View source for Cearadactylus

{{Short description|Genus of anhanguerid pterosaur from the Early Cretaceous}}
{{speciesbox
| fossil_range = [[Albian]], {{Fossil range|112}}
| image= Cearadactylus in Japan.jpg
| image_caption = Fossil specimen
| display_parents = 4
| genus = Cearadactylus
| species = atrox
| parent_authority = Leonardi & Borgomanero, 1985
| authority = 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 [[type species|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.

==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]] of northeastern [[Brazil]]. It consists of a single skull with a length of {{convert|57|cm|ft|sp=us}}.<ref name="leonardi&borgomanero">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.</ref> 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.
[[File:Cearadaktyl (Cearadactylus) - JuraPark Baltow (4).JPG|left|thumb|Model in Poland showing how the jaws were 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 [[piscivore|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''.<ref>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</ref>

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, {{convert|403|mm|in|sp=us}} 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 {{convert|6|m|ft|sp=us}}; Kellner, pointing out that the skull is not larger than the ''C. atrox'' holotype, estimated it at {{convert|5|m|ft|sp=us}} 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''.<ref name=UDM02/> In 2005, Steel ''et al.'' suggested that it was a ''[[Coloborhynchus]] ligabuei''.<ref name=Caulkicephalus>{{cite journal|url=https://www.researchgate.net/publication/222821761|author=Steel, L., Martill, D.M., Unwin, D.M. and Winch, J. D.|year=2005|title=A new pterodactyloid pterosaur from the Wessex Formation (Lower Cretaceous) of the Isle of Wight, England.|journal=Cretaceous Research|volume=26|issue=4|pages=686–698|doi=10.1016/j.cretres.2005.03.005|bibcode=2005CrRes..26..686S }}</ref>

==Description==
[[File:Cearadactylus.jpg|thumb|Life restoration]] 
The [[wingspan]] of ''Cearadactylus'' was by the describers estimated to have been around {{convert|4|m|ft|sp=us}},<ref name=EoDP>{{cite book |editor=Palmer, D.|year=1999 |title= The Marshall Illustrated Encyclopedia of Dinosaurs and Prehistoric Animals|publisher= Marshall Editions|location=London|page= 104|isbn= 1-84028-152-9}}</ref> with a weight of perhaps {{convert|15|kg|lb}}.<ref name="leonardi&borgomanero"/> [[Peter Wellnhofer]] in 1991 estimated a wingspan of 5.5 meters (18&nbsp;ft).<ref>{{cite book|last=Wellnhofer|first=P.|year=1991|title=The Illustrated Encyclopedia of Pterosaurs|location=London|publisher=Salamander|pages=125–126|doi=10.1080/02724634.1992.10011439|isbn=9780861015665}}</ref>

==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]].<ref name=UDM02>{{cite journal |last=Unwin |first=D. M. |year=2002 |title=On the systematic relationships of ''Cearadactylus atrox'', an enigmatic Early Cretaceous pterosaur from the Santana Formation of Brazil |journal=Mitteilungen Museum für Naturkunde Berlin, Geowissenschaftlichen Reihe |volume=5 |issue=1 |pages=1239–263|doi=10.5194/fr-5-239-2002 |doi-access=free |bibcode=2002FossR...5..239U }}</ref> 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).<ref name=Prejanopterus>{{Cite journal|author1=Xabier Pereda-Suberbiola |author2=Fabien Knoll |author3=José Ignacio Ruiz-Omeñaca |author4=Julio Company |author5=Fidel Torcida Fernández-Baldor |year=2012|title=Reassessment of ''Prejanopterus curvirostris'', a Basal Pterodactyloid Pterosaur from the Early Cretaceous of Spain|url=https://www.academia.edu/26678951 |journal=Acta Geologica Sinica|volume=86|issue=6|pages=1389–1401|doi=10.1111/1755-6724.12008|bibcode=2012AcGlS..86.1389P |hdl=10651/13364|s2cid=129917510 |hdl-access=free}}</ref> 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.<ref name=kryptodrakon>{{Cite journal | url = https://www.cell.com/current-biology/pdfExtended/S0960-9822(14)00322-4 | doi = 10.1016/j.cub.2014.03.030| title = The Earliest Pterodactyloid and the Origin of the Group| journal = Current Biology| year = 2014| last1 = Andres | first1 = B. | last2 = Clark | first2 = J. | last3 = Xu | first3 = X. | pmid=24768054 | volume=24 | issue = 9| pages=1011–6| doi-access = free | bibcode = 2014CBio...24.1011A}}</ref><ref name=longrichetal2018>Longrich, N.R., Martill, D.M., and Andres, B. (2018). [http://journals.plos.org/plosbiology/article?id=10.1371/journal.pbio.2001663 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|doi-access=free}}</ref><ref name="Pentland2019">{{Cite journal|last1=Pentland|first1=Adele H.|last2=Poropat|first2=Stephen F.|last3=Tischler|first3=Travis R.|last4=Sloan|first4=Trish|last5=Elliott|first5=Robert A.|last6=Elliott|first6=Harry A.|last7=Elliott|first7=Judy A.|last8=Elliott|first8=David A.|date=December 2019|title=''Ferrodraco lentoni'' gen. et sp. nov., a new ornithocheirid pterosaur from the Winton Formation (Cenomanian–lower Turonian) of Queensland, Australia|journal=Scientific Reports|volume=9|issue=1|pages=13454|doi=10.1038/s41598-019-49789-4|pmid=31582757|pmc=6776501|bibcode=2019NatSR...913454P|issn=2045-2322}}</ref> 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.<ref name="Pentland2019"/> 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]]''.<ref name="Holgado2019">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}}</ref><ref name="Mimodactylus">{{cite journal |last1=Kellner |first1=Alexander W. A. |last2=Caldwell |first2=Michael W. |last3=Holgado |first3=Borja |last4=Vecchia |first4=Fabio M. Dalla |last5=Nohra |first5=Roy |last6=Sayão |first6=Juliana M. |last7=Currie |first7=Philip J. |title=First complete pterosaur from the Afro-Arabian continent: insight into pterodactyloid diversity |journal=Scientific Reports |date=2019 |volume=9 |issue=1 |page=17875 |doi=10.1038/s41598-019-54042-z|pmid=31784545 |pmc=6884559 |bibcode=2019NatSR...917875K |doi-access=free }}</ref><ref name="Kellner2019">{{Cite journal|last1=Kellner|first1=Alexander W. A.|last2=Weinschütz|first2=Luiz C.|last3=Holgado|first3=Borja|last4=Bantim|first4=Renan A. M.|last5=Sayão|first5=Juliana M.|date=19 August 2019|title=A new toothless pterosaur (Pterodactyloidea) from Southern Brazil with insights into the paleoecology of a Cretaceous desert|journal=Anais da Academia Brasileira de Ciências|language=en|volume=91|issue=suppl 2|pages=e20190768|doi=10.1590/0001-3765201920190768|pmid=31432888|issn=0001-3765|doi-access=free}}</ref><ref name="Targaryendraco">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]]</ref> The cladogram on the right shows a topology made by Borja Holgado and Rodrigo Pêgas in 2020.<ref name="holgado2020">{{cite journal |last1=Holgado |first1=B. |last2=Pêgas |first2=R.V. |year=2020 |title=A taxonomic and phylogenetic review of the anhanguerid pterosaur group Coloborhynchinae and the new clade Tropeognathinae |journal=Acta Palaeontologica Polonica |volume=65 |doi=10.4202/app.00751.2020|doi-access=free }}</ref>

