Many of the unresolved debates in palaeoanthropology regarding evolution of particular locomotor or manipulative behaviours are founded in differing opinions about the functional significance of the ...preserved external fossil morphology. However, the plasticity of internal bone morphology, and particularly trabecular bone, allowing it to respond to mechanical loading during life means that it can reveal greater insight into how a bone or joint was used during an individual's lifetime. Analyses of trabecular bone have been commonplace for several decades in a human clinical context. In contrast, the study of trabecular bone as a method for reconstructing joint position, joint loading and ultimately behaviour in extant and fossil non‐human primates is comparatively new. Since the initial 2D studies in the late 1970s and 3D analyses in the 1990s, the utility of trabecular bone to reconstruct behaviour in primates has grown to incorporate experimental studies, expanded taxonomic samples and skeletal elements, and improved methodologies. However, this work, in conjunction with research on humans and non‐primate mammals, has also revealed the substantial complexity inherent in making functional inferences from variation in trabecular architecture. This review addresses the current understanding of trabecular bone functional adaptation, how it has been applied to hominoids, as well as other primates and, ultimately, how this can be used to better interpret fossil hominoid and hominin morphology. Because the fossil record constrains us to interpreting function largely from bony morphology alone, and typically from isolated bones, analyses of trabecular structure, ideally in conjunction with that of cortical structure and external morphology, can offer the best resource for reconstructing behaviour in the past.
For several decades, it was largely assumed that stone tool use and production were abilities limited to the genus Homo. However, growing palaeontological and archaeological evidence, comparative ...extant primate studies, as well as results from methodological advancements in biomechanics and morphological analyses, have been gradually accumulating and now provide strong support for more advanced manual manipulative abilities and tool-related behaviours in pre-Homo hominins than has been traditionally recognized. Here, I review the fossil evidence related to early hominin dexterity, including the recent discoveries of relatively complete early hominin hand skeletons, and new methodologies that are providing a more holistic interpretation of hand function, and insight into how our early ancestors may have balanced the functional requirements of both arboreal locomotion and tool-related behaviours.
Modelling a muscle key to flexing the thumb, a new study suggests that the powerful opposability that characterises the dextrous human hand evolved in some of our fossil relatives about two million ...years ago — a time when tool use became more important.
Modelling a muscle key to flexing the thumb, a new study suggests that the powerful opposability that characterises the dextrous human hand evolved in some of our fossil relatives about two million years ago — a time when tool use became more important.
In 1994, Hunt published the ‘postural feeding hypothesis’—a seminal paper on the origins of hominin bipedalism—founded on the detailed study of chimpanzee positional behavior and the functional ...inferences derived from the upper and lower limb morphology of the Australopithecus afarensis A.L. 288-1 partial skeleton. Hunt proposed a model for understanding the potential selective pressures on hominins, made robust, testable predictions based on Au. afarensis functional morphology, and presented a hypothesis that aimed to explain the dual functional signals of the Au. afarensis and, more generally, early hominin postcranium. Here we synthesize what we have learned about Au. afarensis functional morphology and the dual functional signals of two new australopith discoveries with relatively complete skeletons (Australopithecus sediba and StW 573 ‘Australopithecus prometheus’). We follow this with a discussion of three research approaches that have been developed for the purpose of drawing behavioral inferences in early hominins: (1) developments in the study of extant apes as models for understanding hominin origins; (2) novel and continued developments to quantify bipedal gait and locomotor economy in extant primates to infer the locomotor costs from the anatomy of fossil taxa; and (3) novel developments in the study of internal bone structure to extract functional signals from fossil remains. In conclusion of this review, we discuss some of the inherent challenges of the approaches and methodologies adopted to reconstruct the locomotor modes and behavioral repertoires in extinct primate taxa, and notably the assessment of habitual terrestrial bipedalism in early hominins.
Morphological variation in the hominoid capitate has been linked to differences in habitual locomotor activity due to its importance in movement and load transfer at the midcarpal joint proximally ...and carpometacarpal joints distally. Although the shape of bones and their articulations are linked to joint mobility, the internal structure of bones has been shown experimentally to reflect, at least in part, the loading direction and magnitude experienced by the bone. To date, it is uncertain whether locomotor differences among hominoids are reflected in the bone microarchitecture of the capitate. Here, we apply a whole‐bone methodology to quantify the cortical and trabecular architecture (separately and combined) of the capitate across bipedal (modern Homo sapiens), knuckle‐walking (Pan paniscus, Pan troglodytes, Gorilla sp.), and suspensory (Pongo sp.) hominoids (n = 69). It is hypothesized that variation in bone microarchitecture will differentiate these locomotor groups, reflecting differences in habitual postures and presumed loading force and direction. Additionally, it is hypothesized that trabecular and cortical architecture in the proximal and distal regions, as a result of being part of mechanically divergent joints proximally and distally, will differ across these portions of the capitate. Results indicate that the capitate of knuckle‐walking and suspensory hominoids is differentiated from bipedal Homo primarily by significantly thicker distal cortical bone. Knuckle‐walking taxa are further differentiated from suspensory and bipedal taxa by more isotropic trabeculae in the proximal capitate. An allometric analysis indicates that size is not a significant determinate of bone variation across hominoids, although sexual dimorphism may influence some parameters within Gorilla. Results suggest that internal trabecular and cortical bone is subjected to different forces and functional adaptation responses across the capitate (and possibly other short bones). Additionally, while separating trabecular and cortical bone is normal protocol of current whole‐bone methodologies, this study shows that when applied to carpals, removing or studying the cortical bone separately potentially obfuscates functionally relevant signals in bone structure.
