Tupandactylus

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Tupandactylus
Temporal range: Early Cretaceous, 112  Ma
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Tupandactylus skeleton - Museu Nacional, Rio de Janeiro.jpg
Reconstructed T. imperator skeleton, National Museum of Brazil
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Order: Pterosauria
Suborder: Pterodactyloidea
Family: Tapejaridae
Subfamily: Tapejarinae
Tribe: Tapejarini
Genus: Tupandactylus
Kellner & Campos, 2007
Type species
Tapejara imperator
Campos & Kellner, 1997
Species
  • Tupandactylus imperator
    (Campos & Kellner, 1997)
  • Tupandactylus navigans
    (Frey, Martill & Buchy, 2003)
Synonyms
List of synonyms
  • Genus synonymy
      • Ingridia
        Unwin & Martill, 2007
    Synonyms of T. imperator
      • Tapejara imperator
        Campos & Kellner, 1997
      • Ingridia imperator
        (Campos & Kellner, 1997)
    Synonyms of T. navigans
      • Tapejara navigans
        Frey, Martill & Buchy, 2003
      • Ingridia navigans
        (Frey, Martill & Buchy, 2003)

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

T. navigans skeleton showing soft tissue crest impression Tupandactylus navigans skeleton.PNG
T. navigans skeleton showing soft tissue crest impression

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. [1] It was initially described as a species of Tapejara , [2] 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. [3] 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. [4]

Description

Life restoration showing distribution of filaments preserved in a T. imperator specimen Filaments of Tupandactylus.jpg
Life restoration showing distribution of filaments preserved in a T. imperator specimen

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.

Profile concepts of Tupandactylus imperator (C), Tupandactylus navigans (B), and Tapejara wellnhoferi (A) Tapejarines mmartyniuk.png
Profile concepts of Tupandactylus imperator (C), Tupandactylus navigans (B), and Tapejara wellnhoferi (A)

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. [5]

T. imperator is estimated to have had a wingspan about 3 to 4 meters (9.8 to 13.1 ft), while T. navigans is smaller, with a wingspan of 2.7 meters (8.9 ft). [4] [5]

A 2022 study reported vaned feathers near the base of the crest of a T. cf. imperator specimen. [6]

Classification

T. imperator skull model Cast of Tupandactylus imperator - Pterosaurs Flight in the Age of Dinosaurs.jpg
T. imperator skull model

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. [3] 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. [7] 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. [8] 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. [5]

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. [9]

  Azhdarchoidea  

Paleobiology

Artist's reconstruction of T. navigans Tapejara weln DB2.jpg
Artist's reconstruction 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. [10] [11]

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. [12] 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." "[W]e'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." [13]

The accuracy of these studies has been contested by paleontologist 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, [14] [15] and Sankar Chatterjee seemingly ignored more recent aerodynamic studies in pterosaurs for these conclusions. [16]

See also

Related Research Articles

<i>Tapejara wellnhoferi</i> Genus of tapejarid pterosaur from the Early Cretaceous

Tapejara is a genus of Brazilian pterosaur from the Cretaceous Period. Tapejara crests consisted of a semicircular crest over the snout, and a bony prong which extended back behind the head. It was a small pterosaur, with a wingspan of approximately 1.23–1.3 metres (4.0–4.3 ft).

<i>Tupuxuara</i> Genus of azhdarchoid pterosaur from the Early Cretaceous

Tupuxuara is a genus of large, crested, and toothless pterodactyloid pterosaur from the Early Cretaceous period of what is now the Romualdo Formation of the Santana Group, Brazil, about 125 to 112 million years ago. Tupuxuara is a close relative of Thalassodromeus, and both form a group that is either called Thalassodrominae or Thalassodromidae.

<i>Tropeognathus</i> Genus of anhanguerid pterosaur from the Early Cretaceous

Tropeognathus is a genus of large pterosaurs from the late Early Cretaceous of South America. This genus is considered to be a member of the family Anhangueridae, however, several studies have also recovered it within another family called Ornithocheiridae. Both of these families are diverse groups of pterosaurs known for their keel-tipped snouts and large size. Tropeognathus is regarded as the largest pterosaur found in the Southern Hemisphere, only rivaled by the huge azhdarchids. The type and only species is Tropeognathus mesembrinus. Fossil remains of Tropeognathus have been recovered from the Romualdo Formation, which is a Lagerstätte located in the Santana Group of the Araripe Basin in northeastern Brazil.

