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Temporal range: Albian, 112 Ma
Cearadactylus in Japan.jpg
Fossil specimen
Scientific classification edit
Kingdom: Animalia
Phylum: Chordata
Order: Pterosauria
Suborder: Pterodactyloidea
Clade: Anhangueria
Family: Anhangueridae
Subfamily: Anhanguerinae
Genus: Cearadactylus
Leonardi & Borgomanero, 1985
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.

Model in Germany, showing how the jaws where originally reconstructed

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.[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]


Life restoration

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]

See also[edit]


  1. ^ a b 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.
  2. ^ Bruno C. Vila Nova, Alexander W.A. Kellner, Juliana M. Sayão, 2010, "Short Note on the Phylogenetic Position of Cearadactylus Atrox, and Comments Regarding Its Relationships to Other Pterosaurs", Acta Geoscientica Sinica 31 Supp.1: 73–75
  3. ^ a b Unwin, D. M. (2002). "On the systematic relationships of Cearadactylus atrox, an enigmatic Early Cretaceous pterosaur from the Santana Formation of Brazil". Mitteilungen Museum für Naturkunde Berlin, Geowissenschaftlichen Reihe. 5: 1239–263.
  4. ^ Steel, L., Martill, D.M., Unwin, D.M. and Winch, J. D. (2005). "A new pterodactyloid pterosaur from the Wessex Formation (Lower Cretaceous) of the Isle of Wight, England". Cretaceous Research. 26 (4): 686–698. doi:10.1016/j.cretres.2005.03.005.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  5. ^ Palmer, D., ed. (1999). The Marshall Illustrated Encyclopedia of Dinosaurs and Prehistoric Animals. London: Marshall Editions. p. 104. ISBN 1-84028-152-9.
  6. ^ Wellnhofer, P. (1991). The Illustrated Encyclopedia of Pterosaurs. London: Salamander. pp. 125–126. doi:10.1080/02724634.1992.10011439. ISBN 9780861015665.
  7. ^ Xabier Pereda-Suberbiola; Fabien Knoll; José Ignacio Ruiz-Omeñaca; Julio Company; Fidel Torcida Fernández-Baldor (2012). "Reassessment of Prejanopterus curvirostris, a Basal Pterodactyloid Pterosaur from the Early Cretaceous of Spain". Acta Geologica Sinica. 86 (6): 1389–1401. doi:10.1111/1755-6724.12008. hdl:10651/13364. S2CID 129917510.
  8. ^ Andres, B.; Clark, J.; Xu, X. (2014). "The Earliest Pterodactyloid and the Origin of the Group". Current Biology. 24 (9): 1011–6. doi:10.1016/j.cub.2014.03.030. PMID 24768054.
  9. ^ 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. doi:10.1371/journal.pbio.2001663
  10. ^ a b c Pentland, Adele H.; Poropat, Stephen F.; Tischler, Travis R.; Sloan, Trish; Elliott, Robert A.; Elliott, Harry A.; Elliott, Judy A.; Elliott, David A. (December 2019). "Ferrodraco lentoni gen. et sp. nov., a new ornithocheirid pterosaur from the Winton Formation (Cenomanian–lower Turonian) of Queensland, Australia". Scientific Reports. 9 (1): 13454. Bibcode:2019NatSR...913454P. doi:10.1038/s41598-019-49789-4. ISSN 2045-2322. PMC 6776501. PMID 31582757.
  11. ^ 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 doi:10.1038/s41598-019-41280-4
  12. ^ Kellner, Alexander W. A.; Caldwell, Michael W.; Holgado, Borja; Vecchia, Fabio M. Dalla; Nohra, Roy; Sayão, Juliana M.; Currie, Philip J. (2019). "First complete pterosaur from the Afro-Arabian continent: insight into pterodactyloid diversity". Scientific Reports. 9 (1): 17875. Bibcode:2019NatSR...917875K. doi:10.1038/s41598-019-54042-z. PMC 6884559. PMID 31784545.
  13. ^ Kellner, Alexander W. A.; Weinschütz, Luiz C.; Holgado, Borja; Bantim, Renan A. M.; Sayão, Juliana M. (19 August 2019). "A new toothless pterosaur (Pterodactyloidea) from Southern Brazil with insights into the paleoecology of a Cretaceous desert". Anais da Academia Brasileira de Ciências. 91 (suppl 2): e20190768. doi:10.1590/0001-3765201920190768. ISSN 0001-3765. PMID 31432888.
  14. ^ 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
  15. ^ a b Holgado, B.; Pêgas, R.V. (2020). "A taxonomic and phylogenetic review of the anhanguerid pterosaur group Coloborhynchinae and the new clade Tropeognathinae". Acta Palaeontologica Polonica. 65. doi:10.4202/app.00751.2020.

Further reading[edit]

  • Dalla Vecchia, F. M. (1993), "Cearadactylus? ligabuei, nov. sp., a new Early Cretaceous (Aptian) pterosaur from Chapada do Araripe (Northeastern Brazil)", Bolletini della Societa Paleontologica Italiano, 32: 401–409