Normannognathus | |
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Scientific classification | |
Kingdom: | Animalia |
Phylum: | Chordata |
Order: | † Pterosauria |
Clade: | † Monofenestrata |
Genus: | † Normannognathus Buffetaut, LePage & LePage, 1998 |
Species: | †N. wellnhoferi |
Binomial name | |
†Normannognathus wellnhoferi Buffetaut, LePage & LePage, 1998 | |
Normannognathus is a genus of pterosaur from the Kimmeridgian-age Upper Jurassic Argiles d'Octeville Formation of France. Initially, Normannognathus was classified to the family Germanodactylidae, sister taxon to Germanodactylus , however, many recent analysis have recovered Normannognathus in different phylogenetic positions, and depending on different authors, Normannognathus is either found as a basal member of the Dsungaripteroidea, as an indeterminate monofenestratan, or as the sister taxon of Cycnorhamphus within the family Gallodactylidae.
In 1993, Jean-Jacques Lepage on the Normandy coast at the Cap de la Hève, near Ecqueville, Octeville-sur-Mer, Seine-Maritime, Normandy, found a ten centimetres long fossil of a pterosaur in a marine claystone layer.
In 1998, Eric Buffetaut et al. named a separate genus for it. The type species is Normannognathus wellnhoferi. The genus name is derived from Normannia, the Medieval Latin name for Normandy, and Greek gnathos, "jaw". The specific name honors Peter Wellnhofer. The genus is based on holotype Musée Géologique Cantonal de Lausanne 59'583, the left front portion of a skull and the associated, but not articulated to it, lower jaws.
The snout is low and pointed, and curves upward. Only the part in front of the nostrils has remained. On the back top of this part a very tall bony crest is present. It abruptly juts out from the premaxillae, formed like a crested wave, having a concave leading margin. After its rounded tip it gradually curves downwards again towards the skull top; its further shape is unknown because at this point the fossil ends. The crest is flat, running down the midline of the upper jaw and shows a fibrous texture that could be indicative of some covering, such as a horn sheath.
The teeth are robust, and not very elongated. They continue to be present until the very tip of the jaws. The tooth count is five per premaxilla; the number is at least nine for the maxilla, and at least fourteen per dentary: no reliable estimates can be given of the last two totals because the back of the head has been lost.
The describers assigned Normannognathus to the Germanodactylidae because it was most similar to Germanodactylus , with the exception that it resembles Dsungaripterus in having an abrupt beginning to a crest larger than that in Germanodactylus and possessing an upward pointing snout. [1] David Unwin in 2006 considered it a basal member of the Dsungaripteroidea. In 2015 however, Witton et al. concluded that it is an indeterminate monofenestratan, noting that it shares some characters with ctenochasmatoids, yet shares other characters with non-pterodactyloid monofenestratans. [2] In 2018 however, Longrich et al. found Normannognathus to be the sister taxon of Cycnorhamphus within the family Gallodactylidae, which in turn was within the Ctenochasmatoidea. Their cladogram is shown below. [3]
Archaeopterodactyloidea |
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Pterodaustro is a genus of ctenochasmatid pterodactyloid pterosaur from South America. Its fossil remains dated back to the Early Cretaceous period, about 105 million years ago. The most distinctive characteristic that separates Pterodaustro from other ctenochasmatids is its bristle-like teeth, a feature not seen in any other pterosaur.
Germanodactylus is a genus of germanodactylid pterodactyloid pterosaur from Upper Jurassic-age rocks of Germany, including the Solnhofen limestone. Its specimens were long thought to pertain to Pterodactylus. The head crest of Germanodactylus is a distinctive feature.
Pterodactyloidea is one of the two traditional suborders of pterosaurs, and contains the most derived members of this group of flying reptiles. They appeared during the middle Jurassic Period, and differ from the basal rhamphorhynchoids by their short tails and long wing metacarpals. The most advanced forms also lack teeth, and by the late Cretaceous, all known pterodactyloids were toothless. Many species had well-developed crests on the skull, a form of display taken to extremes in giant-crested forms like Nyctosaurus and Tupandactylus. Pterodactyloids were the last surviving pterosaurs when the order became extinct at the end of the Cretaceous Period, together with the non-avian dinosaurs and most marine reptiles.
