lesser gluteal muscles among extant hominoids has been suggested to be associated, in part, with biomechanical neck length of the femur. Beyond the hominin lineage, the relationship between biomechanical neck length and locomotor performance remains unclear due, in part, to the limited number of primate taxa directly compared and the need to examine species characterized by a wider range of locomotor diversity. Measurements were taken on the proximal femora of 28 extant taxa, with each species being assigned to a locomotor and phylogenetic category. Pairwise comparisons and phylogenetic generalized least-squares analysis were performed to examine the impact of phylogeny and locomotor adaptation on relative biomechanical neck length. Arboreal quadrupeds that perform varying proportions of climbing/clambering versus leaping were characterized by different biomechanical neck lengths, whereas semi-terrestrial anthropoids that perform either knuckle-walking or palmigrade/digitigrade quadrupedalism were found to have similar relative neck lengths. Samples categorized as either orthograde clamberers or bipeds were distinct from all other anthropoid samples and characterized by the relatively shortest and longest biomechanical neck lengths, respectively. Results of additional analyses that included prosimian primates suggest that relatively long biomechanical necks characterize species adapted to hind limb-dominated forms of locomotion (e.g., vertical clinging and leaping and bipedalism). Thus, biomechanical neck length is useful for signaling reliance on bipedalism (as performed by humans) or leaping, including subtle variation in leaping performance among arboreal quadrupeds. Furthermore, this trait is informative regarding reliance on irregular gait clambering as performed by orangutans.

A Geometric Morphometric Examination of Hominoid Third Metacarpal Shape and Its Implications for Inferring the Precursor to Terrestrial Bipedalism.

The third metacarpal has been a focus of study when examining questions surrounding early hominin locomotion since this bone is adapted to the diverse range of positional behaviors performed by extant hominoids. The shape of this bone is potentially under strong selective pressure related to the biomechanical demands of terrestrial knuckle-walking, arboreal clambering, and brachiation performed by extant hominoids since the hand directly interacts with the substrate during the performance of these movements. The objective of the present study was to explore shape variation of the third metacarpal and examine how different parts of the bone discriminated between hominoid genera that perform these different locomotor behaviors. In addition to examining general interspecies variation, shape analysis was applied to testing the knuckle-walking hypothesis for human evolution. Fourteen 3D landmark coordinates were collected on hominoid third metacarpals, and principal component analysis and Procrustes distances were used to examine metacarpal shape. Comparable measurements were collected on fossilized third metacarpals of Australopithecus afarensis as an early hominin test case for examining the knuckle-walking hypothesis. Analyses that included landmarks collected on both ends of the bone distinguished humans from great apes and presented a strong functional signal related to suspensory locomotion among nonhuman hominoids, whereas the distal articular surface provided the strongest knuckle-walking signal. The shapes of Australopithecus afarensis metacarpals examined in the current study did not provide evidence for a trajectory of shape change in early hominin evolution that started from a metacarpal adapted for terrestrial knuckle-walking. Anat Rec, 302:983-998, 2019. © 2018 Wiley Periodicals, Inc.

Adaptation to suspensory locomotion in Australopithecus sediba.

Australopithecus sediba is represented by well-preserved fossilized remains from the locality of Malapa, South Africa. Recent work has shown that the combination of features in the limb skeleton of A. sediba was distinct from that of earlier species of Australopithecus, perhaps indicating that this species moved differently. The bones of the arm and forearm indicate that A. sediba was adapted to suspensory and climbing behaviors. We used a geometric morphometric approach to examine ulnar shape, potentially identifying adaptations to forelimb suspensory locomotion in A. sediba. Results indicated suspensory capabilities in this species and a stronger forelimb suspensory signal than has been documented in Australopithecus afarensis. Our study confirms the adaptive significance of functional morphological traits for arboreal movements in the locomotor repertoire of A. sediba and provides important insight into the diversity and mosaic nature of locomotor adaptations among early hominins.

Inferring the use of forelimb suspensory locomotion by extinct primate species via shape exploration of the ulna.

Uncovering links between skeletal morphology and locomotor behavior is an essential component of paleobiology because it allows researchers to infer the locomotor repertoire of extinct species based on preserved fossils. In this study, we explored ulnar shape in anthropoid primates using 3D geometric morphometrics to discover novel aspects of shape variation that correspond to observed differences in the relative amount of forelimb suspensory locomotion performed by species. The ultimate goal of this research was to construct an accurate predictive model that can be applied to infer the significance of these behaviors. We studied ulnar shape variation in extant species using principal component analysis. Species mainly clustered into phylogenetic groups along the first two principal components. Upon closer examination, the results showed that the position of species within each major clade corresponded closely with the proportion of forelimb suspensory locomotion that they have been observed to perform in nature. We used principal component regression to construct a predictive model for the proportion of these behaviors that would be expected to occur in the locomotor repertoire of anthropoid primates. We then applied this regression analysis to Pliopithecus vindobonensis, a stem catarrhine from the Miocene of central Europe, and found strong evidence that this species was adapted to perform a proportion of forelimb suspensory locomotion similar to that observed in the extant woolly monkey, Lagothrix lagothricha.

