Irritator

Irritator is a genus of spinosaurid dinosaur that lived during the Early Cretaceous Period (Albian stage), around 110 million years ago. Current estimations indicate a length of 8 meters (26 feet) and a height of 3 meters (9 feet). It was found in Brazil. Irritator was a theropod with an unusually shaped crest at the rear of its head, and probably ate fish.

So far the only fossil that has been found was an 80 centimeter long fossil skull in the Romualdo Member, a layer member of the Brazilian Santana formation. This skull strongly resembles the skulls of Suchomimus and Spinosaurus. The genus is often regarded today as identical (synonymous) with Angaturama, which lived in the same time and the same place as Irritator.

Discovery
Irritator was first scientifically described in 1996 by paleontologists Martill, Cruikshank, Frey, Small and Clarke. Its only known fossil, an 80 cm skull discovered in eastern Brazil, was badly obscured by plaster which was added by the commercial fossil-collecting fossil-poachers who illegally sold it (the trade of fossils is prohibited by law in Brazil), in hopes of making the fossil look more complete and valuable. The buyers were not aware of the modifications to the illegally collected specimen, and it required them a great deal of work to reconstruct the original features — hence the name.

It is probably synonymous with Angaturama limai, another spinosaurid from the same time and place, whose remains curiously seem to complete Irritator's skull, meaning that they could belong to the same specimen.

The type species is I. challengeri. It honors the character of Professor Challenger in Arthur Conan Doyle's The Lost World.

Material
Material of I. challengeri, not counting that of A. limai, hails from the Romualdo Member of the Santana Formation in Brazil. The holotype is SMNS 58022, from the Stuttgart State Museum of the Natural Sciences, and it consists of an incomplete skull, lacking the anterior (front) portion.

The skull was recovered nearly complete and is considered the most complete head find of a spinosaurid. It is characterized particularly by its unusual length and curved lip region, which is strongly compressed laterally. The overall length of the complete head is estimated at approximately 84 centimeters. It possesses a clear Sagittal crest; such a comb is found also with some other dinosaurs. The teeth exhibit a single embedding of the strongly extended and straight teeth with conical tooth crowns, which indicates a continual tooth change, as new teeth were pushed up between the old ones. The teeth exhibit lengths from 6 to approximately 40 millimeters.

In 2004, parts of a spinal column were discovered in the Santana Formation. These have been  assigned, due to their structure, to the Spinosauridae. With very high probability these fossils belong to Irritator, since this is the so far the only so far well-known Spinosaurid of the formation.

Angaturama limai, from the same time and place as Irritator challengeri, was described by A. Kellner and Diogenes A. Campos in February 1996 on the basis of a fossil from the Santana formation. Today most sources consider Angaturama a synonym of Irritator. Some scientists have even speculated that the two partial fossil skulls come from the same individual. Waiters and Campos (2000) as well as Machado and Waiters (2005) hold however the opinion that the fossils come from two different genera and that Angaturama had a clearly higher and laterally more flattened head than Irritator. If Angaturama and Irritator are actually regarded as a member of the same genus, Irritator challengeri would be the valid scientific name under the priority rules.

The fossil consists only of the front part of the head, which is characterized by the fact that it is very narrow and carries a premaxillary sagittal crest. In the premaxilla a broken-off tooth with partial tooth crown was recovered which corresponds to that of Irritator. Altogether the premaxilla had seven teeth; the third tooth was the largest. The fossil is kept today under the number USP GP/2T-5 in the Universidade de São Paulo.

Systematics
I. challengeri was a member of the Spinosauridae, more specifically the subfamily Spinosaurinae. It shares a close relationship with Spinosaurus and possibly Siamosaurus, though this last genus is not well-known from fossil material.

Irritator was originally described as a Maniraptoran within the Tetanurae. It was assigned to the family Baryonychidae, along with Angaturama, Baryonyx, Suchomimus and Spinosaurus by Oliver Rauhut in 2003. Holtz et al. (2004) considered the Baryonychidae synonymous with the family Spinosauridae, and placed these genera within that family. Most later revisions have upheld these classifications.

Paleobiology
The exact discovery site of the Irritator fossil skull is unknown; the fossil was collected by illegal fossil dealers. The skull and the matrix have been assigned to a rock of the Romualdo Member of the Santana formation due to lithological characteristics; this classification was confirmed by microfossils of the ostracod Pattersoncypris. A questioning of the local fossil dealers resulted in the identification of the discovery site near the village of Buxexé close Santana Do Cariri at the flank of the Chapada do Araripe, at a height of approximately 650 meters. Since rock from the Romualdo Member is indeed exposed there, this discovery site is regarded as very probable for the fossil.

The Romualdo Member of the Santana formation is generally assigned to the Albian and thus the last section of the Early Cretaceous. The layers are aged to approximately 110 million years and to have come from a time in which the continents of Africa and South America were still connected with one another in the northern part of Brazil.

The horizon of the Santana formation, in which both fossils were found, resulted with very high probability from sedimentation in a flat lake, which was filled with fresh or brackish water. The fossil finds made so far create an ambivalent picture. The fossil insects which have been recovered are an indication for fresh water; the find of the turtle Santanachelys, which was adapted to seawaters, indicate a saltwater environment. One theory is that the site was a brackish lagoon, which was connected to the sea. The climate was tropical and corresponded to today's climate in Brazil to a large extent.

Irritator probably nourished itself on fish, like the pterosaurs found in large number in the Santana formation. Irritator was probably, like today's crocodiles, a food generalist, eating all other animals that it could catch besides fish. A tooth belonging to Irritator still inserted into a fossil neck vertebral column of a pterosaur, indicates that Irritator ate pterosaurs as well, although it is not known if it actively hunted these animals, or simply scavenged the remains.

All Spinosaurids had very narrow jaws with relatively homogeneous pointed teeth. This arrangement is particularly found in crocodiles such as the Sunda Gavial. The long conical teeth, which do not possess serrated edges, are suitable particularly to grabbing and holding of prey. They differ from teeth of carnivores, which must tear or cut off the seized body parts. Particularly with Irritator and Suchomimus a convergence with crocodiles is regularly discussed in the literature. Individual fossils belonging to the Spinosauridae were regarded in the past as crocodile fossils. For example, Baryonyx fossils from Portugal were originally described as Suchosaurus and only in 2007 were the recognized as those of a spinosaurid.

The nostrils of Irritator were shifted far to the rear of the skull, and the secondary palate make respiration possible even if the majority of the jaw was under water or held prey. In particular, the sagittal crest of Irritator is an indication for a pronounced neck musculature, which would have been necessary in order to pull the jaw closed quickly against water resistance and withdraw the head fast. Sues et al. (2002) point out, however, that there would be no reason to assume that the Spinosauridae specialized completely in fishing. They stress rather that this head morphology indicates a generalistic feeding, particularly on small prey animals. In fact, portions of a young Iguanodon, a landliving herbivore, were found inside the fossil skeleton of one Baryonyx. Naish et al. (2004) support the theory that Irritator hunted both aquatic and terrestrial animals as a generalist within the coastal area and in addition probably also searched for carrion.