The orbitotemporal region and the mandibular joint in the skull of shrews (Soricidae, Mammalia)

Modern phylogenetics place the Soricidae (shrews) into the order Lipotyphla, which belongs to the relatively new superorder clade Laurasiatheria. Their most derived skull feature is the unusual position and shape of the jaw articulation: Whereas in all other mammals the glenoid region of the squamosum is more or less tightly attached to the otic capsule or petrosal, respectively, in the soricids it is attached to the nasal capsule. This new position of the jaw articulation becomes possible by the posterior extension of the nasal capsule and the rostral shift of the glenoid fossa. By the study of dated postnatal ontogenetic stages of Crocidura russula and Sorex araneus, we show that the glenoid part of the squamosal becomes fixed to the nasal capsule by the ossified alae orbitalis and temporalis. The ala orbitalis is displaced laterally by the expanded cupula nasi posterior; this posterior expansion is well documented by the lamina terminalis, which incorporates parts of the palatinum and alisphenoid. Both alae consist largely of 'Zuwachsknochen' ('appositional bone') and are then named orbitosphenoid and alisphenoid. By the forward move of the pars glenoidea and of the alisphenoid, the foramen lacerum medium ('fenestra piriformis') also expands rostrally. Functionally, the forward shift of the jaw joint helps to keep the incisal biting force high. Biomechanically the jaws can be considered as a tweezer, and the rostral position of the jaw joints makes the interorbital pillar and the shell-like walls of the facial skull a lever for the highly specialized incisal dentition.