{{col-begin|width=95%}}
{{col-2}}
'''Topology 1''': Pentland ''et al.'' (2019).<ref name="Pentland2019"/>
{{clade| style=font-size:90%;line-height:90%
                      |label1=[[Anhangueria]]
                        |1={{clade
                           |1=''[[Guidraco]]''
                           |2={{clade
|1={{clade
                              |1=''[[Brasileodactylus]]''
                              |2=''[[Ludodactylus]]''
}}
|2={{clade
|1='''''Cearadactylus'''''
                                 |label2=[[Ornithocheirae]]
                                 |2={{clade
                                    |label1=[[Anhangueridae]]
|1={{clade
|1=''[[Liaoningopterus]]''
|2=''[[Anhanguera (pterosaur)|Anhanguera]]''
}}
                                    |label2=[[Ornithocheiridae]]
|2={{clade
|1=''[[Tropeognathus]]''
|label2=[[Ornithocheirinae]]
|2={{clade
|1=''[[Coloborhynchus]]''
|2={{clade
|1=''[[Ornithocheirus]]''
|2={{clade
|1=''[[Ferrodraco]]''
|2=''[[Mythunga]]''
}} }} }} }} }} }} }} }} }}
{{col-2}}
'''Topology 2:''' Holgado & Pêgas (2020).<ref name="holgado2020"/>
{{clade|style=font-size:90%; line-height:90%
| label1=[[Anhangueridae]]
| 1={{clade
| label1=[[Tropeognathinae]]
| 1={{clade
| 1={{clade
| 1=''[[Siroccopteryx]]''
| 2=''[[Tropeognathus]]''
}}
| 2={{clade
| 1=''[[Mythunga]]''
| 2=''[[Ferrodraco]]''
}}
}}
| label2=[[Coloborhynchinae]]
| 2={{clade
| 1=''[[Aerodraco]]''
| 2={{clade
| 1=''[[Coloborhynchus]]''
| 2={{clade
| 1=''[[Nicorhynchus]]''
| 2=''[[Uktenadactylus]]''
}}
}}
}}
| label3=[[Anhanguerinae]]
| 3={{clade
| 1={{clade
| 1=''[[Caulkicephalus]]''
| 2=''[[Guidraco]]''
| 3=''[[Ludodactylus]]''
}}
| 2={{clade
| 1=''[[Liaoningopterus]]''
| 2={{clade
| 1='''''Cearadactylus'''''
| 2=''[[Maaradactylus]]''
}}
| 3=''[[Anhanguera (pterosaur)|Anhanguera]]''
}}
}}
}}
}}
{{col-end}}

In a phylogenetic analysis performed in 2025, Cearadactylus is found to be a junior synonym of [[Brasileodactylus]].<ref>{{cite journal|last1=Pêgas|first1=R.V.|year=2025|title=On the systematics and phylogenetic nomenclature of the Ornithocheiriformes (Pterosauria, Pteranodontoidea)|journal=Palaeontologica Electronica|volume=28|issue=2|at=a25|doi=10.26879/20}}</ref>

== See also ==
* [[List of pterosaur genera]]
* [[Timeline of pterosaur research]]

== References ==
{{Reflist}}

== Further reading ==
* 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
{{Pterosauria|P.}}
{{Taxonbar|from=Q133271}}
{{Portal bar|Paleontology|Brazil}}

[[Category:Pteranodontoidea]]
[[Category:Early Cretaceous pterosaurs of South America]]
[[Category:Aptian life]]
[[Category:Cretaceous Brazil]]
[[Category:Fossils of Brazil]]
[[Category:Romualdo Formation]]
[[Category:Fossil taxa described in 1985]]