''Normannognathus'' is a
genus
Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
pterosaur
Pterosaurs are an extinct clade of flying reptiles in the order Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 million to 66 million years ago). Pterosaurs are the earli ...
from the
Kimmeridgian
In the geologic timescale, the Kimmeridgian is an age in the Late Jurassic Epoch and a stage in the Upper Jurassic Series. It spans the time between 154.8 ±0.8 Ma and 149.2 ±0.7 Ma (million years ago). The Kimmeridgian follows the Oxfordian ...
-age
Upper Jurassic
The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 161.5 ± 1.0 to 143.1 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987.
In European lithostratigraphy, the name ...
Argiles d'Octeville Formation of
France
France, officially the French Republic, is a country located primarily in Western Europe. Overseas France, Its overseas regions and territories include French Guiana in South America, Saint Pierre and Miquelon in the Atlantic Ocean#North Atlan ...
. Initially, ''Normannognathus'' was classified to the family
Germanodactylidae
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 define ...
, sister taxon to ''
Germanodactylus
''Germanodactylus'' ("German finger") is a genus of germanodactylid pterodactyloidea, pterodactyloid pterosaur from Upper Jurassic-age rocks of Germany, including the Solnhofen Limestone. Its specimens were long thought to pertain to ''Pterodacty ...
'', however, many recent analysis have recovered ''Normannognathus'' in different
phylogenetic
In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...
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
''Cycnorhamphus'' (meaning "swan beak") is a genus of gallodactylid ctenochasmatoid pterosaur from the Late Jurassic period of France and Germany, about 152 million years ago. It is synonymous with the genus ''Gallodactylus''.
History
In 1855, ...
'' within the family
Gallodactylidae
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 pre ...
.
Discovery and naming
In 1993, Jean-Jacques Lepage on the Normandy coast at the
Cap de la Hève, near
Ecqueville,
Octeville-sur-Mer
Octeville-sur-Mer (, literally ''Octeville on Sea'') is a commune in the Seine-Maritime department, Normandy, northern France. It is twinned with Bourne End (Bucks) in United Kingdom since 2003 and with Furci Siculo (Sicily) in Italy since 2010. ...
,
Seine-Maritime
Seine-Maritime () is a department of France in the Normandy region of northern France. It is situated on the northern coast of France, at the mouth of the Seine, and includes the cities of Rouen and Le Havre. Until 1955 it was named Seine-Inf� ...
,
Normandy
Normandy (; or ) is a geographical and cultural region in northwestern Europe, roughly coextensive with the historical Duchy of Normandy.
Normandy comprises Normandy (administrative region), mainland Normandy (a part of France) and insular N ...
, found a ten centimetres long
fossil
A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserve ...
of a pterosaur in a marine claystone layer.
In 1998,
Eric Buffetaut
The given name Eric, Erich, Erikk, Erik, Erick, Eirik, or Eiríkur is derived from the Old Norse name ''Eiríkr'' (or ''Eríkr'' in Old East Norse due to monophthongization).
The first element, ''ei-'' may be derived from the older Proto-Nor ...
''et al.'' named a separate genus for it. The
type species
In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...
is ''Normannognathus wellnhoferi''. The genus name is derived from ''Normannia'', the Medieval Latin name for Normandy, and Greek ''gnathos'', "jaw". The
specific name Specific name may refer to:
* in Database management systems, a system-assigned name that is unique within a particular database
In taxonomy, either of these two meanings, each with its own set of rules:
* Specific name (botany), the two-part (bino ...
honors
Peter Wellnhofer
Peter Wellnhofer (born Munich, 1936) is a German paleontologist at the Bayerische Staatssammlung fur Paläontologie in Munich. He is best known for his work on the various fossil specimens of ''Archaeopteryx'' or "Urvogel", the first known bird. ...
. The genus is based on
holotype
A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
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
premaxilla
The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammals h ...
e, formed like a crested wave, having a
concave
Concave or concavity may refer to:
Science and technology
* Concave lens
* Concave mirror
Mathematics
* Concave function, the negative of a convex function
* Concave polygon
A simple polygon that is not convex is called concave, non-convex or ...
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
A tooth (: teeth) is a hard, calcified structure found in the jaws (or mouths) of many vertebrates and used to break down food. Some animals, particularly carnivores and omnivores, also use teeth to help with capturing or wounding prey, tea ...
count is five per
premaxilla
The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammals h ...
; the number is at least nine for the
maxilla
In vertebrates, the maxilla (: maxillae ) is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The two maxil ...
, and at least fourteen per
dentary
In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla).
The jawbone ...
: 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
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 define ...
because it was most similar to ''
Germanodactylus
''Germanodactylus'' ("German finger") is a genus of germanodactylid pterodactyloidea, pterodactyloid pterosaur from Upper Jurassic-age rocks of Germany, including the Solnhofen Limestone. Its specimens were long thought to pertain to ''Pterodacty ...
'', with the exception that it resembles ''
Dsungaripterus
''Dsungaripterus'' is a genus of dsungaripterid pterosaur which lived during the Early Cretaceous in what is now China and possibly South Korea. Its fossils come from the Wuerho Pterosaur Fauna group (WPF) of the Tugulu Group, comprising sectio ...
'' in having an abrupt beginning to a crest larger than that in ''Germanodactylus'' and possessing an upward pointing snout.
[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.] David Unwin
David (; , "beloved one") was a king of ancient Israel and Judah and the third king of the United Monarchy, according to the Hebrew Bible and Old Testament.
The Tel Dan stele, an Aramaic-inscribed stone erected by a king of Aram-Damas ...
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.
In 2018 however, Longrich ''et al.'' found ''Normannognathus'' to be the sister taxon of ''
Cycnorhamphus
''Cycnorhamphus'' (meaning "swan beak") is a genus of gallodactylid ctenochasmatoid pterosaur from the Late Jurassic period of France and Germany, about 152 million years ago. It is synonymous with the genus ''Gallodactylus''.
History
In 1855, ...
'' within the family
Gallodactylidae
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 pre ...
, which in turn was within the Ctenochasmatoidea. Their cladogram is shown below.
[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. . PMC 5849296 Freely accessible. ]
See also
*
List of pterosaur genera
This list of pterosaurs is a comprehensive listing of all Genus, genera that have ever been included in the order Pterosauria, excluding purely vernacular terms. The list includes all commonly accepted genera, but also genera that are now considere ...
*
Timeline of pterosaur research
This timeline of pterosaur research is a chronologically ordered list of important fossil discoveries, controversies of interpretation, and Biological taxonomy, taxonomic revisions of pterosaurs, the famed flying reptiles of the Mesozoic Era (ge ...
References
External links
''Normannognathus''in The Pterosauria
{{Portal bar, Paleontology, France
Monofenestrata
Late Jurassic pterosaurs of Europe
Fossil taxa described in 1998
Taxa named by Éric Buffetaut