Evolution and ecology of feeding in elasmobranchs.

Paleozoic chondrichthyans had a large gape, numerous spike-like teeth, limited cranial kinesis, and a non-suspensory hyoid, suggesting a feeding mechanism dominated by bite and ram. Modern sharks are characterized by a mobile upper jaw braced by a suspensory hyoid arch that is highly kinetic. In batoids, the upper jaw is dissociated from the cranium permitting extensive protrusion of the jaws. Similar to actinopterygians, the evolution of highly mobile mandibular and hyoid elements has been correlated with extensive radiation of feeding modes in elasmobranchs, particularly that of suction. Modern elasmobranchs possess a remarkable variety of feeding modes for a group containing so few species. Biting, suction or filter-feeding may be used in conjunction with ram to capture prey, with most species able to use a combination of behaviors during a strike. Suction-feeding has repeatedly arisen within all recent major elasmobranch clades and is associated with a suite of morphological and behavioral specializations. Prey capture in a diverse assemblage of purported suction-feeding elasmobranchs is investigated in this study. Drop in water pressure measured in the mouth and at the location of the prey shows that suction inflow drops off rapidly with distance from the predator's mouth. Elasmobranchs specializing in suction-feeding may be limited to bottom associated prey and because of their small gape may have a diet restricted to relatively small prey. Behavior can affect performance and overcome constraints imposed by the fluid medium. Suction performance can be enhanced by proximity to a substrate or by decreasing distance from predator to prey using various morphological and/or behavioral characteristics. Benthic suction-feeders benefit by the increased strike radius due to deflection of water flow when feeding close to a substrate, and perhaps require less accuracy when capturing prey. Suction and ram-suction-feeding elasmobranchs can also use suction inflow to draw prey to them from a short distance, while ram-feeding sharks must accelerate and overtake the prey. The relationship between feeding strategy and ecology may depend in part on ecological, mechanistic or evolutionary specialization. Mechanistic suction-feeding specialist elasmobranchs are primarily benthic, while most epibenthic and pelagic elasmobranchs are generalists and use ram, suction, and biting to catch a diversity of prey in various habitats. Some shark species are considered to be ecological specialists in choosing certain kinds of prey over others. Batoids are evolutionary specialists in having a flattened morphology and most are generalist feeders. Filter-feeding elasmobranchs are ecological, mechanistic, and evolutionary specialists.