surface to travel long distances without flapping, and without the aid of thermals (which do not occur over the open ocean the same way they do over land). While most of a Pteranodon flight would have depended on soaring, like long-winged seabirds, it probably required an occasional active, rapid burst of flapping, and studies of Pteranodon wing loading (the strength of the wings vs. the weight of the body) indicate that they were capable of substantial flapping flight, contrary to some earlier suggestions that they were so big they could only glide. However, a more recent study suggests that it relied on thermal soaring, unlike modern seabirds but much like modern continental flyers and the extinct Pelagornis. Like other pterosaurs, Pteranodon probably took off from a standing, quadrupedal position. Using their long forelimbs for leverage, they would have vaulted themselves into the air in a rapid leap. Almost all of the energy would have been generated by the forelimbs. The upstroke of the wings would have occurred when the animal cleared the ground followed by a rapid down-stroke to generate additional lift and complete the launch into the air. Terrestrial locomotion Historically, the terrestrial locomotion of Pteranodon, especially whether it was bipedal or quadrupedal, has been the subject of debate. Today, most pterosaur researchers agree that pterosaurs were quadrupedal, thanks largely to the discovery of pterosaur trackways. The possibility of aquatic locomotion via swimming has been discussed briefly in several papers (Bennett 2001, 1994, and Bramwell & Whitfield 1974). Diet The diet of Pteranodon is known to have included fish; fossilized fish bones have been found in the stomach area of one Pteranodon, and a fossilized fish bolus has been found between the jaws of another Pteranodon, specimen AMNH 5098. Numerous other specimens also preserve fragments of fish scales and vertebrae near the torso, indicating that fish made up a majority of the diet of Pteranodon (though they may also have taken invertebrates). Traditionally, most researchers have suggested that Pteranodon would have taken fish by dipping their beaks into the water while in low, soaring flight. However, this was probably based on the assumption that the animals could not take off from the water surface. It is more likely that Pteranodon could take off from the water, and would have dipped for fish while swimming rather than while flying. Even a small, female Pteranodon could have reached a depth of