Conserved patterns and locomotor-related evolutionary constraints in the hominoid vertebral column.

The evolution of the hominoid lineage is characterized by pervasive homoplasy, notably in regions such as the vertebral column, which plays a central role in body support and locomotion. Few isolated and fewer associated vertebrae are known for most fossil hominoid taxa, but identified specimens indicate potentially high levels of convergence in terms of both form and number. Homoplasy thus complicates attempts to identify the anatomy of the last common ancestor of hominins and other taxa and stymies reconstructions of evolutionary scenarios. One way to clarify the role of homoplasy is by investigating constraints via phenotypic integration, which assesses covariation among traits, shapes evolutionary pathways, and itself evolves in response to selection. We assessed phenotypic integration and evolvability across the subaxial (cervical, thoracic, lumbar, sacral) vertebral column of macaques (n = 96), gibbons (n = 77), chimpanzees (n = 92), and modern humans (n = 151). We found a mid-cervical cluster that may have shifted cranially in hominoids, a persistent thoracic cluster that is most marked in chimpanzees, and an expanded lumbosacral cluster in hominoids that is most expanded in gibbons. Our results highlight the highly conserved nature of the vertebral column. Taxa appear to exploit existing patterns of integration and ontogenetic processes to shift, expand, or reduce cluster boundaries. Gibbons appear to be the most highly derived taxon in our sample, possibly in response to their highly specialized locomotion.

A shared pattern of midfacial bone modelling in hominids suggests deep evolutionary roots for human facial morphogenesis.

Midfacial morphology varies between hominoids, in particular between great apes and humans for which the face is small and retracted. The underlying developmental processes for these morphological differences are still largely unknown. Here, we investigate the cellular mechanism of maxillary development (bone modelling, BM), and how potential changes in this process may have shaped facial evolution. We analysed cross-sectional developmental series of gibbons, orangutans, gorillas, chimpanzees and present-day humans (n = 183). Individuals were organized into five age groups according to their dental development. To visualize each species's BM pattern and corresponding morphology during ontogeny, maps based on microscopic data were mapped onto species-specific age group average shapes obtained using geometric morphometrics. The amount of bone resorption was quantified and compared between species. Great apes share a highly similar BM pattern, whereas gibbons have a distinctive resorption pattern. This suggests a change in cellular activity on the hominid branch. Humans possess most of the great ape pattern, but bone resorption is high in the canine area from birth on, suggesting a key role of canine reduction in facial evolution. We also observed that humans have high levels of bone resorption during childhood, a feature not shared with other apes.

Shape variation in modern human upper premolars.

Morphological variation in modern human dentition is still an open field of study. The understanding of dental shape and metrics is relevant for the advancement of human biology and evolution and is thus of interest in the fields of dental anthropology, as well as human anatomy and medicine. Of concern is also the variation of the inner aspects of the crown which can be investigated using the tools and methods of virtual anthropology. In this study, we explored inter- and intra-population morphometric variation of modern humans' upper third and fourth premolars (P3s and P4s, respectively) considering both the inner and outer aspects of the crown, and discrete traits. We worked by means of geometric morphometrics on 3D image data from a geographically balanced sample of human populations from five continents, to analyse the shape of the dentinal crown, and the crown outline in 78 P3s and 76 P4s from 85 individuals. For the study of dental traits, we referred to the Arizona State University Dental Anthropology System integrated with more recent classification systems. The 3D shape variation of upper premolar crowns varied between short and mesio-distally broad, and tall and mesio-distally narrow. The observed shape variation was independent from the geographical origin of the populations, and resulted in extensive overlap. We noted a high pairwise correlation (r1 = 0.83) between upper P3s and P4s. We did not find any significant geographic differences in the analysed non-metric traits. Our outcomes thus suggest that geographical provenance does not play a determinant role in the shaping of the dental crown, whose genesis is under strict genetic control.

Naming Homo erectus: A review.

Following the discovery of hominin fossils at Trinil (Java, Indonesia) in 1891 and 1892, Eugène Dubois named a new species, now known as Homo erectus. Although the main historical events are well-known, there appears to be no consensus regarding two important aspects of the naming of the species, including what constitutes the original publication of the name, and what is the name-bearing type specimen. These issues are addressed in this paper with reference to original sources and the International Code of Zoological Nomenclature. Our review confirms earlier studies that cite the published quarterly fieldwork report covering the 3rd quarter of 1892 as the original publication naming the species erectus. However, until recently, the correct publication year of 1893 has consistently been cited as 1892, and it has rarely been recognized that the author of the publication was anonymous, even though the author of the species is specifically named. Importantly, Dubois assigns all three hominin fossils found at Trinil up to that moment to the new species, explicitly stating that they belong to a single individual. The three fossils, a molar, a calotte, and a femur, therefore jointly constitute the original holotype. However, the femur most likely derives from younger strata than the other hominins and shows fully modern human-like morphology, unlike subsequently discovered H. erectus femora. Moreover, there is no consensus over the affinities of the molar, and if it is H. erectus rather than an extinct ape, there is no evidence that it belongs to the same individual as the calotte. Excluding these two fossils from the holotype, the calotte is the appropriate fossil to retain the role as name-bearing specimen.

