Knuckle-walking in Sahelanthropus? Locomotor inferences from the ulnae of fossil hominins and other hominoids.

Because the ulna supports and transmits forces during movement, its morphology can signal aspects of functional adaptation. To test whether, like extant apes, some hominins habitually recruit the forelimb in locomotion, we separate the ulna shaft and ulna proximal complex for independent shape analyses via elliptical Fourier methods to identify functional signals. We examine the relative influence of locomotion, taxonomy, and body mass on ulna contours in Homo sapiens (n = 22), five species of extant apes (n = 33), two Miocene apes (Hispanopithecus and Danuvius), and 17 fossil hominin specimens including Sahelanthropus, Ardipithecus, Australopithecus, Paranthropus, and early Homo. Ulna proximal complex contours correlate with body mass but not locomotor patterns, while ulna shafts significantly correlate with locomotion. African apes' ulna shafts are more robust and curved than Asian apes and are unlike other terrestrial mammals (including other primates), curving ventrally rather than dorsally. Because this distinctive curvature is absent in orangutans and hylobatids, it is likely a function of powerful flexors engaged in wrist and hand stabilization during knuckle-walking, and not an adaptation to climbing or suspensory behavior. The OH 36 (purported Paranthropus boisei) and TM 266 (assigned to Sahelanthropus tchadensis) fossils differ from other hominins by falling within the knuckle-walking morphospace, and thus appear to show forelimb morphology consistent with terrestrial locomotion. Discriminant function analysis classifies both OH 36 and TM 266 with Pan and Gorilla with high posterior probability. Along with its associated femur, the TM 266 ulna shaft contours and its deep, keeled trochlear notch comprise a suite of traits signaling African ape-like quadrupedalism. While implications for the phylogenetic position and hominin status of S. tchadensis remain equivocal, this study supports the growing body of evidence indicating that S. tchadensis was not an obligate biped, but instead represents a late Miocene hominid with knuckle-walking adaptations.

Earliest axial fossils from the genus Australopithecus.

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 plesiomorphic 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.

Neck function in early hominins and suspensory primates: Insights from the uncinate process.

OBJECTIVES: Uncinate processes are protuberances on the cranial surface of subaxial cervical vertebrae that assist in stabilizing and guiding spinal motion. Shallow uncinate processes reduce cervical stability but confer an increased range of motion in clinical studies. Here we assess uncinate processes among extant primates and model cervical kinematics in early fossil hominins. MATERIALS AND METHODS: We compare six fossil hominin vertebrae with 48 Homo sapiens and 99 nonhuman primates across 20 genera. We quantify uncinate morphology via geometric morphometric methods to understand how uncinate process shape relates to allometry, taxonomy, and mode of locomotion. RESULTS: Across primates, allometry explains roughly 50% of shape variation, as small, narrow vertebrae feature the relatively tallest, most pronounced uncinate processes, whereas larger, wider vertebrae typically feature reduced uncinates. Taxonomy only weakly explains the residual variation, however, the association between Uncinate Shape and mode of locomotion is robust, as bipeds and suspensory primates occupy opposite extremes of the morphological continuum and are distinguished from arboreal generalists. Like humans, Australopithecus afarensis and Homo erectus exhibit shallow uncinate processes, whereas A. sediba resembles more arboreal taxa, but not fully suspensory primates. DISCUSSION: Suspensory primates exhibit the most pronounced uncinates, likely to maintain visual field stabilization. East African hominins exhibit reduced uncinate processes compared with African apes and A. sediba, likely signaling different degrees of neck motility and modes of locomotion. Although soft tissues constrain neck flexibility beyond limits suggested by osteology alone, this study may assist in modeling cervical kinematics and positional behaviors in extinct taxa.

The cervical spine of Australopithecus sediba.

Cervical vertebrae are rare in the early hominin fossil record, presenting a challenge for understanding the evolution of the neck and head carriage in hominin evolution. Here, we examine the cervical vertebrae of Australopithecus sediba, which unlike other South African taxa is known from associated cervical vertebrae. The A. sediba cervical vertebrae exhibit human-like values for wedging, pedicle cross-sectional areas, and articular facet heights, indicating reduced ventral loading relative to African apes. These features combine with a pattern of vertebral body bone distribution and caudally progressive size expansion suggesting a mode of cervical lordosis, load mitigation, and head carriage similar to humans and distinct from the cantilevered mode of head carriage of the extant African great apes. Yet these derived features in A. sediba are accompanied by ape-like vertebral body and dorsal pillar sizes, articular facet orientation, and uncinate process morphology signaling reduced lateral and rotational coupled movements between vertebral elements and indicate a considerably stiffer neck than in humans. A primitively long and horizontally-oriented C7 spinous process is likely related to a prognathic viscerocranium, although the complimentary C3 spinous process is short, implying large moments emanating from scapular and shoulder elevators rather than large muscles of head stabilization. Cross-sectional spinous process shape and robust anterior tubercles similarly signal increased arm elevation consistent with climbing behavior in corroboration with arboreal signatures previously observed in the shoulder, arms, and hand of A. sediba. Spinal canal shape and size suggests that A. sediba lacked the cervical spinal cord enlargement of Homo that confers humans with enhanced motor control to the upper limbs. The cervical spine of A. sediba thus presents a mosaic of primitive and derived characters, with anatomical features relating to neck posture and head carriage mirroring humans juxtaposed with most other aspects of functional anatomy that resemble chimpanzees.

