Tupuxuara Temporal range: Albian, | |
---|---|
Reconstructed skeleton of T. leonardii, American Museum of Natural History | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Order: | † Pterosauria |
Suborder: | † Pterodactyloidea |
Family: | † Thalassodromidae |
Genus: | † Tupuxuara Kellner & Campos, 1988 |
Type species | |
†Tupuxuara longicristatus Kellner & Campos, 1988 | |
Other species | |
|
Tupuxuara is a genus of thalassodromid pterosaur that lived during the Albian age of the Early Cretaceous, about 112 million years ago. Its remains were found in what is now the Romualdo Formation of the Santana Group in Brazil. Tupuxuara was named in 1988 by paleontologists Alexander Kellner and Diógenes de Almeida Campos. The name Tupuxuara means "familiar spirit" referring to a familiar spirit in the mythology of the Tupi people in Brazil. Two species have been named, T. longicristatus, the type species, and T. leonardii. An additional species has been named in 2013, T. deliradamus. However, the validity of this species has been put into question and it may not even belong to Tupuxuara.
Tupuxuara was quite large in size. It had a big crest at the back portion of its head, stemming from the snout. This crest is most likely an indicator of sexual maturity, given that it appears more prominently in mature individuals. The beak of Tupuxuara was toothless.
The closest relative of Tupuxuara is Thalassodromeus , both of which form a family called Thalassodromidae, which, depending on the study, has been found to be within either the group Tapejaromorpha, closely related to the family Tapejaridae, or within the group Neoazhdarchia, closely related to the family Dsungaripteridae.
Early studies had suggested that Tupuxuara was piscivorous (a fish eater), with some even suggesting a fruit-based diet. However, based off of its azhdarchoid lineage, Tupuxuara most likely had been a terrestrial omnivore or carnivore.
Tupuxuara was named and described by Brazilian paleontologists Alexander Kellner and Diógenes de Almeida Campos in 1988. The type species is Tupuxuara longicristatus. The generic name Tupuxuara means "familiar spirit", referring to a familiar spirit from the mythology of the Tupi people in Brazil. The specific name longicristatus means "long-crested" in Latin. [1]
The holotype specimen, MN 6591-V, was found in the Santana Formation (now known as the Romualdo Formation) of Brazil, a rock formation dating back to the Albian stage of the Early Cretaceous period. The holotype consists of a snout and some partial wing bones. [1]
In 1994, Kellner and Campos named a second species, Tupuxuara leonardii. The specific name honors Giuseppe Leonardi, an Italian geologist and paleontologist. The holotype specimen is MN 6592-V, a fragmentary skull with a more rounded crest than the type species T. longicristatus. Other similar material has been referred to T. leonardii as well. [2]
In 2009, British paleontologist Mark Witton named a purported third species, Tupuxuara deliradamus. The holotype is SMNK PAL 6410, a skull. Another skull is the paratype: KPMNH DL 84. The specific name is derived from Latin delirus, "insane" or "crazy", and adamas, "invincible", but also the word from which "diamond" is derived. The species has a distinctive diamond-shaped skull opening and low eye sockets. The name is a tribute to the song "Shine On You Crazy Diamond" by Pink Floyd, one of Witton's favorite bands. [3] However, this species has been considered as a nomen dubium (dubious name) by Kellner in 2013. [4] A 2023 study also classified T. deliradamus as a nomen dubium, specifically an indeterminate member of the subfamily Tapejarinae and a sister species of Caupedactylus , which challenges its initial classification within the Thalassodrominae, the group which Tupuxuara belongs to. [5]
Tupuxuara was a fairly large pterosaur in terms in size. The most distinguishing characteristic of Tupuxuara is the large crest that it possessed, which protruded from the back portion its head and originated from its snout. This crest consisted primarily of bone. [6] [7] This feature was more prominent in mature individuals. More recently found fossil material show considerable variation in morphology. Some researchers explain this as intra-specific variability, being caused by a difference in age or sex. Others even assume that it is due to different species being present in the discoveries. [8]
In his 2022 pterosaur book, American paleontologist Gregory S. Paul indicated that Tupuxuara had a wingspan measuring 4.7 m (15 ft), a total body length of 2 m (6.6 ft) and a body mass of 25 kg (55 lb). Paul noted that Tupuxuara had long wings relative to its body mass. [8] Tupuxuara was slightly smaller compared to its closest relative, Thalassodromeus , and it bore a skull that was less heavily built due to its crest having a smaller size. [9] [10] A key difference between the two species of Tupuxuara is the structure of the crest. The type species, T. longicristatus, featured a more elongated crest, as its name suggests, while T. leonardii featured a more rounded crest in terms of shape. [2]
