The auditory region of dermoptera: Morphology and function relative to other living mammals

The dermopteran basicranium combines a primitively constructed and oriented auditory bulla formed by ectotympanic, rostral entotympanic, and tubal cartilage with derived features of the middle ear transformer and internal carotid circulation. Living dermopterans possess a primitive eutherian auditory region that has been structurally modified to perceive a lower frequency sound spectrum than probably was utilized by ancestral Mesozoic therians. Perception of the low to midfrequency range is enhanced in Dermoptera by reducing stiffness in the mechanical transformer while maintaining low mass of the component parts. Stiffness has been reduced by (1) development of an epitympanic sinus about four times the volume of the middle ear cavity proper, (2) detachment of the anterior process of the malleus from the ectotympanic, and (3) by delicate suspension of the ear ossicles within the middle ear. We apply to dermopterans a measure of hearing efficiency derived from recent functional studies of the mammalian middle ear that regards the middle ear mechanism as an impedance matching transformer. Calculation of the impedance transformer ratio for Dermoptera suggests that these mammals are relatively efficient in comparison to other eutherians in their ability to match the impedance of cochlear fluids to that of air at the eardrum. Dermopterans theoretically are capable of using over 90% of incident sound energy striking the eardrum at the resonant or natural frequency. Mechanical impedance of the middle ear transformer exerts a minimal influence on hearing efficiency due to low mass, little stiffness, and little frictional resistance. Analysis of measurements of the middle ear transformer published by Gerald Fleischer and integration of these data with current theory on the peripheral hearing mechanism in mammals allow us to propose a model that describes the structural and functional evolution of the mammalian middle ear transformer. Structural changes appear to be correlated with alteration in function from primitive small mammals with stiff middle ear transformers and high frequency dominated hearing to mammals with a wider range in body size with more mobile middle ear transformers and a greater range of frequency perception, often including improved sensitivity to lower frequencies. Mammals employ different anatomical strategies in attainment of increased hearing efficiency and sensitivity. Efficiency is improved by adjustment of lever and areal ratios of the middle ear transformer to achieve an optimum impedance match of external air and cochlear fluids. Sensitivity over a broad frequency spectrum is attained by minimizing mass, stiffness, and frictional resistance of the transformer. The morphology of the auditory region of both living and fossil mammals seems explicable in terms of selection pressure directed toward these ends.