The Role Of The Intramandibular Joint, Symphyseal Tissues, And Wrapping Muscles On Theropod Dinosaur Mandibular Function

Sauropsids, unlike mammals, possess an intramandibular joint (IMJ) separating the dentary and postdentary bones, with different lineages either rigidifying (turtles & crocodilians) or maintaining compliance (lepidosaurs & birds) about the joint. IMJ construction and its role on mandibular performance is unclear, impeding our understanding of its function in extinct animals with extreme feeding behaviors like Tyrannosaurus rex. Sauropsids like T. rex pose a particular biomechanical paradox; feeding traces, coprolites, and their robust crania and teeth indicate that they regularly bit through and ingested bone, but their dorsoventrally-tall and mediolaterally-thin hemimandibles, IMJ, and patent mandibular symphyses suggest their mandibles were ill-suited for bone-crushing bites. Extant sauropsids also exhibit wrapping intramandibularis (mIRA) and pterygoideus ventralis (mPTv) muscles, whose effect on mandibular performance and interaction with the IMJ are unknown. Here we model the effect of the IMJ, symphyseal tissue properties, and wrapping muscle orientation on T. rex mandible biomechanical performance in order to give insight into the biomechanical constraints faced by sauropsids, and the link between their mandibular form and function. Joint tissue histology, gross dissection, and iodine contrast data of extant sauropsids were performed to inform model tissue properties, IMJ construction and muscle orientation. We imported a 3D STL model of the T. rex “STAN” and modeled the IMJ by differing material properties of modeled joint tissues to test the influence of linkage materials on mandibular performance. Muscles were mapped onto the model, and their corresponding muscle moments calculated from area centroids, muscle architecture, and volumes. Wrapping mIRA and mPTv muscles were simulated by directing their force vectors to appropriately positioned dummy points using extant sauropsids as a guide. The mandible was then constrained at the jaw joint and in a series of points along the tooth row from rostral to caudal, simulating both unilateral and bilateral bites. We find that while strains are quite high (>6000 µstrains) about the intramandibular joint, mediolateral bending stresses and strain are markedly reduced by the prearticular, which exhibits greater strain than the surrounding bone. Our results suggest the prearticular of T. rex acted as a strain sink to counteract bending about the IMJ and thereby rigidifying the mandible during feeding. We hypothesize that differences in IMJ articulation and prearticular construction may differentially facilitate or impede prearticular streptognathy, and thus intramandibular kinesis about the IMJ, in sauropsids. Forthcoming models will simulate other theropods within and without Tyrannosauridae to test this hypothesis.