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Cimoliopterus Temporal range: Albian–Cenomanian ~112–94 Ma PreꞒ Ꞓ O S D C P T J K Pg N
Holotype snout tip of C. cuvieri shown from the right side and below
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Order:	†Pterosauria
Suborder:	†Pterodactyloidea
Clade:	†Ornithocheirae
Clade:	†Targaryendraconia
Family:	†Cimoliopteridae
Genus:	†Cimoliopterus Rodrigues & Kellner, 2013
Type species
†Pterodactylus cuvieri Bowerbank, 1851
Species
†C. cuvieri (Bowerbank, 1851) †C. dunni Myers, 2015
Synonyms
Synonyms of C. cuvieri Pterodactylus cuvieri Bowerbank, 1851 Ornithocheirus cuvieri Seeley, 1870 Coloborhynchus cuvieri Owen, 1874 Anhanguera cuvieri Bakhurina & Unwin, 1995 Ornithocheirus brachyrhinus? Seeley, 1870 Pterodactylus fittoni? Owen, 1859 Ornithocheirus fittoni? Seeley, 1870 Anhanguera fittoni? Unwin, 2001

Cimoliopterus is a genus of pterodactyloid pterosaur from the Cretaceous of England, United Kingdom and Texas, United States.

The type species, Cimoliopterus cuvieri, was previously considered parts of several different genera depending on author, but received its own genus in 2013.

History of discovery

The history of Cimoliopterus and related pterosaurs from the UK is considered confusing, since many of them were described from very fragmentary remains, and named using convoluted nomenclatural schemes of the Victorian era. In 1851, the British naturalist James Scott Bowerbank described a large pterosaur snout he had obtained, which was found in a pit in the Chalk Formation at Burham, Kent, in South East England. Pterosaur fossils had earlier been discovered in the same pit, including the front part of some jaws Bowerbank had used as the basis for the species Pterodactylus giganteus in 1846, as well as other bones. Based on the new snout, Bowebank named the new species Pterodactylus cuvieri; at this time, the genus Pterodactylus was used formany different pterosaur species which are now thought to be distantly related to each other. The specific name honours the French palaeontologist Georges Cuvier, who recognised that pterosaurs were flying reptiles.

The holotype snout, today catalogued as NHMUK PV 39409 at the Natural History Museum in London, consists of the front 18 cm (7 in) of a partial upper jaw, represented entirely by the premaxillae, the frontmost bones of the upper jaw, including part of a premaxillary crest. Eleven alveoli (tooth sockets) are preserved on each side. It was originally reported to preserve a single tooth in the right first alveolus (tooth socket at the front of the snout), but this had disappeared when the holotype was examined in 2007 and 2009. Two complete teeth were also originally reported to be preserved in the same block of chalk as the snout. Bowerbank also believed some large bones in three other collections may either have belonged to the same species, to P. giganteus, or to a third possible species. The British biologist Richard Owen assigned one long bone to P. cuvieri based on its size and the character of its surface, and identified it as a possible phalanx bone of the elongated wing-finger.

In the 1850s the British artist Benjamin Waterhouse Hawkins created full-sized sculptures of prehistoric animals for the Crystal Palace in London, under the supervision of Owen. Among them were two species of pterosaur, including two P. cuvieri sculptures made of dark stone, and two smaller statues of Pterodactylus bucklandi. The larger statues are today in disrepair due to their delicate nature and vandalism, and the smaller ones have been lost. The British palaeontologist Mark Witton stated in 2019 that the anatomy of the sculptures was probably based on the by then more completely known Pterodactylus antiquus. They were depicted with scales, though pterosaurs are now known to have been covered in hair-like pycnofibres, and they have rather bird-like proportions (as was customary at the time), such as small head and large torsos.

In 1869, Harry Govier Seeley placed P. cuvieri in the new genus Ptenodactylus along with other British pterosaurs mainly known from upper jaws, in an index of specimens in the Woodwardian Museum. He noted that these names were provisional, only intended for the conveniewce of students using the museum, and not necessarily as names for these species. In 1870, Seeley placed the species in the genus Ornithocheirus, which he had originally used for other species in his 1869 index, and figured a dentary bone that he listed as belonging to O. cuvieri. In 1874, Owen assigned the species to the genus Coloborhynchus. The British palaeontologist Reginald Walter Hooley used the name O. cuvieri in his 1914 revision of Ornithocheirus, using Seeley's terminology.

