Tupandactylus: Difference between revisions

* {{extinct}}'''''Tupandactylus imperator''''' <br/><small>(Campos & Kellner, 1997)</small>

* {{extinct}}'''''Tupandactylus navigans''''' <br/><small>(Frey, Martill & Buchy, 2003)</small>

* ''Ingridia'' <br/><small>Unwin & Martill, 2007</small>

* ''[[Tapejara (pterosaur)|Tapejara]] imperator'' <br/><small>Campos & Kellner, 1997</small>

* ''Ingridia imperator'' <br/><small>(Campos & Kellner, 1997)</small>

* ''Tapejara navigans'' <br/><small>Frey, Martill & Buchy, 2003</small>

* ''Ingridia navigans'' <br/><small>(Frey, Martill & Buchy, 2003)</small>

'''''Tupandactylus''''' (meaning "Tupan finger", in reference to the [[Tupi people|Tupi]] thunder god) is a [[genus]] of [[tapejaridae|tapejarid]] [[pterodactyloid]] [[pterosaur]] from the [[Early Cretaceous]] [[Crato Formation]] of [[Brazil]].

''Tupandactylus imperator'' is known from four nearly complete skulls. The [[holotype]] specimen is [[MuseuEarth de Ciências daSciences TerraMuseum|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)wellnhoferi|Tapejara]]'',<ref name="CK97">{{cite journal |lastlast1=Campos |firstfirst1=D.A. |author2last2=Kellner, |first2=A.W.A. |author-link2=Alexander Kellner |year=1997 |title=Short note on the first occurrence of Tapejaridae in the Crato Member (Aptian), Santana Formation, Araripe Basin, Northeast Brazil |journal=Anais da Academia Brasileira de Ciências |volume=69 |issue=1 |pages=83–87 }}</ref> but later research has indicated it warrants its own genus. The skull was toothless and had a prominent sagittal crest, only the base of which was bony: the front of the crest featured a tall bony rod extending up and back, and the rear of the crest had a long prong of bone projecting behind it. The bulk of the crest was made up of soft tissue similar to [[keratin]], supported by the two bony struts.<ref name="KC07">{{cite journal |lastlast1=Kellner |firstfirst1=A.W.A. |author2author-link=Alexander Kellner |last2=Campos, |first2=D.A. |year=2007 |title=Short note on the ingroup relationships of the Tapejaridae (Pterosauria, Pterodactyloidea |journal=Boletim do Museu Nacional |volume=75 |pages=1–14 }}</ref> An additional skull described in 2011, specimen CPCA 3590, preserved more of the lower jaw, showing that like ''Tapejara'', ''T. imperator'' had a large, asymmetrical "keel"-like crest on the underside of the lower jaw tip.

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>

''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="pinheiroetal2011Beccari&co2021" /><ref name="Beccari&co2021pinheiroetal2011" />

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=1–5684–688 |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>

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 |title=Archived copy |access-date=November 15, 2008 |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>

[[File:Tapejara weln DB2.jpg|250px|thumb|Artist's conceptionreconstruction of ''T. navigans'']]

''Tupandactylus navigans'' may have largely been a terrestrial forager. Examination of the specimen GP/2E 9266 suggests that the pterosaur was capable of flight, but seemingly spent much of its time on the ground thanks to its large crest, longer forelimbs and neck, only taking short flights to possibly escape from predators. Simultaneously, it was not adapted to the same terrestrial stalking lifestyle as [[azhdarchids]] are believed to have utilized.<ref>{{Cite journal|url=https://journals.plos.org/plosone/article?id=10.1371/journal.pone.0254789|title=Osteology of an exceptionally well-preserved tapejarid skeleton from Brazil: Revealing the anatomy of a curious pterodactyloid clade|first1=Victor|last1=Beccari|first2=Felipe Lima|last2=Pinheiro|first3=Ivan|last3=Nunes|first4=Luiz Eduardo|last4=Anelli|first5=Octávio|last5=Mateus|first6=Fabiana Rodrigues|last6=Costa|date=August 25, 2021|journal=PLOS ONE|volume=16|issue=8|pages=e0254789|via=PLoS Journals|doi=10.1371/journal.pone.0254789|pmid=34432814 |pmc=8386889|bibcode=2021PLoSO..1654789B |doi-access=free }}</ref><ref>{{Cite journal|url=https://www.nature.com/articles/d41586-021-02283-2|title=A plundered pterosaur reveals the extinct flyer’sflyer's extreme headgear|date=August 25, 2021|journal=Nature|viavolume=www.nature.com597 |issue=7874 |page=10 |doi=10.1038/d41586-021-02283-2|bibcode=2021Natur.597R..10. |doi-access=free}}</ref>

