Tinkering with elephants’ feet

December 24, 2011 • 10:14 am

by Greg Mayer

One of the most important lessons of comparative anatomy is that evolution usually proceeds by the modification of pre-existing structures (or, stated more precisely, the modification of the pre-existing developmental programs that produce those structures). Certain changes are easier to evolve because the developmental system can be modified to produce them—evolution follows the developmental path of least resistance. In terms of the skeleton of vertebrates, this means that most evolutionary changes are reduction, fusion, loss, lengthening, shortening, thickening,  and narrowing of bones. Evolution uses what’s already there, and rarely do wholly new structures arise.

An interesting example of this principle is in a paper published in yesterday’s issue of Science (BBC story here.)

Right front foot of elephant. The dark blue bone is the metacarpal of the first digit (the thumb in primates); the dark green bone is the sixth 'digit' or prepollex, a sesamoid bone (a bone developed within a tendon). (J.R. Hutchinson, via BBC)

A paper by John R. Hutchinson and colleagues of the Royal Veterinary College (see also his neat elephant lab website) reports the presence of a sixth ‘digit’ in the fatty pad of the feet of elephants (oddly, they don’t say which species of elephant, at least not in the published paper). Primitively, mammals are flat footed, but as they become more cursorial (i.e., adapted for running), they move to running on their toes, and finally, as in horses, on the tips of their toes.

Elephants are midway between walking on their toes and toe tips (a condition known as subunguligrade), and have a large fatty pad at the bottom of their feet to support their great weight. One of the neatest aspect of this paper is that the relatively good fossil record of elephants allows Hutchinson and colleagues to trace when (about 40 mya) and in what groups this evolutionary change toward subunguligrade locomotion and in the foot skeleton occurred. The sixth ‘digit’ is an enlarged sesamoid bone medial to the first true digit. (A sesamoid is a bone that develops within a tendon.) It is not a true toe, which would have a meta carpal/tarsal and one or more phalanges. Rather, a different bone altogether has been pressed into service to approximate a digit.

Such jury rigged adaptations were made famous by Stephen Jay Gould, who wrote about another modification of a sesamoid bone, the panda’s thumb. The panda uses its enlarged sesamoid on the front limb to aid in stripping leaves off of bamboo. A true thumb, as in primates and certain marsupials, is the first digit. The panda, being a bear, has all five of its flat footed digits, and the sesamoid is pressed into service as a thumb substitute.

Darwin and Gould used such jury-rigged adaptations as powerful evidence for descent with modification, because their imperfection and evident relation to structures of different use in other organisms (what we would now recognize as pre-existing structures) show them to be traces of history, not paragons of design. The ‘one bone-two bones-many bones’ structure of the tetrapod limb, and its manifold modifications for walking, running, digging, grasping, swimming, flying, etc., are the most familiar example of this. This tetrapod unity of type embraces similarities beyond what might be called for for functional reasons, and instead is a marker of ancestry. The limbs undergo a sequence of adaptations to various modes of life, but all are modifications of the immediately ancestral limb.

One of my favorite examples of the sequential adaptation of limbs is Richard Swann Lull’s account of the hind limb in marsupials. Primitive marsupials are arboreal (i.e., living in trees), while more derived forms are quite varied, including the speedy kangaroos. Primitive marsupials have a grasping great toe (digit I), while kangaroos have the elongated limbs and feet and reduced side toes typical of cursorial or saltatorial runners; they have lost the great toe altogether. Lull calls these primary and secondary adaptations. Tree kangaroos have become arboreal again. While it might be nice to have a grasping toe, ancestral kangaroos lacked this toe, and so they made do with what they had: the hind foot becomes broadened and shortened for use on tree limbs, and the claws are used for digging in.

As the Zen poet Donald Rumsfeld might have put it, “You become arboreal with the feet you have, not the feet you want.”

Sequential adaptation in marsupials. Hind feet of a climbing opossum (A), a saltatorial kangaroo (B) and a climbing tree kangaroo (C). Toe I is the 'opossum's thumb' (actually it's big toe), which is lacking in the kangaroos. (From Lull, 1917)


Gould, S.J. 1978. The panda’s peculiar thumb. Natural History 87 (November): 20, 24-30.

Hutchinson J.R., C. Delmer, C.E. Miller, T. Hildebrandt, A.A. Pitsillides, and A. Boyde. 2011. From flat foot to fat foot: structure, ontogeny, function, and evolution of elephant “sixth toes”. Science 334:1699-1703.

Lull, R.S. 1917. Organic Evolution. Macmillan, New York.

5 thoughts on “Tinkering with elephants’ feet

  1. Thanks for the alert – evolution is great at shoving the foot into the shoe as far as it will go.

    “they don’t say which species of elephant” – that is a bit of a boob from the editors surely!

    I thought this would be an ‘exaptation’ though?

    1. “Exaptation” is an unnecessary term in the science of form. It confuses more than it clarifies. Lull’s notion of primary and secondary adaptations, or Gregory’s habitus and heritage, are much better ways of looking at changes of function. I’ve attributed the phrase “sequential adaptation” for a long time to Lull, but after a recent rereading I don’t think he used it; it was my term for summarizing his views.

      Here’s an example. Feathers probably evolved in dinosaurs as insulation. They later were modified to form an airfoil. Are feathers an “exaptation”? No. Feathers, qua feathers, are an adaptation for thermoregulation. The aerodynamic form of feathers (vane near the leading edge), and their arrangement on the forelimb, are adaptations for aerial locomotion. They are sequential adaptations. The error of “exaptation” is to think of traits or features of organisms as unanalysed wholes: ‘feathers’ are not merely feathers, but a whole suite of features of the organism. If “exaptation” means that later adaptations are based on the pre-existing structures, then the term is vacuous– all adaptations are then exaptations.

      Another example. Bird’s wings may, in shorthand, be said to be adaptations for flight. (It’s shorthand because it leaves out the long history of sequential adaptation of the many parts of the wing. The vertebrate forelimb was for walking before flying, swimming before walking, and so on, and there are a large number of adaptations to changing function along the way.) Some herons now use their wings to shade the water, thus blocking glare, so that they may more easily see fish in the water. How are we to understand this? As sequential adaptation. The second adaptation in this case is behavioral, and, like all adaptations arises form the pre-existing features of the organism. If, in some future time, wings evolve to be larger, or more opaque, or in some other way to aid the shading function, then these features may be said to be adaptations for shade fishing. The concept of “exaptation” never comes into play.


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