Variation of thoracolumbar vertebral morphology in anthropoid primates.

OBJECTIVES: Morphological variation among extant primates in the lumbar vertebral column is well studied. However, knowledge concerning the thoracic spine, an important region responsible for supporting and facilitating movement in the upper trunk, remains relatively scarce. Consequently, our comprehension of the functional differentiation exhibited throughout the thoracolumbar vertebral column among various primate species remains constrained. In this study, we examined patterns of morphological variation in the thoracolumbar vertebral column of extant hominoids, cercopithecoids, and Ateles. MATERIALS AND METHODS: We collected external shape data on 606 thoracic and lumbar vertebrae from Homo sapiens, Pan troglodytes, Hylobates lar, Macaca fuscata, Chlorocebus aethiops, Colobus guereza, Ateles geoffroyi, and A. belzebuth. Forty-four landmarks were obtained on the three-dimensional surface. Geometric morphometrics was used to quantify the centroid size and variation of the shapes of thoracic and lumbar vertebrae. RESULTS: Cercopithecoids exhibited greater variation in the size and shape of their thoracic and lumbar vertebrae compared to hominoids and Ateles. Although many vertebral features contributed to the observed variation throughout the thoracolumbar vertebral column within the taxon, the transverse and spinous processes exhibited relatively major contributions. DISCUSSION: Our results suggest that quadrupedal locomotion requires the functional differentiation between thoracic and lumbar vertebrae, and for hominoids, functional adaptation to orthograde posture necessitates a relatively more uniform shape of thoracic and lumbar vertebrae.

Evidence for the smallest fossil Pongo in southern China.

The rarity of Pongo fossils with precise absolute dating from the Middle Pleistocene hampers our understanding of the taxonomy and spatiotemporal distribution of Quaternary orangutans in southern China. Here, we report a newly discovered sample of 113 isolated teeth of fossil Pongo from Zhongshan Cave in the Bubing Basin, Guangxi, southern China. We describe the Pongo specimens from Zhongshan Cave and compare them metrically to other samples of fossil Pongo species (i.e., Pongo weidenreichi, Pongo devosi, Pongo duboisi, Pongo palaeosumatrensis, Pongo javensis, and Pongo sp.) and to extant orangutans (i.e., Pongo pygmaeus and Pongo abelii). The Zhongshan Pongo assemblage is dated using U-series and coupled electron spin resonance/U-series methods. Our results reasonably constrain the Zhongshan Pongo assemblage to 184 ± 16 ka, which is consistent with the biostratigraphic evidence. The Zhongshan Pongo teeth are only 6.5% larger on average than those of extant Pongo. The Zhongshan teeth are smaller overall than those of Pongo from all other cave sites in southern China, and they currently represent the smallest fossil orangutans in southern China. Based on their dental size, and the presence of a well-developed lingual pillar and lingual cingulum on the upper and lower incisors, an intermediate frequency of lingual cingulum remnants on the upper molars, and a higher frequency of moderate to heavy wrinkling on the upper and lower molars, we provisionally assign the Zhongshan fossils to P. devosi. Our results confirm earlier claims that P. weidenreichi is replaced by a smaller species in southern China, P. devosi, by the late Middle Pleistocene. The occurrence of P. devosi in Zhongshan Cave further extends its spatial and temporal distribution. The Pongo specimens from Zhongshan provide important new evidence to demonstrate that the dental morphological features of Pongo in southern China changed substantially during the late Middle Pleistocene.

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.

On the genetic basis of tail-loss evolution in humans and apes.