In 1919 and 1922, the Austrian naturalist Gustav von Arthaber lamented that the scientific literature had accepted the many Ornithocheirus names that had only been mentioned in a catalogue made by Seeley for the benefit of students. In his opinion names were worthless without an illustration of the specimens they were based on, or better still a complete restoration of the relevant skeleton. For this reason, he provided a drawing of the skull of Ornithocheirus cuvieri, one of the few species for which the known jaw material proved its validity.

In 1987, the German palaeontologist Peter Wellnhofer described the new crested pterosaur Tropeognathus from Brazil and noted the similarities between it and other newly described Brazilian taxa such as Anhanguera to British taxa that were based on fragmentary snouts, such as the various species assigned to Ornithocheirus. He concluded that while the appearance of the British taxa had long been a puzzle, the discovery of the related Brazilian species made this clearer. In 1991, Wellnhofer stated that the genus Ornithocheirus had become something of a wastebin of species from different countries, assigned to it on the basis of insufficient characters, and needed clearer defitition, and perhaps included several distinct genera.

In 1995, the Russian palaeontologist Natalia N. Bakhurina and the British palaeontologist David M. Unwin assigned a partial mandibular symphysis (the part connecting the two halves of the lower jaw) from Russia to Anhanguera cf. Anhanguera cuvieri (possibly belonging to this or a related animal), thereby using a new combination. In 2000, assigned pterosaur skeletal elements from the Toolebuc Formation of Australia to Anhanguera ?cuvieri. In 2001, Unwin also used the combination A. cuvieri, and considered it the most common pterosaur of the Cambridge Greensand, represented by 23 jaw fragments. He concluded that the species O. brachyrhinus, O. colorhinus, O. dentatus, O. denticulatus, O. enchorhynchus, O. scaphorhynchus and O. xyphorhynchus were junior synonyms of A. cuvieri. He also considered another Greensand species, originally named Pterodactylus fittoni by Owen in 1859, as a member of Anhanguera. In 2011, the Brazilian palaeontologists Alexander W. Kellner, Taissa Rodrigues, and Fabiana R. Costa made the Australian material that had previously been assigned to A. ?cuvieri the basis of a new genus, Aussiedraco, and stated that they saw no ground to place O. cuvieri in Anhanguera.

New genus and assigned species

In 2013, Rodrigues and Kellner reviewed the contents of the genus Ornithocheirus and other British Cretaceous pterosaurs. They concluded that A. cuvieri differed enough from its relatives to be placed in a new, monotypic genus; Cimoliopterus. This generic name combines the Greek words Kimolia (Κιμωλία), which means "chalk", and pteron (πτερόν) which means "wing". Rodrigues and Kellner also found that while the species P. fittoni, O. brachyrhinus, and O. enchorhynchus had various features in common with C. cuvieri, and could therefore not be excluded from that genus, they were too fragmentary to be definitely assigned to it, and were considered nomina dubia (dubious names). Other species that were assigned to A. cuvieri by Unwin in 2001 were also instead considered dubious or to belong to other taxa by them. They also considered the Russian specimen assigned to A. cf. A. cuvieri in 1995 unrelated.

In 2013, a pterosaur snout fragment was discovered by the American amateur fossil collector Brent Dunn in the Britton Formation near Lewisville Lake, northwest of Dallas, in Texas, USA. Dunn donated the specimen to the Shuler Museum of Paleontology of Southern Methodist University. In 2015, the specimen (catalogued as SMU 76892) was described by the American palaeontologist Timothy S. Myers who made it the holotype of the new species Cimoliopterus dunni; the specific name honours Dunn, who died in 2013. The holotype specimen consists of the front 18.5 cm (7.3 in) of the snout, comprising the premaxilla and maxilla (including a largely intact premaxillary crest), broken off just behind the 13th pair of alveoli. The fossil was initially covered by a thin layer of red clay, which was prepared away. The three-dimensionally preserved fossil does not appear to have suffered the extreme crushing often seen in pterosaur fossils, though some of the hind part was distorted by compaction.

