XROMM Analysis of Feeding Mechanics in Toads: Interactions of the Tongue, Hyoid, and Pectoral Girdle

During feeding in many terrestrial vertebrates, the tongue acts in concert with the hyoid and pectoral girdle. In frogs, these three elements are interconnected by musculature. While the feeding mechanics of the anuran tongue are well-studied, little is known of how the motions of the tongue relate to the movements of the skeleton or how buccal structures move following closure of the mouth. Although features such as the pectoral girdle and hyoid are not externally visible in frogs, their motions can be tracked in X-ray video. We used XROMM (X-ray Reconstruction of Moving Morphology) techniques to track the 3D movements of the tongue, hyoid apparatus, pectoral girdle, skull, and jaw during the feeding cycle of the cane toad, Rhinella marina. We show how the movements of these elements are integrated during tongue protrusion and prey capture, as well as during prey transport, swallowing, and recovery. Our findings suggest that the hyoid apparatus is important both for prey manipulation and swallowing. The tongue consistently stretches posterior to the skull during swallowing, often more than it stretches during protrusion to reach the prey. Feeding kinematics are similar between individuals, and the kinematics of unsuccessful strikes generally resemble those of successful strikes. Our data also provide a new perspective on the potential role of the pectoral girdle, an element with a predominant locomotor function, during feeding events. This work raises new questions about the evolution of feeding in frogs, as well as how the diversity of pectoral and buccal anatomy observed across anurans may influence feeding kinematics.