Reappraisal of the envenoming capacity of Euchambersia mirabilis (Therapsida, Therocephalia) using μCT-scanning techniques.

Euchambersia mirabilis is an iconic species of Permo-Triassic therapsid because of its unusually large external maxillary fossa linked through a sulcus to a ridged canine. This anatomy led to the commonly accepted conclusion that the large fossa accommodated a venom gland. However, this hypothesis remains untested so far. Here, we conducted a μCT scan assisted reappraisal of the envenoming capacity of Euchambersia, with a special focus on the anatomy of the maxillary fossa and canines. This study shows that the fossa, presumably for the venom-producing gland, is directly linked to the maxillary canal, which carries the trigeminal nerve (responsible for the sensitivity of the face). The peculiar anatomy of the maxillary canal suggests important reorganisation in the somatosensory system and that a ganglion could possibly have been present in the maxillary fossa instead of a venom gland. Nevertheless, the venom gland hypothesis is still preferred since we describe, for the first time, the complete crown morphology of the incisiform teeth of Euchambersia, which strongly suggests that the complete dentition was ridged. Therefore Euchambersia manifests evidence of all characteristics of venomous animals: a venom gland (in the maxillary fossa), a mechanism to deliver the venom (the maxillary canal and/or the sulcus located ventrally to the fossa); and an apparatus with which to inflict a wound for venom delivery (the ridged dentition).