Description

The size of Cimoliopterus is hard to estimate since it is currently only knwon from fragments of the skull. Extrapolating from the remains of another species, P. giganteus, from the same formation as Cimoliopterus cuvieri (the type species of Cimoliopterus), Bowerbank, the original describer of C. cuvieri, estimated a wingspan for P. cuvieri of about 5.1 metres (16.6 ft), making it the largest flying animal, extinct or extant, known in 1851. This claim at the time was met with considerable scepticism, though today, larger pterosaurs are known to have existed. In 2001, Unwin, comparing the remains of C. cuvieri with more complete remains of the pterosaur Anhanguera from the Santana Formation of Brazil, estimated a wingspan of about 3.5 metres (11 ft), smaller than what Bowerbank estimated it. An assigned specimen, CAMSM B54.431, was estimated to reach a winspan measuring up to 5 metres (16 ft), however, this specimen was reassigned to the species Camposipterus colorhinus in 2013. As a pterosaur, Cimoliopterus would have been covered in hair-like pycnofibres, and had extensive wing-membranes, which were distended by a long wing-finger.

The two species of Cimoliopterus, C. cuvieri and C. dunni, do not show the pronounced lateral front expansion (a spoon-like expansion) at the tip of the rostrum, this feature is a key characteristic of a lot of other toothed pteranodontoids. In both species, this lateral rostral expansion is developed so weakly that it had even been considered to be absent once, and in comparison to the pronounced rostral expansions of other pterosaurs such as those of Anhanguera, Coloborhynchus, and Ornithocheirus, the rostral expansions seen in both C. cuvieri and C. dunni bear so little resemblance to them. This is also why C. cuvieri can be distinguished from the three former genera; it has a low rostrum compared to the genus Ornithocheirus, it also possess a forward-facing first pair of alveoli, unlike Ornithocheirus; C. cuvieri can furthermore be distinguished from the genus Anhanguera because it does not possess an anterior expansion of the rostrum, as mentioned before, which is seen in members of the family Anhangueridae, it also does not have the fourth and fifth alveoli at a smaller size than the third and sixth, which is a key feature in Anhanguera. Due to the more posteriorly positioned premaxillary crest of Cimoliopterus cuvieri, unlike the premaxillary crests seen in anhanguerids, it may perhaps indicate that the crests that C. cuvieri bore had evolved in a separate way.

C. cuvieri

The holotype of the species Cimoliopterus cuvieri consists of an incomplete upper jaw, which is narrow in the preserved portion. The crest of C. cuvieri is, on the premaxilla (the frontmost cranial bone), placed far back, at the level of the seventh tooth pair, however, it begins before the nasoantorbital fenestra (the skull opening where the antorbital fenestra and the bony nostril are combined). In the anterior (front) part of the jaw of C. cuvieri there are three alveoli (tooth sockets) per 3 centimetres (1.2 in) of jaw margin, and two alveoli per 3 centimetres (1.2 in) of jaw margin at the back of the jaw. The front portion of the ridge on the midline of the palate (upper part of the mouth) reaches the level of the third tooth pair, the palate itself is curved upwards and there is no expansion of the jaw at the front. The second tooth pair is about the same size as the third, and both of them are larger than the fourth tooth pair. To the rear part of the jaw, the distance between the teeth gradually increases.

Other pterosaur species have also been tentatively assigned to Cimoliopterus cuvieri due to the possession of similar features and characteristics, however, most of them were too fragmental to be unquestionably assigned to C. cuvieri. The species Pterodactylus fittoni for example, shares with C. cuvieri a low snout tip, the absence of an anterior expansion at the end of the snout, a dorsally curved palate, as well as a palatal ridge extending back to the third pair of tooth sockets, the lattermost feature however, would only be shared with C. cuvieri if the first preserved pair of alveoli in P. fittoni is actually its second pair. The height of P. fittoni's snout can be differentiated from C. cuvieri's in being, and its tip is wider than higher; the latter difference could be due to the fracture of the snout tip of P. fittoni. Due to its fragmentary remains, P. fittoni cannot be definitely assigned to C. cuvieri. Another species, Ornithocheirus brachyrhinus, shares a number of features with C. cuvieri, this includes its palate being curved, the front end being higher than wider, an anterior expansion of the snout, and the absence of an anteriorly positioned crest. The structure of O. brachyrhinus' snout tip perfectly corresponds to the structure of C. cuvieri's snout tip, these characteristics suggest that O. brachyrhinus is referable to C. cuvieri. However, CAMSM B54443 (the holotype of O. brachyrhinus), is fragmentary and thus it is unfeasible to establish if O. brachyrhinus had a crest that is located posteriorly, or if it had the size variation of tooth sockets that is diagnostic in Cimoliopterus cuvieri.

