Cortical bone architecture of hominid intermediate phalanges reveals functional signals of locomotion and manipulation.

OBJECTIVES: Reconstruction of fossil hominin manual behaviors often relies on comparative analyses of extant hominid hands to understand the relationship between hand use and skeletal morphology. In this context, the intermediate phalanges remain understudied. Thus, here we investigate cortical bone morphology of the intermediate phalanges of extant hominids and compare it to the cortical structure of the proximal phalanges, to investigate the relationship between cortical bone structure and inferred loading during manual behaviors. MATERIALS AND METHODS: Using micro-CT data, we analyze cortical bone structure of the intermediate phalangeal shaft of digits 2-5 in Pongo pygmaeus (n = 6 individuals), Gorilla gorilla (n = 22), Pan spp. (n = 23), and Homo sapiens (n = 23). The R package morphomap is used to study cortical bone distribution, cortical thickness and cross-sectional properties within and across taxa. RESULTS: Non-human great apes generally have thick cortical bone on the palmar shaft, with Pongo only having thick cortex on the peaks of the flexor sheath ridges, while African apes have thick cortex along the entire flexor sheath ridge and proximal to the trochlea. Humans are distinct in having thicker dorsal shaft cortex as well as thick cortex at the disto-palmar region of the shaft. DISCUSSION: Variation in cortical bone distribution and properties of the intermediate phalanges is consistent with differences in locomotor and manipulative behaviors in extant great apes. Comparisons between the intermediate and proximal phalanges reveals similar patterns of cortical bone distribution within each taxon but with potentially greater load experienced by the proximal phalanges, even in knuckle-walking African apes. This study provides a comparative context for the reconstruction of habitual hand use in fossil hominins and hominids.

Evaluating the muscle attachment hypothesis for sagittal cresting in Gorilla and Pongo.

Primate mandibular morphology is often associated with jaw functionality of the masticatory complex in the context of variation in diets. Recent research into the disparities between the diet and jaw functionality in male and female hominoids is inconclusive and suggests that sexual dimorphism in the mandible may be influenced by external factors such as temporalis and masseter muscle morphology, which in turn may be influenced by sexual selection. As the muscles associated with mastication (i.e., the type of chewing exhibited by primates and other mammals) encompass the mandible as well as the neurocranium, including the sagittal crest among some individuals, this study investigates sex-specific associations between regions of the mandibular ramus and neurocranium associated with mastication in a dentally mature sample of Gorilla and Pongo. A total of four cranial and mandibular variables were measured in two Gorilla taxa (Gorilla gorilla gorilla and Gorilla beringei graueri) and one Pongo taxon (Pongo pygmaeus pygmaeus) (n = 220). For all three taxa, we investigate (a) whether the degree of sexual dimorphism in cranial regions associated with sagittal cresting (sagittal crest size (SCS) and temporalis muscle attachment area (TMAA)) is proportional to the degree of mandibular ramus area (MRA) and coronoid process height (CPH) sexual dimorphism, (b) whether there are sex differences in scaling relationships between TMAA and MRA, and (c) whether there are sex differences in the strength of association between TMAA and CPH. We show that for G. g. gorilla, variables associated with sagittal cresting show higher sexual dimorphism values than our two mandibular ramus variables, which is not the case for G. b. graueri or for P. p. pygmaeus. All three taxa show similar sex-specific scaling relationships between TMAA and MRA, where for males this relationship does not diverge from isometry, and for females there is a negative allometric relationship. Our findings also show intraspecific sex differences in allometric slopes between MRA and TMAA for all three taxa. Only G. g. gorilla shows a significant association between TMAA and CPH, which is observed in both sexes. Although there are some statistical associations between the cranial and mandibular regions associated with mastication, our results show that among male gorillas and orangutans, patterns of variation in the sagittal crest, TMAA, mandibular ramus and the coronoid process cannot be explained by the muscle attachment hypothesis alone. These findings have implications surrounding the associations between social behaviour and the morphology of the craniofacial complex.

Comparative genomic analyses provide new insights into evolutionary history and conservation genomics of gorillas.

