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Knuckle-Walking and the Origin of Human Bipedalism

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Abstract

Functional analysis of fossil great apes and humans indicate that no known fossil taxon was a habitual knuckle-walker. However, the phylogenetic relations among hominoids suggests that the last common ancestor of the African apes and humans was in fact a knuckle-walker. Anatomical, fossil and molecular evidence of relations among the Hominoidea strongly suggest than Pan and Homo share a common ancestor not shared by any other living taxon. If this is correct, then knuckle-walking must have evolved once in the common ancestor of Pan, Gorilla and Homo (in which it was lost), or twice, independently in each African ape lineage. In addition to being less parsimonious, most multiple origins hypotheses for knuckle-walking also fail to account for characters shared by African apes and humans that are plausibly functionally related to increased terrestriality. These include vertebral, limb and intra-limb proportions, and limb long bone, carpal, tarsal and metapodial, and phalangeal morphology. In theory, knuckle-walking and obligate bipedalism could have evolved from an unknown type of terrestrial quadrupedalism, possibly associated with high frequencies of facultative bipedality. This would also account for characters and positional behavior shared by African apes and humans, the differences between Pan and Gorilla, and the apparent retention of primitive features of the trunk in humans. However, it implies a substantial increase in homoplasy as well as a hypothetical ancestral morphotype unkown in any group or out group. Functional anatomy and phylogeny together continue to suggest that humans are most likely to have evolved from a knuckle-walker. However, a functionally plausible, though less parsimonious facultative bipedality hypothesis is also possible. In this regard, hints of the unique functional morphology of Ardipithecus ramidus suggest that this taxon would serve to test these two alternatives.

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Begun, D.R. (2004). Knuckle-Walking and the Origin of Human Bipedalism. In: Meldrum, D.J., Hilton, C.E. (eds) From Biped to Strider. Springer, Boston, MA. https://doi.org/10.1007/978-1-4419-8965-9_2

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