Other species, including O. dentatus and O. enchorhynchus have also been referred to C. cuvieri. However, the former differs by the presence of smaller alveoli that are placed closer together, while the latter, even though quite similar to C. cuvieri (sharing features including the absence of anteriorly positioned crest, the palate dorsally curved, the first pair of teeth facing forward, as well as the absence of an anterior rostral expansion), is, like O. brachyrhinus, fragmentary, which makes it hard to compare to other pterosaur species such as C. cuvieri. The species O. scaphorhynchus has also been tentatively assigned to Cimoliopterus cuvieri, but again, the incompleteness of the species makes it difficult to be referred with certainty to any known genus. The margin of the tooth sockets in O. scaphorhynchus is poorly preserved, but it does not seem to fit the spacing pattern present in the holotype specimen of C. cuvieri.

C. dunni

The holotype of the species C. dunni, SMU 76892, is composed of 18.5 centimetres (7.3 in) of the anterior portion of the rostrum (snout), and comprises both the premaxilla and the maxilla, which appear to have been broken until the thirteeth pair of alveoli. The total number of alveoli found in the specimen is twenty-six. The premaxillary crest of C. dunni begins just above the fourth alveoli, and it slightly curves upward, this forms a front edge that is concave. The upper edge of the crest seems to descend moderately just before a broken portion, this suggests that the premaxillary crest of C. dunni was, in a lateral view, symmetrical. Supposing that the pterosaur's crest was symmetrical in shape, the complete crest of it would have had a length that was approximately 15 to 16 centimetres (5.9 to 6.3 in). The maximum height of the premaxillary crest of C. dunni is 38 millimetres (1.5 in), the region of the crest where this height is reached is located above the ninth and tenth alveoli. The cortical bone (the hard outer layer of bones) is well-preserved in C. dunni, however, on the left side of the crest, the conjoinment between premaxilla and maxilla is not visible, and on the right side there are several regions where cortical bone is either damaged or missing. The tip of the snout in C. dunni is blunt, and the rostrum has an angle of 45° respect to the anterior part of the palate.

An upward angle of 8 degrees relative to the flat area of the rear part of the palate can be assumed based on its front portion being dorsally reflected, and an inflection point (the point where a curve changes from convex to concave) close to the level of the eighth alveoli can also be seen. At less than 1 millimetre (0.039 in) high, a narrow palatal ridge extends anteriorly from the broken portion on posterior end of the premaxillary crest. In palatal view of the palate, the anterior end of the rostrum of C. dunni posteriorly expands to a maximum width of 1.6 centimetres (0.63 in) above the third alveoli, narrowing to a minimum width of 1.5 centimetres (0.59 in) at the level of the fourth alveoli. The width of the rostrum continues to increase afterwards, until it reaches a maximum of 1.8 to 1.9 centimetres (0.71 to 0.75 in) at the broken part of the posterior edge of the crest. The spaces between the alveoli of C. dunni gradually increase backward along the row of teeth of the pterosaur, the minimum between the spaces measuring 1.6 millimetres (0.063 in), and the maximum measuring 11.5 millimetres (0.45 in). In total, there are thirteen pairs of alveoli, this is indicates a minimum of twenty-six teeth in the upper jaw.

Classification

Cimoliopterus was by Rodrigues and Kellner (and Myers in 2015) assigned to the Pteranodontoidea, in an uncertain position (incertae sedis). Cladistic analyses indicated a position in the evolutionary tree above Istiodactylus but below a polytomy formed by more derived forms.

In 2018, Jacobs et al. performed a phylogenetic analysis where they placed Cimoliopterus within the family Ornithocheiridae, as the sister taxon of Camposipterus. They published their analysis in 2019. In the same year however, Pêgas et al. reassigned Cimoliopterus to the clade Targaryendraconia, more specifically to its own family, the Cimoliopteridae, and sister taxon to both Aetodactylus and Camposipterus.

