Morphology of the tongue in the tuatara, Sphenodon punctatus (Reptilia: Lepidosauria), with comments on function and phylogeny

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Abstract
The morphology and histology of the tongue in Sphenodon punctatus are described and used to infer function and to determine character state polarities in lepidosaurs. The tongue lacks an anterior notch and is covered with filamentous papillae, including specialized gustatory papillae containing taste buds. Lingual glands are restricted to mucocytes covering the papillae. Three intrinsic tongue muscles are identified and shown to be discrete fiber systems and not merely elaborations of the M. hyoglossus. These muscles interact with a connective tissue skeleton, particularly three septal planes, to cause changes in tongue shape. Tongue protrusion is probably caused by hyoid protraction and contraction of posterior genioglossus fibers; retraction by hyoid retraction, hyoglossus contraction, and contraction of anterior genioglossus fibers.It is argued that taste is important in prey discrimination and possibly in courtship. Vomeronasal function is probably mediated by inhalation and not tongue movement.Insertion of genioglossus fibers into the buccal floor is a derived feature of lepidosaur tongues. Derived features of squamate tongues include an anterior bifurcation, a divided genioglossus comprising medial and lateral portions, ventral transverse and circular muscle fiber systems around the hyoglossus, and the presence of a median septum. The tongue of the squamate family Iguanidae shares many plesiomorphic features with Sphenodon.

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