The Stapes of Gomphodont Cynodonts: Insights into the Middle Ear Structure of Non-Mammaliaform Cynodonts

The stapes is known in several non-mammaliaform cynodonts, although it has only been cursorily studied. Here we thoroughly analyze the stapedial anatomy of several basal cynodonts in a phylogenetic framework. Our study shows that the stapedial anatomy is more variable than previously thought. The morphological variation of the stapes led to the recognition of 11 phylogenetic characters that were included in a total evidence data matrix centered in the analysis of gomphodont cynodonts. Stapes morphology does not provide evidence to suggest a direct connection between the stapes and a postquadrate tympanic membrane (if present) and the hypothesis of a dorsal process as the site of attachment of a small ligament or the stapedial muscle is supported. The re-evaluation of the theories concerning the position of the tympanic membrane in non-mammaliaform cynodonts allowed us to conclude that the hypothetical postquadrate tympanic membrane associated with the squamosal sulcus is at best relictual and most likely non-functional (not connected with the stapes). The sound waves were most likely transmitted to the stapes from a postdentary tympanic membrane through the quadrate. Our analysis results in a better understanding of the auditory system in basal cynodonts and its evolution, highlighting the variability of the stapedial anatomy.