page_title: Irritator
section_title: Classification
hierarchical_section_title: Irritator / Classification
caption_attribution_description: English: A, Premaxillae and maxillae of Baryonyx walkeri. B, Partial rostrum (MNN GDF501) referred to Suchomimus tenerensis. C, Specimen MSNM V4047. D, Reconstructed skull of Irritator challengeri. All specimens are aligned based on the position of the fourth maxillary tooth, whereas the last maxillary tooth (or alveolus) in each is indicated by a black star. Abbreviations for teeth follow Hendrickx et al. [58]. Additional abbreviations: n.c, nasal contact; ao.f, antorbital fenestra; e.n, external naris.
context_page_description: Irritator is a genus of spinosaurid dinosaur that lived in what is now Brazil during the Albian stage of the Early Cretaceous Period, about 110 million years ago. It is known from a nearly complete skull found in the Romualdo Formation of the Araripe Basin. Fossil dealers had acquired this skull and illegally sold it to the State Museum of Natural History Stuttgart. In 1996, the specimen became the holotype of the type species Irritator challengeri. The genus name comes from the word "irritation", reflecting the feelings of paleontologists who found the skull had been heavily damaged and altered by the collectors. The species name is an homage to the fictional character Professor Challenger from Arthur Conan Doyle's novels.
  Some paleontologists regard Angaturama limai—known from a snout tip that was described later in 1996—as a potential junior synonym of Irritator. Both animals hail from the same stratigraphic units of the Araripe Basin. It was also previously proposed that Irritator and Angaturama's skull parts belonged to the same specimen. Although this has been cast into doubt, more overlapping fossil material is needed to confirm whether they are the same animal or not.
context_section_description: Martill and his team originally classified Irritator as a maniraptoran dinosaur in the clade Bullatosauria (a group no longer considered monophyletic), as a close relative to the feathered ornithomimosaurs and troodontids. Given that its dental morphology, particularly long snout, and assumed fin-shaped crest were features unknown in "other" maniraptorans, the researchers erected the new family Irritatoridae within the clade. They recognized Irritator's affinities with Spinosaurus, in that they both had similarly shaped and unserrated teeth, but noted that the latter's mandible would not conform with the front of Irritator's upper jaw, and that other non-avian dinosaurs like Compsognathus and Ornitholestes also bore no serrations on some or all of their teeth. Some of these claims were questioned in 1996 by Kellner who found that Irritator's skull lacked the one autapomorphy (distinguishing feature) diagnosed in maniraptorans at the time, which was having its jugal (cheek) bone forming part of the antorbital fenestra. He also pointed out that since Irritator challengeri's holotype lacked the tip of its snout, it would not be possible to know if Spinosaurus's dentary could complement it or not. Based on comparisons with Spinosaurus, Kellner resolved Irritator as a spinosaurid and synonymized Irritatoridae with that family. Irritator was then assigned to the Baryonychidae along with Angaturama, Baryonyx, Suchomimus, and Spinosaurus by Oliver W.M. Rauhut in 2003. Thomas Holtz and colleagues in 2004 considered the Baryonychidae synonymous with Spinosauridae, and moved these genera to the latter family. Most later revisions have upheld these classifications. As spinosaurids, Irritator and Angaturama are placed within the superfamily Megalosauroidea, with Spinosauridae being a possible sister taxon to the Megalosauridae.
  In 1998, Sereno and colleagues defined two subfamilies within the Spinosauridae based on craniodental (skull and tooth) characteristics. They were Spinosaurinae, where they placed Spinosaurus and Irritator; and Baryonychinae, to which they assigned Baryonyx, Suchomimus, and Cristatusaurus. Spinosaurines were distinguished by their unserrated, straighter, and more widely spaced teeth, as well as smaller first teeth of the premaxilla. In 2005, Dal Sasso and colleagues assumed Irritator's nostrils as being located above the middle of the maxillary tooth row; more posteriorly than in baryonychines, but less so than in Spinosaurus. Sales and Schultz in 2017 found that Irritator's 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 limai's 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, Angaturama's 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)
caption: Diagram illustrating skull elements from Irritator (bottom) and other spinosaurids, comparing the relative positions of the external nares, or bony nostrils, (labeled e.n.)
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