Genome sequencing is a powerful tool to understand species evolutionary history, uncover genes under selection, which could be informative of local adaptation, and infer measures of genetic diversity, inbreeding and mutational load that could be used to inform conservation efforts. Gorillas, critically endangered primates, have received considerable attention and with the recently sequenced Bwindi mountain gorilla population, genomic data is now available from all gorilla subspecies and both mountain gorilla populations. Here, we reanalysed this rich dataset with a focus on evolutionary history, local adaptation and genomic parameters relevant for conservation. We estimate a recent split between western and eastern gorillas of 150,000-180,000 years ago, with gene flow around 20,000 years ago, primarily between the Cross River and Grauer's gorilla subspecies. This gene flow event likely obscures evolutionary relationships within eastern gorillas: after excluding putatively introgressed genomic regions, we uncover a sister relationship between Virunga mountain gorillas and Grauer's gorillas to the exclusion of Bwindi mountain gorillas. This makes mountain gorillas paraphyletic. Eastern gorillas are less genetically diverse and more inbred than western gorillas, yet we detected lower genetic load in the eastern species. Analyses of indels fit remarkably well with differences in genetic diversity across gorilla taxa as recovered with nucleotide diversity measures. We also identified genes under selection and unique gene variants specific for each gorilla subspecies, encoding, among others, traits involved in immunity, diet, muscular development, hair morphology and behavior. The presence of this functional variation suggests that the subspecies may be locally adapted. In conclusion, using extensive genomic resources we provide a comprehensive overview of gorilla genomic diversity, including a so-far understudied Bwindi mountain gorilla population, identify putative genes involved in local adaptation, and detect population-specific gene flow across gorilla species.

Morphological integration and shape covariation between the trapezium and first metacarpal among extant hominids.

OBJECTIVES: The shape of the trapezium and first metacarpal (Mc1) markedly influence thumb mobility, strength, and the manual abilities of extant hominids. Previous research has typically focused solely on trapezium-Mc1 joint shape. Here we investigate how morphological integration and shape covariation between the entire trapezium (articular and non-articular surfaces) and the entire Mc1 reflect known differences in thumb use in extant hominids. MATERIALS AND METHODS: We analyzed shape covariation in associated trapezia and Mc1s across a large, diverse sample of Homo sapiens (n = 40 individuals) and other extant hominids (Pan troglodytes, n = 16; Pan paniscus, n = 13; Gorilla gorilla gorilla, n = 27; Gorilla beringei, n = 6; Pongo pygmaeus, n = 14; Pongo abelii, n = 9) using a 3D geometric morphometric approach. We tested for interspecific significant differences in degree of morphological integration and patterns of shape covariation between the entire trapezium and Mc1, as well as within the trapezium-Mc1 joint specifically. RESULTS: Significant morphological integration was only found in the trapezium-Mc1 joint of H. sapiens and G. g. gorilla. Each genus showed a specific pattern of shape covariation between the entire trapezium and Mc1 that was consistent with different intercarpal and carpometacarpal joint postures. DISCUSSION: Our results are consistent with known differences in habitual thumb use, including a more abducted thumb during forceful precision grips in H. sapiens and a more adducted thumb in other hominids used for diverse grips. These results will help to infer thumb use in fossil hominins.

"Untying the knot": The primitive orofacial muscle architecture in the gorilla (Gorilla gorilla) as a key to the evolution of hominin facial movement.

The anatomical underpinnings of primate facial expressions are essential to exploring their evolution. Traditionally, it has been accepted that the primate face exhibits a "scala natura" morphocline, ranging from primitive to derived characteristics. At the primitive end, the face consists of undifferentiated muscular sheets, while at the derived end there is greater complexity with more muscles and insertion points. Among these, the role of the human modiolus ("knoten" in German) has been emphasized. Recent studies have challenged this view by revealing significant complexity in the faces of several non-human primates, thereby rejecting the linear notion of facial evolution. However, our knowledge of the facial architecture in gorillas, the second closest living relatives to modern humans, remains a significant gap in the literature. Here, we present new findings based on dissection and histological analysis of one gorilla craniofacial specimen, alongside 30 human hemifaces. Our results indicate that while the number and overall arrangement of facial muscles in the gorilla are comparable to those of chimpanzees and modern humans, several orofacial features distinguish the gorilla's anatomy from that of hominins. Among these are the absence of a modiolus, the continuity of muscular fibers over the region of the mouth corner, the flat (uncurving) sheet of the orbicularis oris muscle, and the insertion of direct labial tractors both anterior and posterior to it. Collectively, the anatomical characteristics observed in the gorilla suggest that the complex anatomy of the hominin face should be considered synapomorphic (shared-derived) within the Pan-Homo clade.

