Cearadactylus Temporal range: Albian, | |
---|---|
Fossil specimen | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Order: | † Pterosauria |
Suborder: | † Pterodactyloidea |
Clade: | † Anhangueria |
Family: | † Anhangueridae |
Subfamily: | † Anhanguerinae |
Genus: | † Cearadactylus Leonardi & Borgomanero, 1985 |
Species: | †C. atrox |
Binomial name | |
†Cearadactylus atrox Leonardi & Borgomanero, 1985 | |
Cearadactylus is a genus of large anhanguerid pterodactyloid pterosaur from the Romualdo Formation of Brazil, South America. Fossil remains of Cearadactylus dated back to the Albian stage of the Early Cretaceous period, about 112 million years ago. The only known species is C. atrox, described and named in 1985 by Giuseppe Leonardi and Guido Borgomanero. The name refers to the Brazilian state Ceará, and combines this with Greek daktylos, "finger", a reference to the wing finger of pterosaurs. The Latin atrox means "frightful", a reference to the fearsome dentition of the species.
The holotype of Cearadactylus is MN 7019-V (formerly CB-PV-F-O93), which was discovered in the Romualdo Formation of the Santana Group in the Araripe plateau of northeastern Brazil. It consists of a single skull with a length of 57 centimeters (1.87 ft). [1] It was traded to Italy in 1983 and bought by Borgomanero for his collection. The skull is severely damaged, especially on the top, and was perhaps reconstructed by the fossil dealer.
As shown by a later preparation by the Brazilian Museu Nacional, in the first preparation many serious mistakes were made. The fronts of the snout and of the lower jaws were confused leading to a reconstruction in which the anterior part of the head was upside down. The teeth were extensively restored and enlarged until the wider front of the jaws showed very large and robust teeth projecting outwards, forming a sort of "rosette". This kinked upper jaw and its interlocking teeth suggested a piscivorous diet, allowing the animal to keep hold of slippery fish. No crests seemed to be present. The new preparation made clear that a crest was present on the snout and that the rosette was a lot smaller. Many details were discovered that were useful in determining the phylogenetic position of Cearadactylus. [2]
In 1993, Fabio Marco Dalla Vecchia named a second species, Cearadactylus ligabuei. The specific name honors Giancarlo Ligabue, the director of the Centro Studi Ricerche Ligabue in Venice. It is based on holotype CCSRL 12692/12713, again a heavily damaged crestless skull, 403 millimeters (15.9 in) long. The skull consists of two pieces, the front and the back part, glued together by fossil traders; it is uncertain whether they belong to the same individual or indeed to the same species. Dalla Vecchia was himself not convinced the new species in fact belonged to Cearadactylus, but the skull was not sufficiently unique to base its own genus on yet still too different from known species to be assigned to them, so he created a new species for the genus the fossil most resembled. Later authors have consistently denied the identity referring to the taxon as "Cearadactylus" ligabuei.
Dalla Vecchia estimated the wingspan of "C." ligabuei at 6 meters (20 ft); Kellner, pointing out that the skull is not larger than the C. atrox holotype, estimated it at 5 meters (16 ft) at the most. Dalla Vecchia assigned C. ligabuei to the Cearadactylidae. Kellner concluded it was probably a member of Anhangueridae; Unwin in 2002 even named it Anhanguera ligabuei. [3] In 2005, Steel et al. suggested that it was a Coloborhynchus ligabuei. [4]
The wingspan of Cearadactylus was by the describers estimated to have been around 4 meters (13 ft), [5] with a weight of perhaps 15 kilograms (33 lb). [1] Peter Wellnhofer in 1991 estimated a wingspan of 5.5 meters (18 ft). [6]
Leonardi did abstain from assigning the genus to a family. Wellnhofer created a special family called Cearadactylidae, but this concept is no longer used. In 2000, Alexander Kellner concluded that it was related to, but lacking a crest not part of, the Anhangueridae within the larger group Pteranodontoidea sensu Kellner. In 2002 however, David Unwin stated it was a highly deviant member of the family Ctenochasmatidae. [3] In 2010, Kellner entered the new information into three existing databases of pterosaur features, to calculate through cladistic analysis the position of Cearadactylus in the phylogenetic tree. Although the three resulting trees differed, all had in common that Cearadactylus was close to the Anhangueridae. In 2012, Pereda-Suberbiola et al. found Cearadactylus within the group Ctenochasmatoidea, part of a polytomy that also comprises Gnathosaurus and Ardeadactylus (identified as Pterodactylus longicolum in the analysis). [7] Subsequent recent analysis however, have found Cearadactylus as a member of the group Anhangueria, and depending on the different analyses, Cearadactylus is either placed in a more derived or basal position. [8] [9] [10] A phylogenetic analysis by Pentland et al. in 2019 for example, had found Cearadactylus in a derived position within the Anhangueria, just outside the Ornithocheirae, which, by their definition, is the clade that contains the families Ornithocheiridae and Anhangueridae. Their cladogram is shown on the left. [10] Other studies however, have concluded that Cearadactylus is a member of the family Anhangueridae, more specifically a member of the subfamily Anhanguerinae, sister taxon to Maaradactylus . [11] [12] [13] [14] The cladogram on the right shows a topology made by Borja Holgado and Rodrigo Pêgas in 2020. [15]
Topology 1: Pentland et al. (2019). [10] | Topology 2: Holgado & Pêgas (2020). [15] |
Caulkicephalus is a genus of anhanguerid pterosaur from the Isle of Wight off the coast of England. It lived during the Early Cretaceous period, about 130 million years ago.
