Tupuxuara Temporal range: Albian, | |
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Reconstructed skeleton of T. leonardii, American Museum of Natural History | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Order: | † Pterosauria |
Suborder: | † Pterodactyloidea |
Subfamily: | † Thalassodrominae |
Genus: | † Tupuxuara Kellner & Campos, 1988 |
Type species | |
†Tupuxuara longicristatus Kellner & Campos, 1988 | |
Other species | |
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Tupuxuara is a genus of large, crested, and toothless pterodactyloid pterosaur from the Early Cretaceous period (Albian stage) of what is now the Romualdo Formation of the Santana Group, Brazil, about 125 to 112 million years ago. Tupuxuara is a close relative of Thalassodromeus , and both form a group that is either called Thalassodrominae (if placed within the family Tapejaridae) or Thalassodromidae (if placed within the clade Neoazhdarchia).
The genus was named and described by Alexander Kellner and Diógenes de Almeida Campos in 1988. [1] The type species is Tupuxuara longicristatus. The generic name refers to a familiar spirit from the mythology of the Tupi. The specific name means "long-crested" in Latin.
The holotype, MN 6591-V, was found in the Early Cretaceous (Albian) Santana Formation (now known as the Romualdo Formation) of Brazil. It consists of a snout and some partial wing bones. Mature individuals of T. longicristatus had a back-swept crest arising from the snout. Much more fossil material has later been found, showing considerable variation in morphology. Some researchers explain this as intra-specific variability, being caused by a difference in age or sex. Others, however, assume there are different species present. The largest specimens indicate a wingspan of 4.7 m (15 ft), body length of 2 m (6.6 ft) and body msss of 25 kg (55 lb). [2]
In 1994, Kellner and Campos named a second species, Tupuxuara leonardii. The specific name honors Giuseppe Leonardi. [3] The holotype is MN 6592-V, a fragmentary skull with a more rounded crest. Other such material has been referred to T. leonardii.
In 2009, Mark Witton named a purported third species, Tupuxuara deliradamus. The holotype is SMNK PAL 6410, a skull. Another skull is the paratype: KPMNH DL 84. The specific name is derived from Latin delirus, "insane" or "crazy", and adamas, "invincible", but also the word from which "diamond" is derived. The species has a distinctive diamond-shaped skull opening and low eye sockets. The name is a tribute to the song "Shine On You Crazy Diamond" by Pink Floyd, one of Witton's favorite bands. [4] However, this species has been considered as a nomen dubium by Alexander Kellner in 2013. [5] A 2023 study also classified T. deliradamus as a nomen dubium , specifically an indeterminate tapejarine instead of thalassodromine, and a sister species of Caupedactylus . [6]
Tupuxuara is a member of the group Azhdarchoidea. Kellner assigned it to the Tapejaridae within Azhdarchoidea. According to some analyses however, Tupuxuara is closer to the Azhdarchidae (the group that includes the giant Texan form Quetzalcoatlus ) than to Tapejara and its relatives.
The cladogram below follows the 2011 analysis of Felipe Pinheiro and colleagues. [7]
Azhdarchoidea |
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Below is a cladogram showing the phylogenetic placement of Tupuxuara within Neoazhdarchia from Andres and Myers (2013). [8]
Neoazhdarchia |
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It was once been suggested that Tupuxuara was a fish eater at the coasts of South America, while some deviant hypotheses include the possibility it was a fruit eater. However, based on its azhdarchoid affinities, it was most likely a terrestrial omnivore or carnivore. The closely related Thalassodromeus was specialized for larger prey, while both Tupuxuara species lacked such specializations. [9]
A subadult described by David Martill and Darren Naish from the University of Portsmouth in 2006 had not yet fully developed its crest, which supports the suggestion that the crest was a marker for sexual maturity.
Comparisons between the scleral rings of Tupuxuara and modern birds and reptiles suggest that it may have been diurnal. [10]
Tapejara is a genus of Brazilian pterosaur from the Cretaceous Period. Tapejara crests consisted of a semicircular crest over the snout, and a bony prong which extended back behind the head. It was a small pterosaur, with a wingspan of approximately 1.23–1.3 metres (4.0–4.3 ft).
Tropeognathus is a genus of large pterosaurs from the late Early Cretaceous of South America. This genus is considered to be a member of the family Anhangueridae, however, several studies have also recovered it within another family called Ornithocheiridae. Both of these families are diverse groups of pterosaurs known for their keel-tipped snouts and large size. Tropeognathus is regarded as the largest pterosaur found in the Southern Hemisphere, only rivaled by the huge azhdarchids. The type and only species is Tropeognathus mesembrinus. Fossil remains of Tropeognathus have been recovered from the Romualdo Formation, which is a Lagerstätte located in the Santana Group of the Araripe Basin in northeastern Brazil.
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", 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 claim that Banguela is a valid genus. Another species was described in 2015 based on a supposed crest fragment, but this was later shown to be part of a turtle shell.
Anhanguera is a genus of pterodactyloid pterosaur known from the Early Cretaceous Romualdo Formation of Brazil and the Late Cretaceous Kem Kem Group of Morocco. This pterosaur is closely related to Ornithocheirus, but belongs in the family Anhangueridae. The generic name comes from the Tupi words añanga, meaning "spirit protector of the animals" + wera "bygone".
Tapejaridae are a family of pterodactyloid pterosaurs from the Cretaceous period. Members are currently known from Brazil, England, Hungary, Morocco, Spain, the United States, and China. The most primitive genera were found in China, indicating that the family has an Asian origin.
Eopteranodon is a genus of tapejarid pterosaur from the Aptian-age Lower Cretaceous Yixian Formation of Beipiao City, Liaoning, China. The genus was named in 2005 by paleontologists Lü Junchang and Zhang Xingliao. The type species is Eopteranodon lii.
