Ludodactylus Temporal range: Aptian ~ | |
---|---|
Fossil skull of Ludodactylus with yucca leaf | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Order: | † Pterosauria |
Suborder: | † Pterodactyloidea |
Family: | † Anhangueridae |
Subfamily: | † Anhanguerinae |
Genus: | † Ludodactylus Frey et al. 2003 |
Type species | |
†Ludodactylus sibbicki Frey et al. 2003 |
Ludodactylus (meaning "play finger") is a genus of anhanguerid pterodactyloid pterosaur from the Early Cretaceous period (Aptian stage) of what is now the Crato Formation of the Araripe Basin in Ceará, Brazil. The type and only species is L. sibbicki. The generic name Ludodactylus refers to the fact that the animal had the combination of teeth and a Pteranodon -like head crest, similar to many toy pterosaurs, and no such creature was known to exist until the discovery of Ludodactylus. However, Ludodactylus is not the only pterosaur known to possess these features, its very close relative Caulkicephalus is another example. [1]
The genus was named by Eberhard Frey et al. in 2003 and contains one known species, Ludodactylus sibbicki. The name is derived from Latin ludus, "play" and Greek daktylos, "finger". Ludus refers to the fact, long lamented by paleontologists, that many toy pterosaurs combined teeth with a Pteranodon -like head crest, while no such creature was known to exist — however, Ludodactylus shows exactly this combination of features. "Dactylus", in reference to the characteristic long wing finger, has been a common element in the names of pterosaurs, since the first known genus was named Pterodactylus . The specific name "sibbicki" is an homage to the paleoartist John Sibbick. [2]
Ludodactylus is based on holotype SMNK PAL 3828, a skull missing part of the head crest, that was removed from the plate before the fossil was illegally sold. Unlike other ornithocheirids (the group of which Ludodactylus was originally assigned to), it had no premaxillary crest on the snout, but did have a crest at the back of the skull. Frey et al. interpreted the deep mandible as a crest on the lower jaws. Trapped between the rami of the mandible is a yucca leaf; Frey suggested that the animal got it caught in its beak and unsuccessfully tried to dislodge it (the edge of the leaf is frayed), and then possibly died from starvation or a complication of starving. The skull would have been more than 66 centimeters (26 in) long. [2]
In 2003, Frey et al. classified Ludodactylus in the family Ornithocheiridae. In 2007, Frey had reconsidered the validity of Ludodactylus, suggesting that it may represent the same animal as Brasileodactylus , and therefore a junior synonym. In 2014 however, Brian Andres and colleagues, in a large cladistic analysis of pterosaurs, found Ludodactylus to be slightly more closely related to ornithocheirids and anhanguerids than to Brasileodactylus. In the analysis of Andres and colleagues, Ludodactylus is classified just outside Ornithocheiridae and Anhangueridae as a derived member of the more inclusive group Anhangueria. Their cladogram is shown on the left. [3] In 2020, a different topology by Borja Holgado and Rodrigo Pêgas recovered Ludodactylus within the family Anhangueridae, more specifically to the subfamily Anhanguerinae, sister taxon to both Caulkicephalus and Guidraco . This proves that it is indeed very closely related to the anhanguerids. Their cladogram is shown on the right. [4]
Topology 1: Andres et al. (2014). [3] | Topology 2: Holgado & Pêgas (2020). [4] |
Caulkicephalus is a genus of anhanguerid pterosaur from the Isle of Wight off the coast of England. It lived during the Early Cretaceous period, about 130 million years ago.
Cearadactylus is a genus of large anhanguerid pterodactyloid pterosaur from the Romualdo Formation of Brazil, South America. Fossil remains of Cearadactylus dated back to the Albian stage of the Early Cretaceous period, about 112 million years ago. The only known species is C. atrox, described and named in 1985 by Giuseppe Leonardi and Guido Borgomanero. The name refers to the Brazilian state Ceará, and combines this with Greek daktylos, "finger", a reference to the wing finger of pterosaurs. The Latin atrox means "frightful", a reference to the fearsome dentition of the species.
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.
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.
Brasileodactylus a genus of pterosaur from the Aptian-age lower Santana formation of Chapada do Araripe, Ceará, Brazil.
