Pteranodon
Pteranodon is a genus of pterosaur that included some of the largest known flying reptiles, with P. longiceps having a wingspan of over. They lived during the late Cretaceous geological period of North America in present-day Kansas, Nebraska, Wyoming, South Dakota and Alabama. More fossil specimens of Pteranodon have been found than any other pterosaur, with about 1,200 specimens known to science, many of them well preserved with nearly complete skulls and articulated skeletons. It was an important part of the animal community in the Western Interior Seaway.
When the first fossils of Pteranodon were found, they were assigned to toothed pterosaur genera, such as Ornithocheirus and Pterodactylus. In 1876, Othniel Charles Marsh recognised it as a genus of its own, making particular note of its complete lack of teeth, which at the time was unique among pterosaurs. Over the decades, multiple species would be assigned to Pteranodon, though today, only two are recognised: P. longiceps, the type species, and P. sternbergi. A third species, P. maiseyi, may also exist. Some researchers have suggested the latter two as a genus of their own, Geosternbergia, though this is the subject of some debate. Another genus split from Pteranodon, Dawndraco, may be synonymous with Geosternbergia if that genus is valid, or with Pteranodon if it is not.
Pteranodon is part of the family Pteranodontidae, part of the clade Pteranodontia, which also includes nyctosaurids. Pteranodontians form a larger clade, Pteranodontoidea, alongside ornithocheiromorphs, and that clade falls under the suborder Pterodactyloidea. While not dinosaurs, pterosaurs such as Pteranodon form a clade closely related to dinosaurs as both fall within the clade Avemetatarsalia.
Male and female Pteranodon differed in size and crest shape. Males attained wingspans of ; females were smaller, averaging. The crests of males were far larger than those of females. In P. longiceps, they were long and backswept, whereas in P. sternbergi, they were tall and upright. Females also had wider pelvises than males.
Discovery and history
First fossils
Pteranodon was the first pterosaur found outside of Europe. Its fossils first were found by Othniel Charles Marsh in 1871, in the Late Cretaceous Smoky Hill Chalk deposits of western Kansas. These chalk beds were deposited at the bottom of what was once the Western Interior Seaway, a large shallow sea over what now is the midsection of the North American continent. These first specimens, YPM 1160 and YPM 1161, consisted of partial wing bones, as well as a tooth from the prehistoric fish Xiphactinus, which Marsh mistakenly believed to belong to this new pterosaur. In 1871, Marsh named the find Pterodactylus oweni, assigning it to the well-known European genus Pterodactylus. Marsh also collected more wing bones of the large pterosaur in 1871. Realizing that the name he had chosen had already been used for Harry Seeley's European pterosaur species Pterodactylus oweni in 1864, Marsh renamed his giant North American pterosaur Pterodactylus occidentalis, meaning "Western wing finger," in his 1872 description of the new specimen. He named two additional species, based on size differences: Pterodactylus ingens, and Pterodactylus velox.Meanwhile, Marsh's rival Edward Drinker Cope had unearthed several specimens of the large North American pterosaur. Based on these specimens, Cope named two new species, Ornithochirus umbrosus and Ornithochirus harpyia, in an attempt to assign them to the large European genus Ornithocheirus, though he misspelled the name. Cope's paper naming his species was published in 1872, just five days after Marsh's paper. This resulted in a dispute, fought in the published literature, over whose names had priority in what obviously were the same species. Cope conceded in 1875 that Marsh's names did have priority over his, but maintained that Pterodactylus umbrosus was a distinct species from any that Marsh had named previously. Re-evaluation by later scientists has supported Marsh's case, refuting Cope's assertion that P. umbrosus represented a larger, distinct species.
