This study asks whether there are discernable links between precision gripping, tool behaviors, and hand morphology in modern hominoids, which may guide functional interpretation of early hominid hand morphology. Findings from a three-pronged investigation answer this question in the affirmative, as follows: (1) Experimental manufacture of early prehistoric tools provides evidence of connections between distinctive human precision grips and effective tool making. (A connection is not found between the "fine" thumb/index finger pad precision grip and early tool making.) (2) Manipulative behavior studies of chimpanzees, hamadryas baboons, and human show that human precision grips are distinguished by the greater force with which objects may be secured by the thumb and fingers of one hand (precision pinching) and the ability to adjust the orientation of gripped objects through movements at joints distal to the wrist (precision handling). (3) Morphological studies reveal eight featured distinctive of modern humans which facilitate use of these grips. Among these features are substantially larger moment arms for intrinsic muscles that stabilize the proximal thumb joints. Examination of evidence for these reveals that three of the eight features occur in Australopithecus afarensis, but limited thumb mobility would have compromised tool making. Also, Olduvai hand morphology strongly suggests a capacity for stone tool making. However, functional and behavioral implications of Sterkfontein and Swartkrans hand morphology are less clear. At present, no single skeletal feature can be safely relied upon as an indicator of distinctively human capabilities for precision gripping or tool making in fossil hominids.
Molecular evidence indicates that the last common ancestor of the genus Pan and the hominin clade existed between 8 and 4 million years ago (Ma). The current fossil record indicates the Pan-Homo last common ancestor existed at least 5 Ma and most likely between 6 and 7 Ma. Together, the molecular and fossil evidence has important consequences for interpreting the evolutionary history of the hand within the tribe Hominini (hominins). Firstly, parsimony supports the hypothesis that the hand of the last common ancestor most likely resembled that of an extant great ape overall ( Pan , Gorilla , and Pongo ), and that of an African ape in particular. Second, it provides a context for interpreting the derived changes to the hand that have evolved in various hominins. For example, the Australopithecus afarensis hand is likely derived in comparison with that of the Pan -Homo last common ancestor in having shorter fingers relative to thumb length and more proximo-distally oriented joints between its capitate, second metacarpal, and trapezium. This evidence suggests that these derived features evolved prior to the intensification of stone tool-related hominin behaviors beginning around 2.5 Ma. However, a majority of primitive features most likely present in the Pan -Homo last common ancestor are retained in the hands of Australopithecus , Paranthropus /early Homo , and Homo floresiensis . This evidence suggests that further derived changes to the hands of other hominins such as modern humans and Neandertals did not evolve until after 2.5 Ma and possibly even later than 1.5 Ma, which is currently the earliest evidence of Acheulian technology. The derived hands of modern humans and Neandertals may indicate a morphological commitment to tool-related manipulative behaviors beyond that observed in other hominins, including those (e.g. H. floresiensis ) which may be descended from earlier tool-making species.
Was stone tool making a factor in the evolution of human hand morphology? Is it possible to find evidence in fossil hominin hands for this capability? These questions are being addressed with increasingly sophisticated studies that are testing two hypotheses; (i) that humans have unique patterns of grip and hand movement capabilities compatible with effective stone tool making and use of the tools and, if this is the case, (ii) that there exist unique patterns of morphology in human hands that are consistent with these capabilities. Comparative analyses of human stone tool behaviours and chimpanzee feeding behaviours have revealed a distinctive set of forceful pinch grips by humans that are effective in the control of stones by one hand during manufacture and use of the tools. Comparative dissections, kinematic analyses and biomechanical studies indicate that humans do have a unique pattern of muscle architecture and joint surface form and functions consistent with the derived capabilities. A major remaining challenge is to identify skeletal features that reflect the full morphological pattern, and therefore may serve as clues to fossil hominin manipulative capabilities. Hominin fossils are evaluated for evidence of patterns of derived human grip and stress-accommodation features.
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