Ирритатор: Бразилиялық спинозаврид динозавры

Spinosaurid theropod dinosaur genus from the Early Cretaceous Period

Irritator is a genus of spinosaurid dinosaur that lived in what is now Brazil during the Albian stage of the Early Cretaceous Period, about 113 to 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 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 a 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 Angaturamas 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. Other spinosaurid skeletal material, some of which could belong to Irritator or Angaturama, was retrieved from the Romualdo Formation, allowing for a replica skeleton to be made and mounted for display at the National Museum of Rio de Janeiro in 2009. Estimated at between in length, Irritator weighed around , making it one of the smallest spinosaurids known. Its long, shallow and slender snout was lined with straight and unserrated conical teeth. Lengthwise atop the head ran a thin sagittal crest, to which powerful neck muscles were likely anchored. The nostrils were positioned far back from the tip of the snout, and a rigid secondary palate on the roof of the mouth would have strengthened the jaw when feeding. Belonging to a subadult, Irritator challengeris holotype remains the most completely preserved spinosaurid skull yet found. The Angaturama snout tip expanded to the sides in a rosette like shape, bearing long teeth and an unusually tall crest. One possible skeleton indicates it, like other spinosaurids, had enlarged first finger claws and a sail running down its back. Irritator had been mistaken initially for a pterosaur, and later a maniraptoran dinosaur. In 1996, the animal was identified as a spinosaurid theropod. The holotype skull was thoroughly prepared before being redescribed in 2002, confirming this classification. Both Irritator and Angaturama belong to the Spinosaurinae subfamily. A generalist diet—like that of today's crocodilians—has been suggested; Irritator might have preyed mainly on fish and any other small prey animals it could catch. Fossil evidence is known of an individual that ate a pterosaur, either from hunting or scavenging it. Irritator may have had semiaquatic habits, and inhabited the tropical environment of a coastal lagoon surrounded by dry regions. It coexisted with other carnivorous theropods as well as turtles, crocodyliforms, and a large number of pterosaur and fish species.

The holotype of Irritator was excavated from a chalk concretion containing the rear of a large skull with lower jaws near the town of Santana do Cariri in northeastern Brazil. This fossil was acquired by dealers who sold it to Rupert Wild of the State Museum of Natural History Stuttgart, Germany.]] The skull was flattened sideways somewhat and, as is common with fossils, was partly crushed. The right side was well preserved, while the left was extensively damaged during collection. Some of the skull's hindmost upper surface had eroded, and the lower jaw lacked its front end, both owing to breakage during fossilization. Parts of the specimen were also cracked due to being part of a septarian concretion. The tip of the upper jaw was also missing. Since there were no signs of erosion, it had most probably broken off during or after the fossil's collection. Evident corrosion on certain bones indicates acid preparation had been attempted. A vertical fracture was present across the middle of the skull, which had apparently been sealed with car body filler. In hopes of making it look more complete and valuable, the fossil traders had severely obscured the skull beneath plaster; The buyers were unaware of the modifications to the specimen until it was sent to universities in the United Kingdom for CT scan imaging. This revealed the collectors had tried to reconstruct the skull by grafting parts of the maxilla (main upper jaw bone) onto the front of the rostrum (snout). When Martill and colleagues first described Irritator challengeri, the holotype was still extensively encased in calcareous rock matrix. Under the supervision of American paleontologist Hans Dieter Sues, technician Diane M. Scott from the University of Toronto at Mississauga assumed the task of fully extracting the skull bones from the rock, allowing for a detailed redescription in 2002. Published by Sues, Frey, Martill, and Scott, this inspection of the now fully prepared specimen negated many of Martill and colleagues' original observations, which were based on misinterpretations of the damaged and largely concealed skull. The estimated length of the complete skull was shorter than previously proposed. What was originally thought to be a prominent head crest proved to be an unattached, indeterminate bone fragment. As in the previous study, Sues and colleagues regarded the African genus Spinosaurus as the most similar taxon to Irritator, because they shared many dental features, including mostly straight conical tooth crowns, thin enamel, well defined edges with no , and lengthwise fluting. Since little was known of Spinosauruss skull at the time, these similarities were enough for the authors to suggest a possible junior synonymy of Irritator with Spinosaurus. Sues and colleagues noted that more overlapping skull material was needed for further diagnosis. Although the site of discovery is uncertain, the specimen most probably stems from the Romualdo Formation (previously designated the Romualdo Member of the then Santana Formation). In 1997, British paleontologists Alan J. Charig and Angela C. Milner considered Angaturama a likely junior synonym of Irritator, noting that both genera had retracted nostrils, long jaws, and characteristic spinosaurid dentition. Paul Sereno and colleagues in 1998 agreed with this possibility, and additionally observed that the holotype of Angaturama seems to complete that of Irritator (meaning that they could belong to the same specimen). Authors including Éric Buffetaut and Mohamed Ouaja in 2002, Cristiano Dal Sasso and colleagues in 2005, Darren Naish in 2013, In their redescription of Irritator, Sues and colleagues pointed out that both holotypes are equally as narrow, and share transversely round teeth with defined yet unserrated edges. They also noted that a sagittal crest on Angaturamas premaxillae may correspond with that of Irritators nasal bones. A review of both fossils by the Brazilian paleontologists Marcos A. F. Sales and Cesar L. Schultz in 2017 noted that the specimens also differ in other aspects of their preservation: the Irritator specimen is brighter in color and is affected by a vertical crack, while the Angaturama specimen bears many cavities; the damage to the teeth of the Irritator challengeri holotype is also much less severe. Sales and Schultz also identified a possible point of overlap, the third left maxillary tooth, and observed that the skull of Angaturama could have been larger than that of Irritator based on the proportions of the closely related genus Baryonyx. They therefore concluded that the two specimens do not belong to the same individual, Sales and Schultz noted that synonymy at the genus level would need to be verified by more extensively overlapping remains. If Angaturama and Irritator are regarded as a member of the same genus, the latter would be the valid scientific name under rules of priority, since it was named almost a month earlier.