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.
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.
Brasileodactylus a genus of pterosaur from the Aptian-age lower Santana formation of Chapada do Araripe, Ceará, Brazil.
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, but its validity has been questioned.
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.
Ornithocheiroidea is a group of pterosaurs within the extinct suborder Pterodactyloidea. They were typically large pterosaurs that lived from the Early to Late Cretaceous periods, with fossil remains found all over the world except Antarctica.
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.
Ornithocheiridae is a group of pterosaurs within the suborder Pterodactyloidea. These pterosaurs were among the last to possess teeth. Members that belong to this group lived from the Early to Late Cretaceous periods, around 140 to 90 million years ago.
Hongshanopterus is a genus of pterodactyloid pterosaur from the Lower Cretaceous Jiufotang Formation of Liaoning, China.
Uktenadactylus is a genus of anhanguerid pterodactyloid pterosaurs from the Lower Cretaceous Paw Paw Formation of Texas, United States and the Wessex Formation on the Isle of Wight, England. Fossil remains of Uktenadactylus dated back to the Early Cretaceous period, from about 125 to 100 million years ago.
Anhangueridae is a group of pterosaurs within the suborder Pterodactyloidea. They were among the last pterosaurs to possess teeth. A recent study discussing the group considered the Anhangueridae to be typified by a premaxillary crest and a lateral expansion in the distal rostrum. The same study presented a cladistic analysis, for which an "agreement subtree" was calculated. The Anhangueridae was found to be sister taxon to the large crested Tropeognathus.
Guidraco is an extinct genus of toothed pterodactyloid pterosaur known from the Early Cretaceous of Liaoning Province, northeast China. According to many recent studies, Guidraco is a member of the group Anhanguerinae, a subfamily belonging to the larger group Anhangueridae.
Ornithocheirae is an extinct clade of pteranodontoid pterosaurs from the Early Cretaceous to the Late Cretaceous of Asia, Europe, North America and South America. It was named by Harry Seeley in 1870 as a family that contains Ornithocheirus and its relatives. The name was emended to Ornithocheiridae, to match the requirements of the ICZN Code that a family-ranked clade should end with an -idae suffix. Brian Andres (2010) in his review of pterosaur phylogeny, defined the name Ornithocheirae phylogenetically, as a node-based taxon consisting of the last common ancestor of Anhanguera and Ornithocheirus and all its descendants. Thus Ornithocheirae is defined to include two families, the Anhangueridae and the Ornithocheiridae, following the opinion of Alexander Kellner and Andres that these families should not be synonymized based on their original phylogenetical definitions. However, subsequent studies in 2019 have found Ornithocheirae to be a more inclusive group containing both Anhangueria and Targaryendraconia.
Anhangueria is a group of pterosaurs belonging to the clade Pteranodontoidea. Fossil remains of this group date back from the Early to Late Cretaceous periods, around 140 to 92.5 million years ago. Anhangueria was named by paleontologists Taissa Rodrigues and Alexander Kellner in a review of Ornithocheirus species in 2013, they defined the clade as a branch-based taxon consisting of all pteranodontoids more closely related to Anhanguera blittersdorffi than to Istiodactylus latidens and Cimoliopterus cuvieri.
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.
Nicorhynchus is a genus of anhanguerid pterosaur from the Cretaceous period. It contains two species, the type species, N. capito, from the Cambridge Greensand of England, and N. fluviferox from the Kem Kem Group of Morocco. These species were previously assigned to Coloborhynchus.
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