Tupuxuara was initially assigned to the family Tapejaridae in its description by Kellner, closely related to the pterosaur Tapejara . [2] Within Tapejaridae, its closest relative was Thalassodromeus, indicated by their bony crest, a characteristic only shared by them in the family. Meanwhile, other tapejarids bore a crest mostly consisting of soft tissue. [6] [7] In 2007, Kellner and Campos divided Tapejaridae into the subfamilies Tapejarinae and Thalassodrominae, Tupuxuara being a member of the latter, alongside Thalassodromeus. Tapejaridae itself was recovered within the superfamily Azhdarchoidea. [7] This arrangement would later be supported by many different studies. [11] [10] [12] [13] [14]
However, conflicting studies also arose. A new group had been named in 2003 by paleontologist David Unwin, the Neoazhdarchia, containing Tupuxuara and the family Azhdarchidae. This new group, like the Tapejaridae, would be inside Azhdarchoidea. In 2006, this arrangement would be further supported by British paleontologists David Martill and Darren Naish, who found Tapejaridae to be paraphyletic (unnatural), and found both Tupuxuara and Thalassodromeus as sister taxa to the Azhdarchidae. [15] [16] Later, in a study by Chinese paleontologist Lü Junchang and colleagues in 2008, the name "Tupuxuaridae" would be used to include Tupuxuara and Thalassodromeus. [17] However, in 2009, Witton stated that it had never been a validly established name, arguing that Thalassodrominae should be the proper name for the group containing Tupuxuara and Thalassodromeus. Witton supported the concept of the group Neoazhdarchia, so he further converted the subfamily Thalassodrominae into its own family called Thalassodromidae, and included it within said group. [3] A few subsequent studies would support this model and would further elaborate by grouping Thalassodromidae alongside the family Dsungaripteridae within a group named Dsungaripteromorpha, which itself was nested within Neoazhdarchia. [18] [19]
Considering that the group containing Tupuxuara and Thalassodromeus has received two different denominations throughout the years, Thalassodromidae and Thalassodrominae, in 2023, paleontologist Rodrigo Pêgas and colleagues argued that despite the disagreements between the position of said group within Azhdarchoidea, the species contained within it have almost always been the same ones. Therefore, they deemed the difference in naming pattern undesirable. They favored the denomination Thalassodromidae, in order to have consistency with other studies that used the same name. In their analysis, they supported a close relationship between the thalassodromids and the family Tapejaridae, following the classification model established by Kellner. They included both families within the larger group Tapejaromorpha (defined as the most inclusive clade containing Tapejara wellnhoferi but not Azhdarcho lancicollis ). [20] Subsequent studies using Kellner's model would also employ the denomination Thalassodromidae instead of Thalassodrominae. [21] [22]
Below are two cladograms showing different conflicting studies regarding the position of Thalassodromidae. The first one is from a phylogenetic analysis by paleontologist Nicholas Longrich and colleagues in 2018. It showcases the interrelationships within Azhdarchoidea, in which Thalassodromidae was recovered as the sister taxon of Dsungaripteridae. Aside from Thalassodromeus, the pterosaurs Alanqa and Aerotitan were also found as thalassodromids. [18] The second cladogram is based on the phylogenetic analysis conducted by Pêgas and colleagues in 2023. In this study, Thalassodromidae consisted of the two species of Tupuxuara (T. longicristatus and T. leonardii), Thalassodromeus, and Kariridraco . Both Thalassodromidae and Tapejaridae were found as sister taxa within the larger Tapejaromorpha, corroborating the relationship between the two families. [20]
Topology 1: Longrich and colleagues (2018).
| Topology 2: Pêgas and colleagues (2023).
|
It has once been suggested that Tupuxuara had a piscivorous (a fish-based) diet, preying in the coasts of modern day South America, while some deviant hypotheses even include the possibility that it was a fruit eater. However, basing off of its azhdarchoid affinities, it was most likely a terrestrial omnivore or carnivore. The closely related Thalassodromeus was specialized for larger prey, while both Tupuxuara species lacked such specializations. [9]
A subadult described by Martill and Naish from the University of Portsmouth in 2006 had not yet fully developed its crest, which supports the suggestion that the crest was a marker for sexual maturity. [16]
Comparisons between the scleral rings of Tupuxuara and modern birds and reptiles suggest that it may have been diurnal. [23]
Azhdarchidae is a family of pterosaurs known primarily from the Late Cretaceous Period, though an isolated vertebra apparently from an azhdarchid is known from the Early Cretaceous as well. Azhdarchids are mainly known for including some of the largest flying animals discovered, but smaller cat-size members have also been found. Originally considered a sub-family of Pteranodontidae, Nesov (1984) named the Azhdarchinae to include the pterosaurs Azhdarcho, Quetzalcoatlus, and Titanopteryx. They were among the last known surviving members of the pterosaurs, and were a rather successful group with a worldwide distribution. Previously it was thought that by the end of the Cretaceous, most pterosaur families except for the Azhdarchidae disappeared from the fossil record, but recent studies indicate a wealth of pterosaurian fauna, including pteranodontids, nyctosaurids, tapejarids and several indeterminate forms.
