found that Irritators nostrils were in fact positioned closer to the front of the jaw, as in Baryonyx and Suchomimus; this more forward nostril placement was typically considered characteristic of baryonychines. Nevertheless, Irritator also bore unserrated teeth, a trait associated with spinosaurines. Sales and Schultz thus noted that the Araripe Basin spinosaurids Irritator and Angaturama might represent intermediate forms between the earlier baryonychines and later spinosaurines, and that further research may eventually render the former a paraphyletic (unnatural) group. Irritator is further distinguished from Baryonyx, Suchomimus, and Cristatusaurus by having slightly over half as many teeth in the maxilla, and from Spinosaurus due to its comparatively larger and more forwardly positioned nostril openings, which, unlike in Spinosaurus, are also formed by the premaxilla. The narrow sagittal crest, which ends in a knob-like process above the frontals, is another autapomorphy separating Irritator from other spinosaurids. Although Angaturama limais snout is generally narrower than in other spinosaurids, this might be due to damage sustained by the fossil; the holotype appears partly crushed and broken on its lower margin, with some of the preserved teeth having been sectioned off along their length. Therefore, Angaturamas only valid autapomorphy is its sagittal crest, which extends farther forwards on the rostrum and is more exaggerated than in other known spinosaurid skulls. Topology A: Benson and colleagues (2009) Topology B: Sales and Schultz (2017) Paleobiology Diet and feeding In 1996, Martill and colleagues theorized that Irritator challengeri, with its elongated snout and unserrated conical teeth, likely had at least a partly piscivorous (fish-eating) diet. Although much of the holotype's morphology turned out to be greatly different than they thought, later studies supported these observations. Spinosaurids had very narrow and elongated jaws with relatively homogeneous pointed teeth, an arrangement found particularly in animals like the Indian gharial—the most piscivorous extant crocodilian. The long conical teeth, which in spinosaurines did not possess serrated edges, were suitable for grabbing and holding prey. They differed from teeth of other theropods, which seemed geared towards tearing or cutting off seized body parts. Irritator shared with crocodilians a stiff secondary palate and reduced antorbital fenestrae. In 2007, a finite-element analysis study by British paleontologist Emily J. Rayfield and colleagues found that these attributes, present in other spinosaurids as well, made the skull more resistant to torsion from prey item loads when feeding. The authors pointed out that in contrast, most theropods lacked secondary