C. dunni dates to the Cenomanian stage, of the early Late Cretaceous. This find extends the distribution of the genus to North America and shows that the pterosaur faunas from Europe and North America still had contact during the middle Cretaceous. The phylogenetic analysis included in the description of this species indicates that Cimoliopterus is a basal pteranodontoid closely related with Aetodactylus, which has also been found in Cenomanian strata from Texas.

Palaeobiology

Diet

The tooth-arrangement seen in the group Cimoliopterus belongs to has been referred to as "fish-grabs".

Wellnhofer stated in 1987 that while the crests of pterosaurs such as Pteranodon and Dsungaripterus were probably used as aerodynamic rudders or as areas for muscle attachment, the crests on the front ends of the jaws of Tropeognathus and its relatives could not serve this purpose. He instead proposed that the terminal crests have functioned as the keel of a boat, to stabilise the jaws in water while skimming and catching fish with the jaws submerged at high speed. The shape of the keel would reduce flow-resistance, keeping the head in a stable position, and would therefore have a hydronynamic rather than aerodynamic function.

Palaeoenvironment

The holotype of C. cuvieri, NHMUK PV 39409, had been found in the Grey Chalk Subgroup dating from the Cenomanian-Turonian.

See also

• List of pterosaur genera
• Timeline of pterosaur research

References

1. ^ Myers, Timothy S. (2015). "First North American occurrence of the toothed pteranodontoid pterosaur Cimoliopterus". Journal of Vertebrate Paleontology. 35 (6): e1014904. doi:10.1080/02724634.2015.1014904. S2CID 86099117.
2. ^ Rodrigues, T.; Kellner, A. (2013). "Taxonomic review of the Ornithocheirus complex (Pterosauria) from the Cretaceous of England". ZooKeys (308): 1–112. doi:10.3897/zookeys.308.5559. PMC 3689139. PMID 23794925.{{cite journal}}: CS1 maint: unflagged free DOI (link)
3. ^ Witton 2013, pp. 152–163.
4. ^ Unwin, D. M. (2001). "An overview of the pterosaur assemblage from the Cambridge Greensand (Cretaceous) of Eastern England". Mitteilungen aus dem Museum für Naturkunde in Berlin, Geowissenschaftliche Reihe. 4: 189–221. doi:10.1002/mmng.20010040112.
5. ^ Bowerbank, J. S. (1851). "On the pterodactyles of the Chalk Formation". Proceedings of the Zoological Society of London. 19: 14–20. doi:10.1111/j.1096-3642.1851.tb01125.x.
6. Witton 2013, p. 98.
7. Martill, D. M. (2010). "The early history of pterosaur discovery in Great Britain". Geological Society, London, Special Publications. 343 (1): 287–311. Bibcode:2010GSLSP.343..287M. doi:10.1144/SP343.18. S2CID 130116778.
8. Jobling, J. A. (2010). The Helm Dictionary of Scientific Bird Names. London: Christopher Helm. p. 125. ISBN 978-1-4081-2501-4.
9. ^ Owen, R. (1849–1884). A History of British Fossil Reptiles. Vol. 1. London: Cassell & Company Limited. pp. 242–258. doi:10.5962/bhl.title.7529.
10. Owen, R. (1854). Geology and Inhabitants of the Ancient World. Vol. 8. London: Crystal Palace library. pp. 5–7, 11–13. ISBN 978-1166913045.
11. Witton, M. (2019). "Pterosaur – information about the Crystal Palace statues". cpdinosaurs.org. Retrieved 16 February 2021.
12. Witton, M. (2019). "The science of the Crystal Palace Dinosaurs, part 2: Teleosaurus, pterosaurs and Mosasaurus". Mark Witton.com Blog. Retrieved 16 February 2021.