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 2015, and assigned back to Thalassodromeus as the species T. oberlii in 2018, though other researchers consider it a valid genus. Another species was described in 2015 based on a supposed crest fragment, but this was later shown to be part of a turtle shell.
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.
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. A second species, E. yixianensis, was named in 2023, but its validity has been questioned.
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.
Jidapterus is a genus of chaoyangopterid pterosaur from the Aptian-age Lower Cretaceous Jiufotang Formation of Chaoyang, Liaoning, China. The genus was in 2003 named by Dong Zhiming, Sun Yue-Wu and Wu Shao-Yuan. The type species is Jidapterus edentus. The genus name is derived from Jílín Dàxué or "Jilin University" and a Latinized Greek pteron, "wing". The specific name means "toothless" in Latin.
Lonchodectes was a genus of lonchodectid pterosaur from several formations dating to the Turonian of England, mostly in the area around Kent. The species belonging to it had been assigned to Ornithocheirus until David Unwin's work of the 1990s and 2000s. Several potential species are known; most are based on scrappy remains, and have gone through several other generic assignments. The genus is part of the complex taxonomy issues surrounding Early Cretaceous pterosaurs from Brazil and England, such as Amblydectes, Anhanguera, Coloborhynchus, and Ornithocheirus.
Lonchodectidae or Lonchodraconidae is a group of pterosaurs within the clade Pterodactyloidea. It has variously been considered to be within Ctenochasmatoidea, Azhdarchoidea and Pteranodontoidea. They are notable for their high, conical tooth sockets and raised alveolar margins.
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. Other notable pterosaur families belonging to the Azhdarchoidea are the Tapejaridae, the Thalassodromidae, and the Chaoyangopteridae. This group contains some of the most advanced members of the pterosaur order.
Tupandactylus is a genus of tapejarid pterodactyloid pterosaur from the Early Cretaceous Crato Formation of Brazil.
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.
Chaoyangopteridae is a family of pterosaurs within the larger group Azhdarchoidea. Chaoyangopterids lived mostly during the Early Cretaceous period, though possible members, Microtuban, Xericeps and Argentinadraco, may extend the fossil range to the Late Cretaceous.
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 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.
Dsungaripteridae is a group of pterosaurs within the suborder Pterodactyloidea. They were robust pterosaurs with good terrestrial abilities and flight honed for inland settings, and were commonly interpreted as durophagous and possibly piscivorous pterosaurs. Fossils have been discovered from Early Cretaceous deposits in Asia, South America and possibly Europe.
Banguela is a genus of azhdarchoid pterosaur from the Early Cretaceous period of what is now Brazil. Only one species is known, Banguela oberlii.
Tapejaroidea is a group of pterosaurs belonging to the clade Ornithocheiroidea. Tapejaroids lived from the Early to Late Cretaceous periods, with one possible member, Tendaguripterus, extending the fossil range to the Late Jurassic period. Tapejaroidea contains two groups, the Dsungaripteridae and the Azhdarchoidea, which in turn includes the azhdarchids, the group that contains some of the largest flying animals. The group was named by Brazilian paleontologist Alexander Wilhelm Armin Kellner in 1996.
Tapejaromorpha is a group of pterosaurs within the clade Azhdarchoidea. The fossil remains of tapejaromorphs dated back to the Cretaceous period.
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 have been discovered worldwide except Antarctica, though most genera have been 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 pteranodontians 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.
Argentinadraco is an extinct genus of azhdarchoid pterosaur from the Late Cretaceous Portezuelo Formation of Argentina. It contains a single species, A. barrealensis, named in 2017 by Alexander Kellner and Jorge Calvo. Argentinadraco is unusual for bearing a bottom jaw with a concave bottom edge, as well as a pair of ridges and depressions on the top surface. These features distinguish it from all other azhdarchoid groups, complicating its assignment, but it may belong to the Chaoyangopteridae. The ridges on the lower jaw may have been used to feed on small invertebrates in loose sediment within the system of lakes and rivers that it resided in.
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.