The variability of inner ear orientation in saurischian dinosaurs: testing the use of semicircular canals as a reference system for comparative anatomy

The vestibular system of the inner ear houses three semicircular canals-oriented on three nearly-orthogonal planes-that respond to angular acceleration stimuli. In recent years, the orientation of the lateral semicircular canal (LSC) has been regularly used to determine skull orientations for comparative purposes in studies of non-avian dinosaurs. Such orientations have been inferred based on fixing the LSC to a common set of coordinates (parallel to the Earth's horizon), given that the orientation to gravity of this sensory system is assumed constant among taxa. Under this assumption, the LSC is used as a baseline (a reference system) both to estimate how the animals held their heads and to describe craniofacial variation among dinosaurs. However, the available data in living birds (extant saurischian dinosaurs) suggests that the orientation of the LSC in non-avian saurischian dinosaurs could have been very variable and taxon-specific. If such were the case, using the LSC as a comparative reference system would cause inappropriate visual perceptions of craniofacial organization, leading to significant descriptive inconsistencies among taxa. Here, we used Procrustes methods (Geometric Morphometrics), a suite of analytical tools that compares morphology on the basis of shared landmark homology, to show that the variability of LSC relative to skull landmarks is large (ca. 50°) and likely unpredictable, thus making it an inconsistent reference system for comparing and describing the skulls of saurischian (sauropodomorph and theropod) dinosaurs. In light of our results, the lateral semicircular canal is an inconsistent baseline for comparative studies of craniofacial morphology in dinosaurs.