Gallodactylidae is a group of pterosaurs within the suborder Pterodactyloidea. Gallodactylids differed from other related pterosaurs in several distinct features, including fewer than 50 teeth present only in the jaw tips, and rounded crests present on the rear portion of the skull and jaws but not near the ends of their snouts. At least some species possessed jaw flanges, possibly used to bissect hard-shelled prey.
Boreopterus is a genus of boreopterid pterodactyloid pterosaur from the Barremian-Aptian-age Lower Cretaceous Yixian Formation of Dalian, Liaoning, China.
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.
Cycnorhamphus is a genus of gallodactylid ctenochasmatoid pterosaur from the Late Jurassic period of France and Germany, about 152 million years ago. It is probably synonymous with the genus Gallodactylus.
Feilongus is an extinct genus of ctenochasmatid pterodactyloid pterosaur from the Barremian–Aptian-age Lower Cretaceous Yixian Formation of Beipiao, Liaoning, China.
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 Pteranodontia. They are notable for their high, conical tooth sockets and raised alveolar margins.
Ctenochasmatoidea is a group of early pterosaurs within the suborder Pterodactyloidea. Their remains are usually found in what were once coastal or lake environments. They generally had long wings, long necks, and highly specialized teeth.
Dsungaripteroidea is a group of pterosaurs within the suborder Pterodactyloidea.
The Argiles d'Octeville is a geological formation in Normandy, France. It dates back to the Kimmeridgian stage of the Late Jurassic. It is equivalent to the Kimmeridge Clay in England and predominantly consists of claystone, with some limestone. It is well exposed in cliff section at Cap de la Hève
Ctenochasmatidae is a group of pterosaurs within the suborder Pterodactyloidea. They are characterized by their distinctively looking teeth, which is thought to have been used for filter-feeding. Ctenochasmatids lived from the Late Jurassic to the Early Cretaceous periods.
Alanqa is a genus of pterodactyloid pterosaur from the Late Cretaceous period of what is now the Kem Kem Beds of southeastern Morocco. The name Alanqa comes from the Arabic word العنقاءal-‘Anqā’, for a mythical bird of Arabian culture.
Euctenochasmatia is an extinct group of pterodactyloid pterosaurs. It was named by David Unwin in 2003 as the group that contains the most recent common ancestor of Pterodactylus and Ctenochasma, and all their descendants.
Boreopteridae is a group of pterodactyloid pterosaurs from the Aptian-age Lower Cretaceous Yixian Formation of Liaoning, China.
Germanodactylidae is a controversial group of pterosaurs within the suborder Pterodactyloidea. It was first named by Yang Zhongjian in 1964, and given a formal phylogenetic definition in 2014 by Brian Andres, James Clark, and Xu Xing. They defined it as the least inclusive clade containing Germanodactylus cristatus and Normannognathus wellnhoferi, which they considered to be close relatives at the time. However, more recent studies by the same researchers have found that these pterosaurs may be only distantly related.
Cuspicephalus is an extinct genus of monofenestratan pterosaur known from Dorset in England. Its fossil remains date back to the Late Jurassic period.
Archaeopterodactyloidea is an extinct clade of pterodactyloid pterosaurs that lived from the middle Late Jurassic to the latest Early Cretaceous periods of Africa, Asia, Europe and North America. It was named by Alexander Wilhelm Armin Kellner in 1996 as the group that contains Germanodactylus, Pterodactylus, the Ctenochasmatidae and the Gallodactylidae. In 2003, Kellner defined the clade as a node-based taxon consisting of the last common ancestor of Pterodactylus, Ctenochasma and Gallodactylus and all its descendants. Although phylogenetic analyses that based on David Unwin's 2003 analysis do not recover monophyletic Archaeopterodactyloidea, phylogenetic analyses that based on Kellner's analyses, or the analyses of Brian Andres recover monophyletic Archaeopterodactyloidea at the base of the Pterodactyloidea.