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Abstract Complex prey processing requires the repositioning of food between the teeth, as modulated by a soft tissue appendage like a tongue or lips. In this study, we trace the evolution of lips and ligaments, which are used during prey capture and prey processing in an herbivorous group of fishes. Pacus (Serrasalmidae) are Neotropical freshwater fishes that feed on leaves, fruits, and seeds. These prey are hard or tough, require high forces to fracture, contain abrasive or caustic elements, or deform considerably before failure. Pacus are gape‐limited and do not have the pharyngeal jaws many bony fishes use to dismantle and/or transport prey. Despite their gape limitation, pacus feed on prey larger than their mouths, relying on robust teeth and a hypertrophied lower lip for manipulation and breakdown of food. We used histology to compare the lip morphology across 14 species of pacus and piranhas to better understand this soft tissue. We found that frugivorous pacus have larger, more complex lips which are innervated and folded at their surface, while grazing species have callused, mucus‐covered lips. Unlike mammalian lips or tongues, pacu lips lack any intrinsic skeletal or smooth muscle. This implies that pacu lips lack dexterity; however, we found a novel connection to the primordial ligament which suggests that the lips are actuated by the jaw adductors. We propose that pacus combine hydraulic repositioning of prey inside the buccal cavity with direct oral manipulation, the latter using a combination of a morphologically heterodont dentition and compliant lips for reorienting food. 

Abstract Durophagous predators consume hard‐shelled prey such as bivalves, gastropods, and large crustaceans, typically by crushing the mineralized exoskeleton. This is costly from the point of view of the bite forces involved, handling times, and the stresses inflicted on the predator's skeleton. It is not uncommon for durophagous taxa to display an ontogenetic shift from softer to harder prey items, implying that it is relatively difficult for smaller animals to consume shelled prey. Batoid fishes (rays, skates, sawfishes, and guitarfishes) have independently evolved durophagy multiple times, despite the challenges associated with crushing prey harder than their own cartilaginous skeleton.Potamotrygon leopoldiis a durophagous freshwater ray endemic to the Xingu River in Brazil, with a jaw morphology superficially similar to its distant durophagous marine relatives, eagle rays (e.g.,Aetomylaeus, Aetobatus). We used second moment of area as a proxy for the ability to resist bending and analyzed the arrangement of the mineralized skeleton of the jaw ofP. leopoldiover ontogeny using data from computed tomography (CT) scans. The jaws ofP. leopoldido not resist bending nearly as well as other durophagous elasmobranchs, and the jaws are stiffest nearest the joints rather than beneath the dentition. While second moment has similar material distribution over ontogeny, mineralization of the jaws under the teeth increases with age. Neonate rays have low jaw stiffness and poor mineralization, suggesting thatP. leopoldimay not feed on hard‐shelled prey early in life. These differences in the shape, stiffness and mineralization of the jaws ofP. leopoldicompared to its durophagous relatives show there are several solutions to the problem of crushing shelled prey with a compliant skeleton.