Homology of the “ethmoid complex” of tyrannosaurids and its implications for the reconstruction of the olfactory apparatus of non-avian theropods

ABSTRACT

The “ethmoid complex” is an enigmatic element of the anterior portion of the braincase first described in Tyrannosaurus rex in 1912, which has since been recognized in many non-avian theropods. Because the “ethmoid complex” is a landmark for the reconstruction of the olfactory apparatus of non-avian theropods, we clarify the homology of this structure among archosaurs. The “ethmoid complex” consists of a trough-shaped element that is attached to an anteriorly-located median septum capped by a dorsal plate. Based on anatomical comparisons with the olfactory region of extant birds and crocodylians, the components of the “ethmoid complex” are shown to have cartilaginous or osteological homologues among extant archosaurs: the trough is homologous to the anterior portion of the planum supraseptale of crocodylians and embryonic birds, whereas the median septum and overlying dorsal plate are homologous to the avian mesethmoid and to the nasal septum and tectum nasi of crocodylians. Based on the location and ossification of olfactory region structures in non-avian theropods, the most appropriate terms for elements of the “ethmoid complex” are the sphenethmoid for the trough and the mesethmoid for the median septum and dorsal plate. The olfactory bulbs of nonavian theropods were housed within the sphenethmoid, which restricted the maximum size of the olfactory bulbs to a size smaller than the cerebral hemispheres.