The loss of the tail is among the most notable anatomical changes to have occurred along the evolutionary lineage leading to humans and to the 'anthropomorphous apes'1-3, with a proposed role in contributing to human bipedalism4-6. Yet, the genetic mechanism that facilitated tail-loss evolution in hominoids remains unknown. Here we present evidence that an individual insertion of an Alu element in the genome of the hominoid ancestor may have contributed to tail-loss evolution. We demonstrate that this Alu element-inserted into an intron of the TBXT gene7-9-pairs with a neighbouring ancestral Alu element encoded in the reverse genomic orientation and leads to a hominoid-specific alternative splicing event. To study the effect of this splicing event, we generated multiple mouse models that express both full-length and exon-skipped isoforms of Tbxt, mimicking the expression pattern of its hominoid orthologue TBXT. Mice expressing both Tbxt isoforms exhibit a complete absence of the tail or a shortened tail depending on the relative abundance of Tbxt isoforms expressed at the embryonic tail bud. These results support the notion that the exon-skipped transcript is sufficient to induce a tail-loss phenotype. Moreover, mice expressing the exon-skipped Tbxt isoform develop neural tube defects, a condition that affects approximately 1 in 1,000 neonates in humans10. Thus, tail-loss evolution may have been associated with an adaptive cost of the potential for neural tube defects, which continue to affect human health today.

New craniodental fossils of Paranthropus robustus from Kromdraai, South Africa (2014-2017 excavations).

Since the initial discovery of Paranthropus robustus at the site of Kromdraai in 1938, the hypodigm of this species has been expanded by subsequent work at the localities of Swartkrans and Drimolen, with a few fossils also known from Cooper's D, Gondolin and Sterkfontein Member 5. Beginning in 2014, systematic excavations at Kromdraai uncovered a large and previously unknown fossiliferous area, shedding light on Units O and P in the earliest part of the site's stratigraphic sequence. The aim of this paper is to provide detailed descriptions and illustrations of 30 P. robustus craniodental specimens recovered between 2014 and 2017 within the Unit P deposits at Kromdraai. This new sample predates all prior conspecific specimens found at this site (including the holotype of P. robustus from Kromdraai, TM 1517). Its basic dental morphology dimensions and cranial features are compared in a preliminary analysis with other P. robustus samples. The P. robustus sample from Kromdraai Unit P documents previously unknown portions of the P. robustus juvenile cranium. The new dental and cranial remains aid in the exploration of potential morphological distinctions between site-specific P. robustus samples and are compared favorably in size and morphology with the small P. robustus specimens from Drimolen (e.g., DNH 7). These findings do not support the hypothesis that the specimens from Drimolen belong to a different taxonomic group. Instead, they reinforce the presence of a significant degree of sexual dimorphism within P. robustus. The Kromdraai Unit P specimens also contribute to the biodemographic profile of P. robustus. The notable prevalence of infants (i.e., juvenile individuals before the emergence of their first permanent molars) mirrors the natural mortality profiles observed in wild chimpanzees. This suggests a closer resemblance in the processes of accumulation in Kromdraai Unit P and Drimolen than at Swartkrans.

A lineage perspective on hominin taxonomy and evolution.

An uncritical reliance on the phylogenetic species concept has led paleoanthropologists to become increasingly typological in their delimitation of new species in the hominin fossil record. As a practical matter, this approach identifies species as diagnosably distinct groups of fossils that share a unique suite of morphological characters but, ontologically, a species is a metapopulation lineage segment that extends from initial divergence to eventual extinction or subsequent speciation. Working from first principles of species concept theory, it is clear that a reliance on morphological diagnosabilty will systematically overestimate species diversity in the fossil record; because morphology can evolve within a lineage segment, it follows that early and late populations of the same species can be diagnosably distinct from each other. We suggest that a combination of morphology and chronology provides a more robust test of the single-species null hypothesis than morphology alone.

Technical note: A volumetric method for measuring the longitudinal arch of human tracks and feet.

Fossil footprints (i.e., tracks) were believed to document arch anatomical evolution, although our recent work has shown that track arches record foot kinematics instead. Analyses of track arches can thereby inform the evolution of human locomotion, although quantifying this 3-D aspect of track morphology is difficult. Here, we present a volumetric method for measuring the arches of 3-D models of human tracks and feet, using both Autodesk Maya and Blender software. The method involves generation of a 3-D object that represents the space beneath the longitudinal arch, and measurement of that arch object's geometry and spatial orientation. We provide relevant tools and guidance for users to apply this technique to their own data. We present three case studies to demonstrate potential applications. These include, (1) measuring the arches of static and dynamic human feet, (2) comparing the arches of human tracks with the arches of the feet that made them, and (3) direct comparisons of human track and foot arch morphology throughout simulated track formation. The volumetric measurement tool proved robust for measuring 3-D models of human tracks and feet, in static and dynamic contexts. This tool enables researchers to quantitatively compare arches of fossil hominin tracks, in order to derive biomechanical interpretations from them, and/or offers a different approach for quantifying foot morphology in living humans.