Intraspecific variation of long bone cross-sectional properties in Pan troglodytes troglodytes and Gorilla gorilla gorilla.

OBJECTIVES: Morphological intraspecific variation is due to the balance between skeletal plasticity and genetic constraint on the skeleton. Osteogenic responses to external stimuli, such as locomotion, have been well documented interspecifically across the primate order, but less so at the intraspecific level. Here, we examine the differences in cross-sectional variability of the femur, humerus, radius, and tibia in Pan troglodytes troglodytes versus Gorilla gorilla gorilla. We investigate whether there are sex, species, bone, and trait differences in response to variable body size and locomotion. MATERIALS AND METHODS: Adult male and female P. t. troglodytes and G. g. gorilla long bones from the Cleveland Museum of Natural History were scanned with a peripheral quantitative computer tomography system. Scans were taken at the midshaft of each bone according to functional bone length. Coefficients of variation were used to provide a size-independent measure of variation. We applied a Bonferroni correction to account for the multiple pairwise tests. RESULTS: There were limited significant differences between males and females, however, females tended to be more variable than males. Variation in Gorilla, when significant, was greater than in Pan, although significant differences were limited. There were no differences between bone variability in male and female Gorilla, and female Pan. DISCUSSION: Increased female variability may be due to more variable locomotor behavior, particularly during periods of pregnancy, lactation, and caring for an offspring compared to consistent locomotion over the life course by males. Body size may be a contributing factor to variability; more work is needed to understand this relationship.

Morphological variation of the Pan talus relative to that of Gorilla.

OBJECTIVES: Differences in talar articular morphology relative to locomotion have recently been found within Pan and Gorilla. Whole-bone talar morphology within, and shared variation among, Pan and Gorilla (sub)species, however, has yet to be investigated. Here we separately analyze talar external shape within Pan (P. t. troglodytes, P. t. schweinfurthii, P. t. verus, P. paniscus) and Gorilla (G. g. gorilla, G. b. beringei, G. b. graueri) relative to degree of arboreality and body size. Pan and Gorilla are additionally analyzed together to determine if consistent shape differences exist within the genera. MATERIALS AND METHODS: Talar external shape was quantified using a weighted spherical harmonic analysis. Shape variation both within and among Pan and Gorilla was described using principal component analyses. Root mean square distances were calculated between taxon averages, and resampling statistics conducted to test for pairwise differences. RESULTS: P. t. verus (most arboreal Pan) talar shape significantly differs from other Pan taxa (p < 0.05 for pairwise comparisons) driven by more asymmetrical trochlear rims and a medially-set talar head. P. t. troglodytes, P. t. schweinfurthii, and P. paniscus do not significantly differ (p > 0.05 for pairwise comparisons). All gorilla taxa exhibit significantly different talar morphologies (p < 0.007 for pairwise comparisons). The more terrestrial subspecies of G. beringei and P. troglodytes exhibit a superoinferiorly taller talar head/neck complex. DISCUSSION: P. t. verus exhibits talar morphologies that have been previously related to more frequent arboreality. The adaptations in the more terrestrial G. beringei and P. troglodytes subspecies may serve to facilitate load transmission.

Cortical bone distribution of the proximal phalanges in great apes: implications for reconstructing manual behaviours.

