Cimoliopterus
Cimoliopterus is a genus of pterosaur that lived during the Late Cretaceous in what is now England and the United States. The first known specimen, consisting of the front part of a snout including part of a crest, was discovered in the Grey Chalk Subgroup of Kent, England, and described as the new species Pterodactylus cuvieri in 1851. The specific name cuvieri honoured the palaeontologist George Cuvier, and the genus Pterodactylus was then used for many pterosaurs of species that are no longer thought to be closely related. It was among the first pterosaurs to be depicted as sculptures, in Crystal Palace Park in the 1850s. The species was subsequently assigned to various other genera, including Ornithocheirus and Anhanguera. In 2013, the species was moved to a new genus, as C. cuvieri; the generic name Cimoliopterus is derived from the Greek words for "chalk" and "wing". Other specimens and species have also been assigned to or synonymised with the species with various levels of certainty. In 2015, a snout discovered in the Britton Formation of Texas, US, was named as a new species in the genus, C. dunni. A species from the Cambridge Greensand of England originally assigned to Ornithocheirus was assigned to Cimoliopterus in 2025, as C. colorhinus.
C. cuvieri is estimated to have had a wingspan of, and C. dunni is thought to have been similar to C. cuvieri in size. One C. colorhinus specimen is estimated to have had a wingspan of. Cimoliopterus can be distinguished from related pterosaurs in features such as having a premaxillary crest that begins hindward on the snout, in having a ridge on the palate that extends forwards until the third pair of tooth sockets, and in the spacing and proportions of the tooth sockets. Unlike similar pterosaurs, the tip of the snout is only subtly expanded to the sides. C. cuvieri and C. dunni differ from each other in various details in the configuration of these features; for example, the crest of C. cuvieri begins by the seventh tooth socket, whereas that of C. dunni begins at the fourth. More completely known related genera were fairly large pterosaurs, with proportionally large skulls, long jaws and tooth-rows, often with large, rounded crests at the front of the jaws. The teeth at the front of the jaws were large and recurved; further back, the teeth were smaller, slightly recurved, and well-spaced. As pterosaurs, Cimoliopterus would have been covered in pycnofibres, and had extensive wing-membranes, which were distended by long wing-fingers.
While long considered an ornithocheiran, the affinities of C. cuvieri were unclear due to the fragmentary nature of it and other English pterosaurs, until more complete relatives were reported from Brazil in the 1980s. Cimoliopterus was moved to the family Cimoliopteridae within the clade Targaryendraconia in 2019, with its closest relative being Camposipterus. That C. cuvieri from England and C. dunni from North America were so similar despite living on opposite sides of the North Atlantic Ocean indicates they became less isolated from each other than other animal groups because they could fly. This kind of pterosaur was probably adapted for long-distance oceanic soaring, which is also supported by their fossils mainly being found in marine settings. While they may have been proficient in water, their terrestrial abilities were limited due to their having short hindlimbs but long forelimbs. They possibly fed while flying by dipping their long jaws into water and catching fish with their interlocking teeth. While some have suggested the premaxillary crests were used to stabilise the jaws while submerged in water during feeding, they may have been used as display structures.
