Form and function of the tongue in agamid lizards with comments on its phylogenetic significance

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Abstract
The morphology of the tongue of agamid lizards is reviewed and discussed in the context of its functional and phylogenetic significance. It is shown that in several features, including the development of the central musculature of the tongue into a ring muscle and the presence of a genioglossus internus muscle in adults, the tongue in most agamids is derived relative to that in other squamates. In some features, such as the vertical connective tissue septa, agamids share primitive features with Sphenodon. Some conditions found in agamids are also found in anoline iguanids. Two genera, Uromastyx and Leiolepis, differ significantly from other agamids in intrinsic tongue musculature. The functional significance of the unique tongue morphology is that agamids utilize a different mechanism of tongue protrusion from that of other lizards. This mechanism involves the production of force against the lingual process, leading to an anterior slide of the tongue, and is detailed in this paper. Finally, I discuss the mechanical basis for the transformation series of tongue protrusion mechanisms from agamids to chamaeleonids. It is suggested that the mechanism of tongue protrusion in chamaeleonids is not unique, but is a highly derived state of the condition found in agamids.