Thalassodromeus is a genus of pterosaur that lived in what is now Brazil during the Early Cretaceous period, about a hundred million years ago. The original skull, discovered in 1983 in the Araripe Basin of northeastern Brazil, was collected in several pieces. In 2002, the skull was made the holotype specimen of Thalassodromeus sethi by palaeontologists Alexander Kellner and Diogenes de Almeida Campos. The generic name means "sea runner" (in reference to its supposed mode of feeding), and the specific name refers to the Egyptian god Seth due to its crest being supposedly reminiscent of Seth's crown. Other scholars have pointed out that the crest was instead similar to the crown of Amon. A jaw tip was assigned to T. sethi in 2005, became the basis of the new genus Banguela in 2015, and assigned back to Thalassodromeus as the species T. oberlii in 2018, though other researchers consider it a valid genus. Another species (T. sebesensis) was described in 2015 based on a supposed crest fragment, but this was later shown to be part of a turtle shell. Thalassodromeus had one of the largest known skulls among pterosaurs, around 1.42 m (4 ft 8 in) long, with one of the proportionally largest cranial crests of any vertebrate. Though only the skull is known, the animal is estimated to have had a wingspan of 4.2 to 4.5 m (14 to 15 ft). The crest was lightly built and ran from the tip of the upper jaw to beyond the back of the skull, ending in a unique V-shaped notch. The jaws were toothless, and had sharp upper and lower edges. Its skull had large nasoantorbital fenestrae (opening that combined the antorbital fenestra in front of the eye with the bony nostril), and part of its palate was concave. The lower jaw was blade-like, and may have turned slightly upwards. The closest relative of Thalassodromeus was Tupuxuara; both are grouped in a clade called Thalassodromidae, which, depending on the study, has been placed either within Tapejaromorpha, closely related to the family Tapejaridae, or within Neoazhdarchia, closely related to the family Dsungaripteridae. Several theories have been suggested to explain the function of Thalassodromeus's crest, including thermoregulation and display, but it likely had more than one function. The crests of thalassodromids appear to have developed late in growth (probably correlated with sexual maturity) and they may have been sexually dimorphic (differing according to sex). As the genus name implies, Thalassodromeus was originally proposed to have fed like a modern skimmer bird, by skimming over the water's surface and dipping its lower jaws to catch prey. This idea was later criticised for lack of evidence; Thalassodromeus has since been found to have had strong jaw musculature, and may have been able to kill and eat relatively large prey on the ground. The limb proportions of related species indicate that it may have adapted to fly in inland settings, and would have been efficient at moving on the ground. Thalassodromeus is known from the Romualdo Formation, where it coexisted with many other types of pterosaurs, dinosaurs and other animals.
History of discovery
The first known specimen of this pterosaur (an extinct order of flying reptiles) was collected in 1983 near the town of Santana do Cariri in the Araripe Basin of northeastern Brazil. Found in outcrops of the Romualdo Formation, it was collected over a long period in several pieces. The specimen (catalogued as DGM 1476-R at the Museu de Ciências da Terra) was preserved in a calcareous nodule, and consists of an almost-complete, three-dimensional skull (pterosaur bones are often flattened compression fossils), missing two segments of the bottom of the skull and mandible and the front of the lower jaw. The left jugal region and right mandibular ramus (half of the mandible) are pushed slightly inward. The skull was first reported in a 1984 Italian book, and preliminarily described and figured in 1990 by palaeontologists Alexander Kellner and Diogenes de Almeida Campos. Although the pieces of skull had been divided between museums in South and North America, they were assembled before 2002. In 2002, Kellner and Campos described and named the new genus and species Thalassodromeus sethi, skull DGM 1476-R being the holotype specimen. The generic name is derived from the Ancient Greek words θάλασσα, thálassa, '"sea"' and δρομεύς, dromeús, '"runner"', meaning "sea runner" in reference to the animal's supposed skim-feeding behaviour. The specific name refers to the Egyptian god Seth. The specimen was not fully prepared at the time of this preliminary description. The original describers chose the name sethi because the crest of this pterosaur was supposedly reminiscent of the crown worn by Seth, but the palaeontologists André Jacques Veldmeijer, Marco Signore, and Hanneke J. M. Meijer pointed out in 2005 that the crown (with its two tall plumes) was typically worn by the god Amon (or Amon-Ra) and his manifestations – not by Seth.
In 2006, palaeontologists David M. Martill and Darren Naish suggested that Thalassodromeus was a junior synonym of the related genus Tupuxuara, which was named by Kellner and Campos in 1988 based on fossils from the same formation. In the view of Martill and Naish, the differences between these genera (including two species of Tupuxuara, T. longicristatus and T. leonardii) were due to ontogeny (changes during growth) and compression of the fossils; Thalassodromeus was simply an older, larger, and better-preserved individual. This idea was rejected by Kellner and Campos in 2007, who pointed out these species had differences in features other than their crests. They also noted that one specimen of Tupuxuara had a larger skull than Thalassodromeus (measured from the tip of the premaxilla to the back of the squamosal bone), despite Martill and Naish's contention that the latter was an older individual. Kellner and Campos' view has since been accepted by other researchers, including Martill and Naish.
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