Dental morphology in Homo habilis and its implications for the evolution of early Homo.

The phylogenetic position of Homo habilis is central to debates over the origin and early evolution of the genus Homo. A large portion of the species hypodigm consists of dental remains, but they have only been studied at the often worn enamel surface. We investigate the morphology of the H. habilis enamel-dentine junction (EDJ), which is preserved in cases of moderate tooth wear and known to carry a strong taxonomic signal. Geometric morphometrics is used to characterise dentine crown shape and size across the entire mandibular and maxillary tooth rows, compared with a broad comparative sample (n = 712). We find that EDJ morphology in H. habilis is for the most part remarkably primitive, supporting the hypothesis that the H. habilis hypodigm has more in common with Australopithecus than later Homo. Additionally, the chronologically younger specimen OH 16 displays a suite of derived features; its inclusion in H. habilis leads to excessive levels of variation.

Ulnar shape of extant primates: Functional signals and covariation with triquetrum shape.

OBJECTIVES: In this study, we investigated the shape differences of the distal ulna in a phylogenetic context among a broad range of primate taxa. Furthermore, we evaluated covariation between ulnar and triquetrum shape and a possible association between ulnar shape and locomotor behavior. MATERIALS AND METHODS: We applied 3D geometric morphometrics on a large dataset comprising the distal ulna of 124 anthropoid primate specimens belonging to 12 different genera. For each species, a mean shape was calculated using 11 Procrustes-aligned surface landmarks on the distal ulna. These mean shapes are used in a bgPCA, pPCA, and PACA and 3D morphs were used to visualize more subtle differences between taxa. A p2B-PLS analysis was performed to test the covariance between distal ulnar and triquetrum shape. RESULTS: The results show that more closely related species exhibit a similar distal ulnar shape. Overall, extant hominid ulnae show a shape shift compared to those of extant monkeys and hylobatids. This includes a shortening of the ulnar styloid process and dorspalmarly widening of the ulnar head, shape characteristics that are independent of phylogeny. Within the hominids, Pongo pygmaeus seem to possess the most plesiomorphic distal ulnar shape, while Gorilla and Homo sapiens display the most derived distal ulna. Cercopithecoids, hylobatids, and P. pygmaeus are characterized by a relatively deep ECU groove, which is a shape trait dependent of phylogeny. Although there was no significant covariation between distal ulnar shape and triquetrum shape, the shape differences of the distal ulna between the different primate taxa reveal a possible link with locomotor behavior. CONCLUSIONS: The comparative analyses of this study reveal different shape trends in a phylogenetic context. Highly arboreal primates, such as hylobatids and Ateles fusciceps, show a distal ulnar morphology that appears to be adapted to tensile and torsional forces. In primates that use their wrist under more compressive conditions, such as quadrupedal cercopithecoids and great apes, the distal ulnar morphology seems to reflect increased compressive forces. In modern humans, the distal ulnar shape can be associated to enhanced manipulative skills and power grips. There was no significant covariation between distal ulnar shape and triquetrum shape, probably due to the variation in the amount of contact between the triquetrum and ulna. In combination with future research on wrist mobility in diverse primate taxa, the results of this study will allow us to establish form-function relationships of the primate wrist and contribute towards an evidence-based interpretation of fossil remains.

Captive bonobos (Pan paniscus) apply precision grips when using flaked stone tools.

OBJECTIVES: Current evidence suggests that flaked stone tool technologies did not emerge until ~3.3-2.6 million-years-ago (Ma). It is often hypothesized that early hominin (principally Ardipithecus and early Australopithecus) manual anatomy may have prevented an earlier emergence, as the forceful precision grips essential to flake tool-use may have been ineffectively performed by these species. Marzke, Marchant, McGrew, and Reece (2015) observed potentially forceful pad-to-side precision grips being recruited by wild chimpanzees (Pan troglodytes) during feeding behaviors, indicating that Pan-like manual anatomy, and therefore potentially early hominin anatomy, may be capable of effectively securing flake stone tools during their use. MATERIALS AND METHODS: Here, we report on the grips recruited by four captive, human-trained, bonobos (Pan paniscus) during the use of stone and organic tools, including flake stone tools during cutting behaviors. RESULTS: It is revealed that pad-to-side precision grips are frequently recruited by these bonobos when securing stone flakes during cutting actions. In some instances, high forces could have been resisted and applied by the thumb and fingers. DISCUSSION: While our analyzes are preliminary and limited to captive individuals, and Pan is not suggested to secure flakes with the same efficacy as Homo or Australopithecus, it points to early hominins potentially being able to perform the precision grips required to use flake stone tools. In turn, the ability to gain tangible benefits from the effective use of flake tools (i.e., gain energetic returns from processing food resources) may have been - at least anatomically - possible in early Australopithecus and other pre-Early Stone Age hominin species. In turn, hominin manual anatomy may not be a leading restriction on the emergence of the earliest stone tool technologies.