<i>Thalassodromeus</i> Genus of azhdarchoid pterosaur from the Early Cretaceous

Thalassodromeus is a genus of pterosaur that lived in what is now Brazil during the Early Cretaceous period, about a hundred million years ago. The original skull, discovered in 1983 in the Araripe Basin of northeastern Brazil, was collected in several pieces. In 2002, the skull was made the holotype specimen of Thalassodromeus sethi by palaeontologists Alexander Kellner and Diogenes de Almeida Campos. The generic name means "sea runner", and the specific name refers to the Egyptian god Seth due to its crest being supposedly reminiscent of Seth's crown. Other scholars have pointed out that the crest was instead similar to the crown of Amon. A jaw tip was assigned to T. sethi in 2005, became the basis of the new genus Banguela in 2014, and assigned back to Thalassodromeus as the species T. oberlii in 2018. Another species was described in 2015 based on a supposed crest fragment, but this was later shown to be part of a turtle shell.

Anhanguera is a genus of pterodactyloid pterosaur known from the Early Cretaceous Romualdo Formation of Brazil and the Late Cretaceous Kem Kem Group of Morocco. This pterosaur is closely related to Ornithocheirus, but belongs in the family Anhangueridae. The generic name comes from the Tupi words añanga, meaning "spirit protector of the animals" + wera "bygone".

<span class="mw-page-title-main">Tapejaridae</span> Family of azhdarchoid pterosaurs from the Cretaceous period

Tapejaridae are a family of pterodactyloid pterosaurs from the Cretaceous period. Members are currently known from Brazil, England, Hungary, Morocco, Spain, the United States, and China. The most primitive genera were found in China, indicating that the family has an Asian origin.

<i>Bakonydraco</i> Genus of tapejarid pterosaur from the Late Cretaceous

Bakonydraco is a genus of pterodactyloid pterosaur from the Late Cretaceous period of what is now the Csehbánya Formation of the Bakony Mountains, Iharkút, Veszprém, western Hungary.

Brasileodactylus a genus of pterosaur from the Aptian-age lower Santana formation of Chapada do Araripe, Ceará, Brazil.

<i>Eopteranodon</i> Genus of tapejarid pterosaur from the Early Cretaceous

Eopteranodon is a genus of tapejarid pterosaur from the Aptian-age Lower Cretaceous Yixian Formation of Beipiao City, Liaoning, China. The genus was named in 2005 by paleontologists Lü Junchang and Zhang Xingliao. The type species is Eopteranodon lii.

<i>Feilongus</i> Genus of ctenochasmatid pterosaur from the Early Cretaceous

Feilongus is an extinct genus of ctenochasmatid pterodactyloid pterosaur from the Barremian–Aptian-age Lower Cretaceous Yixian Formation of Beipiao, Liaoning, China.

<i>Sinopterus</i> Genus of tapejarid pterosaur from the Early Cretaceous

Sinopterus is a genus of tapejarid pterodactyloid pterosaur from the Aptian-age Lower Cretaceous Jiufotang Formation of Chaoyang, Liaoning, China. It was first described and named by Wang Xiaolin and Zhou Zhonghe. Historically, there were multiple species attributed to the genus although only one is considered to be valid. Sinopterus is known for its proportionally large skull, which has a birdlike pointed beak, a long bony crest that starts with a tall premaxilla and goes back along the middle of the skull to form a point overhanging the rear of the skull, and its lack of teeth.

<span class="mw-page-title-main">Azhdarchoidea</span> Superfamily of ornithocheiroid pterosaurs

Azhdarchoidea is a group of pterosaurs within the suborder Pterodactyloidea, more specifically within the group Ornithocheiroidea. Pterosaurs belonging to this group lived throughout the Early and Late Cretaceous periods, with one tentative member, Tendaguripterus, that lived in the Late Jurassic period. The largest azhdarchoids include members of the family Azhdarchidae, examples of these are Quetzalcoatlus, Hatzegopteryx, and Arambourgiania. The Azhdarchoidea has been recovered as either closely related to the Ctenochasmatoidea, as the sister taxon of the Pteranodontoidea within the Ornithocheiroidea, or within the Tapejaroidea, which in turn was also within the Ornithocheiroidea.