Primate fingers are typically in direct contact with the environment during both locomotion and manipulation, and aspects of external phalangeal morphology are known to reflect differences in hand use. Since bone is a living tissue that can adapt in response to loading through life, the internal bone architecture of the manual phalanges should also reflect differences in manual behaviours. Here, we use the R package Morphomap to analyse high-resolution microCT scans of hominid proximal phalanges of digits 2-5 to determine whether cortical bone structure reflects variation in manual behaviours between bipedal (Homo), knuckle-walking (Gorilla, Pan) and suspensory (Pongo) taxa. We test the hypothesis that relative cortical bone distribution patterns and cross-sectional geometric properties will differ both among extant great apes and across the four digits due to locomotor and postural differences. Results indicate that cortical bone structure reflects the varied hand postures employed by each taxon. The phalangeal cortices of Pongo are significantly thinner and have weaker cross-sectional properties relative to the African apes, yet thick cortical bone under their flexor sheath ridges corresponds with predicted loading during flexed finger grips. Knuckle-walking African apes have even thicker cortical bone under the flexor sheath ridges, as well as in the region proximal to the trochlea, but Pan also has thicker diaphyseal cortices than Gorilla. Humans display a distinct pattern of distodorsal thickening, as well as relatively thin cortices, which may reflect the lack of phalangeal curvature combined with frequent use of flexed fingered hand grips during manipulation. Within each taxon, digits 2-5 have a similar cortical distribution in Pongo, Gorilla and, unexpectedly, Homo, which suggest similar loading of all fingers during habitual locomotion or hand use. In Pan, however, cortical thickness differs between the fingers, potentially reflecting differential loading during knuckle-walking. Inter- and intra-generic variation in phalangeal cortical bone structure reflects differences in manual behaviours, offering a comparative framework for reconstructing hand use in fossil hominins.

Carpal allometry of African apes among mammals.

OBJECTIVES: Morphological variation in African ape carpals has been used to support the idea that Pan and Gorilla evolved knuckle-walking independently. Little work, however, has focused on the effect of body mass on carpal morphology. Here, we compare carpal allometry in Pan and Gorilla to that of other quadrupedal mammals with similar body mass differences. If allometric trends in Pan and Gorilla carpals mirror those of other mammals with similar body mass variation, then body mass differences may provide a more parsimonious explanation for African ape carpal variation than the independent evolution of knuckle-walking. MATERIALS AND METHODS: Three linear measurements were collected on the capitate, hamate, lunate, and scaphoid (or scapholunate) of 39 quadrupedal species from six mammalian families/subfamilies. Relationships between linear measurements and estimated body mass were analyzed using reduced major axis regression. Slopes were compared to 0.33 for isometry. RESULTS: Within Hominidae, higher body mass taxa (Gorilla) have relatively anteroposteriorly wider, mediolaterally wider, and/or proximodistally shorter capitates, hamates, and scaphoids than low body mass taxa (Pan). These allometric relationships are mirrored in most, but not all, mammalian families/subfamilies included in the analysis. CONCLUSIONS: Within most mammalian families/subfamilies, carpals of high body mass taxa are proximodistally shorter, anteroposteriorly wider, and mediolaterally wider than those of low body mass taxa. These distinctions may be caused by the need to accommodate relatively higher forelimb loading associated with greater body mass. Because these trends occur within multiple mammalian families/subfamilies, some carpal variation in Pan and Gorilla is consistent with body mass differences.

Modularity of the wrist in extant hominines.

Wrist shape varies greatly across primates and previous studies indicate that the numerous morphological differences among them are related to a complex mixture of phylogeny and function. However, little is known about whether the variation in these various anatomical differences is linked and to what extent the wrist bones vary independently. Here, we used 3D geometric morphometrics on a sample of extant hominines (Homo sapiens, Pan troglodytes, Gorilla gorilla, and Gorilla beringei), to find the model that best describes the covariation patterns among four of the eight carpals (i.e., capitate, lunate, scaphoid, and trapezium). For this purpose, 15 modular hypotheses were tested using the Covariance Ratio. Results indicate that there is a covariation structure common to all hominines, which corresponds to stronger covariation within each carpal as compared to the covariation between carpals. However, the results also indicate that that there is a degree of codependence in the variation of some carpals, which is unique in humans, chimpanzees, and gorillas, respectively. In humans there is evidence of associated shape changes between the lunate and capitate, and between the scaphoid and trapezium. This covariation between lunate and capitate is also apparent in gorillas, while chimpanzees display the greatest disassociation among carpals, showing low covariation values in all pairwise comparisons. Our analyses indicate that carpals have an important level of variational independence which might suggest a high degree of independent evolvability in the wrists of hominines, and that although weak, the structure of associated changes of these four carpals varies across genera. To our knowledge this is the first report on the patterns of modularity between these four wrist bones in the Homininae and future studies might attempt to investigate whether the anatomical shape associations among carpals are functionally related to locomotion and manipulation.