13. Seeley, H. G. (1869). Index to the fossil remains of Aves, Ornithosauria, and Reptilia, from the secondary system of strata arranged in the Woodwardian Museum of the University of Cambridge. Cambridge,Deighton, Bell, and co. p. 16. doi:10.5962/bhl.title.159380.
14. Seeley, H.G. (1870). The Ornithosauria: an elementary study of the bones of pterodactyls, made from fossil remains found in the Cambridge Upper Greensand, and arranged in the Woodwardian Museum of the University of Cambridge. Cambridge: Deighton, Bell, and Co. p. 113.
15. Owen, R. (1874). A Monograph on the Fossil Reptilia of the Mesozoic Formations. Vol. 1. London: The Palæontographical society. pp. 6–8. doi:10.1017/CBO9781316151099.003.
16. Hooley, R. W. (1914). "LXI.— On the Ornithosaurian genus Ornithocheirus, with a review of the specimens from the Cambridge Greensand in the Sedgwick Museum, Cambridge". Annals and Magazine of Natural History. 13 (78): 529–557. doi:10.1080/00222931408693521.
17. Von Arthaber, G. (1919). "Studien über Flugsaurier auf Grund der Bearbeitung des Wiener Exemplars von Dorygnathus banthensis Theod Sp". Denkschriften der königlichen Akademie der Wissenschaften. Mathematisch-Naturwissenschaftliche Klasse. 97: 391–464.
18. von Arthaber, G. (1922). "Über Entwicklung, Ausbildung und Absterben der Flugsaurier". Paläontologische Zeitschrift. 4: 1–47. doi:10.1007/BF03041557. S2CID 131644821.
19. ^ Wellnhofer, P. (1987). "New Crested Pterosaurs from the Lower Cretaceous of Brazil". Mitteilungen der Bayerischen Staatssammlung für Paläontologie und Histor. Geologie. 27: 175–186.
20. Wellnhofer, P. (1991). The Illustrated Encyclopedia of Pterosaurs. New York: Crescent Books. pp. 108–112. ISBN 978-0-517-03701-0.
21. Bakhurina, N. N.; Unwin, D. M. (1995). "A survey of pterosaurs from the Jurassic and Cretaceous of the former soviet union and Mongolia". Historical Biology. 10 (3): 197–245. doi:10.1080/10292389509380522.
22. Unwin, D. M.; Lü, J.; Bakhurina, N. N. (2000). "On the systematic and stratigraphic significance of pterosaurs from the Lower Cretaceous Yixian Formation (Jehol Group) of Liaoning, China". Fossil Record. 3 (1): 181–206. doi:10.1002/mmng.20000030109.
23. Kellner, A. W. A.; Rodrigues, T.; Costa, F. R. (2011). "Short note on a Pteranodontoid pterosaur (Pterodactyloidea) from western Queensland, Australia". Anais da Academia Brasileira de Ciências. 83 (1): 301–308. doi:10.1590/S0001-37652011000100018. PMID 21437387.
24. Allen, M. (2013). "North America's newest pterosaur is a Texan — and flying reptile's closest cousin is English". SMU Research. Retrieved 21 March 2021.
25. ^ Pêgas, R.V.; Holgado, B.; Leal, M.E.C. (2019). "On Targaryendraco wiedenrothi gen. nov. (Pterodactyloidea, Pteranodontoidea, Lanceodontia) and recognition of a new cosmopolitan lineage of Cretaceous toothed pterodactyloids". Historical Biology: 1–15. doi:10.1080/08912963.2019.1690482.
26. Witton, M.P.; Martill, D.M.; Loveridge, R.F. (2010). "Clipping the Wings of Giant Pterosaurs: Comments on Wingspan Estimations and Diversity". Acta Geoscientica Sinica. 31: 79–81.
27. Witton 2013, pp. 51–52.
28. ^ Jacobs, M.L.; Martill, D.M.; Ibrahim, N.; Longrich, N. (2019). "A new species of Coloborhynchus (Pterosauria, Ornithocheiridae) from the mid-Cretaceous of North Africa" (PDF). Cretaceous Research. 95: 77–88. doi:10.1016/j.cretres.2018.10.018.

Bibliography

• Witton, M. P. (2013). Pterosaurs: Natural History, Evolution, Anatomy (1st ed.). Princeton and Oxford: Princeton University Press. ISBN 978-0-691-15061-1.