A toothless pterosaur
While the first Pteranodon wing bones were collected by Marsh and Cope in the early 1870s, the first Pteranodon skull was found on May 2, 1876, along the Smoky Hill River in Wallace County, Kansas, USA, by Samuel Wendell Williston, a fossil collector working for Marsh. A second, smaller skull soon was discovered as well. These skulls showed that the North American pterosaurs were different from any European species, in that they lacked teeth and had bony crests on their skulls. Marsh recognized this major difference, describing the specimens as "distinguished from all previously known genera of the order Pterosauria by the entire absence of teeth." Marsh recognized that this characteristic warranted a new genus, and he coined the name Pteranodon in 1876. Marsh reclassified all the previously named North American species from Pterodactylus to Pteranodon. He considered the smaller skull to belong to Pteranodon occidentalis, based on its size. Marsh classified the larger skull, YPM 1117, in the new species Pteranodon longiceps, which he thought to be a medium-sized species in between the small P. occidentalis and the large P. ingens. Marsh also named several additional species: Pteranodon comptus and Pteranodon nanus were named for fragmentary skeletons of small individuals, while Pteranodon gracilis was based on a wing bone that he mistook for a pelvic bone. He soon realized his mistake, and re-classified that specimen again into a separate genus, which he named Nyctosaurus. P. nanus was also later recognized as a Nyctosaurus specimen.In 1892, Samuel Williston examined the question of Pteranodon classification. He noticed that, in 1871, Seeley had mentioned the existence of a partial set of toothless pterosaur jaws from the Cambridge Greensand of England, which he named Ornithostoma. Because the primary characteristic Marsh had used to separate Pteranodon from other pterosaurs was its lack of teeth, Williston concluded that "Ornithostoma" must be considered the senior synonym of Pteranodon. However, in 1901, Pleininger pointed out that "Ornithostoma" had never been scientifically described or even assigned a species name until Williston's work, and therefore had been a nomen nudum and could not beat out Pteranodon for naming priority. Williston accepted this conclusion and went back to calling the genus Pteranodon. However, both Williston and Pleininger were incorrect, because unnoticed by both of them was the fact that, in 1891, Seeley himself had finally described and properly named Ornithostoma, assigning it to the species O. sedgwicki. In the 2010s, more research on the identity of Ornithostoma showed that it was probably not Pteranodon or even a close relative, but may in fact have been an azhdarchoid, a different type of toothless pterosaur.
Revising species
Williston was also the first scientist to critically evaluate all of the Pteranodon species classified by Cope and Marsh. He agreed with most of Marsh's classification, with a few exceptions. First, he did not believe that P. ingens and P. umbrosus could be considered synonyms, which even Cope had come to believe. He considered both P. velox and P. longiceps to be dubious; the first was based on non-diagnostic fragments, and the second, though known from a complete skull, probably belonged to one of the other, previously-named species. In 1903, Williston revisited the question of Pteranodon classification, and revised his earlier conclusion that there were seven species down to just three. He considered both P. comptus and P. nanus to be specimens of Nyctosaurus, and divided the others into small, medium, and large species, based primarily on the shape of their upper arm bones. He thought P. longiceps, the only one known from a skull, could be a synonym of either P. velox or P. occidentalis, based on its size.In 1910, Eaton became the first scientist to publish a more detailed description of the entire Pteranodon skeleton, as it was known at the time. He used his findings to revise the classification of the genus once again based on a better understanding of the differences in pteranodont anatomy. Eaton conducted experiments using clay models of bones to help determine the effects of crushing and flattening on the shapes of the arm bones Williston had used in his own classification. Eaton found that most of the differences in bone shapes could be easily explained by the pressures of fossilization, and concluded that no Pteranodon skeletons had any significant differences from each other besides their size. Therefore, Eaton was left to decide his classification scheme based on differences in the skulls alone, which he assigned to species just as Marsh did, by their size. In the end, Eaton recognized only three valid species: P. occidentalis, P. ingens, and P. longiceps.
The discovery of specimens with upright crests, classified by Harksen in 1966 as the new species Pteranodon sternbergi, complicated the situation even further. prompting another revision of the genus by Halsey W. Miller in 1972. Because it was impossible to determine crest shape for all of the species based on headless skeletons, Miller concluded that all Pteranodon species except the two based on skulls must be considered nomena dubia and abandoned. The skull Eaton thought belonged to P. ingens was placed in the new species Pteranodon marshi, and the skull Eaton assigned to P. occidentalis was re-named Pteranodon eatoni. Miller also recognized another species based on a skull with a crest similar to that of P. sternbergi; Miller named this Pteranodon walkeri. To help bring order to this tangle of names, Miller created three subgenera. P. marshi and P. longiceps were placed in the subgenus Longicepia, though this was later changed to simply Pteranodon due to the rules of priority. P. sternbergi and P. walkeri, the upright-crested species, were given the subgenus Sternbergia, which was later changed to Geosternbergia because Sternbergia was preoccupied. Finally, Miller named the subgenus Occidentalia for P. eatoni, the skull formerly associated with P. occidentalis. Miller further expanded the concept of Pteranodon to include Nyctosaurus as a fourth subgenus. Miller considered these to be an evolutionary progression, with the primitive Nyctosaurus, at the time thought to be crestless, giving rise to small-crested Occidentalia, which in turn gave rise to long-crested Pteranodon, finally leading to tall-crested Geosternbergia. However, Miller made several mistakes in his study concerning which specimens Marsh had assigned to which species, and most scientists disregarded his work on the subject in their later research. In 1984, Robert Milton Schoch published another revision that essentially returned to Marsh's original classification scheme, most notably sinking P. longiceps as a synonym of P. ingens.