Evolution of the Tribosphenic Molar Pattern in Early Mammals, with Comments on the Dual-Origin Hypothesis

Development of the tribosphenic molar was a fundamental event that likely influenced the rise of modern mammals. This multi-functional complex combined shearing and grinding in a single chewing stroke, and provided the base morphology for the later evolution of the myriad dental morphologies employed by mammals today. Here a series of morphotypes are presented that represent stepwise acquisition of characters of the molar crown, in an effort to clarify homologies and functional analogies among molars of tribosphenic and tribosphenic-like mammals, as well as their putative sister groups. This is accomplished by evaluation of wear features, which provide direct evidence of occlusal function, and mapping these features on molars of the various morphotypes demonstrates their utility in determining homology. The original singular lower molar talonid cusp is homologous with the hypoconid, and upper molar cusp C in early mammals is homologous with the metacone (cusp C is a neomorph with variable occurrence). The lingual translation of the metacone to a position more directly distal to the paracone (as in Peramus) creates an embrasure for the lower molar hypoconid, and is accompanied by the development of the hypoconulid and a new shearing surface. Lastly, the Gondwanan radiation of tribosphenic-like mammals, the Australosphenida (including monotremes), is determined to be functionally non-tribosphenic. The Tribosphenida are restricted to Laurasian taxa, with an origin at or just prior to the Jurassic-Cretaceous boundary.