External links

• Southern Methodist University - New toothy Texas pterosaur, with an English cousin – Timothy Myers talks about Cimoliopterus dunni

v t e Pterosauria
Kingdom: Animalia Phylum: Chordata Class: Sauropsida Clade: Archosauria Clade: Avemetatarsalia Clade: Pterosauromorpha Avemetatarsalia see Avemetatarsalia Pterosauria see below↓
Pterosauria Pterosauria Daohugoupterus Preondactylia Austriadactylus Preondactylus Caviramidae? Carniadactylus? Caviramus Raeticodactylus? Austriadraconidae Arcticodactylus? Austriadraco Seazzadactylus Eopterosauria Austriadraco? Peteinosaurus? Preondactylia? Eudimorphodontoidea Raeticodactylidae Caviramus? Pachagnathus Raeticodactylus Yelaphomte Eudimorphodontidae Arcticodactylus? Eudimorphodontinae Carniadactylus? Eudimorphodon Zambellisauria? Peteinosaurus? Macronychoptera Herbstosaurus Dimorphodontidae Allkaruen? Caelestiventus Dimorphodon Parapsicephalus? Peteinosaurus? Rhamphinion? Lonchognatha? Eudimorphodon? Novialoidea Campylognathoididae Bergamodactylus Campylognathoides Breviquartossa Rhamphorhynchidae Dolicorhamphus? Klobiodon Parapsicephalus? Scaphognathinae? Rhamphorhynchinae Bellubrunnus Cacibupteryx Dorygnathus Harpactognathus Nesodactylus Orientognathus Qinglongopterus Rhamphorhynchae Angustinaripterini Angustinaripterus Dearc Harpactognathus? Qinglongopterus? Sericipterus Rhamphorhynchini Cacibupteryx? Nesodactylus? Rhamphorhynchus Digibrevisauria? Scaphognathidae? Fenghuangopterus Jianchangnathus Jianchangopterus Scaphognathus Sordes? Pterodactylomorpha see below↓
Pterodactylomorpha Pterodactylomorpha Allkaruen? Sordes? Monofenestrata Archaeoistiodactylus Melkamter Normannognathus Anurognathidae? Darwinoptera Ceoptera Pterorhynchus Wukongopteridae Kunpengopterus Wukongopterinae Archaeoistiodactylus? Cuspicephalus Darwinopterus Douzhanopterus? Wukongopterus Pterodactyliformes Changchengopterus? Douzhanopterus Skiphosoura Caelidracones Anurognathidae Mesadactylus Anurognathinae Anurognathus Dendrorhynchoides? Jeholopterus? Luopterus Vesperopterylus Batrachognathinae Batrachognathus Cascocauda Dendrorhynchoides? Jeholopterus? Sinomacrops Pterodactyloidea Dermodactylus Eurolimnornis Herbstosaurus? Kryptodrakon Ningchengopterus Pangupterus Propterodactylus Samrukia? Wenupteryx Lophocratia see below↓
Lophocratia Archaeopterodactyloidea Prejanopterus Germanodactylidae Altmuehlopterus? Germanodactylus Normannognathus? Tendaguripterus? Euctenochasmatia Diopecephalus Pterodactylus Ctenochasmatoidea Gallodactylidae Aurorazhdarcho? Cycnorhamphus Normannognathus? Petrodactyle? Moganopterinae? Aurorazhdarchia Aerodactylus Gallodactylidae? Aurorazhdarchidae Ardeadactylus? Aurorazhdarcho Huanhepterus? Ctenochasmatidae Ardeadactylus? Balaenognathus Cathayopterus Cratonopterus Elanodactylus Forfexopterus Gladocephaloideus Kepodactylus Liaodactylus Otogopterus Petrodactyle Pterofiltrus Moganopterinae? Feilongus Moganopterus Gnathosaurinae Gnathosaurus Huanhepterus? Lusognathus Plataleorhynchus Tacuadactylus Ctenochasmatinae Ctenochasma Pterodaustrini Beipiaopterus Eosipterus Gegepterus Pterodaustro Eupterodactyloidea Altmuehlopterus? Ornithocheiroidea see below↓