<span class="mw-page-title-main">Ornithocheiridae</span> Family of ornithocheiran pterosaurs

Ornithocheiridae is a group of pterosaurs within the suborder Pterodactyloidea. These pterosaurs were among the last to possess teeth. Members that belong to this group lived from the Early to Late Cretaceous periods, around 140 to 90 million years ago.

<span class="mw-page-title-main">Thalassodrominae</span> Subfamily of azhdarchoid pterosaurs from the Cretaceous period

Thalassodrominae or Thalassodromidae is a group of azhdarchoid pterosaurs from the Cretaceous period. Its traditional members come from Brazil, however, other possible members also come from other places, including the United States, Morocco, and Argentina. Thalassodrominae is considered either to be a subfamily within the pterosaur family Tapejaridae, or as a distinct family, Thalassodromidae, within the clade Neoazhdarchia, closely related to dsungaripterids or azhdarchids.

<i>Europejara</i> Genus of tapejarid pterosaur from the Early Cretaceous

Europejara is a genus of tapejarid pterosaur from the Early Cretaceous period of Spain. The type and only species known is Europejara olcadesorum.

<i>Caiuajara</i> Genus of tapejarid pterosaur from the Late Cretaceous

Caiuajara is an extinct genus of tapejarid pterosaur from the Early Cretaceous period of Brazil. It is known from a single type species, Caiuajara dobruskii.

<span class="mw-page-title-main">Ornithocheiromorpha</span> Clade of pteranodontoid pterosaurs

Ornithocheiromorpha is a group of pterosaurs within the suborder Pterodactyloidea. Fossil remains of this group date back from the Early to Late Cretaceous periods, around 140 to 92.5 million years ago. Ornithocheiromorphs were discovered worldwide except Antarctica, though most genera were recovered in Europe, Asia and South America. They were the most diverse and successful pterosaurs during the Early Cretaceous, but throughout the Late Cretaceous they were replaced by better adapted and more advanced pterosaur species such the pteranodontids and azhdarchoids. The Ornithocheiromorpha was defined in 2014 by Andres and colleagues, and they made Ornithocheiromorpha the most inclusive clade containing Ornithocheirus, but not Pteranodon.

<i>Aymberedactylus</i> Genus of tapejarid pterosaur from the Early Cretaceous

Aymberedactylus is a genus of tapejarid pterosaur from the Early Cretaceous Crato Formation of Brazil. It contains a single species, A. cearensis.

<i>Wellnhopterus</i> Genus of azhdarchid pterosaur from the Late Cretaceous

Wellnhopterus is an azhdarchid pterosaur recovered from the Late Cretaceous Javelina Formation in Texas that was previously identified as a thalassodromine. It consists of a set of upper and lower jaws, as well as some cervical vertebrae and a fragmentary long bone. In July 2021, the jaws were given the genus name "Javelinadactylus", with the type and only species as "J. sagebieli"; however, this article has now been retracted. In a paper published in December 2021, the complete holotype was independently named Wellnhopterus, with the only species being W. brevirostris. As of 2022, this is the formal name of this pterosaur.

<i>Huaxiadraco</i> Genus of tapejarid pterosaur from the Early Cretaceous period

Huaxiadraco is a genus of tapejarid pterodactyloid pterosaur from the Aptian-age Lower Cretaceous Jiufotang Formation of Chaoyang, Liaoning, China. It is the third valid genus of tapejarid from the Jehol Biota, after Sinopterus and Eopteranodon. It contains one species, Huaxiadraco corollatus, originally assigned to the defunct genus Huaxiapterus.

References

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  2. Campos, D.A.; Kellner, A.W.A. (1997). "Short note on the first occurrence of Tapejaridae in the Crato Member (Aptian), Santana Formation, Araripe Basin, Northeast Brazil". Anais da Academia Brasileira de Ciências. 69 (1): 83–87.
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