Geometric morphometrics and virtual anthropology: advances in human evolutionary studies.

Geometric morphometric methods have been increasingly used in paleoanthropology in the last two decades, lending greater power to the analysis and interpretation of the human fossil record. More recently the advent of the wide use of computed tomography and surface scanning, implemented in combination with geometric morphometrics (GM), characterizes a new approach, termed Virtual Anthropology (VA). These methodological advances have led to a number of developments in human evolutionary studies. We present some recent examples of GM and VA related research in human evolution with an emphasis on work conducted at the University of Tübingen and other German research institutions.

New Neanderthal remains from Mani peninsula, Southern Greece: the Kalamakia Middle Paleolithic cave site.

The Kalamakia cave, a Middle Paleolithic site on the western coast of the Mani peninsula, Greece, was excavated in 1993-2006 by an interdisciplinary team from the Ephoreia of Paleoanthropology and Speleology (Greek Ministry of Culture) and the Muséum national d'Histoire naturelle (Paris). The site is dated to between ca. 100,000 and >39,000 years BP (Before Present) and has yielded Mousterian lithics, a rich fauna, and human remains from several layers. The latter include 10 isolated teeth, a cranial fragment and three postcranial elements. The remains represent at least eight individuals, two of them subadults, and show both carnivore and anthropogenic modifications. They can be identified as Neanderthal on the basis of diagnostic morphology on most specimens. A diet similar to that of Neanderthals from mixed habitat is suggested by our analysis of dental wear (occlusal fingerprint analysis) and microwear (occlusal texture microwear analysis), in agreement with the faunal and palynological analyses of the site. These new fossils significantly expand the Neanderthal sample known from Greece. Together with the human specimens from Lakonis and Apidima, the Kalamakia human remains add to the growing evidence of a strong Neanderthal presence in the Mani region during the Late Pleistocene.

Exploring third metacarpal capitate facet shape in early hominins.

The joint between the capitate and third metacarpal plays an important role in stabilizing the manus during hand use in great apes and humans. Researchers have examined the morphology of this region in humans, our fossil relatives, and other extant primates to try to understand the importance of this joint in human evolution. The first goal of our research was to explore shape variation of the third metacarpal capitate facet across extant anthropoids, including hominoids, cercopithecoids, and platyrrhines. This analysis allowed us to examine the range of variation in the capitate facet and the degree to which locomotor behavior, phylogeny, and size explained shape variation. We also examined capitate facet shape in the early hominin fossil record in order to explore how the shape of this articular surface has changed during early hominin evolution. We captured six landmark coordinates on the edge of the capitate facet in extant anthropoids and fossil specimens to quantify and visualize shape variation in this region. We used principal components analysis, Procrustes distances, and multivariate regression analysis to investigate different possible influences on shape variation. We found that shape variation corresponded to function, phylogeny, and size. With the exception of brachiation, shape variation did not clearly correspond with any specific locomotor behavior. However, we identified a shift in the relative mediolateral breadth of the capitate facet during early hominin evolution, which is most likely one of several adaptations for a more stable joint surface.

Metacarpophalangeal joint orientation in anthropoid manual phalanges.

The proximal articular surface angle of orientation (AO) of proximal phalanges of the hand and foot has been used to infer the locomotor profile of extinct Miocene catarrhines and early hominins. Previous work has found that joint orientation distinguishes quadrupedal from suspensory anthropoids. The purpose of this study is to expand on previous research by examining this feature within and across several primate clades, allowing us to investigate the potential influences of locomotion, substrate usage, hand posture, and phylogeny. We also report AO measurements in human proximal hand phalanges, allowing us to examine human skeletal variation within a wide comparative context. The angle of orientation was measured on manual proximal third phalanges of 21 extant anthropoid species using a Microscribe digitizer. Comparisons were made between locomotor groups within hominoids, platyrrhines, and cercopithecoids. Proximal phalanges of quadrupedal species were characterized by greater dorsal orientation than those of suspensory taxa in hominoids and atelids. In addition, arboreal quadrupeds had greater AO values than terrestrial quadrupeds within the Cercopithecoidea. However, within the terrestrial locomotor group, mean AO values did not differ between palmigrade and digitigrade taxa. Thus, while there appears to be a functional signal related to substrate usage, differences in use of hand postures when moving on the ground were not reflected in proximal joint orientation of the proximal phalanx. Finally, we measured relatively low AO values in human phalanges, which might be related to integration with serially homologous pedal phalanges that are under strong selective pressure related to bipedalism.