Siroccopteryx is an extinct genus of anhanguerid pterodactyloid pterosaur, known from middle Cretaceous sediments in modern-day Morocco. Some researchers, such as David M. Unwin, consider the genus a junior synonym of Coloborhynchus.
Coloborhynchus is a genus of pterodactyloid pterosaur belonging to the family Anhangueridae, though it has also been recovered as a member of the Ornithocheiridae in some studies. Coloborhynchus is known from the Lower Cretaceous of England, and depending on which species are included, possibly the Albian and Cenomanian ages as well. Coloborhynchus was once thought to be the largest known toothed pterosaur, however, a specimen of the closely related Tropeognathus is now thought to have had a larger wingspan.
Lonchodectidae or Lonchodraconidae is a group of pterosaurs within the clade Pterodactyloidea. It has variously been considered to be within Ctenochasmatoidea, Azhdarchoidea and Pteranodontia. They are notable for their high, conical tooth sockets and raised alveolar margins.
Ornithocheiroidea is a group of pterosaurs within the extinct suborder Pterodactyloidea. They were typically large pterosaurs that lived from the Early to Late Cretaceous periods, with fossil remains found all over the world except Antarctica.
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.
Uktenadactylus is a genus of anhanguerid pterodactyloid pterosaurs from the Lower Cretaceous Paw Paw Formation of Texas, United States and the Wessex Formation on the Isle of Wight, England. Fossil remains of Uktenadactylus dated back to the Early Cretaceous period, from about 125 to 100 million years ago.
Anhangueridae is a group of pterosaurs within the suborder Pterodactyloidea. They were among the last pterosaurs to possess teeth. A recent study discussing the group considered the Anhangueridae to be typified by a premaxillary crest and a lateral expansion in the distal rostrum. The same study presented a cladistic analysis, for which an "agreement subtree" was calculated. The Anhangueridae was found to be sister taxon to the large crested Tropeognathus.
Barbosania is an extinct genus of crestless targaryendraconian pterosaur from the Cretaceous Romualdo Formation of the Santana Group of northeastern Brazil, dating to the Aptian to Albian.
Boreopteridae is a group of pterodactyloid pterosaurs from the Aptian-age Lower Cretaceous Yixian Formation of Liaoning, China.
Guidraco is an extinct genus of toothed pterodactyloid pterosaur known from the Early Cretaceous of Liaoning Province, northeast China. According to many recent studies, Guidraco is a member of the group Anhanguerinae, a subfamily belonging to the larger group Anhangueridae.
Ornithocheirae is an extinct clade of pteranodontoid pterosaurs from the Early Cretaceous to the Late Cretaceous of Asia, Europe, North America and South America. It was named by Harry Seeley in 1870 as a family that contains Ornithocheirus and its relatives. The name was emended to Ornithocheiridae, to match the requirements of the ICZN Code that a family-ranked clade should end with an -idae suffix. Brian Andres (2010) in his review of pterosaur phylogeny, defined the name Ornithocheirae phylogenetically, as a node-based taxon consisting of the last common ancestor of Anhanguera and Ornithocheirus and all its descendants. Thus Ornithocheirae is defined to include two families, the Anhangueridae and the Ornithocheiridae, following the opinion of Alexander Kellner and Andres that these families should not be synonymized based on their original phylogenetical definitions. However, subsequent studies in 2019 have found Ornithocheirae to be a more inclusive group containing both Anhangueria and Targaryendraconia.
Anhangueria is a group of pterosaurs belonging to the clade Pteranodontoidea. Fossil remains of this group date back from the Early to Late Cretaceous periods, around 140 to 92.5 million years ago. Anhangueria was named by paleontologists Taissa Rodrigues and Alexander Kellner in a review of Ornithocheirus species in 2013, they defined the clade as a branch-based taxon consisting of all pteranodontoids more closely related to Anhanguera blittersdorffi than to Istiodactylus latidens and Cimoliopterus cuvieri.
Maaradactylus is a genus of anhanguerid pterodactyloid pterosaur known from the Lower Cretaceous period of the Romualdo Formation of northeastern Brazil.
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.
Nicorhynchus is a genus of anhanguerid pterosaur from the Cretaceous period. It contains two species, the type species, N. capito, from the Cambridge Greensand of England, and N. fluviferox from the Kem Kem Group of Morocco. These species were previously assigned to Coloborhynchus.
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