Can skull form predict the shape of the temporomandibular joint? A study using geometric morphometrics on the skulls of wolves and domestic dogs.

The temporomandibular joint (TMJ) conducts and restrains masticatory movements between the mammalian cranium and the mandible. Through this functional integration, TMJ morphology in wild mammals is strongly correlated with diet, resulting in a wide range of TMJ variations. However, in artificially selected and closely related domestic dogs, dietary specialisations between breeds can be ruled out as a diversifying factor although they display an enormous variation in TMJ morphology. This raises the question of the origin of this variation. Here we hypothesise that, even in the face of reduced functional demands, TMJ shape in dogs can be predicted by skull form; i.e. that the TMJ is still highly integrated in the dog skull. If true, TMJ variation in the dog would be a plain by-product of the enormous cranial variation in dogs and its genetic causes. We addressed this hypothesis using geometric morphometry on a data set of 214 dog and 60 wolf skulls. We digitized 53 three-dimensional landmarks of the skull and the TMJ on CT-based segmentations and compared (1) the variation between domestic dog and wolf TMJs (via principal component analysis) and (2) the pattern of covariation of skull size, flexion and rostrum length with TMJ shape (via regression of centroid size on shape and partial least squares analyses). We show that the TMJ in domestic dogs is significantly more diverse than in wolves: its shape covaries significantly with skull size, flexion and rostrum proportions in patterns which resemble those observed in primates. Similar patterns in canids, which are carnivorous, and primates, which are mostly frugivorous imply the existence of basic TMJ integration patterns which are independent of dietary adaptations. However, only limited amounts of TMJ variation in dogs can be explained by simple covariation with overall skull geometry. This implies that the final TMJ shape is gained partially independently of the rest of the skull.