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Normannognathus; Temporal range: Kimmeridgian PreꞒ Ꞓ O S D C P T J K Pg N
Scientific classification
Domain:	Eukaryota
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.

Discovery and naming

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.

Description

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.

Classification

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

Germanodactylidae

	Germanodactylus cristatus

	Germanodactylus rhamphastinus

Euctenochasmatia

Pterodactylus antiquus

Ctenochasmatoidea

Gallodactylidae

	Cycnorhamphus suevicus

	Normannognathus wellnhoferi

Ctenochasmatidae

References

^ Buffetaut, E., Lepage, J.-J., and Lepage, G. (1998). A new pterodactyloid pterosaur from the Kimmeridgian of the Cap de la Hève (Normandy, France). Geological Magazine 135(5):719–722.
^ Witton, Mark P.; O’Sullivan, Michael; Martill, David M. (2015-05-08). Laurin, M. (ed.). "The relationships of Cuspicephalus scarfi Martill and Etches, 2013 and Normannognathus wellnhoferi Buffetaut et al., 1998 to other monofenestratan pterosaurs". Contributions to Zoology. 84 (2): 115–127. doi:10.1163/18759866-08402002. ISSN 1383-4517.
^ Longrich, N.R.; Martill, D.M.; Andres, B.B. (2018). "Late Maastrichtian pterosaurs from North Africa and mass extinction of Pterosauria at the Cretaceous-Paleogene boundary". PLOS Biology. 16 (3): e2001663. doi:10.1371/journal.pbio.2001663. PMC 5849296 Freely accessible. PMID 29534059