A research team consisting of paleontologist [[Sankar Chatterjee]] of [[Texas Tech University]], aeronautical engineer Rick Lind of the [[University of Florida]], and their students Andy Gedeon and Brian Roberts sought to mimic the physical and biological characteristics of this pterosaur—skin, blood vessels, muscles, tendons, nerves, cranial plate, skeletal structure, and more &ndash; tomore—to develop an [[unmanned aerial vehicle]] that not only flies but also walks and sails just like the original, to be called a Pterodrone.<ref>{{cite web |urldate=October 2, https://www.sciencedaily.com/releases/2008/10/081002103649.htm |title= Pterodactyl-Inspired Robot To Master Air, Ground And Sea |url=https://www.sciencedaily.com/releases/2008/10/081002103649.htm |access-date=July October 21, 20082012 |work= Geological Society of America (2008, October 2)|publisher= ScienceDaily|access-date= July 1, 2012}}</ref>

The large, thin rudder-like sail on its head functioned as a sensory organ that acted similarly to a flight computer in a modern-day aircraft and also helped with the animal's turning agility. “These"These animals take the best parts of bats and birds,”" Chatterjee said. “They"They had the maneuverability of a bat, but could glide like an albatross. Nothing alive today compares to the performance and agility of these animals. They lived for 160 million years, so they were not stupid animals. The skies were darkened by flocks of them. They were the dominant flying animals of their time.”" “"[W]e’vee've found they could actually sail on the wind for very long periods as they flew over the oceans.... By raising their wings like sails on a boat, they could use the slightest breeze in the same way a catamaran moves across water. They could take off quickly and fly long distances with little effort.” "<ref>{{cite web |urldate=October 13, https://www.sciencedaily.com/releases/2008/10/081013140010.htm |title= Ancient Airways: Flying Drone Design Based On Prehistoric Flying Reptile |url=https://www.sciencedaily.com/releases/2008/10/081013140010.htm |access-date=July October 131, 20082012 |work= Texas Tech University (2008, October 13)|publisher= ScienceDaily|access-date= July 1, 2012}}</ref>

The accuracy of these studies has been atcontested timesby contestedpaleontologist Mark Witton, however. It has been noted that tapejarids had short wings, about as suited for soaring as those of [[Galliformes]], which are indeed consistent with adaptations for terrestriality and climbing. Likewise, no evidence for an aerodynamic function of the crest has been perceived,<ref>{{Cite web|url=http://pterosaur-net.blogspot.com/2011/01/what-despair-pterosaurs-and-david.html|title=Pterosaur.net Blog: What despair, pterosaurs and David Attenborough have in common|first=Mark|last=Witton|date=January 25, 2011}}</ref><ref>Witton, Mark P. ''Pterosaurs: Natural History, Anatomy, Evolution'', Mark P. Witton</ref> and [[Sankar Chatterjee]] seemingly ignored more recent aerodynamic studies in pterosaurs for these conclusions.<ref>{{Cite web|url=http://pterosaur-net.blogspot.com/2012/11/how-giant-pterosaurs-are-struggling-to.html|title=Pterosaur.net Blog: How giant pterosaurs are struggling to take off from the sinking ship of science journalism|first=Mark|last=Witton|date=November 11, 2012}}</ref>

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