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Hominin evolution featured shifts from a trunk shape suitable for climbing and housing a large gut to a trunk adapted to bipedalism and higher quality diets. Our knowledge regarding the tempo, mode, and context in which these derived traits evolved has been limited, based largely on a small-bodied Australopithecus partial skeleton (A.L. 288-1; "Lucy") and a juvenile Homo erectus skeleton (KNM-WT 15000; "Turkana Boy"). Two recent discoveries, of a large-bodied Australopithecus afarensis (KSD-VP-1/1) and two Australopithecus sediba partial skeletons (MH1 and MH2), have added to our understanding of thorax evolution; however, little is known about thorax morphology in early Homo. Here we describe hominin vertebrae, ribs, and sternal remains from the Dinaledi chamber of the Rising Star cave system attributed to Homo naledi. Although the remains are highly fragmented, the best-preserved specimensdtwo lower thoracic vertebrae and a lower ribdwere found in association and belong to a small-bodied individual. A second lower rib may belong to this individual as well. All four of these individual elements are amongst the smallest known in the hominin fossil record. H. naledi is characterized by robust, relatively uncurved lower ribs and a relatively large spinal canal. We expect that the recovery of additional material from Rising Star Cave will clarify the nature of these traits and shed light on H. naledi functional morphology and phylogeny.

Spinal Evolution, 2019

In this chapter, we summarize vertebral remains from early Pleistocene Homo, including H. erectus, as well as H. naledi and H. floresiensis fossils from the Middle and Late Pleistocene, respectively. Two partial immature H. erectus skeletons where vertebrae are well represented are KNM-WT 15000 (“Turkana boy”) and the D2700 individual from Dmanisi. Vertebrae from H. naledi are also considered here, including those from the LES1 partial skeleton (“Neo”), despite their younger date to the Middle Pleistocene. We review the fossil record of presacral vertebrae in early Homo, and summarize work on the functional morphology, metameric patterning, and postcranial neuroanatomy of early Homo, comparing and contrasting the presacral spine with their putative australopith forbears and extant apes and humans. Based on the current evidence, the vertebral column of H. erectus possessed a modal number of twelve thoracic and five lumbar segments, as is the case in australopiths, as well as modern humans. The spine of H. erectus reveals key changes relative to earlier hominins, with an expanded thoracolumbar spinal canal offering increased neurovascular capacities, and a ventral pillar (formed by the vertebral bodies) better equipped to mitigate compressive loads and provide energy return. These biological developments are germane to understanding the advent of derived human behaviors, including efficient long-range locomotion and the first hominin expansion out of Africa.

The vertebral column plays a central role in the evolution and performance of positional behaviors, including upright posture and bipedal locomotion in the human lineage. The lumbar column, in particular, is associated with locomotor function. As such, its numerical composition has been a major source of contention in the paleoanthropological literature. Ever since description and interpretation of the nearly complete thoracolumbar vertebral column of Sts 14 (Australopithecus africanus), researchers have, with few exceptions, consistently stated that early hominins possessed six lumbar vertebrae (

The species Homo naledi was recently named from specimens recovered from the Dinaledi Chamber of the Rising Star cave system in South Africa. This large skeletal sample lacks associated faunal material and currently does not have a known chronological context. In this paper, we present comprehensive descriptions and metric comparisons of the recovered cranial and mandibular material. We describe 41 elements attributed to Dinaledi Hominin (DH1eDH5) individuals and paratype U.W. 101-377, and 32 additional cranial fragments. The H. naledi material was compared to Plio-Pleistocene fossil hominins using qualitative and quantitative analyses including over 100 linear measurements and ratios. We find that the Dinaledi cranial sample represents an anatomically homogeneous population that expands the range of morphological variation attributable to the genus Homo. Despite a relatively small cranial capacity that is within the range of australopiths and a few specimens of early Homo, H. naledi shares cranial characters with species across the genus Homo, including Homo habilis, Homo rudolfensis, Homo erectus, and Middle Pleistocene Homo. These include aspects of cranial form, facial morphology, and mandibular anatomy. However, the skull of H. naledi is readily distinguishable from existing species of Homo in both qualitative and quantitative assessments. Since H. naledi is currently undated, we discuss the evolutionary implications of its cranial morphology in a range of chronological frameworks. Finally, we designate a sixth Dinaledi Hominin (DH6) individual based on a juvenile mandible.

