The scaling of locomotor performance in predator-prey encounters: from fish to killer whales.

During predator-prey encounters, a high locomotor performance in unsteady manoeuvres (i.e. acceleration, turning) is desirable for both predators and prey. While speed increases with size in fish and other aquatic vertebrates in continuous swimming, the speed achieved within a given time, a relevant parameter in predator-prey encounters, is size independent. In addition, most parameters indicating high performance in unsteady swimming decrease with size. Both theoretical considerations and data on acceleration suggest a decrease with body size. Small turning radii and high turning rates are indices of maneuverability in space and in time, respectively. Maneuverability decreases with body length, as minimum turning radii and maximum turning rates increase and decrease with body length, respectively. In addition, the scaling of linear performance in fish locomotion may be modulated by turning behaviour, which is an essential component of the escape response. In angelfish, for example, the speed of large fish is inversely related to their turning angle, i.e. fish escaping at large turning angles show lower speed than fish escaping at small turning angles. The scaling of unsteady locomotor performance makes it difficult for large aquatic vertebrates to capture elusive prey by using whole-body attacks, since the overall maneuverability and acceleration of small prey is likely to be superior to that of large predators. Feeding strategies in vertebrate predators can be related to the predator-prey length ratios. At prey-predator ratios higher than approximately 10(-2), vertebrate predators are particulate feeders, while at smaller ratios, they tend to be filter feeders. At intermediate ratios, large aquatic predators may use a variety of feeding methods that aid, or do not involve, whole body attacks. Among these are bubble curtains used by humpback whales to trap fish schools, and tail-slapping of fish by delphinids. Tail slapping by killer whales is discussed as an example of these strategies. The speed and acceleration achieved by the flukes of killer whales during tail slaps are higher and comparable, respectively, to those that can be expected in their prey, making tail-slapping an effective predator behaviour.