| Cycnorhamphus | |
|---|---|
 | |
| Juvenile specimen | |
Scientific classification | |
| Domain: | Eukaryota |
| Kingdom: | Animalia |
| Phylum: | Chordata |
| Order: | †Pterosauria |
| Suborder: | †Pterodactyloidea |
| Clade: | †Ctenochasmatoidea |
| Family: | †Gallodactylidae |
| Genus: | †Cycnorhamphus Seeley, 1870 |
| Species: | †C. suevicus |
| Binomial name | |
| †Cycnorhamphus suevicus (Quenstedt, 1855) | |
| Synonyms | |
Genus synonymy
Species synonymy
| |
Cycnorhamphus (meaning "swan beak") is agenus ofgallodactylidctenochasmatoidpterosaur from theLate Jurassic period ofFrance andGermany, about 152 million years ago.[1] It is synonymous with the genusGallodactylus.
In 1855, afossil in a plate ofshale from theKimmeridgian, found nearNusplingen inWürttemberg,holotype GPIT "Orig. Quenstedt 1855, Taf. 1" orGPIT 80, was namedPterodactylus suevicus byFriedrich August Quenstedt.[2][3][4] The specific name refers to the tribal area ofSuevia. Quenstedt had earlier mentioned the find in a letter to ProfessorHeinrich Georg Bronn, which was published in 1854. In it he used the namePterodactylus württembergicus.[5] In 1855 and 1858,Christian Erich Hermann von Meyer adopted this older species name[6] but it would be forgotten afterwards. The publication in 1854 was not meant to be a nomenclatural act.[7] According toPeter Wellnhofer,Pterodactylus württembergicus should be considered anomen oblitum.[8]
In 1858,Johann Andreas Wagner described a second specimen consisting of the wings, a shinbone and a foot. He named itPterodactylus (Ornithocephalus) eurychirus, "the broad-handed", but later in the same publication used the namePterodactylus suevicus eurychirus as if it were asubspecies.[9] This is today considered a junior synonym. The specimen was acquired by theBayerische Staatssammlung für Paläontologie und Geologie but was lost in April 1944 during theMunich bombardment.[8]
In 1870,Harry Govier Seeley assignedP. suevicus to a new genus:Cycnorhamphus.[10] The name is derived from Greek κύκνος,kyknos, "swan" and ράμφος,ramphos, "beak", in reference to the snout shape. Thetype species of this genus isPterodactylus suevicus, thecombinatio nova isCycnorhamphus suevicus.
In 1878,Oscar Fraas referred a specimen ofPterodactylus longicollum, the laterExemplar Nr 58, toPterodactylus suevicus.[11] Seeley made this thetype specimen of yet another species:Cycnorhamphus Fraasii, in 1891.[12]
In 1907 however,Felix Plieninger rejected the split betweenPterodactylus andCycnorhamphus and denied the validity ofC. fraasii.[13] 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 nearAiguines. ALagerstätte 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, byLéonard Ginsburg and Guy Mennessier.[14] In 1974, French paleontologistJacques Fabre based on this specimen, MNHN CNJ-71, named a new speciesGallodactylus canjuersensis.[15] The generic name combines a reference toGallia with Greekdaktylos, "finger". He concluded it was the same genus asP. suevicus, but did not reviveCycnorhamphus, judging that the latter name was unavailable because of mistakes in the diagnosis by Seeley, already pointed uit by Plieninger.P. suevicus thus becameGallodactylus suevicus. In 1976, Fabre again named the species, describing it in greater detail but not mentioning the earlier publication.[16] 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.[7] In 1983, the Ghirardis sold their entire collection to theNational Museum of Natural History, France.
However, in 1996, Christopher Bennett pointed out that such mistakes do not invalidate a name and that thereforeCycnorhamphus has priority, makingGallodactylus canjuersensisC. canjuersensis.[4] 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 synonymizedC. canjuersensis andC. suevicus.[7]
Cycnorhamphus had historically been assumed to have had long jaws with teeth at the very tip, akin to those ofPterodactylus antiquus. However, recent work on a specimen nicknamed "The Painten Pelican"[7] 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,[17] but they are more obvious and well developed in adult animals. It has been speculated that the jaws functioned similar to those ofopenbill storks, allowing the animal to hold hard invertebrates like mollusks and either crush or bisect them.[18]
As illustrated below, the results of atopology are based on a phylogenetic analysis made by Longrich, Martill, and Andres in 2018. They placed theCycnorhamphus within the cladeEuctenochasmatia, more precisely within the familyGallodactylidae, sister taxon toNormannognathus.[19]