The correspondence between proximal phalanx morphology and locomotion: implications for inferring the locomotor behavior of fossil catarrhines.

Phalanges are considered to be highly informative in the reconstruction of extinct primate locomotor behavior since these skeletal elements directly interact with the substrate during locomotion. Variation in shaft curvature and relative phalangeal length has been linked to differences in the degree of suspension and overall arboreal locomotor activities. Building on previous work, this study investigated these two skeletal characters in a comparative context to analyze function, while taking evolutionary relationships into account. This study examined the correspondence between proportions of suspension and overall substrate usage observed in 17 extant taxa and included angle of curvature and relative phalangeal length. Predictive models based on these traits are reported. Published proportions of different locomotor behaviors were regressed against each phalangeal measurement and a size proxy. The relationship between each behavior and skeletal trait was investigated using ordinary least-squares, phylogenetic generalized least-squares (pGLS), and two pGLS transformation methods to determine the model of best-fit. Phalangeal curvature and relative length had significant positive relationships with both suspension and overall arboreal locomotion. Cross-validation analyses demonstrated that relative length and curvature provide accurate predictions of relative suspensory behavior and substrate usage in a range of extant species when used together in predictive models. These regression equations provide a refined method to assess the amount of suspensory and overall arboreal locomotion characterizing species in the catarrhine fossil record.

Skeletal correlates of quadrupedalism and climbing in the anthropoid forelimb: implications for inferring locomotion in Miocene catarrhines.

Several well-known Miocene catarrhines, including Proconsul heseloni, have been inferred to combine quadrupedal walking in an arboreal substrate with a significant amount of climbing during locomotion. The degree to which some of these species were adapted to perform these behaviors is not fully understood due to the mosaic of 'ape-like' and 'monkey-like' traits identified in the forelimb. Given these unique combinations of forelimb features in the fossils, we report on forelimb traits that should be emphasized when investigating skeletal adaptation to quadrupedalism (defined in this manuscript as symmetrical gait movement on horizontal supports, excluding knuckle-walking) and climbing (including both vertical climbing and clambering). We investigate the correspondence between: 1) quadrupedalism and two well-known forelimb traits, humeral torsion and olecranon process length, and 2) climbing and phalangeal curvature. We also test the degree of phylogenetic signal in these relationships using phylogenetic generalized least-squares and branch length transformation methods in order to determine the models of best-fit. We present models that can be used to predict proportions of quadrupedalism and climbing in extant and extinct anthropoid taxa. Each trait-behavior correlation is significant and characterized by an absence of phylogenetic signal. Thus, we employ models assuming a star phylogeny to predict locomotor proportions. The climbing model based on phalangeal curvature and a proxy for size provides the most accurate predictions of behavior across anthropoids. The two quadrupedalism models are less accurate, but distinguish highly quadrupedal species from those that are not. Predictive equations based on these traits support the inference that P. heseloni performed a high proportion of quadrupedalism with a significant climbing component. The degree of phalangeal curvature measured in Pliopithecus vindobonensis predicts that this Miocene catarrhine species performed a proportion of climbing similar to Proconsul, while humeral torsion and olecranon process length provide conflicting inferences of quadrupedal locomotion in this species.

Brain structure variation in great apes, with attention to the mountain gorilla (Gorilla beringei beringei).

This report presents data regarding the brain structure of mountain gorillas (Gorilla beringei beringei) in comparison with other great apes. Magnetic resonance (MR) images of three mountain gorilla brains were obtained with a 3T scanner, and the volume of major neuroanatomical structures (neocortical gray matter, hippocampus, thalamus, striatum, and cerebellum) was measured. These data were included with our existing database that includes 23 chimpanzees, three western lowland gorillas, and six orangutans. We defined a multidimensional space by calculating the principal components (PCs) from the correlation matrix of brain structure fractions in the well-represented sample of chimpanzees. We then plotted data from all of the taxa in this space to examine phyletic variation in neural organization. Most of the variance in mountain gorillas, as well as other great apes, was contained within the chimpanzee range along the first two PCs, which accounted for 61.73% of the total variance. Thus, the majority of interspecific variation in brain structure observed among these ape taxa was no greater than the within-species variation seen in chimpanzees. The loadings on PCs indicated that the brain structure of great apes differs among taxa mostly in the relative sizes of the striatum, cerebellum, and hippocampus. These findings suggest possible functional differences among taxa in terms of neural adaptations for ecological and locomotor capacities. Importantly, these results fill a critical gap in current knowledge regarding great ape neuroanatomical diversity.