''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 The Early Cretaceous ( geochronological name) or the Lower Cretaceous (chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145 Ma to 100.5 Ma. Geology Pro ...

period, about 112 million years ago. The only known species is ''C. atrox'', described and named in 1985 by

Giuseppe Leonardi Giuseppe Leonardi (born 31 July 1996) is an Italian sprinter, selected to be part of the Italian athletics team for the Tokyo 2020 Olympics, as a possible member of the relay team A relay race is a racing competition where members of a team ...

and

Guido Borgomanero Guido is a given name Latinised from the Old High German name Wido. It originated in Medieval Italy. Guido later became a male first name in Austria, Germany, the Low Countries, Scandinavia, Spain, Portugal, Latin America and Switzerland. The mea ...

. The name refers to the Brazilian state

Ceará Ceará (, pronounced locally as or ) is one of the 26 states of Brazil, located in the northeastern part of the country, on the Atlantic coast. It is the eighth-largest Brazilian State by population and the 17th by area. It is also one of the ...

, 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.

Discovery

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 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 flo ...

of northeastern Brazil. 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, 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 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 Fabio is a given name descended from Latin ''Fabius'' and very popular in Italy and Latin America (due to Italian migration). Its English equivalent is Fabian. The name is written without an accent in Italian and Spanish, but is usually accented ...

named a second species, ''Cearadactylus ligabuei''. The specific name honors

Giancarlo Ligabue Giancarlo Ligabue (30 October 1931 – 25 January 2015) was an Italian paleontologist, scholar, politician and businessman. Born in Venice, Ligabue graduated in Economics at the Ca' Foscari University of Venice and in Geology at la Sorbonne, and ...

, the director of the

Centro Studi Ricerche Ligabue Centro may refer to: Places Brazil * Centro, Santa Maria, a neighborhood in Santa Maria, Rio Grande do Sul, Brazil * Centro, Porto Alegre, a neighborhood of Porto Alegre, Rio Grande do Sul, Brazil *Centro (Duque de Caxias), a neighborhood of Du ...

Venice Venice ( ; it, Venezia ; vec, Venesia or ) is a city in northeastern Italy and the capital of the Veneto Regions of Italy, region. It is built on a group of 118 small islands that are separated by canals and linked by over 400 ...

. 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 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 ''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 ...

, 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 Ornithocheirae is an extinct clade of pteranodontoid pterosaurs from the Early Cretaceous to the Late Cretaceous (Valanginian to Turonian stages) of Asia, Europe, North America and South America. It was named by Harry Seeley in 1870 as a family ...

, 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 ''Maaradactylus'' is a genus of anhanguerid pterodactyloid pterosaur known from the Lower Cretaceous period (Aptian to Albian stages) of the Romualdo Formation of northeastern Brazil. Discovery ''Maaradactylus'' is based on the Museu Paleonto ...

''.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