Chaoyangopterus is a genus of chaoyangopterid pterosaur known from a partial skeleton found in Liaoning, China. Chaoyangopterus was found in rocks dating back to the Aptian-age Lower Cretaceous Jiufotang Formation of Dapingfang, Chaoyang.
Jidapterus is a genus of chaoyangopterid pterosaur from the Aptian-age Lower Cretaceous Jiufotang Formation of Chaoyang, Liaoning, China. The genus was in 2003 named by Dong Zhiming, Sun Yue-Wu and Wu Shao-Yuan. The type species is Jidapterus edentus. The genus name is derived from Jílín Dàxué or "Jilin University" and a Latinized Greek pteron, "wing". The specific name means "toothless" in Latin.
Santanadactylus was a genus of pterodactyloid pterosaur from the Albian-age Romualdo Member of the Upper Cretaceous Santana Formation, of Barra do Jardim, Araripe Plateau, Ceará State, Brazil. Four species have been named, but today are no considered congeneric with each other. It was a rather large pterosaur.
Azhdarchoidea is a group of pterosaurs within the suborder Pterodactyloidea, more specifically within the group Ornithocheiroidea. Pterosaurs belonging to this group lived throughout the Early and Late Cretaceous periods, with one tentative member, Tendaguripterus, that lived in the Late Jurassic period. The largest azhdarchoids include members of the family Azhdarchidae, examples of these are Quetzalcoatlus, Hatzegopteryx, and Arambourgiania. The Azhdarchoidea has been recovered as either closely related to the Ctenochasmatoidea, as the sister taxon of the Pteranodontoidea within the Ornithocheiroidea, or within the Tapejaroidea, which in turn was also within the Ornithocheiroidea.
Tupandactylus is a genus of tapejarid pterodactyloid pterosaur from the Early Cretaceous Crato Formation of Brazil.
Lacusovagus is a genus of azhdarchoid pterodactyloid pterosaur from the Lower Cretaceous of Brazil. It is based on SMNK PAL 4325, a partial upper jaw comprising sections of the skull in front of the eyes. This specimen was found in rocks of the Early Cretaceous-age Nova Olinda Member of the Crato Formation. The skull was long, and unusually wide. The section in front of the combined nasal-antorbital fenestra was relatively short. Also unusual was the combination of its toothless jaws and no bony head crest. Lacusovagus was described in 2008 by Mark Witton. The type species is L. magnificens, meaning "grand lake wanderer", in reference to its large size—it is currently the largest pterosaur known from the Crato Formation with an estimated wingspan of approximately 4 metres (13 ft) and a body mass of 30 kg (66 lb).
Thalassodrominae or Thalassodromidae is a group of azhdarchoid pterosaurs from the Cretaceous period. Its traditional members come from Brazil, however, other possible members also come from other places, including the United States, Morocco, and Argentina. Thalassodrominae is considered either to be a subfamily within the pterosaur family Tapejaridae, or as a distinct family, Thalassodromidae, within the clade Neoazhdarchia, closely related to dsungaripterids or azhdarchids.
Caupedactylus is an extinct genus of tapejarid azhdarchoid pterosaur known from the Early Cretaceous period of what is now the Romualdo Formation of the Araripe Basin in northeastern Brazil. The type and only species of Caupedactylus is Caupedactylus ybaka.
Banguela is a genus of azhdarchoid pterosaur from the Early Cretaceous period of what is now Brazil. Only one species is known, Banguela oberlii.
Tapejaroidea is a group of pterosaurs belonging to the clade Ornithocheiroidea. Tapejaroids lived from the Early to Late Cretaceous periods, with one possible member, Tendaguripterus, extending the fossil range to the Late Jurassic period. Tapejaroidea contains two groups, the Dsungaripteridae and the Azhdarchoidea, which in turn includes the azhdarchids, the group that contains some of the largest flying animals. The group was named by Brazilian paleontologist Alexander Wilhelm Armin Kellner in 1996.
Argentinadraco is an extinct genus of azhdarchoid pterosaur from the Late Cretaceous Portezuelo Formation of Argentina. It contains a single species, A. barrealensis, named in 2017 by Alexander Kellner and Jorge Calvo. Argentinadraco is unusual for bearing a bottom jaw with a concave bottom edge, as well as a pair of ridges and depressions on the top surface. These features distinguish it from all other azhdarchoid groups, complicating its assignment, but it may belong to the Chaoyangopteridae. The ridges on the lower jaw may have been used to feed on small invertebrates in loose sediment within the system of lakes and rivers that it resided in.
Aratasaurus is an extinct genus of basal coelurosaurian theropod dinosaur from the Early Cretaceous (Aptian) Romualdo Formation of Brazil. The genus contains a single species, A. museunacionali, known from a partial right leg. Aratasaurus represents the only tetrapod fossil known from the lower levels of the Romualdo Formation.
Wellnhopterus is an azhdarchid pterosaur recovered from the Late Cretaceous Javelina Formation in Texas that was previously identified as a thalassodromine. It consists of a set of upper and lower jaws, as well as some cervical vertebrae and a fragmentary long bone. In July 2021, the jaws were given the genus name "Javelinadactylus", with the type and only species as "J. sagebieli"; however, this article has now been retracted. In a paper published in December 2021, the complete holotype was independently named Wellnhopterus, with the only species being W. brevirostris. As of 2022, this is the formal name of this pterosaur.
Kariridraco is a genus of tapejarid pterosaur from the Early Cretaceous-aged Romualdo Formation, Brazil. The genus contains a single species, Kariridraco dianae.