Ontogeny and cranial morphology of the tympanic region of the Tupaiidae, with special reference to Ptilocercus

The structure of the tympanic region of the skull of Ptilocercus lowii was studied in an embryo of 30 mm crown-rump length and in 5 osteocrania. As in Tupaia, the anterior wall of the bulla of Ptilocercus is not completed by a tympanic process of the alisphenoid, contrary to earlier reports. Ptilocercus resembles Tupaia in the following derived characters. The ventral wall of the tympanic cavity is formed by a rostral entotympanic and by a caudal tympanic process of the petrosal. The entotympanic develops in primary connection with the tubal cartilage. The tympanic aperture of the auditory tube is bordered by the entotympanic. The ring-shaped tympanicum is covered by the entotympanicum and is aphaneric. The musculus tensor tympani is lacking. Among mammals, these characters can be regarded as synapomorphic for the Tupaiidae, that is, to have been present in the common ancestor of the two subfamilies. From the evidence of the tympanic region, the Tupaiidae, therefore, form a monophyletic group. Besides these synapomorphies, there are remarkable differences between Ptilocercus and Tupaia in the structure of the bulla. In Ptilocercus the bulla is smaller and less pneumatized than in Tupaia. An anterior intrabullar septum, present in Tupaia, is lacking in Ptilocercus. The epitympanic wing of the alisphenoid is smaller in Ptilocercus than in Tupaia. A lateral prefacial commissure of the tegmen tympani is present in Ptilocercus, but absent in Tupaia. The caudal tympanic process of the petrosal is larger in Ptilocercus than in Tupaia. These characters are autapomorphic for the Ptilocercinae and for the Tupaiinae, respectively. They demonstrate that the auditory bulla of Ptilocercus and that of Tupaia have evolved independently to a considerable extent. An early phylogenetic separation of their respective ancestors seems likely. The tympanic region of the skull provides no evidence for close relationships of the tree shrews to the primates or to any other eutherians. The classification of the Tupaiidae in a separate order, Scandentia, is supported.