The vertebrae and ribs of Homo naledi.

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 specimens-two lower thoracic vertebrae and a lower rib-were 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 cord evolution in early Homo.

The discovery at Nariokotome of the Homo erectus skeleton KNM-WT 15000, with a narrow spinal canal, seemed to show that this relatively large-brained hominin retained the primitive spinal cord size of African apes and that brain size expansion preceded postcranial neurological evolution. Here we compare the size and shape of the KNM-WT 15000 spinal canal with modern and fossil taxa including H. erectus from Dmanisi, Homo antecessor, the European middle Pleistocene hominins from Sima de los Huesos, and Pan troglodytes. In terms of shape and absolute and relative size of the spinal canal, we find all of the Dmanisi and most of the vertebrae of KNM-WT 15000 are within the human range of variation except for the C7, T2, and T3 of KNM-WT 15000, which are constricted, suggesting spinal stenosis. While additional fossils might definitively indicate whether H. erectus had evolved a human-like enlarged spinal canal, the evidence from the Dmanisi spinal canal and the unaffected levels of KNM-WT 15000 show that unlike Australopithecus, H. erectus had a spinal canal size and shape equivalent to that of modern humans. Subadult status is unlikely to affect our results, as spinal canal growth is complete in both individuals. We contest the notion that vertebrae yield information about respiratory control or language evolution, but suggest that, like H. antecessor and European middle Pleistocene hominins from Sima de los Huesos, early Homo possessed a postcranial neurological endowment roughly commensurate to modern humans, with implications for neurological, structural, and vascular improvements over Pan and Australopithecus.

Lucy's back: Reassessment of fossils associated with the A.L. 288-1 vertebral column.

The Australopithecus afarensis partial skeleton A.L. 288-1, popularly known as "Lucy" is associated with nine vertebrae. The vertebrae were given provisional level assignments to locations within the vertebral column by their discoverers and later workers. The continuity of the thoracic series differs in these assessments, which has implications for functional interpretations and comparative studies with other fossil hominins. Johanson and colleagues described one vertebral element (A.L. 288-1am) as uniquely worn amongst the A.L. 288-1 fossil assemblage, a condition unobservable on casts of the fossils. Here, we reassess the species attribution and serial position of this vertebral fragment and other vertebrae in the A.L. 288-1 series. When compared to the other vertebrae, A.L. 288-1am falls well below the expected size within a given spinal column. Furthermore, we demonstrate this vertebra exhibits non-metric characters absent in hominoids but common in large-bodied papionins. Quantitative analyses situate this vertebra within the genus Theropithecus, which today is solely represented by the gelada baboon but was the most abundant cercopithecoid in the KH-1s deposit at Hadar where Lucy was discovered. Our additional analyses confirm that the remainder of the A.L. 288-1 vertebral material belongs to A. afarensis, and we provide new level assignments for some of the other vertebrae, resulting in a continuous articular series of thoracic vertebrae, from T6 to T11. This work does not refute previous work on Lucy or its importance for human evolution, but rather highlights the importance of studying original fossils, as well as the efficacy of the scientific method.

Inferring lumbar lordosis in Neandertals and other hominins.

Lumbar lordosis is a key adaptation to bipedal locomotion in the human lineage. Dorsoventral spinal curvatures enable the body's center of mass to be positioned above the hip, knee, and ankle joints, and minimize the muscular effort required for postural control and locomotion. Previous studies have suggested that Neandertals had less lordotic (ventrally convex) lumbar columns than modern humans, which contributed to historical perceptions of postural and locomotor differences between the two groups. Quantifying lower back curvature in extinct hominins is entirely reliant upon bony correlates of overall lordosis, since the latter is significantly influenced by soft tissue structures (e.g. intervertebral discs). Here, we investigate sexual dimorphism, ancestry, and lifestyle effects on lumbar vertebral body wedging and inferior articular facet angulation, two features previously shown to be significantly correlated with overall lordosis in living individuals, in a large sample of modern humans and Neandertals. Our results demonstrate significant differences between postindustrial cadaveric remains and archaeological samples of people that lived preindustrial lifestyles. We suggest these differences are related to activity and other aspects of lifestyle rather than innate population (ancestry) differences. Neandertal bony correlates of lumbar lordosis are significantly different from all human samples except preindustrial males. Therefore, although Neandertals demonstrate more bony kyphotic wedging than most modern humans, we cast doubt on proposed locomotor and postural differences between the two lineages based on inferred lumbar lordosis (or lack thereof), and we recommend future research compare fossils to modern humans from varied populations and not just recent, postindustrial samples.

