Molar Occlusion in Late Triassic Mammals

Summary 1. A new genus and species of late Triassic mammal, Megazostrodon rudnerae , from Lesotho in southern Africa is described. The molars are similar to those of the British Eozostrodon parvus except that they are slightly larger and the upper molars have a large external cingulum supporting well‐developed cusps. 2. Molar occlusion is discussed in two groups of late Triassic mammals: Eoxostrodon and the closely related Megazostrodon on one the hand and the unnamed primitive symmetrodonts on the other. It is shown that in Eoxostrodon the upper and lower molars did not have matching occlusal surfaces upon eruption but that wear produced matching occlusal surfaces. These surfaces are confined to the internal surface of the upper molars and the external surface of the lower molars and form a series of wide‐angled triangles. The main cusp of an upper molar occluded between the main and posterior subsidiary cusp of the lower molar and the main cusp of the lower molar occluded between the main and anterior subsidiary cusp of the upper molar, 3. It is shown that the molars of Docodon and HaIdanodon were possibly derived from those of a primitive mammal such as Eozostrodon . The transition involved the development on the upper molars of an internal extension which, as it increased in size, established contact with the dorsal surfaces of two adjacent lower molars. The process involved is fundamentally different from that leading to tribosphenic molars. 4. In Megaxostrodon the main cusp of the upper molars occluded between the posterior and anterior subsidiary cusps of two adjacent lower molars, i.e. more posteriorly than in Eozostrodon . Primitive Rhaetic symmetrodonts were derived from mammals which had this type of occlusion and which were also closely related to Eoxostrodon and Megaxostrodon . The transition involved a rotation of the subsidiary cusps of the upper molars externally and those of the lower molars internally. This rotation increased the shearing surfaces between occluding upper and lower molars. Cusp rotation was carried further in the acute‐angled symmetrodonts ( Peralestes and Spalacotherium ) and pantotheres. It appears that marked cusp rotation was coupled with the acquisition of transverse movements of the lower jaw during mastication. Transverse movement was apparently not possible in cynodonts, in Eoxostrodon (and related forms) and in Docodon . 5. The evolution of therian molars involves cusp rotation as originally proposed by the Cope—Osborn theory. Criticisms of the Cope—Osborn theory are re‐evaluated in light of the new late Triassic material. 6. In Rhaetic symmetrodonts, molar wear produces matching occlusal facets, but the amount of attrition necessary to produce these facets was considerably less than in Eoxostrodon . In acute‐angled symmetrodonts and in pantotheres, the molars erupt with more precise occlusal surfaces and attrition was not necessary to produce matching surfaces. 7. On the basis of the structure of the molar teeth it was concluded that Eozostrodon, Megazostrodon and Erythrotherium were closely related to the Rhaetic symmetrodonts. Slightly different occlusal relationships between upper and lower molars indicated that in these early mammals constant occlusal relations were being established. 8. Primitive cynodonts, such as Thrinaxodon , are characterized by alternate tooth replacement; there is a total lack of a constant occlusal relationship between upper and lower postcanine teeth. In Thrinaxodon individual postcanines were replaced several times. The crown structures of successive generations of postcanines were different so that a freshly erupted postcanine tooth had a crown structure quite distinct from the tooth which it replaced. It has been shown that the crown structure of one of the generations of postcanine teeth of Thrinaxodon is almost identical to that of Eozostrodon except that Thrinaxodon postcanines have a single root, On the basis of this similarity and the over‐all structure of the primitive cynodont skull, it was concluded that Rhaetic mammals (excluding ictidosaurs and haramyids) could be derived from primitive cynodonts. 9. All the orders of Jurassic mammals (with the possible exception of multituber‐culates) were probably derived from late Triassic mammals. The apparent close relationship of late Triassic mammals is evidence of a monophyletic origin of this class.