Biomechanical energetics of terrestrial locomotion in California sea lions (Zalophus californianus).

Pinnipedia, an order of semi-aquatic marine mammals, adapted a body design that allows for efficient aquatic locomotion but limited terrestrial locomotion. Otariids, like the California sea lion (Zalophus californianus), have enlarged forelimbs and can bring their hindlimbs under the body to locomote quadrupedally on land, but phocids (true seals) have reduced forelimbs and are unable to bring their hindlimbs beneath them during terrestrial locomotion. Because of these differences, phocids are expected to have greater energetic costs when moving on land compared with otariids. The mechanical costs of transport (COT) and power outputs of terrestrial locomotion were first obtained from one male and two female adult California sea lions through video recording locomotion sequences across a level runway. The center of mass, along with six other anatomical points, were digitized to obtain variables such as velocity (V), amplitude of heave (A) and the frequency (f) of oscillations during the locomotion cycle. These variables represent the principal parameters of a biomechanical model that computes the power output of individuals. The three California sea lions in this study averaged a power output of 112.04 W and a COT of 0.63 J kg-1 m-1. This footage was compared against video footage previously recorded of three phocid species (harbor seal, gray seal and northern elephant seal). Power output and mechanical COT were compared for all four pinniped species by tracking the animals' center of mass. The quadrupedal gait of sea lions showed lower vertical displacements of the center of mass, and higher velocities compared with the terrestrial gait of phocids. Northern elephant seals, gray seals and harbor seals showed significantly higher COT and power outputs than the sea lions. California sea lions locomote with lower energetic costs, and thus higher efficiency compared with phocids, proving that they are a mechanically intermediate species on land between terrestrial mammals and phocids. This study provides novel information on the mechanical energy exerted by pinnipeds, particularly California sea lions, to then be used in future research to better understand the limitations of these aquatic mammals.

The role of California sea lion (Zalophus californianus) hindflippers as aquatic control surfaces for maneuverability.

California sea lions (Zalophus californianus) are a highly maneuverable species of marine mammal. During uninterrupted, rectilinear swimming, sea lions oscillate their foreflippers to propel themselves forward without aid from the collapsed hindflippers, which are passively trailed. During maneuvers such as turning and leaping (porpoising), the hindflippers are spread into a delta-wing configuration. There is little information defining the role of otarrid hindflippers as aquatic control surfaces. To examine Z. californianus hindflippers during maneuvering, trained sea lions were video recorded underwater through viewing windows performing porpoising behaviors and banking turns. Porpoising by a trained sea lion was compared with sea lions executing the maneuver in the wild. Anatomical points of reference (ankle and hindflipper tip) were digitized from videos to analyze various performance metrics and define the use of the hindflippers. During a porpoising bout, the hindflippers were considered to generate lift when surfacing with a mean angle of attack of 14.6±6.3 deg. However, while performing banked 180 deg turns, the mean angle of attack of the hindflippers was 28.3±7.3 deg, and greater by another 8-12 deg for the maximum 20% of cases. The delta-wing morphology of the hindflippers may be advantageous at high angles of attack to prevent stalling during high-performance maneuvers. Lift generated by the delta-shaped hindflippers, in concert with their position far from the center of gravity, would make these appendages effective aquatic control surfaces for executing rapid turning maneuvers.