This research examines the trabecular and cortical architecture of the capitate of great apes. It is concluded that the internal structure of the human capitate is differentiated from nonhuman apes by a combination of thin cortical bone in the distal segment and more anisotropic trabeculae in the proximal. Further, there are significant differences in the proximal and distal segments of the capitate which indicates variable loading and functional adaptation across the small bone in hominoids.
La main humaine a changé de forme et de fonction tout au long de notre évolution en raison, entre autres, de la bipédie terrestre, de l’utilisation d’outils et de l’asymétrie directionnelle telle que ...la préférence manuelle. Cependant, nous manquons d’informations sur la variabilité potentielle des facteurs clés pouvant affecter la force de préhension et la dextérité, en particulier la taille, la forme et la fonction quotidienne de la main. Nous avons mesuré la force de préhension par pincemen...
Differences in grip techniques used across primates are usually attributed to variation in thumb-finger proportions and muscular anatomy of the hand. However, this cause-effect relationship is not ...fully understood because little is known about the biomechanical functioning and mechanical loads (e.g., muscle or joint forces) of the non-human primate hand compared to that of humans during object manipulation. This study aims to understand the importance of hand proportions on the use of different grip strategies used by humans, extant great apes (bonobos, gorillas and orangutans) and, potentially, fossil hominins (Homo naledi and Australopithecus sediba) using a musculoskeletal model of the hand. Results show that certain grips are more challenging for some species, particularly orangutans, than others, such that they require stronger muscle forces for a given range of motion. Assuming a human-like range of motion at each hand joint, simulation results show that H. naledi and A. sediba had the biomechanical potential to use the grip techniques considered important for stone tool-related behaviors in humans. These musculoskeletal simulation results shed light on the functional consequences of the different hand proportions among extant and extinct hominids and the different manipulative abilities found in humans and great apes.
Bipedalism is a defining trait of the hominin lineage, associated with a transition from a more arboreal to a more terrestrial environment. While there is debate about when modern human-like ...bipedalism first appeared in hominins, all known South African hominins show morphological adaptations to bipedalism, suggesting that this was their predominant mode of locomotion. Here we present evidence that hominins preserved in the Sterkfontein Caves practiced two different locomotor repertoires. The trabecular structure of a proximal femur (StW 522) attributed to Australopithecus africanus exhibits a modern human-like bipedal locomotor pattern, while that of a geologically younger specimen (StW 311) attributed to either Homo sp. or Paranthropus robustus exhibits a pattern more similar to nonhuman apes, potentially suggesting regular bouts of both climbing and terrestrial bipedalism. Our results demonstrate distinct morphological differences, linked to behavioral differences between Australopithecus and later hominins in South Africa and contribute to the increasing evidence of locomotor diversity within the hominin clade.
Objectives
The dexterity of fossil hominins is often inferred by assessing the comparative manual anatomy and behaviors of extant hominids, with a focus on the thumb. The aim of this study is to test ...whether trabecular structure is consistent with what is currently known about habitually loaded thumb postures across extant hominids.
Materials and methods
We analyze first metacarpal (Mc1) subarticular trabecular architecture in humans (Homo sapiens, n = 10), bonobos (Pan paniscus, n = 10), chimpanzees (Pan troglodytes, n = 11), as well as for the first time, gorillas (Gorilla gorilla gorilla, n = 10) and orangutans (Pongo sp., n = 1, Pongo abelii, n = 3 and Pongo pygmaeus, n = 5). Using a combination of subarticular and whole‐epiphysis approaches, we test for significant differences in relative trabecular bone volume (RBV/TV) and degree of anisotropy (DA) between species.
Results
Humans have significantly greater RBV/TV on the radiopalmar aspects of both the proximal and distal Mc1 subarticular surfaces and greater DA throughout the Mc1 head than other hominids. Nonhuman great apes have greatest RBV/TV on the ulnar aspect of the Mc1 head and the palmar aspect of the Mc1 base. Gorillas possessed significantly lower DA in the Mc1 head than any other taxon in our sample.
Discussion
These results are consistent with abduction of the thumb during forceful “pad‐to‐pad” precision grips in humans and, in nonhuman great apes, a habitually adducted thumb that is typically used in precision and power grips. This comparative context will help infer habitual manipulative and locomotor grips in fossil hominins.
PDF