Morphological integration in the hominid midfoot.

The calculation of morphological integration across living apes and humans may provide important insights into the potential influence of integration on evolutionary trajectories in the hominid lineage. Here, we quantify magnitudes of morphological integration among and within elements of the midfoot in great apes and humans to examine the link between locomotor differences and trait covariance. We test the hypothesis that the medial elements of the great ape foot are less morphologically integrated with one another compared to humans based on their abducted halluces, and aim to determine how adaptations for midfoot mobility/stiffness and locomotor specialization influence magnitudes of morphological integration. The study sample is composed of all cuneiforms, the navicular, the cuboid, and metatarsals 1-5 of Homo sapiens (n = 80), Pan troglodytes (n = 63), Gorilla gorilla (n = 39), and Pongo sp. (n = 41). Morphological integration was quantified using the integration coefficient of variation of interlandmark distances organized into sets of a priori-defined modules. Magnitudes of integration across these modules were then compared against sets of random traits from the whole midfoot. Results show that all nonhuman apes have less integrated medial elements, whereas humans have highly integrated medial elements, suggesting a link between hallucal abduction and reduced levels of morphological integration. However, we find considerable variation in magnitudes of morphological integration across metatarsals 2-5, the intermediate and lateral cuneiform, the cuboid, and navicular, emphasizing the influence of functional and nonfunctional factors in magnitudes of integration. Lastly, we find that humans and orangutans show the lowest overall magnitudes of integration in the midfoot, which may be related to their highly specialized functions, and suggest a link between strong diversifying selection and reduced magnitudes of morphological integration.

Morphological variation of the maxilla in modern humans and African apes.

Differences in morphology among modern humans and African apes are frequently used when assessing whether hominin fossils should be attributed to a single species or represent evidence for taxic diversity. A good understanding of the degree and structure of the intergeneric, interspecific, and intraspecific variation, including aspects such as sexual dimorphism and age, are key in this context. Here we explore the variation and differences shown by the maxilla of extant hominines, as maxillary morphology is central in the diagnosis of several hominin taxa. Our sample includes adults of all currently recognized hominine species and subspecies, with a balanced species sex ratio. In addition, we compared the adults with a small sample of late juveniles. The morphology of the maxillae was captured using three-dimensional landmarks, and the size and shape were analyzed using geometric morphometric methods. Key observations are that 1) the maxillae of all extant hominine species and subspecies show statistically significant differences, but complete separation in shape is only seen at the genus level; 2) the degree of variation is not consistent between genera, with subspecies of Gorilla being more different from each other than are species of Pan; 3) the pattern of sexual shape dimorphism is different in Pan, Gorilla, and Homo, often showing opposite trends; and 4) differentiation between maxillary shapes is increased after adjustment for static intraspecific allometry. These results provide a taxonomically up-to-date comparative morphological framework to help interpret the hominin fossil record, and we discuss the practical implications in that context.

Three-dimensional polygonal muscle modelling and line of action estimation in living and extinct taxa.

