Durophagous biting in sea otters (Enhydra lutris) differs kinematically from raptorial biting of other marine mammals.

Sea otters represent an interesting model for studies of mammalian feeding evolution. Although they are marine mammals, sea otters returned to the sea relatively recently and feed at the surface. Therefore, they represent a transitional stage of aquatic adaptation. Currently no feeding performance studies of sea otters have been conducted. The main objective of this study was to characterize the feeding kinematic profile in sea otters. It was hypothesized that sea otters would exhibit a terrestrial feeding behavior and that they forcefully crush hard prey at large gapes. As a result, biting kinematics would be congruent with biting behavior reported for their terrestrial ancestors, thus providing additional evidence that raptorial biting is a conserved behavior even in recently aquatic mammals. Sea otters consistently used a durophagous raptorial biting mode characterized by large gapes, large gape angles and lack of lateral gape occlusion. The shorter skulls and mandibles of sea otters, along with increased mechanical advantages of the masseter and increased bite force, form a repertoire of functional traits for durophagy. Here we consider durophagy to be a specialized raptorial biting feeding mode. A comparison of feeding kinematics of wild versus captive sea otters showed no significant differences in lateral kinematic profiles, and only minor differences in three frontal kinematic profiles, which included a slower maximum opening gape velocity, a slower maximum gape opening velocity, and a slower maximum closing gape velocity in captive sea otters. Data indicate functional innovations for producing large bite forces at wide gape and gape angles.

Divergent Skull Morphology Supports Two Trophic Specializations in Otters (Lutrinae).

Variation in terrestrial mammalian skull morphology is known to constrain feeding performance, which in turn influences dietary habits and ultimately fitness. Among mustelids, otters have evolved two feeding specializations: underwater raptorial capture of prey (mouth-oriented) and capture of prey by hand (hand-oriented), both of which have likely associations with morphology and bite performance. However, feeding biomechanics and performance data for otters are sparse. The first goal of this study was to investigate the relationships between feeding morphology and bite performance among two mouth-oriented piscivores (Pteronura brasiliensis and Lontra canadensis) and two hand-oriented invertebrate specialists (Enhydra lutris and Aonyx cinerea). Since other vertebrate taxa that are mouth-oriented piscivores tend to possess longer skulls and mandibles, with jaws designed for increased velocity at the expense of biting capability, we hypothesized that mouth-oriented otters would also possess long, narrow skulls indicative of high velocity jaws. Conversely, hand-oriented otters were expected to possess short, blunt skulls with adaptations to increase bite force and crushing capability. Concomitant with these skull shapes we hypothesized that sea otters would possess a greater mandibular bluntness index, providing for a greater mechanical advantage compared to other otter species investigated. A second goal was to examine morphological variation at a finer scale by assessing variation in cranial morphology among three sea otter subspecies. Since diet varies among these subspecies, and their populations are isolated, we hypothesized that the magnitude of mandibular bluntness and concomitant mechanical advantage, as well as occlusal surface area would also vary within species according to their primary food source (fish versus hard invertebrates). Functional expectations were met for comparisons among and within species. Among species the phylogeny suggests a deeply rooted transition to alternative foraging types. Yet within foraging types alternative species were also strongly variable, suggesting either selective differences in the extent or nature of realized foraging mode, or an accumulation of non-adaptive changes during the long independent evolutionary history. At the finest scale, variation among subspecies indicates that trophic adaptation occurred rapidly, making it interesting that we happened to find both deeply and shallowly-rooted transformations associated with diet type in otter species and subspecies.