Nature Ecology & Evolution

2019

The abundant femoral assemblage of Homo naledi found in the Dinaledi Chamber provides a unique opportunity to test hypotheses regarding the taxonomy, locomotion, and loading patterns of this species. Here we describe neck and shaft cross-sectional structure of all the femoral fossils recovered in the Dinaledi Chamber and compare them to a broad sample of fossil hominins, recent humans, and extant apes. Cross-sectional geometric (CSG) properties from the femoral neck (base of neck and midneck) and diaphysis (subtrochanteric region and midshaft) were obtained through CT scans for H. naledi and through CT scans or from the literature for the comparative sample. The comparison of CSG properties of H. naledi and the comparative samples shows that H. naledi femoral neck is quite derived with low superoinferior cortical thickness ratio and high relative cortical area. The neck appears superoinferiorly elongated because of two bony pilasters on its superior surface. Homo naledi femoral shaft shows a relatively thick cortex compared to the other hominins. The subtrochanteric region of the diaphysis is mediolaterally elongated resembling early hominins while the midshaft is anteroposteriorly elongated, indicating high mobility levels. In term of diaphyseal robusticity, the H. naledi femur is more gracile that other hominins and most apes. Homo naledi shows a unique combination of characteristics in its femur that undoubtedly indicate a species committed to terrestrial bipedalism but with a unique loading pattern of the femur possibly consequence of the unique postcranial anatomy of the species.

2019

The early hominin (Ardipithecus and Australopithecus) fossil record contains over 100 preserved vertebral elements (n = 107; approximately half of which are well-preserved), ~65% of which have not been described since the turn of the millennium. Many are fragments, some for which detailed descriptions are pending (e.g., those of Australopithecus anamensis). Australopithecus afarensis and Australopithecus sediba are known from cervical, thoracic, and lumbar vertebrae, whereas Australopithecus africanus is known from thoracic and lumbar vertebrae but not cervical vertebrae. A partial skeleton from Member 4 of Sterkfontein, StW 573, preserves vertebrae from all presacral regions, but its species designation is debated and not yet formalized in the literature. Other early hominin species, such as Sahelanthropus tchadensis, Orrorin tugenensis, Ardipithecus kadabba, Australopithecus deyiremeda, Australopithecus bahrelghazali, and Australopithecus garhi, do not preserve vertebrae. Vertebra...

Reconstructing the morphology of the Neanderthal rib cage not only provides information about the general evolution of human body shape but also aids understanding of functional anatomy and energetics. Despite this paleobiological importance there is still debate about the nature and extent of variations in the size and shape of the Neandertal thorax. The El Sidr
on Neandertals can be used to contribute to this debate, providing new costal remains ranging from fully preserved and undistorted ribs to highly fragmented elements. Six first ribs are particularly well preserved and offer the opportunity to analyze thorax morphology in Neandertals. The aims of this paper are to present this new material, to compare the ontogenetic trajectories of the first ribs between Neandertals and modern humans, and, using geometric morphometrics, to test the hypothesis of morphological integration between the first rib and overall thorax morphology. The first ribs of the El Sidr
on adult Neandertals are smaller in centroid size and tend to be less curved when compared with those of modern humans, but are similar to Kebara 2. Our results further show that the straightening of the first ribs is significantly correlated with a straightening of the ribs of the upper thorax (R ¼ 0.66; p < 0.0001) in modern humans, suggesting modularity in the upper and lower thorax units as reported in other hominins. It also supports the hypothesis that the upper thorax of Neandertals differs in shape from modern humans with more anteriorly projecting upper ribs during inspiration. These differences could have biomechanical consequences and account for stronger muscle attachments in Neandertals. Different upper thorax shape would also imply a different spatial arrangement of the shoulder girdle and articulation with the humerus (torsion) and its connection to the upper thorax. Future research should address these inferences in the context of Neandertal overall body morphology.