Biomechanical models and simulations of musculoskeletal function rely on accurate muscle parameters, such as muscle masses and lines of action, to estimate force production potential and moment arms. These parameters are often obtained through destructive techniques (i.e., dissection) in living taxa, frequently hindering the measurement of other relevant parameters from a single individual, thus making it necessary to combine multiple specimens and/or sources. Estimating these parameters in extinct taxa is even more challenging as soft tissues are rarely preserved in fossil taxa and the skeletal remains contain relatively little information about the size or exact path of a muscle. Here we describe a new protocol that facilitates the estimation of missing muscle parameters (i.e., muscle volume and path) for extant and extinct taxa. We created three-dimensional volumetric reconstructions for the hindlimb muscles of the extant Nile crocodile and extinct stem-archosaur Euparkeria, and the shoulder muscles of an extant gorilla to demonstrate the broad applicability of this methodology across living and extinct animal clades. Additionally, our method can be combined with surface geometry data digitally captured during dissection, thus facilitating downstream analyses. We evaluated the estimated muscle masses against physical measurements to test their accuracy in estimating missing parameters. Our estimated muscle masses generally compare favourably with segmented iodine-stained muscles and almost all fall within or close to the range of observed muscle masses, thus indicating that our estimates are reliable and the resulting lines of action calculated sufficiently accurately. This method has potential for diverse applications in evolutionary morphology and biomechanics.

Relationships between the hard and soft dimensions of the nose in Pan troglodytes and Homo sapiens reveal the positions of the nasal tips of Plio-Pleistocene hominids.

By identifying homogeneity in bone and soft tissue covariation patterns in living hominids, it is possible to produce facial approximation methods with interspecies compatibility. These methods may be useful for producing facial approximations of fossil hominids that are more realistic than currently possible. In this study, we conducted an interspecific comparison of the nasomaxillary region in chimpanzees and modern humans with the aim of producing a method for predicting the positions of the nasal tips of Plio-Pleistocene hominids. We addressed this aim by first collecting and performing regression analyses of linear and angular measurements of nasal cavity length and inclination in modern humans (Homo sapiens; n = 72) and chimpanzees (Pan troglodytes; n = 19), and then performing a set of out-of-group tests. The first test was performed on four subjects that belonged to the same genus as the training sample, i.e., Homo (n = 2) and Pan (n = 2), and the second test, which functioned as an interspecies compatibility test, was performed on Pan paniscus (n = 1), Gorilla gorilla (n = 3), Pongo pygmaeus (n = 1), Pongo abelli (n = 1), Symphalangus syndactylus (n = 3), and Papio hamadryas (n = 3). We identified statistically significant correlations in both humans and chimpanzees with slopes that displayed homogeneity of covariation. Prediction formulae combining these data were found to be compatible with humans and chimpanzees as well as all other African great apes, i.e., bonobos and gorillas. The main conclusion that can be drawn from this study is that our set of regression models for approximating the position of the nasal tip are homogenous among humans and African apes, and can thus be reasonably extended to ancestors leading to these clades.

Ontogenetic changes to metacarpal trabecular bone structure in mountain and western lowland gorillas.

The trabecular bone morphology of adult extant primates has been shown to reflect mechanical loading related to locomotion. However, ontogenetic studies of humans and other mammals suggest an adaptive lag between trabecular bone response and current mechanical loading patterns that could result in adult trabecular bone morphology reflecting juvenile behaviours. This study investigates ontogenetic changes in the trabecular bone structure of the third metacarpal of mountain gorillas (Gorilla beringei beringei; n = 26) and western lowland gorillas (Gorilla gorilla gorilla; n = 26) and its relationship to expected changes in locomotor loading patterns. Results show that trabecular bone reflects predicted mechanical loading throughout ontogeny. Bone volume fraction, trabecular thickness and trabecular number are low at birth and increase with age, although degree of anisotropy remains relatively stable throughout ontogeny. A high concentration of bone volume fraction can be observed in the distopalmar region of the third metacarpal epiphysis in early ontogeny, consistent with the high frequency of climbing, suspensory and other grasping behaviours in young gorillas. High trabecular bone concentration increases dorsally in the epiphysis during the juvenile period as terrestrial knuckle-walking becomes the primary form of locomotion. However, fusion of the epiphysis does not take place until 10-11 years of age, and overall trabecular structure does not fully reflect the adult pattern until 12 years of age, indicating a lag between adult-like behaviours and adult-like trabecular morphology. We found minimal differences in trabecular ontogeny between mountain and western lowland gorillas, despite presumed variation in the frequencies of arboreal locomotor behaviours. Altogether, ontogenetic changes in Gorilla metacarpal trabecular structure reflect overall genus-level changes in locomotor behaviours throughout development, but with some ontogenetic lag that should be considered when drawing functional conclusions from bone structure in extant or fossil adolescent specimens.

