Cycnorhamphus
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''Cycnorhamphus'' (meaning "
swan Swans are birds of the family Anatidae within the genus ''Cygnus''. The swans' closest relatives include the geese and ducks. Swans are grouped with the closely related geese in the subfamily Anserinae where they form the tribe Cygnini. Som ...
beak") is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
of gallodactylid ctenochasmatoid
pterosaur Pterosaurs (; from Greek ''pteron'' and ''sauros'', meaning "wing lizard") is 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 ...
from the
Late Jurassic The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 163.5 ± 1.0 to 145.0 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987. In European lithostratigraphy, the name ...
period of
France France (), officially the French Republic ( ), is a country primarily located in Western Europe. It also comprises of Overseas France, overseas regions and territories in the Americas and the Atlantic Ocean, Atlantic, Pacific Ocean, Pac ...
and
Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwee ...
, about 152 million years ago. It is probably synonymous with the genus ''Gallodactylus''.


History

In 1855, a
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 preserved ...
in a plate of
shale Shale is a fine-grained, clastic sedimentary rock formed from mud that is a mix of flakes of clay minerals (hydrous aluminium phyllosilicates, e.g. kaolin, Al2 Si2 O5( OH)4) and tiny fragments (silt-sized particles) of other minerals, especiall ...
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 157.3 ± 1.0 Ma and 152.1 ± 0.9 Ma (million years ago). The Kimmeridgian follows the Oxford ...
, found near Nusplingen in
Württemberg Württemberg ( ; ) is a historical German territory roughly corresponding to the cultural and linguistic region of Swabia. The main town of the region is Stuttgart. Together with Baden and Hohenzollern, two other historical territories, Württ ...
,
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of seve ...
GPIT "Orig. Quenstedt 1855, Taf. 1" or GPIT 80, was named '' Pterodactylus Suevicus'' by
Friedrich August Quenstedt Friedrich August von Quenstedt (July 10, 1809 – December 21, 1889), was a German geologist and palaeontologist. Life Von Quenstedt was born at Eisleben in Saxony, and educated at the Humboldt University of Berlin. After a period as assistant ...
. The specific name refers to the tribal area of Suevia. Quenstedt had earlier mentioned the find in a letter to Professor
Heinrich Georg Bronn Heinrich Georg Bronn (3 March 1800 – 5 July 1862) was a German geologist and paleontologist. He was the first to translate Charles Darwin's ''On the Origin of Species'' into German in 1860, although not without introducing his own interpretation ...
, which was published in 1854. In it he used the name ''Pterodactylus Württembergicus''. In 1855 and 1858,
Christian Erich Hermann von Meyer Christian Erich Hermann von Meyer (3 September 1801 – 2 April 1869), known as Hermann von Meyer, was a German palaeontologist. He was awarded the 1858 Wollaston medal by the Geological Society of London. Life He was born at Frankfurt am ...
adopted this older species name but it would be forgotten afterwards. The publication in 1854 was not meant to be a nomenclatural act. According to Peter Wellnhofer, ''Pterodactylus württembergicus'' should be considered a ''
nomen oblitum In zoological nomenclature, a ''nomen oblitum'' (plural: ''nomina oblita''; Latin for "forgotten name") is a disused scientific name which has been declared to be obsolete (figuratively 'forgotten') in favour of another 'protected' name. In its p ...
''.Wellnhofer, P. 1970. "Die Pterodactyloidea (Pterosauria) der Oberjura-Plattenkalke Süddeutschlands", ''Bayerische Akademie der Wissenschaften, mathematisch-wissenschaftlichen Klasse, Abhandlungen'', 141 : 1-133 In 1858, Johann Andreas Wagner described a second specimen consisting of the wings, a shinbone and a foot. He named it ''Pterodactylus (Ornithocephalus) eurychirus'', "the broad-handed", but later in the same publication used the name ''Pterodactylus suevicus eurychirus'' as if it were a
subspecies In biological classification, subspecies is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics ( morphology), but that can successfully interbreed. Not all specie ...
. This is today considered a junior synonym. The specimen was acquired by the ''Bayerische Staatssammlung für Paläontologie und Geologie'' but was lost in April 1944 during the Munich bombardment. In 1870, Harry Govier Seeley assigned ''P. suevicus'' to a new genus: ''Cycnorhamphus''. The name is derived from Greek κύκνος, ''kyknos'', "swan" and ράμφος, ''ramphos'', "beak", in reference to the snout shape. The
type species In 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 species that contains the biological type specim ...
of this genus is ''Pterodactylus suevicus'', the ''
combinatio nova ''Combinatio nova'', abbreviated ''comb. nov.'' (sometimes ''n. comb.''), is Latin for "new combination". It is used in taxonomic biology literature when a new name is introduced based on a pre-existing name. The term should not to be confused w ...
'' is ''Cycnorhamphus suevicus''. In 1878,
Oscar Fraas Oscar Friedrich von Fraas (17 January 1824, in Lorch (Württemberg) – 22 November 1897, in Stuttgart) was a German clergyman, paleontologist and geologist. He was the father of geologist Eberhard Fraas (1862–1915). Biography He studied ...
referred a specimen of ''
Pterodactylus longicollum ''Ardeadactylus'' (from ''Ardea'' – meaning "heron", and also a name of a genus of herons – and ''dactylus'', meaning "finger") is an extinct genus of ctenochasmatoid pterosaur known from the Late Jurassic Solnhofen limestone of Bavaria, ...
'', the later ''Exemplar Nr 58'', to ''Pterodactylus suevicus''. Seeley made this the
type specimen In biology, a type is a particular specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally attached. In other words, a type is an example that serves to anchor or centralizes th ...
of yet another species: ''Cycnorhamphus Fraasii'', in 1891. In 1907 however, Felix Plieninger rejected the split between ''Pterodactylus'' and ''Cycnorhamphus'' and denied the validity of ''C. fraasii''. This would be the standard interpretation, shared by most paleontologists, for over sixty years. During the late 1960s, the Ghirardi family began to exploit the chalkstone quarries of Les Besson, located on the French army base of Canjuers near
Aiguines Aiguines (; oc, Aiguina) is a commune in the Var department in the Provence-Alpes-Côte d'Azur region in southeastern France. Geography Climate Aiguines has a hot-summer Mediterranean climate ( Köppen climate classification ''Csa'') closely b ...
. A ''
Lagerstätte A Lagerstätte (, from '' Lager'' 'storage, lair' '' Stätte'' 'place'; plural ''Lagerstätten'') is a sedimentary deposit that exhibits extraordinary fossils with exceptional preservation—sometimes including preserved soft tissues. These f ...
'' proved to be present from which many high quality fossils could be collected. One of these was a slab showing a pterosaur. The precise time and location of this discovery are unknown. It was first reported in the scientific literature in 1971, by Léonard Ginsburg and Guy Mennessier. In 1974, French paleontologist Jacques Fabre based on this specimen, MNHN CNJ-71, named a new species ''Gallodactylus canjuersensis''. The generic name combines a reference to Gallia with Greek ''daktylos'', "finger". He concluded it was the same genus as ''P. suevicus'', but did not revive ''Cycnorhamphus'', judging that the latter name was unavailable because of mistakes in the diagnosis by Seeley, already pointed uit by Plieninger. ''P. suevicus'' thus became ''Gallodactylus suevicus''. In 1976, Fabre again named the species, describing it in greater detail but not mentioning the earlier publication. This confused later researchers who mistakenly assumed that 1976 was the formal naming date. In fact the 1974 paper contains a sufficient description and the species was validly named that year. In 1983, the Ghirardis sold their entire collection to the
National Museum of Natural History, France The French National Museum of Natural History, known in French as the ' (abbreviation MNHN), is the national natural history museum of France and a ' of higher education part of Sorbonne Universities. The main museum, with four galleries, is lo ...
. However, in 1996, Christopher Bennett pointed out that such mistakes do not invalidate a name and that therefore ''Cycnorhamphus'' has priority, making ''Gallodactylus canjuersensis'' ''C. canjuersensis''. In 2010 and 2012, Bennett published further re-studies of the fossils, concluding that the differences between the two species could be explained by age, sex or individual variation, and formally synonymized ''C. canjuersensis'' and ''C. suevicus''.


