palates and had large antorbital fenestrae, exchanging strength for lighter skull builds. The nostrils of Irritator were shifted far back from the snout tip; this, along with the secondary palate, which separated the animal's nasal passages from the inside of its mouth, made respiration possible even if most of the jaw was underwater or held prey. In particular, the sagittal crest of Irritator is an indication for pronounced neck musculature, which would have been necessary to close the jaws quickly against water resistance and withdraw the head rapidly. In 2015, German paleontologist Serjoscha W. Evers and colleagues found evidence for similar adaptations in the African spinosaur Sigilmassasaurus. The neck vertebrae of this genus have a heavily furrowed undersurface. This is consistent with the attachment of powerful neck muscles for use in fishing or rapidly snatching small prey, a trait also observed in extant crocodilians and birds. Sales and Schultz in 2017 found that Irritator and baryonychines might have relied more on their sense of smell for hunting than Spinosaurus did, since they had larger, less retracted nostrils and more room in their skulls for the nasal cavity. Spinosaurus itself probably made heavier use of senses like vision or the mechanoreceptors on the tip of its snout, like those used by crocodilians to sense prey moving in the water. Another trait spinosaurs shared with gharials was an enlarged snout tip that wielded a rosette-shaped array of interlocking teeth, adept for impaling and catching fish. Although to a lesser degree than most known spinosaurs, this feature was also present in the Angaturama limai holotype. In 2002, Sues and colleagues pointed out, however, that there would be no reason to assume that the Spinosauridae specialized completely in fishing. They stressed rather that this head morphology indicates a generalistic feeding, particularly on small prey animals. In fact, portions of a young Iguanodon, a terrestrial herbivore, were found inside the fossil skeleton of one Baryonyx. Naish and colleagues in 2004 supported the theory that Irritator hunted both aquatic and terrestrial animals as a generalist within the coastal area and in addition probably searched for carrion. A tooth belonging to Irritator was discovered still inserted into the fossil neck vertebral column of an ornithocheirid pterosaur, likely with a wingspan of . This indicates that Irritator ate pterosaurs as well, although it is not known if it actively hunted these animals or simply scavenged the remains. In