More convergence and arboreality: Knuckle-walking and the African apes

As long as we're on the topic of homoplasy, a recent study suggests that knuckle-walking evolved independently in chimpanzees (Pan) and gorillas (Gorilla). If true, this suggests that hominins did not evolve from a knuckle-walking ancestor. Interesting.

Take-home points from the paper include:

• Many purported 'knuckle-walking' features of the hominoid wrist might rather indicate arboreal wrist postures
  
• Knuckle-walking in Pan and Gorilla are biomechanically distinct, and may thus have evolved independently in each lineage
• More tentatively: Humans may not have evolved from a knuckle-walking ancestor, lending further credence to the idea that Pan is not a great model for the Pan-human common ancestor
• This may be another example of one of Futuyma's Principles of Evolution: HOMOPLASY IS COMMON IN EVOLUTION

"Knuckle-walking" refers to the mode of locomotion employed by most Pan and Gorilla when on the ground. Whereas most terrestrially quadrupdal primates use either the palmar surfaces of their 'fingers' or their palms to contact the ground, Pan and Gorilla's hands contact the ground with the back surface of the middle of their fingers (their intermediate phalanges, in technical terms). It is a very unusual posture--so far as I know, among all animals it is unique to these apes. So, it is perfectly sensible to assume that that knuckle-walking in chimpanzees and gorillas is homologous, represents the ancestral posture in African apes, and that humans evolved from a knuckle-walking ancestor.

But Tracey Kivell and Dan Schmitt present evidence from the wrist that suggests knuckle-walking in Pan and Gorilla are biomechanically and developmentally distinct. They point to several features of the wrist bones (carpals) that have traditionally been assumed to reflect knuckle-walking behavior. The expression of these features does not fit expectations given size and maturation differences between the two African apes. In fact, most of the features are more common/pronounced in Pan, and sometimes even other primates, more so than in Gorilla. The authors thus posit that many of the hitherto-knuckle-walking features of the wrist are actually indicative of arboreal wrist postures, and not knuckle-walking.

That authors acknowledge that it is possible that the wrist differences between Pan and do not necessarily preclude the possibility that knuckle-walking in the two apes has a common, ancestral origin, and that the differences accumulated after the evolutionary split between Gorilla on the one hand and Pan-humans on the other. That is to say, the behavior in the apes is homologous (as in common ancestry) but non-identical. Another possibility, which would also indicate that humans did evolve from a knuckle-walking ancestor, is that the behavior evolved separately in the Gorilla lineage, and in the Pan-hominin lineage before the split between Pan on the one hand and hominins on the other. The only way to test such a hypothesis is with fossils, fossils which so far as I know we do not have (yet).

Reference
Kivell T and Schmitt D. Independent evolution of knuckle-walking in African apes shows that humans did not evolve from a knuckle-walking ancestor. Proceedings of the National Academy of Sciences, in press.