Functional morphology of the “tongue‐bite” in the osteoglossomorph fish Notopterus

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Abstract
Osteoglossomorph fishes are characterized by having three sets of jaws: a mandibular jaw apparatus (MJA) anteriorly, a pharyngeal jaw apparatus (PJA) posteriorly, and a tongue‐bite apparatus (TBA) associated with basihyal and parasphenoid teeth. The TBA is a novel complex feature of the head that characterizes osteoglossomorph fishes and provides a case study in the origin of novel functions and roles in the vertebrate musculoskeletal system.The function of the tongue‐bite in the osteoglossomorph fish Notopterus was characterized by using high‐speed cinematography and electromyography. The tongue‐bite is used during intraoral prey processing to shred and disable prey. Two distinct uses of the TBA were defined on the basis of kinematic and electromyographic profiles: raking and open‐mouth chewing. During raking behavior, the prey is held fixed in the MJA, the neurocranium is elevated, and the pectoral girdle is retracted. The adductor mandibulae, hypaxialis, epaxialis, and posterior intermandibularis muscles are all highly active, but only very low activity is observed in the sternohyoideus muscle. During open‐mouth chewing behavior, the prey is located within the oral cavity, posterior pectoral girdle rotation is less than during raking, and the levator operculi muscle shows relatively high activity.We propose that a shearing action of the basihyal (moved anteroposteriorly by the posterior intermandibularis and hypaxial muscles) with respect to the neurocranium (elevated by epaxial muscles) is the critical aspect of the tongue‐bite in Notopterus. The body muscles (epaxials and hypaxials) provide the main power for the tongue‐bite. We hypothesize that lack of sternohyoideus activity during intraoral prey processing, posterior pectoral girdle rotation, and a long‐fibered posterior intermandibularis muscle are novel structures associated with the tongue‐bite apparatus within osteoglossomorph fishes.