Functional and phylogenetic interpretation of enamel microstructure in rhinoceroses

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Abstract
Worn enamel surfaces of the cheek teeth in fossil and recent rhinoceroses are characterized by fine, parallel ridges aligned perpendicular to the enamel-dentin interface. We show that these ridges result from an unusual enamel ultrastructure in which a primitively horizontal layering of the prisms has become vertical. The new structure apparently appeared between early and middle Eocene, at the time when the superfamilies of perissodactyls were rapidly diverging. Similar modifications of the enamel structure occurred in certain parts of the cheek teeth in tapiroids, chalicotherioids and brontotherioids, but hardly at all in the equids. The modified enamel structure, where it occurs in groups other than rhinocerotoids, is associated with lophs but not cusps. Experimental evidence shows that the modified enamel is more resistant to wear than the unmodified enamel. The consistent association with thin lophs rather than cusps suggests that the modified enamel evolved to prolong the life of the lophs, where occlusal pressures are highest and attrition greatest. The dominance of modified enamel in rhinocerotoids correlates with the higher degree of compression of the cusps and extreme lophodonty in this group. The absence of modified enamel structure in the equids, even in the ectoloph, correlates with the lesser importance of the ectoloph in equids relative to brontotherioids, chalicotherioids and rhinocerotoids.

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