Description

''Cycnorhamphus'' had historically been assumed to have had long jaws with teeth at the very tip, akin to those of '' Pterodactylus antiquus''. However, recent work on a specimen nicknamed "The Painten Pelican" has revealed that the animal possesses a very unusual jaw anatomy, with peg-like teeth at the jaw tips - blunter and stouter in older individuals -, jaw curvatures behind said teeth that form angled arcs away from the biting surface, forming thus an opening, and two poorly understood soft tissue structures occupying this opening from the upper jaw, showing mineralization. The purpose of these adaptations is unknown, but they are more obvious and well developed in adult animals. It has been speculated that the jaws functioned similar to those of openbill storks, allowing the animal to hold hard invertebrates like mollusks and either crush or bisect them.Witton, Mark P. (2013), ''Pterosaurs: Natural History, Evolution, Anatomy''


Classification

As illustrated below, the results of a
topology In mathematics, topology (from the Greek words , and ) is concerned with the properties of a geometric object that are preserved under continuous deformations, such as stretching, twisting, crumpling, and bending; that is, without closing ...
are based on a phylogenetic analysis made by Longrich, Martill, and Andres in 2018. They placed the ''Cycnorhamphus'' within the clade Euctenochasmatia, more precisely within the family Gallodactylidae, sister taxon to ''
Normannognathus ''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'', ...
''.


See also

* List of pterosaur genera * Timeline of pterosaur research


References

{{Portal bar, Paleontology, Germany, France Late Jurassic pterosaurs of Europe Ctenochasmatoids Taxa named by Harry Seeley Fossil taxa described in 1870