New fossils of Australopithecus sediba reveal a nearly complete lower back.

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 possessed a lower back consistent with 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'). These results contrast with some 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 overall shape but its vertebral body is somewhat intermediate in shape between modern humans and great apes. Additionally, it bears long, cranially and ventrally oriented costal (transverse) processes, implying powerful trunk musculature. We interpret this combination of features to indicate that A. sediba used its lower back in both bipedal and arboreal positional behaviors, as previously suggested based on multiple lines of evidence from other parts of the skeleton and reconstructed paleobiology of A. sediba.

One of the defining features of humans is our ability to walk comfortably on two legs. To achieve this, our skeletons have evolved certain physical characteristics. For example, the lower part of the human spine has a forward curve that supports an upright posture; whereas the lower backs of chimpanzees and other apes ­ which walk around on four limbs and spend much of their time in trees ­ lack this curvature. Studying the fossilized back bones of ancient human remains can help us to understand how we evolved these features, and whether our ancestors moved in a similar way. Australopithecus sediba was a close-relative of modern humans that lived about two million years ago. In 2008, fossils from an adult female were discovered at a cave site in South Africa called Malapa. However, the fossils of the lower back region were incomplete, so it was unclear whether the female ­ referred to as Malapa Hominin 2 (MH2) ­ had a forward-curving spine and other adaptations needed to walk on two legs. Here, Williams et al. report the discovery of new A. sediba fossils from Malapa. The new fossils are mainly bones from the lower back, and they fit together with the previously discovered MH2 fossils, providing a nearly complete lower spine. Analysis of the fossils suggested that MH2 would have had an upright posture and comfortably walked on two legs, and the curvature of their lower back was similar to modern females. However, other aspects of the bones' shape suggest that as well as walking, A. sediba probably spent a significant amount of time climbing in trees. The findings of Williams et al. provide new insights in to our evolutionary history, and ultimately, our place in the natural world around us. Our lower back is prone to injury and pain associated with posture, pregnancy and exercise (or lack thereof). Therefore, understanding how the lower back evolved may help us to learn how to prevent injuries and maintain a healthy back.

Late subadult ontogeny and adult aging of the human thorax reveals divergent growth trajectories between sexes.

Sexual dimorphism is an important feature of adult thorax morphology, but when and how sex-related differences in the ribcage arise during ontogeny is poorly known. Previous research proposed that sex-related size differences in the nasal region arise during puberty. Therefore, we explore whether ribcage sexual dimorphism also arises at that time and whether this sexual dimorphism is maintained until old age. We measured 526 (semi)landmarks on 80 CT-based human ribcage reconstructions, on individuals ranging from 7 to 65 year-old. The 3D coordinates were submitted to the Procrustes superimposition and analyzed. Our results show that the trajectories of thorax size and shape between sexes diverge at around 12 years of age, and continue slightly diverging until old age. The differential ontogenetic trends cause adult male ribcages to become deeper, shorter, and wider than female. Our results are consistent with the evidence from the cranial respiratory system, with the development of sexual dimorphism probably related to changes in body composition during puberty combined with changes in the reproductive system.

New fossil remains of Homo naledi from the Lesedi Chamber, South Africa.

The Rising Star cave system has produced abundant fossil hominin remains within the Dinaledi Chamber, representing a minimum of 15 individuals attributed to Homo naledi. Further exploration led to the discovery of hominin material, now comprising 131 hominin specimens, within a second chamber, the Lesedi Chamber. The Lesedi Chamber is far separated from the Dinaledi Chamber within the Rising Star cave system, and represents a second depositional context for hominin remains. In each of three collection areas within the Lesedi Chamber, diagnostic skeletal material allows a clear attribution to H. naledi. Both adult and immature material is present. The hominin remains represent at least three individuals based upon duplication of elements, but more individuals are likely present based upon the spatial context. The most significant specimen is the near-complete cranium of a large individual, designated LES1, with an endocranial volume of approximately 610 ml and associated postcranial remains. The Lesedi Chamber skeletal sample extends our knowledge of the morphology and variation of H. naledi, and evidence of H. naledi from both recovery localities shows a consistent pattern of differentiation from other hominin species.

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

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