Comparative ontogeny of functional aspects of human cervical vertebrae.

OBJECTIVES: Differences between adult humans and great apes in cervical vertebral morphology are well documented, but the ontogeny of this variation is still largely unexplored. This study examines patterns of growth in functionally relevant features of C1, C2, C4, and C6 in extant humans and apes to understand the development of their disparate morphologies. MATERIALS AND METHODS: Linear and angular measurements were taken from 530 cervical vertebrae representing 146 individual humans, chimpanzees, gorillas, and orangutans. Specimens were divided into three age-categories based on dental eruption: juvenile, adolescent, and adult. Inter- and intraspecific comparisons were evaluated using resampling methods. RESULTS: Of the eighteen variables examined here, seven distinguish humans from apes at the adult stage. Human-ape differences in features related to atlantoaxial joint function tend to be established by the juvenile stage, whereas differences in features related to the nuchal musculature and movement of the subaxial elements do not fully emerge until adolescence or later. The orientation of the odontoid process-often cited as a feature that distinguishes humans from apes-is similar in adult humans and adult chimpanzees, but the developmental patterns are distinct, with human adultlike morphology being achieved much earlier. DISCUSSION: The biomechanical consequences of the variation observed here is poorly understood. Whether the differences in growth patterns represent functional links to cranial development or postural changes, or both, requires additional investigation. Determining when humanlike ontogenetic patterns evolved in hominins may provide insight into the functional basis driving the morphological divergence between extant humans and apes.

A review of the distal femur in Australopithecus.

In 1938, the first distal femur of a fossil Australopithecus was discovered at Sterkfontein, South Africa. A decade later, another distal femur was discovered at the same locality. These two fossil femora were the subject of a foundational paper authored by Kingsbury Heiple and Owen Lovejoy in 1971. In this paper, the authors discussed functionally relevant anatomies of these two fossil femora and noted their strong affinity to the modern human condition. Here, we update this work by including eight more fossil Australopithecus distal femora, an expanded comparative dataset, as well as additional linear measurements. Just as Heiple and Lovejoy reported a half-century ago, we find strong overlap between modern humans and cercopithecoids, except for inferiorly flattened condyles and a high bicondylar angle, both of which characterize modern humans and Australopithecus and are directly related to striding bipedalism. All other measured aspects of the femora are by-products of these key morphological traits. Additional fossil material from the early Pliocene will help to inform the evolution of the hominin distal femur and its condition in the Pan-Homo common ancestor that preceded bipedal locomotion.

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.

Covariation between the shape and mineralized tissues of the rib cross section in Homo sapiens, Pan troglodytes and Sts 14.

OBJECTIVES: Studying rib torsion is crucial for understanding the evolution of the hominid ribcage. Interestingly, there are variables of the rib cross section that could be associated with rib torsion and, consequently, with the morphology of the thorax. The aim of this research is to conduct a comparative study of the shape and mineralized tissues of the rib cross section in different hominids to test for significant differences and, if possible, associate them to different thoracic morphotypes. MATERIALS AND METHODS: The sample consists of the rib cross sections at the midshaft taken from 10 Homo sapiens and 10 Pan troglodytes adult individuals, as well as from A. africanus Sts 14. The shape of these rib cross sections was quantified using geometric morphometrics, while the mineralized tissues were evaluated using the compartmentalization index. Subsequently, covariation between both parameters was tested by a Spearman's ρ test, a permutation test and a linear regression. RESULTS: Generally, P. troglodytes individuals exhibit rib cross sections that are rounder and more mineralized compared to those of H. sapiens. However, the covariation between both parameters was only observed in typical ribs (levels 3-10). Although covariation was not found in the rib cross sections of Sts 14, their parameters are closer to P. troglodytes. DISCUSSION: On the one hand, the differences observed in the rib cross sections between H. sapiens and P. troglodytes might be related to different degrees of rib torsion and, consequently, to different thoracic 3D configurations. These findings can be functionally explained by considering their distinct modes of breathing and locomotion. On the other hand, although the rib cross sections belonging to Sts 14 are more similar to those of P. troglodytes, previous publications determined that their overall morphology is closer to modern humans. This discrepancy could reflect a diversity of post-cranial adaptations in Australopithecus.