Tupandactylus imperator

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

History

''Tupandactylus imperator'' is known from four nearly complete skulls. The holotype specimen is 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.Martill, D.M., Bechly, G. and Loveridge, R.F. (2007). ''The Crato fossil beds of Brazil: window into an ancient world.'' Cambridge University Press. , It was initially described as a species of '' Tapejara'', 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. 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.

Description

''Tupandactylus'' is notable for its large cranial crest, composed partly of bone and partly of soft tissue. The genus ''Tupandactylus'' possibly contains two species, both bearing differently sized/shaped crests that may have been used to signal and display for other ''Tupandactylus'', much as toucans use their bright bills to signal to one another.

''Tupandactylus'' crests consisted of a semicircular crest over the snout, and in the case of the type species ''T. imperator'', a bony prong which extended back behind the head. A second species, ''T. navigans'', lacked this prong, and had a much more vertical crest.

Soft tissue impressions also show that the small bony crests were extended by a much larger structure made of a keratinous material. The complete crest of ''T. navigans'' rose in a sharp, sail-like "dome" high above the rest of the skull.

Some ''Tupandactylus'' specimens preserve evidence of a keratinous beak at the jaw tips. However, this was restricted to the crested portion of the lower jaw, as one specimen also preserves pycnofibres (simple feather-like filaments) covering the jaws further back.

''T. imperator'' is estimated to have had a wingspan about , while ''T. navigans'' is smaller, with a wingspan of .

A 2022 study reported vaned feathers near the base of the crest of a ''T.'' cf. ''imperator'' specimen.

Classification

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 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''. 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''.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. 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''.Naish, D. (2008). "Crato Formation fossils and the new tapejarids." Weblog entry. ''Tetrapod Zoology''. January 18, 2008. Accessed January 31, 2008 (). 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.

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.

Paleobiology

''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.

Pterodrone unmanned aerial vehicle

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 – to develop an unmanned aerial vehicle that not only flies but also walks and sails just like the original, to be called a Pterodrone.
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 animals take the best parts of bats and birds,” Chatterjee said. “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.” “ ’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.”

The accuracy of these studies has been at times contested, 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, and Sankar Chatterjee seemingly ignored more recent aerodynamic studies in pterosaurs for these conclusions.

See also

* List of pterosaur genera
* Timeline of pterosaur research

References

{{Portal bar, Paleontology, Cretaceous, Brazil

Tapejaromorphs
Early Cretaceous pterosaurs of South America
Aptian life
Cretaceous Brazil
Fossils of Brazil
Crato Formation
Fossil taxa described in 2007
Taxa named by Alexander Kellner