Besides the skull, the snout fragment, and some isolated teeth, the Romualdo Formation has also yielded l remains that may belong to spinosaurids, many of which are hitherto undescribed, and all of them pertaining to the Spinosaurinae subfamily. In 2004, parts of a spinal column (MN 4743 V) were unearthed at the formation. Brazilian paleontologist Jonathas Bittencourt and Kellner assigned these, due to their structure, to the Spinosauridae. It is uncertain whether this specimen can be referred to Irritator or Angaturama, given that both are based only on skull material. In 2007, Machado and Kellner tentatively referred a rib fragment (MN 7021 V) to the Spinosauridae. The skeleton was fully described in 2010 in an as of yet unpublished master's thesis by Machado. In 2018, Tito Aureliano and his team described LPP PV 0042, part of a left tibia from a particularly large individual. Some of the Romualdo Formation postcrania were used as the basis for the creation of a replica Angaturama skeleton, later mounted at the Federal University owned Museu Nacional do Rio de Janeiro (National Museum of Rio de Janeiro). The skeleton depicted the animal carrying an anhanguerid pterosaur in its jaws. In press releases of the exhibit's opening, Kellner informally implied MN 4819 V as belonging to Angaturama. In September 2018, a fire broke out at the palace housing the Museu Nacional, largely destroying the fossil collections and possibly the exhibited Angaturama skeleton and fossil elements. The holotype of Oxalaia quilombensis, which was stored in the same building, may also have been destroyed.

The taphonomy (changes between death and fossilization) of the Irritator challengeri holotype specimen has been discussed by some researchers. The skull was found lying on its side. Preceding fossilization, several bones from the back of the braincase, as well as the dentary, splenial, coronoid, and right angular bones from the lower jaw, were lost. Other bones, mostly from the skull rear, had become and displaced towards alternate regions of the head before burial. Naish and colleagues in 2004 asserted that the Romualdo Formation dinosaur fauna is represented by animals that died near shorelines or rivers before being carried out to sea, where their floating remains were eventually fossilized. In 2018, Aureliano and colleagues argued against this scenario, stating that the Irritator challengeri holotype's mandible was preserved in articulation with the rest of the skull, whereas it would have likely detached had the carcass been floating at sea. They also noted that the corpse would have quickly sunk due to the osteosclerosis of the skeleton. The researchers, therefore, concluded that fossils from the Santana Group represent organisms that were buried in their natural habitat, instead of having been deposited allochthonously (other than at their present position).