Banguela Temporal range: | |
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Illustration of the holotype jaw fragment, with cross-section | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Order: | † Pterosauria |
Suborder: | † Pterodactyloidea |
Clade: | † Azhdarchoidea |
Genus: | † Banguela Headden & Campos, 2015 |
Species: | †B. oberlii |
Binomial name | |
†Banguela oberlii Headden & Campos, 2015 | |
Synonyms | |
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Banguela is a genus of azhdarchoid pterosaur from the Early Cretaceous period (Albian stage) of what is now Brazil. Only one species is known, Banguela oberlii. [1]
The Swiss collector Urs Oberli acquired a pterosaur jaw fragment from the Chapada do Araripe. In 2005, this was described by André Jacques Veldmeijer and colleagues, and referred to Thalassodromeus sethi. [2]
In 2015, it was named and described by Jaime Headden and Hebert Bruno Nascimento Campos as a separate genus Banguela, with the type species Banguela oberlii. The generic name is a Brazilian Portuguese word for "toothless one", especially used as an affectionate term for elderly women. The specific name honors Oberli. [1]
The holotype, NMSG SAO 251093, was probably found in the Romualdo Formation, also known as the Romualdo Member of the Santana Formation, dating from the Albian. It consists of the symphysis, fused front end, of the lower jaws. [1]
Banguela has an estimated skull length of about 61 centimeters (2 ft) and wingspan of over 3.7 meters (12 ft). The symphysis, with a preserved length of 273 millimeters (10.7 in), curves upwards and has a relatively short depression at its upper rear end. The front upper edge of the symphysis is sharp. The front bottom edge is sharp too but lacks a true crest. There are no teeth or tooth sockets present in the fragment. [1]
In 2005, Veldmeijer had already noted similarities to Dsungaripterus , but considered the available data to be insufficient to draw any conclusions from this. [2] In 2015, Headden & Campos placed Banguela in the family Dsungaripteridae, in a basal position. The cladogram of their analysis is shown below: [1]
If Banguela were a dsungaripterid pterosaur, it would have been unique among the group due to the presumed total absence of teeth. Other pterosaur groups, such as pteranodontids, nyctosaurids and azhdarchoids, have also lost their teeth, indicating that toothloss might have independently occurred at least four times among pterosaurs. However, because dsungaripterids are occasionally recovered as derived azhdarchoids, it is possible that toothloss has occurred more often, if as an instance of Dollo's Law azhdarchoids should be originally toothless. [1] [3] If there was a large number of cases, Banguela suggests how it developed in most of these: the development of horned rhamphothecae in the jawtips, with progressive tooth rarification until they cease to be useful. [1] Since dsungaripteroids have some of the most specialized teeth of all sauropsids, [4] the toothlessness of Banguela could indicate some degree of divergent specialization.
However, its identity as a dsungaripterid was disputed by other researchers, since no unambiguous traits of dsungaripterids were found in the type specimen. [5] [6] [7] In 2018, a study suggested that Banguela was a species of Thalassodromeus (T. oberlii), and thus it is assigned to the Thalassodrominae, a subfamily within the clade Tapejaridae. [6] In 2020, McPhee and colleagues considered Banguela as a valid genus, and claimed that it belongs to the Chaoyangopteridae. [7]
Tupuxuara is a genus of large, crested, and toothless pterodactyloid pterosaur from the Early Cretaceous period 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 or Thalassodromidae.
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 consider it 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.
Bennettazhia is a genus of tapejaromorph pterosaur which lived during the Albian stage of the Early Cretaceous from what is now the Hudspeth Formation of the state of Oregon in the United States. Although originally identified as a species of the pteranodontoid pterosaur Pteranodon, Bennettazhia is now thought to have been a different animal. The type and only species is B. oregonensis.
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. A second species, E. yixianensis, was named in 2023.
Coloborhynchus is a genus of pterodactyloid pterosaur belonging to the family Anhangueridae, though it has also been recovered as a member of the Ornithocheiridae in some studies. Coloborhynchus is known from the Lower Cretaceous of England, and depending on which species are included, possibly the Albian and Cenomanian ages as well. Coloborhynchus was once thought to be the largest known toothed pterosaur, however, a specimen of the closely related Tropeognathus is now thought to have had a larger wingspan.
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.
Chaoyangopteridae is a family of pterosaurs within the larger group Azhdarchoidea. Chaoyangopterids lived mostly during the Early Cretaceous period, though possible members, Microtuban, Xericeps and Argentinadraco, may extend the fossil range to the Late Cretaceous.
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.
Dsungaripteridae is a group of pterosaurs within the suborder Pterodactyloidea. They were robust pterosaurs with good terrestrial abilities and flight honed for inland settings, and were commonly interpreted as durophagous and possibly piscivorous pterosaurs. Fossils have been discovered from Early Cretaceous deposits in Asia, South America and possibly Europe.
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.
Maaradactylus is a genus of anhanguerid pterodactyloid pterosaur known from the Lower Cretaceous period of the Romualdo Formation of northeastern Brazil.
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.
Ornithocheiromorpha is a group of pterosaurs within the suborder Pterodactyloidea. Fossil remains of this group date back from the Early to Late Cretaceous periods, around 140 to 92.5 million years ago. Ornithocheiromorphs were discovered worldwide except Antarctica, though most genera were recovered in Europe, Asia and South America. They were the most diverse and successful pterosaurs during the Early Cretaceous, but throughout the Late Cretaceous they were replaced by pteranodontians and azhdarchoids. The Ornithocheiromorpha was defined in 2014 by Andres and colleagues, and they made Ornithocheiromorpha the most inclusive clade containing Ornithocheirus, but not Pteranodon.
Aymberedactylus is a genus of tapejarid pterosaur from the Early Cretaceous Crato Formation of Brazil. It contains a single species, A. cearensis.
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.
Apatorhamphus is an extinct genus of azhdarchoid pterosaur from the Kem Kem Group of Morocco. It might have been part of the Chaoyangopteridae. It is only known from a few snout fragments and it likely had a wingspan of between 3–7 metres (9.8–23.0 ft)
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.