Independent evolution of knuckle-walking in African apes shows that humans did not evolve from a knuckle-walking ancestor

  1. Tracy L. Kivella,1,2 and
  2. Daniel Schmitta
  1. aDepartment of Evolutionary Anthropology, Duke University, P.O. Box 90383 Science Drive, Durham, NC 27708-0383
  • 2Present address: Max Planck Institute for Evolutionary Anthropology, Department of Human Evolution, Deutscher Platz 6, D-04103 Leipzig, Germany.

  1. Edited by Alan Walker, Pennsylvania State University, University Park, PA, and approved June 30, 2009 (received for review February 5, 2009)

Abstract

Despite decades of debate, it remains unclear whether human bipedalism evolved from a terrestrial knuckle-walking ancestor or from a more generalized, arboreal ape ancestor. Proponents of the knuckle-walking hypothesis focused on the wrist and hand to find morphological evidence of this behavior in the human fossil record. These studies, however, have not examined variation or development of purported knuckle-walking features in apes or other primates, data that are critical to resolution of this long-standing debate. Here we present novel data on the frequency and development of putative knuckle-walking features of the wrist in apes and monkeys. We use these data to test the hypothesis that all knuckle-walking apes share similar anatomical features and that these features can be used to reliably infer locomotor behavior in our extinct ancestors. Contrary to previous expectations, features long-assumed to indicate knuckle-walking behavior are not found in all African apes, show different developmental patterns across species, and are found in nonknuckle-walking primates as well. However, variation among African ape wrist morphology can be clearly explained if we accept the likely independent evolution of 2 fundamentally different biomechanical modes of knuckle-walking: an extended wrist posture in an arboreal environment (Pan) versus a neutral, columnar hand posture in a terrestrial environment (Gorilla). The presence of purported knuckle-walking features in the hominin wrist can thus be viewed as evidence of arboreality, not terrestriality, and provide evidence that human bipedalism evolved from a more arboreal ancestor occupying the ecological niche common to all living apes.

Footnotes

  • 1To whom correspondence should be addressed. E-mail: tracy.kivell{at}duke.edu
  • Author contributions: T.L.K. designed research; T.L.K. performed research; T.L.K. and D.S. analyzed data; and T.L.K. and D.S. wrote the paper.

  • The authors declare no conflict of interest.

  • This article is a PNAS Direct Submission.

  • This article contains supporting information online at www.pnas.org/cgi/content/full/0901280106/DCSupplemental.

  • * For clarity in this paper, we use “neutral” when referring to a posture in which the wrist in held in line with the radius and ulna of the forearm. Deviations from neutral in which the angle of the wrist relative to the anterior forearm is greater than 180 degrees is referred to as “extension” (see Fig. 3).

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