Journal of Human Evolution, 2019

Substantial differences among the pelves of anthropoids have been central to interpretations of the selection pressures that shaped extant hominoids, yet the evolution of the hominoid pelvis has been poorly understood due to the scarcity of fossil material. A recently discovered partial hipbone attributed to the 10 million-year-old fossil ape Rudapithecus hungaricus from Rudab anya, Hungary, differs from the hipbones of cercopithecids and earlier apes in functionally significant ways. Comparisons were made to extant and other fossil anthropoids using combination of non-landmark-based and linear metrics. Measurements were taken on 3D polygonal models of hipbones collected using laser scans. These metrics capture functionally relevant morphology given the incomplete preservation of the Rudapithecus specimen. This fossil displays features that reflect changes in spinal musculature and torso structure found only in extant great and lesser apes among hominoids. Rudapithecus has an expanded cranial acetabular lunate surface related to orthograde positional behaviors, a shallow acetabulum and relatively short ischium like orangutans and hylobatids. It displays evidence of moderately coronally-oriented iliac blades as in all extant apes and Ateles, and flaring iliac blade shape of siamangs and great apes, associated with some level of spinal stiffness. However, this fossil lacks the long lower ilium that characterizes chimpanzees, gorillas and orangutans, associated with their reduction of the number of lumbar vertebrae. The R. hungaricus pelvis demonstrates that the extreme elongation of the lower ilium seen in extant great apes does not necessarily accompany adaptation to orthograde posture and forelimb-dominated arboreal locomotion in hominoid evolution. Lower iliac elongation appears to have occurred independently in each lineage of extant great apes, supporting the hypothesis that the last common ancestor of Pan and Homo may have been unlike extant great apes in pelvic length and lower back morphology.

Journal of human …, 2009

First ribs – the first or most superior ribs in the thorax – are rare in the hominin fossil record, and when found, have the potential to provide information regarding the upper thorax shape of extinct hominins. Here, we describe a partial first rib from Member 4 of the Sterkfontein Caves, South Africa. The rib shaft is broken away, so only the head and neck are preserved. The rib is small, falling closest to small-bodied Australopithecus first ribs (AL 288-1 and MH1). Given that it was recovered near the StW 318 femur excavation, which also represents a small individual, we suggest that the two may be associated. Three-dimensional geometric morphometric analyses were used to quantify the rib fragment morphology and compare it to extant hominoid and other fossil hominin ribs. While only the proximal end is preserved, our analyses show that South African Australopithecus share derived features of the proximal first rib more closely resembling A. afarensis and later hominins than great apes.

Journal of Human Evolution, 2019

Australopitheus anamensis fossils demonstrate that craniodentally and postcranially the taxon was more primitive than its evolutionary successor Australopithecus afarensis. Postcranial evidence suggests habitual bipedality combined with primitive upper limbs and an inferred significant arboreal adaptation. Here we report on A. anamensis fossils from the Assa Issie locality in Ethiopia's Middle Awash area dated to ~4.2 Ma, constituting the oldest known Australopithecus axial remains. Because the spine is the interface between major body segments, these fossils can be informative on the adaptation, behavior and our evolutionary understanding of A. anamensis. The atlas, or first cervical vertebra (C1), is similar in size to Homo sapiens, with synapomorphies in the articular facets and transverse processes. Absence of a retroglenoid tubercle suggests that, like humans, A. anamensis lacked the atlantoclavicularis muscle, resulting in reduced capacity for climbing relative to the great apes. The retroflexed C2 odontoid process and long C6 spinous process are reciprocates of facial prognathism, a long clivus and retroflexed foramen magnum, rather than indications of locomotor or postural behaviors. The T1 is derived in shape and size as in Homo with an enlarged vertebral body epiphyseal surfaces for mitigating the high-magnitude compressive loads of full-time bipedality. The full costal facet is unlike the extant great ape demifacet pattern and represents the oldest evidence for the derived univertebral pattern in hominins. These fossils augment other lines of evidence in A. anamensis indicating habitual bipedality despite some plesio-morphic vertebral traits related to craniofacial morphology independent of locomotor or postural behaviors (i.e., a long clivus and a retroflexed foramen magnum). Yet in contrast to craniodental lines of evidence, some aspects of vertebral morphology in A. anamensis appear more derived than its descendant A. afarensis.