Ornithocheiroidea Ornithocheiroidea Piksi? Tapejaroidea Dsungaripteridae Banguela? Lonchognathosaurus? Noripterus Ordosipterus Puntanipterus? Tendaguripterus? Dsungaripterinae Domeykodactylus Dsungaripterus Azhdarchoidea Inabtanin Ornithostoma Tapejaromorpha Bennettazhia Thalassodromidae? Alanqa? Argentinadraco? Banguela? Kariridraco Leptostomia? Thalassodromeus Tupuxuara Xericeps? Tapejariformes Caupedactylia? Aymberedactylus Caupedactylus Tapejaridae Sinopterinae Afrotapejara Bakonydraco? Eopteranodon? Huaxiadraco Nemicolopterus Sinopterus Wightia Tapejarinae Aymberedactylus? Caupedactylus? Keresdrakon? Lacusovagus? Vectidraco Tapejarini Bakonydraco? Europejara Tapejara Tupandactylus Caiuajarina Caiuajara Torukjara Neoazhdarchia Dsungaripteromorpha? Microtuban Dsungaripteridae Thalassodromidae? Azhdarchomorpha Keresdrakon? Neopterodactyloidea Chaoyangopteridae Apatorhamphus? Eoazhdarcho Lacusovagus? Meilifeilong Microtuban? Chaoyangopterinae Chaoyangopterus Jidapterus Shenzhoupterus Azhdarchiformes Montanazhdarcho Radiodactylus Alanqidae? Alanqa Argentinadraco? Leptostomia? Montanazhdarcho? Xericeps? Azhdarchidae Alanqa? Navajodactylus? Palaeocursornis Azhdarchinae Aerotitan? Albadraco? Azhdarcho Mistralazhdarcho? Quetzalcoatlinae Aerotitan? Albadraco? Aralazhdarcho Arambourgiania Cryodrakon Eurazhdarcho? Hatzegopteryx Mistralazhdarcho? Nipponopterus Phosphatodraco? Quetzalcoatlus Thanatosdrakon Wellnhopterus? Zhejiangopterus? Pteranodontoidea see below↓
Pteranodontoidea Pteranodontoidea Santanadactylus Pteranodontia Pteranodontidae Bogolubovia Dawndraco Pteranodon Tethydraco Volgadraco? Nyctosauromorpha Alamodactylus Cretornis Volgadraco? Aponyctosauria Alcione Epapatelo Simurghia Nyctosauridae Barbaridactylus Muzquizopteryx Nyctosaurus Ornithocheiromorpha Aussiedraco Serradraco Unwindia Lonchodectidae Hongshanopterus? Ikrandraco? Lonchodectes Lonchodraco? Targaryendraco? Lanceodontia Draigwenia Lonchodraconidae Ikrandraco Lonchodraco Istiodactyliformes Hongshanopterus? Linlongopterus Yixianopterus Lonchodectidae? Mimodactylidae Haopterus Linlongopterus Mimodactylus Istiodactylidae Lingyuanopterus Longchengpterus Luchibang Nurhachius Istiodactylinae Istiodactylus Liaoxipterus Ornithocheiriformes Barbosania Brasileodactylus Cearadactylus Hamipterus Boreopteridae Boreopterus Zhenyuanopterus Ornithocheirae Ornithocheiridae Araripesaurus Arthurdactylus Camposipterus? Caulkicephalus? Cimoliopterus? Haliskia? Tropeognathus? Ornithocheirinae Aetodactylus? Camposipterus? Coloborhynchus? Draigwenia? Ferrodraco? Guidraco? Ludodactylus? Mythunga? Ornithocheirus Siroccopteryx? Uktenadactylus? Targaryendraconia? Cimoliopteridae Aetodactylus Camposipterus? Cimoliopterus Targaryendraconidae Aussiedraco? Barbosania? Targaryendraco Anhangueria Brasileodactylus? Ornithocheiridae? Hamipteridae? Hamipterus Iberodactylus Anhangueridae Coloborhynchinae? Aerodraco Coloborhynchus Nicorhynchus Siroccopteryx? Uktenadactylus Anhanguerinae Anhanguera Caulkicephalus? Cearadactylus? Guidraco Liaoningopterus Ludodactylus Maaradactylus Tropeognathinae? Tropeognathini Akharhynchus Amblydectes? Siroccopteryx? Tropeognathus Mythungini Ferrodraco? Haliskia? Mythunga Thapunngaka

• Paleontology
• United States
• United Kingdom

• Pteranodontoids
• Albian life
• Cenomanian life
• Early Cretaceous pterosaurs of Europe
• Late Cretaceous pterosaurs of Europe
• Cretaceous England
• Fossils of England
• Late Cretaceous pterosaurs of North America
• Cretaceous Texas
• Paleontology in Texas
• Fossil taxa described in 2013
• Taxa named by Alexander Kellner