''Cearadactylus'' is a

genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...

of large

anhanguerid 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 late ...

pterodactyloid Pterodactyloidea (derived from the Greek words ''πτερόν'' (''pterón'', for usual ''ptéryx'') "wing", and ''δάκτυλος'' (''dáktylos'') "finger" meaning "winged finger", "wing-finger" or "finger-wing") is one of the two traditional ...

pterosaur Pterosaurs (; from Greek ''pteron'' and ''sauros'', meaning "wing lizard") is an extinct clade of flying reptiles in the order, Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 ...

from the

Romualdo Formation The Romualdo Formation is a geologic Konservat-Lagerstätte in northeastern Brazil's Araripe Basin where the states of Pernambuco, Piauí and Ceará come together. The geological formation, previously designated as the Romualdo Member of the Sa ...

Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...

South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the sout ...

. Fossil remains of ''Cearadactylus'' dated back to the

Albian The Albian is both an age of the geologic timescale and a stage in the stratigraphic column. It is the youngest or uppermost subdivision of the Early/Lower Cretaceous Epoch/ Series. Its approximate time range is 113.0 ± 1.0 Ma to 100.5 ± 0 ...

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 Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...

''atrox'' means "frightful", a reference to the fearsome dentition of the species.

Discovery

The

holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of sever ...

of ''Cearadactylus'' is MN 7019-V (formerly CB-PV-F-O93), which was discovered in the

of the

Santana Group The Santana Group is a geologic group, formerly included as the middle part of the Araripe Group, in the Araripe Basin of northeastern Brazil. The group comprises the Crato, Ipubi and Romualdo Formations and is dated to the Aptian to Albian stag ...

in the

Araripe plateau The Chapada do Araripe, also known as the Serra do Araripe, is a ''chapada'' (plateau) in northeastern Brazil. The chapada forms the boundary of Ceará and Pernambuco states, and forms the watershed between the Jaguaribe River of Ceará, which flow ...

of northeastern

. It consists of a single skull with a length of .Leonardi, G. & Borgomanero, G. (1985). "''Cearadactylus atrox'' nov. gen., nov. sp.: novo Pterosauria (Pterodactyloidea) da Chapada do Araripe, Ceara, Brasil." ''Resumos dos communicaçoes VIII Congresso bras. de Paleontologia e Stratigrafia'', 27: 75–80. 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. Cearadactylus model

As shown by a later preparation by the Brazilian

Museu Nacional The National Museum of Brazil ( pt, Museu Nacional) is the oldest scientific institution of Brazil. It is located in the city of Rio de Janeiro, where it is installed in the Paço de São Cristóvão (Saint Christopher's Palace), which is ins ...

, 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 A tooth ( : teeth) is a hard, calcified structure found in the jaws (or mouths) of many vertebrates and used to break down food. Some animals, particularly carnivores and omnivores, also use teeth to help with capturing or wounding prey, t ...

suggested a piscivore, piscivourous 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''. 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, 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 ; Kellner, pointing out that the skull is not larger than the ''C. atrox'' holotype, estimated it at 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 (pterosaur), Anhanguera ligabuei''. In 2005, Steel ''et al.'' suggested that it was a ''Coloborhynchus ligabuei''.

Description

The wingspan of ''Cearadactylus'' was by the describers estimated to have been around , with a weight of perhaps . Peter Wellnhofer in 1991 estimated a wingspan of 5.5 meters (18 ft).

Classification

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. 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). 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.Longrich, N.R., Martill, D.M., and Andres, B. (2018)
Late Maastrichtian pterosaurs from North Africa and mass extinction of Pterosauria at the Cretaceous-Paleogene boundary.
''PLoS Biology'', 16(3): e2001663. 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. 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''.Borja Holgado, Rodrigo V. Pêgas, José Ignacio Canudo, Josep Fortuny, Taissa Rodrigues, Julio Company & Alexander W.A. Kellner, 2019, "On a new crested pterodactyloid from the Early Cretaceous of the Iberian Peninsula and the radiation of the clade Anhangueria", ''Scientific Reports'' 9: 4940 Pêgas, R.V., Holgado, B., Leal, M.E.C., 2019. "On ''Targaryendraco wiedenrothi'' gen. nov. (Pterodactyloidea, Pteranodontoidea, Lanceodontia) and recognition of a new cosmopolitan lineage of Cretaceous toothed pterodactyloids", ''Historical Biology'', 1–15. doi:10.1080/08912963.2019.1690482 The cladogram on the right shows a topology made by Borja Holgado and Rodrigo Pêgas in 2020. Topology 1: Pentland ''et al.'' (2019). Topology 2: Holgado & Pêgas (2020).

Discovery

Description

Classification

See also

References

Further reading