Covariation between the cranium and the cervical vertebrae in hominids.

The analysis of patterns of integration is crucial for the reconstruction and understanding of how morphological changes occur in a taxonomic group throughout evolution. These patterns are relatively constant; however, both patterns and the magnitudes of integration may vary across species. These differences may indicate morphological diversification, in some cases related to functional adaptations to the biomechanics of organisms. In this study, we analyze patterns of integration between two functional and developmental structures, the cranium and the cervical spine in hominids, and we quantify the amount of divergence of each anatomical element through phylogeny. We applied these methods to three-dimensional data from 168 adult hominid individuals, summing a total of more than 1000 cervical vertebrae. We found the atlas (C1) and axis (C2) display the lowest covariation with the cranium in hominids (Homo sapiens, Pan troglodytes, Pan paniscus, Gorilla gorilla, Gorilla beringei, Pongo pygmaeus). H. sapiens show a relatively different pattern of craniocervical correlation compared with chimpanzees and gorillas, especially in variables implicated in maintaining the balance of the head. Finally, the atlas and axis show lower magnitude of shape change during evolution than the rest of the cervical vertebrae, especially those located in the middle of the subaxial cervical spine. Overall, results suggest that differences in the pattern of craniocervical correlation between humans and gorillas and chimpanzees could reflect the postural differences between these groups. Also, the stronger craniocervical integration and larger magnitude of shape change during evolution shown by the middle cervical vertebrae suggests that they have been selected to play an active role in maintaining head balance.

3D geometric morphometrics analysis of mandibular fragments of Ouranopithecus macedoniensis from the late Miocene deposits of Central Macedonia, Greece.

OBJECTIVES: To explore mandibular shape differences between Ouranopithecus macedoniensis and a comparative sample of extant great apes using three-dimensional (3D) geometrics morphometrics. Other objectives are to assess mandibular shape variation and homogeneity within Ouranopithecus, explore the effects of size on mandibular shape, and explore the degree of mandibular sexual size dimorphism in Ouranopithecus. MATERIALS AND METHODS: The comparative sample comprises digitized mandibles from adult extant great apes. The 3D analysis includes three datasets: one with landmarks registered on the mandibular corpus and symphysis of mandibles preserving both sides, one on hemimandibles only, and one focused on the ramus and gonial area. Multivariate statistical analyses were conducted, such as ordination analyses (PCA), intra-specific Procrustes distances pairs, pairwise male-female centroid size differences, and correlation analyses. RESULTS: The male and female specimens of Ouranopithecus have mandibular shapes that are quite similar, although differences exist. The Procrustes distances results suggest more shape variation in Ouranopithecus than in the extant great apes. Ouranopithecus shows some similarities in mandibular shape to the larger great apes, Gorilla and Pongo. Moreover, the degree of sexual dimorphism in the small Ouranopithecus sample is greater than any of the great apes. Based on our correlation analyses of principal components (PC) with size, some PCs are significantly correlated with size, with correlation varying from moderate to substantial. DISCUSSION: This study attempted to understand better the variation within the mandibles of O. macedoniensis and the expression of sexual dimorphism in this taxon in more detail than has been done previously. The overall mandibular morphology of Ouranopithecus shows some similarities to those of the larger great apes, which likely reflects similarities in size. Compared to Gorilla and Pongo, O. macedoniensis shows an elevated degree of morphological variation, although limitations relating to sample size apply. Sexual dimorphism in the mandibles of O. macedoniensis appears to be relatively high, seemingly greater than in Gorilla and high even in comparison to Pongo, but this again is possibly in part an artifact of a small sample size.

Body proportions and environmental adaptation in gorillas.

