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==History==
[[File:Tupandactylus navigans skeleton.PNG|thumb|left|''T. navigans'' skeleton showing soft tissue crest impression]]
''Tupandactylus imperator'' is known from four nearly complete skulls. The [[holotype]] specimen is [[Earth Sciences Museum|MCT]] 1622-R, a skull and partial lower jaw, found in the [[Crato Formation]], dating to the boundary of the [[Aptian]]-[[Albian]] stages of the early [[Cretaceous]] period, about 112 Ma ago.<ref name=martilletal2007>Martill, D.M., Bechly, G. and Loveridge, R.F. (2007). ''The Crato fossil beds of Brazil: window into an ancient world.'' Cambridge University Press. {{ISBN|0-521-85867-4}}, {{ISBN|978-0-521-85867-0}}</ref> It was initially described as a [[species]] of ''[[Tapejara (pterosaur)|Tapejara]]'',<ref name="CK97">{{cite journal |
A 2021 study describing a very complete ''T. navigans'' specimen suggested that the two species might represent different sexes of one [[sexually dimorphic]] species, but cautioned that further study was needed to test this.<ref name="Beccari&co2021">{{cite journal |last1=Beccari |first1=Victor |last2=Pinheiro |first2=Felipe Lima |last3=Nunes |first3=Ivan |last4=Anelli |first4=Luiz Eduardo |last5=Mateus |first5=Octávio |last6=Costa |first6=Fabiana Rodrigues |title=Osteology of an exceptionally well-preserved tapejarid skeleton from Brazil: Revealing the anatomy of a curious pterodactyloid clade |journal=PLOS ONE |date=2021 |volume=16 |issue=8 |pages=e0254789 |pmid=34432814|doi=10.1371/journal.pone.0254789|pmc=8386889 |bibcode=2021PLoSO..1654789B |doi-access=free }}</ref>
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''T. imperator'' is estimated to have had a wingspan about {{convert|3 to 4|m|ft|sp=us}}, while ''T. navigans'' is smaller, with a wingspan of {{convert|2.7|m|ft|sp=us}}.<ref name="Beccari&co2021"/><ref name="pinheiroetal2011"/>
A 2022 study reported vaned feathers near the base of the crest of a ''T.'' cf. ''imperator'' specimen.<ref name="Filaments">{{cite journal |last1=Cincotta |first1=Aude |last2=Nicolaï |first2=Michaël |last3=Campos |first3=Hebert Bruno Nascimento |last4=McNamara |first4=Maria |last5=D’Alba |first5=Liliana |last6=Shawkey |first6=Matthew D. |last7=Kischlat |first7=Edio-Ernst |last8=Yans |first8=Johan |last9=Carleer |first9=Robert |last10=Escuillié |first10=François |last11=Godefroit |first11=Pascal |title=Pterosaur melanosomes support signalling functions for early feathers |journal=Nature |date=2022 |volume=604 |issue=7907 |pages=684–688 |doi=10.1038/s41586-022-04622-3 |pmid=35444275|s2cid=248298392 |doi-access=free |pmc=9046085 |bibcode=2022Natur.604..684C |hdl=1942/37293 |hdl-access=free }}</ref>
==Classification==
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Beginning in 2006, several researchers, including Kellner and Campos (who named ''Tupandactylus''), had found that the three species traditionally assigned to the genus ''Tapejara'' (''T. wellnhofferi'', ''T. imperator'', and ''T. navigans'') are in fact distinct both in anatomy and in their relationships to other tapejarid pterosaurs, and thus needed to be given new [[genus|generic]] names. However, just how the species should be split proved controversial. Kellner and Campos considered only ''T. imperator'' to warrant a new name, creating ''Tupandactylus''.<ref name=KC07/> However, another study published in 2007 by Unwin and Martill found that ''T. navigans'', previously assigned to ''Tapejara'', was actually most closely related to ''T. imperator'' and belonged with it in a new genus separate from ''Tapejara''. In 2007, at a symposium held in honor of renowned pterosaur researcher [[Peter Wellnhofer]], Unwin and Martill announced the new genus name '''''Ingridia''''', in honor of Wellnhofer's late wife Ingrid. However, when they published this name in a 2007 volume, they assigned ''imperator'' as the [[type species]] of their new genus, rather than ''navigans'', which they also included as a species of ''Ingridia''.<ref name="unwin&martill2007">Unwin, D. M. and Martill, D. M. (2007). "Pterosaurs of the Crato Formation." In Martill, D. M., Bechly, G. and Loveridge, R. F. (eds), ''The Crato Fossil Beds of Brazil: Window into an Ancient World.'' Cambridge University Press (Cambridge), pp. 475–524.</ref> Furthermore, Unwin and Martill's paper was not published until several months after the similar paper by Kellner and Campos. Therefore, because both sets of authors used ''imperator'' as the type, ''Ingridia'' is considered a [[junior objective synonym]] of ''Tupandactylus''.<ref name="naishtetzooingridia2008">[[Darren Naish|Naish, D.]] (2008). "Crato Formation fossils and the new tapejarids." Weblog entry. ''Tetrapod Zoology''. January 18, 2008. Accessed January 31, 2008 ({{cite web |title=Tetrapod Zoology : Crato Formation fossils and the new tapejarids |url=http://scienceblogs.com/tetrapodzoology/2008/01/crato_formation_tapejarids.php |url-status=dead |archive-url=https://web.archive.org/web/20080706033647/http://scienceblogs.com/tetrapodzoology/2008/01/crato_formation_tapejarids.php |archive-date=July 6, 2008 |access-date=November 15, 2008}}).</ref> It was not until 2011 that ''T. navigans'' was formally reclassified in the genus ''Tupandactylus'', in a subsequent study supporting the conclusions of Unwin and Martill in 2007.<ref name=pinheiroetal2011>{{cite journal|last1=Pinheiro|first1=Felipe L.|last2=Fortier|first2=Daniel C.|last3=Schultz|first3=Cesar L.|last4=De Andrade|first4=José Artur F.G.|last5=Bantim|first5=Renan A.M. |year=2011 |title=New information on ''Tupandactylus imperator'', with comments on the relationships of Tapejaridae (Pterosauria) |journal=Acta Palaeontologica Polonica|volume=56|issue=3 |pages=567–580 |doi=10.4202/app.2010.0057|doi-access=free}}</ref>
The cladogram below follows the 2014 [[phylogenetic analysis]] by Brian Andres and colleagues. They found both ''T. navigans'' and ''T. imperator'' within the tribe [[Tapejarini]], which in turn was within the larger group [[Tapejaridae]].<ref name=kryptodrakon>{{Cite journal | 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. |volume=24 |issue=9 |pages=1011–1016 | pmid=24768054| doi-access = free | bibcode = 2014CBio...24.1011A}}</ref>
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