South African Journal of Science, 2016

First ribs – the first or most superior ribs in the thorax – are rare in the hominin fossil record, and when found, have the potential to provide information regarding the upper thorax shape of extinct hominins. Here, we describe a partial first rib from Member 4 of the Sterkfontein Caves, South Africa. The rib shaft is broken away, so only the head and neck are preserved. The rib is small, falling closest to small-bodied Australopithecus first ribs (AL 288-1 and MH1). Given that it was recovered near the StW 318 femur excavation, which also represents a small individual, we suggest that the two may be associated. Three-dimensional geometric morphometric analyses were used to quantify the rib fragment morphology and compare it to extant hominoid and other fossil hominin ribs. While only the proximal end is preserved, our analyses show that South African Australopithecus share derived features of the proximal first rib more closely resembling A. afarensis and later hominins than grea...

The phylogeny of the early African hominins has long been confounded by contrasting interpretations of midfacial structure. In particular, the anterior pillar, an externally prominent bony column running vertically alongside the nasal aperture, has been identified as a homology of South African species Australopithecus africanus and Australopithecus robustus. If the anterior pillar is a true synapomorphy of these two species, the evidence for a southern African clade of Australopithecus would be strengthened, and support would be given to the phylogenetic hypothesis of an independent origin for eastern and southern African “robust” australopith clades. Analyses of CT data, however, show that the internal structure of the circumnasal region is strikingly different in the two South African australopith species. In A. africanus the anterior pillar is a hollow column of cortical bone, whereas in A. robustus it is a column of dense trabecular bone. Although Australopithecus boisei usually lacks an external pillar, it has internal morphology identical to that seen in A. robustus. This result supports the monophyly of the “robust” australopiths and suggests that the external similarities seen in the South African species are the result of parallel evolution.

Two partial vertebral columns of Australopithecus sediba grant insight into aspects of early hominin spinal mobility, lumbar curvature, vertebral formula, and transitional vertebra position. Au. sediba likely possessed five non-rib-bearing lumbar vertebrae and five sacral elements, the same configuration that occurs modally in modern humans. This finding contrasts with other interpretations of early hominin regional vertebral numbers. Importantly, the transitional vertebra is distinct from and above the last rib-bearing vertebra in Au. sediba, resulting in a functionally longer lower back. This configuration, along with a strongly wedged last lumbar vertebra and other indicators of lordotic posture, would have contributed to a highly flexible spine that is derived compared with earlier members of the genus Australopithecus and similar to that of the Nariokotome Homo erectus skeleton.

2019

Homo naledi skeletal material described from the Dinaledi Chamber, Rising Star System, in the Cradle of Humankind, South Africa, includes upper limb material with remarkably ape-like morphology occurring in the context of a distinctly modern human-like lower limb, foot, and hand. Here we describe upper limb specimens from a new fossil hominin site within the Rising Star System, the Lesedi Chamber (Site U.W. 102), including an intact clavicle and a well-preserved proximal humerus and proximal ulna. Craniodental remains, in association with the postcranial elements described here, have been attributed to H. naledi. The upper limb material from the Lesedi Chamber is gracile and resembles the upper limb material from the Dinaledi Chamber in overall morphology. The primitive humeral morphology of the Dinaledi material is replicated within the Lesedi material. The fossils from the Lesedi Chamber material also preserve additional humeral morphology not represented in the Dinaledi Chamber, providing new information on humeral form in this species. Three-dimensional geometric morphometric analysis of the Lesedi proximal humerus demonstrates affinities with the humeri of Pan and Australopithecus. The complete clavicle from Lesedi is similarly primitive, supporting previous interpretations of the H. naledi shoulder as being more superiorly positioned than modern humans, and closer to the hypothesized australopith condition. The ulna is gracile, with a mediolaterally narrow olecranon process and anterior-facing trochlear notch similar to the derived state suggested for other fossil hominins. These new findings support the interpretation of overhead reaching and climbing behaviors having continued relevance in the locomotor repertoire of Homo naledi and provide a clearer picture of the upper limb morphology of this species.

