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Columellar Muscle of Neogastropods: Muscle Attachment and the Function of Columellar Folds

Department of Geophysical Sciences, University of Chicago, 5734 S. Ellis Ave., Chicago, Illinois 60637

Malacologists often assume that ornamentation on snail shells is functional, and therefore adaptive. I conducted the first comprehensive test of the widely accepted hypothesis that columellar folds, a type of internal ornamentation, enhance the performance of the columellar muscle, which attaches the snail to its shell. Careful dissections of live, non-relaxed specimens reveal that the physical attachment between the columellar muscle and the columella is not restricted to a small, circular patch located deep within the shell. Instead, the attachment is long and narrow, extending approximately a full whorl along the length of the columella. I developed a novel technique for preparing three-dimensional reconstructions from photographs documenting the dissections. These reconstructions were then used to measure four parameters that describe the muscle: (1) the surface area of the physical attachment between the muscle and columella, (2) the total contact area between the muscle and the columella, (3) the depth of attachment, and (4) the length of attachment. None of these parameters differed significantly between species with and without folds. In light of the biomechanics of muscular hydrostats, values of the first parameter indicate that columellar folds probably do not guide the columellar muscle as the animal moves in and out of its shell. Values of the other parameters indicate that columellar folds neither increase an animal’s ability to maneuver its shell nor facilitate deeper withdrawal. These results, and the fact that folds have evolved convergently several times, might indicate that folds are an easily evolvable solution to many functional problems, none of which are currently understood.

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