Enamel structure in astrapotheres and its functional implications.

Astrapotheres, large extinct ungulates of South America, share with rhinoceroses vertical prism decussation in the cheek tooth enamel. The similarity extends beyond merely the direction of the planes of decussation. The vertical decussation in astrapotheres is confined to the inner part of the enamel and has uniformly well-defined zones in which the prism direction differs by nearly 90 degrees and the zones are separated by narrow transitional borders of intermediate prism direction. The outer enamel consists of predominantly occlusally and outwardly directed prisms. Within the outer enamel is a region of horizontally decussating prisms; here the angle of decussation is usually smaller than that of the inner vertically decussating prisms. Except for the horizontal decussation in the outer enamel, these conditions match structures that have been described for rhinocerotoids. These features, together with the similarity in premortem crack direction and gross shape of the cheek teeth, imply that astrapotheres and rhinocerotoids shared essentially the same system of cheek tooth mechanics. However, the microstructure of the canine enamel in the astrapotheres is distinct. The lower canine enamel of the Oligocene Parastrapotherium exhibits a form of vertical decussation modified by a wavelike bending of prism zones, whereas the decussation in the rhinocerotoid canine is horizontal. The lower canine in Parastrapotherium was subjected to different loading conditions, judging from multiple sets of premortem crack directions. The modified vertical decussation would in theory resist cracking under different directions of tensile stresses. This is confirmed by the sinuous paths of cracks that run in directions differing by up to 90 degrees. That diverse stresses were generated in the enamel during life is confirmed by the pattern of premortem cracks in Parastrapotherium. The enamel in the upper canine of a late Miocene astrapothere lacks decussation but may have resisted cracking under varied loading conditions by virtue of a 3-dimensional wavelike bending of the prisms.