Spinal posture has vast biomechanical, locomotor and pathological implications in hominins. Assessing the curvatures of the spine of fossil hominins can provide important information towards the understanding of their paleobiology. Unfortunately, complete hominin spines are very rarely preserved in the fossil record. The Neanderthal partial skeleton, Kebara 2 from Israel, constitutes a remarkable exception, representing an almost complete spine and pelvis. The aim of this study is, therefore, to create a new 3D virtual reconstruction of the spine of Kebara 2. To build the model, we used the CT scans of the sacrum, lumbar and thoracic vertebrae of Kebara 2, captured its 3D morphology, and, using visualization software (Amira 5.2©), aligned the 3D reconstruction of the original bones into the spinal curvature. First we aligned the sacrum and then we added one vertebra at a time, until the complete spine (T1-S5) was intact. The amount of spinal curvature (lordosis and kyphosis), the sacral orientation, and the coronal plane deviation was determined based on the current literature or measured and calculated specifically for this study based on published methods. This reconstruction provides, for the first time, a complete 3D virtual reconstruction of the spine of an extinct hominin. The spinal posture and spinopelvic alignment of Kebara 2 show a unique configuration compared with that of modern humans, suggesting locomotor and weight-bearing differences between the two. The spinal posture of Kebara 2 also shows slight asymmetry in the coronal plane. Stature estimation of Kebara 2 based on spinal length confirms that the height of Kebara 2 was around 170 cm. This reconstruction can now serve as the basis for a more complete reconstruction of the Kebara 2 specimen, which will include other parts of this remarkable fossil, such as the pelvis, the rib cage and the cervical spine.

The thoracic skeletal morphology of homininae is poorly known and understood. As a result of the representative fossil record of ribs and vertebrae being rare, distorted, fragmentary or unrecognised even when recovered, very little is known about the variability of rib and vertebral morphology when compared to the other cranial and postcranial elements in this lineage. Yet the costal skeleton forms a substantial part of the postcranial skeleton and thus ribs and vertebrae are therefore potentially numerous in the fossil record; but in comparison with other skeletal elements, and for the reasons mentioned above, very little is known about vertebrate and especially hominin rib morphology. The assessment of the structure of the thoracic skeletal elements and its evolutionary and ecological significance, particularly in the Homininae, poses a challenge but is still important as the shape and form of the rib cage has numerous functional and behavioural implications. The present study analysed the ribs of selected primate and non-primate mammalian species by examining fifteen variables, seven indices and eight osteological non-metric features.

bioRxiv, 2021

Adaptations of the lower back to bipedalism are frequently discussed but infrequently demonstrated in early fossil hominins. Newly discovered lumbar vertebrae contribute to a near-complete lower back of Malapa Hominin 2 (MH2), offering additional insights into posture and locomotion in Australopithecus sediba. We show that MH2 demonstrates a lower back consistent with human-like lumbar lordosis and other adaptations to bipedalism, including an increase in the width of intervertebral articular facets from the upper to lower lumbar column (“pyramidal configuration”). This contrasts with recent work on lordosis in fossil hominins, where MH2 was argued to demonstrate no appreciable lordosis (“hypolordosis”) similar to Neandertals. Our three-dimensional geometric morphometric (3D GM) analyses show that MH2’s nearly complete middle lumbar vertebra is human-like in shape but bears large, cranially-directed transverse processes, implying powerful trunk musculature. We interpret this combina...