OBJECTIVES: Limb length and trunk proportions are determined in a large, taxonomically and environmentally diverse sample of gorillas and related to variation in locomotion, climate, altitude, and diet. MATERIALS AND METHODS: The sample includes 299 gorilla skeletons, 115 of which are infants and juveniles, distributed between western lowland (G. gorilla gorilla), low and high elevation grauer (G. beringei graueri), and Virunga mountain gorillas (G. b. beringei). Limb bone and vertebral column lengths scaled to body mass are compared between subgroups by age group. RESULTS: All G. beringei have relatively short 3rd metapodials and manual proximal phalanges compared to G. gorilla, and this difference is apparent in infancy. All G. beringei also have shortened total limb lengths relative to either body mass or vertebral column length, although patterns of variation in individual skeletal elements are more complex, and infants do not display the same patterns as adults. Mountain gorillas have relatively long clavicles, present in infancy, and a relatively long thoracic (but not lumbosacral) vertebral column. DISCUSSION: A variety of environmental factors likely contributed to observed patterns of morphological variation among extant gorillas. We interpret the short hand and foot bones of all G. beringei as genetic adaptations to greater terrestriality in the last common ancestor of G. beringei; variation in other limb lengths to climatic adaptation, both genetic and developmental; and the larger thorax of G. b. beringei to adaptation to reduced oxygen pressure at high altitudes, again as a product of both genetic differences and environmental influences during development.

Morphological differences in the calcaneus among extant great apes investigated by three-dimensional geometric morphometrics.

Investigating the morphological differences of the calcaneus in humans and great apes is crucial for reconstructing locomotor repertories of fossil hominins. However, morphological variations in the calcaneus of the great apes (chimpanzees, gorillas, and orangutans) have not been sufficiently studied. This study aims to clarify variations in calcaneal morphology among great apes based on three-dimensional geometric morphometrics. A total of 556 landmarks and semilandmarks were placed on the calcaneal surface to calculate the principal components of shape variations among specimens. Clear interspecific differences in calcaneal morphology were extracted, corresponding to the degree of arboreality of the three species. The most arboreal orangutans possessed comparatively more slender calcaneal tuberosity and deeper pivot region of the cuboid articular surface than chimpanzees and gorillas. However, the most terrestrial gorillas exhibited longer lever arm of the triceps surae muscle, larger peroneal trochlea, more concave plantar surface, more inverted calcaneal tuberosity, more everted cuboid articular surface, and more prominent plantar process than the orangutans and chimpanzees. These interspecific differences possibly reflect the functional adaptation of the calcaneus to locomotor behavior in great apes. Such information might be useful for inferring foot functions and reconstructing the locomotion of fossil hominoids and hominids.

Morphological integration of the canine region within the hominine alveolar arch.

The early hominin record is characterized by numerous shifts in dental proportions (e.g., canine reduction and megadontia) linked to changes in diet and social behavior. Recent studies suggest that hominins exhibit a reduction in the magnitude of covariation between the anterior and posterior dental components compared with other extant great apes. They point toward, but do not directly test, the relative independence of canine morphology within the hominin alveolar arch. This study focuses specifically on the how the canine region covaries with other regions of the dental arch because the canine region has drastically reduced in size and changed in shape across human evolution. We examine extant primate species most commonly used as a comparative framework for fossil hominin morphology: Gorilla gorilla (n = 27), Pan troglodytes (n = 27), and Homo sapiens (n = 30). We used geometric morphometric methods to test for size and shape covariation between the canine region with other dental regions. We also examined the influence of sexual dimorphism and allometry on intraspecific and interspecific patterns of covariation. The analysis of size and shape covariation between the mandibular canine and other individual tooth regions elucidated complex, species-specific, and sex-specific morphological relationships in the mandibular alveolar arch. There was little evidence to support different patterns of morphological integration between humans on the one hand and nonhuman apes on the other. Canine region morphology was relatively independent from other dental regions across species based on shape and did not significantly covary more with either the incisor or postcanine region in any species. The size correlations between the canine and other dental regions were moderate to high. The species-specific results of this study question the ability to make a priori assumptions about morphological integration in the extant hominin mandibular alveolar arch and its application to the fossil record.