History
The taxonomic history of Cimoliopterus and related pterosaurs from the Cretaceous of England has been described as confusing by modern pterosaur researchers, since many of them were described from very fragmentary remains, and named using convoluted nomenclatural schemes of the Victorian era. Prior to these discoveries, many of the hollow, thin-walled pterosaur bones found in England were thought to have belonged to birds. In 1851, the British naturalist James Scott Bowerbank described a large pterosaur snout he had obtained, which was found in the Lower Culand Pit in what is now called the Grey Chalk Subgroup at Burham, Kent, in South East England. Pterosaur fossils had been discovered earlier 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, Bowerbank named the species Pterodactylus cuvieri; at this time, the genus Pterodactylus was used for many pterosaur species now thought to be only distantly related to each other. The specific name honours the French palaeontologist Georges Cuvier, who had recognised that pterosaurs were flying reptiles. In full, the name can be translated as "Cuvier's wing finger".The snout which Bowerbank made the holotype specimen of P. cuvieri consists of the front of the upper jaw, including part of a premaxillary crest, and is catalogued today as NHMUK PV 39409 at the Natural History Museum in London. It was originally reported to preserve a single tooth in the first right tooth socket, but this had disappeared when the holotype was examined in the 2000s. 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 in 1851 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 Crystal Palace Park in London, under the supervision of Owen. Among them were two species of pterosaur, including two P. cuvieri sculptures made of iron-framed concrete and two smaller statues of Pterodactylus bucklandi. While Owen only identified one of the large sculptures as P. cuvieri in 1854, they are similar enough that they can be assumed to depict the same species. Hawkins' sculptures were the first to depict pterosaurs, and the oldest life reconstructions of giant members of the group. Incidentally, he thought these animals to be the original dragons, such as in the legend of Saint George. The larger P. cuvieri 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 more completely known P. antiquus. They were depicted with scales and bird-like proportions such as small heads and large torsos, as was customary at the time. One sculpture is correctly shown in a quadrupedal pose, and with one foot flat on the ground. While this pterosaur was one of the first known preserved with a crest at the tip of the jaw, this feature is not included in the sculptures, as it seems they were assumed to have been more similar to those of P. antiquus. Witton and the British biologist Ellinor Michel said in 2023 that while it was the largest known pterosaur at the time, its historical significance was overshadowed by the 1870 discovery of the larger Pteranodon, a genus that was widely featured in text and artwork, while the Crystal Palace sculptures remained the only major publicity of P. antiquus.
In 1869, the British palaeontologist Harry Govier Seeley placed P. cuvieri in the new genus "Ptenodactylus" along with other English pterosaurs known mainly from upper jaws, in an index of specimens in the Woodwardian Museum. He noted these names were provisional, only intended for the convenience 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 he listed belonged 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 1922, the Austrian naturalist Gustav von Arthaber lamented that the scientific literature had accepted the many Ornithocheirus names that had only been mentioned in Seeley's catalogue for students. In his opinion names were of no use without an illustration of the specimens they were based on, or better still, a complete reconstruction of the relevant skeleton. For this reason, he provided a drawing of the skull of O. cuvieri, one of the few species for which the known jaw material proved its validity. In 1924, von Arthaber's interpretations were criticised by the Hungarian palaeontologist Franz Nopcsa von Felső-Szilvás, who found some of the skull reconstructions "worthless".
O. cuvieri and many other English pterosaurs were kept in the genus Ornithocheirus for most of the 20th century. In 1987, the German palaeontologist Peter Wellnhofer described the new crested pterosaur Tropeognathus from the Santana Formation of Brazil and noted the similarities between it and other newly described Brazilian taxa such as Anhanguera to English taxa that were based on fragmentary snouts, such as the various species assigned to Ornithocheirus. He concluded that while the appearance of the English taxa had long been a puzzle, the discovery of the related and much better preserved Brazilian species made this clearer, showing that the English species too had premaxillary crests at the end of large, long skulls. When Anhanguera was described in 1985, it was thought to be the only pterosaur with such a crest known until that point. In 1991, Wellnhofer stated that the genus Ornithocheirus had become a "wastebin" of species from different countries, assigned to it on the basis of insufficient characters, and needed clearer definition, and perhaps included several distinct genera.
The British palaeontologist S. Christopher Bennett stated in 1993 that the holotype specimen of O. cuvieri was the right first wing-phalanx bone mentioned by Owen in 1851, and cited him for the name without further explanation. In 1995, the Russian palaeontologist Natalia N. Bakhurina and the British palaeontologist David M. Unwin assigned a partial mandibular symphysis from Russia to cf. Anhanguera cuvieri, thereby using a new combination. Unwin and colleagues assigned pterosaur skeletal elements from the Toolebuc Formation of Australia to Anhanguera ?cuvieri in 2000. In 2001, Unwin also used the combination A. cuvieri, and cited Bowerbank for the name, with the snout as holotype. He assigned 23 jaw fragments from the Cambridge Greensand of England to A. cuvieri and considered it the most common pterosaur of that formation. Unwin 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 a species originally named Pterodactylus fittoni by Owen in 1859 as a member of Anhanguera. In 2011, the Brazilian palaeontologist Alexander W. Kellner and colleagues made the Australian material that had been assigned to A. ?cuvieri the basis of a new genus, Aussiedraco, and stated that they saw no ground for placing O. cuvieri in Anhanguera.