Instruct: Given a question, retrieve passages that answer the question.
Query: How large was the Pteranodon?
Pteranodon (/tɪˈrænədɒn/; from Greek πτερόν ( pteron, "wing") and ἀνόδων (anodon, "toothless") is a genus of pterosaur that included some of the largest known flying reptiles, with wingspans over 8 meters (26 feet). They lived during the late Cretaceous geological period of North America in present-day Kansas, Alabama, Nebraska, Wyoming, and South Dakota. 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.[1]
Many researchers consider there to be at least two species of Pteranodon. However, aside from the differences between males and females described above, the post-cranial skeletons of Pteranodon show little to no variation between species or specimens, and the bodies and wings of all pteranodonts were essentially identical.[6]
After the group escapes, Grant discovers Billy has taken two raptor eggs to use for funding, which provoked the raptor attacks. Grant decides to keep the eggs to ensure the group's survival. The group unknowingly enters a large aviary used to house "Pteranodons", which attack the group and fly away with Eric. Billy rescues Eric using Ben's parasail, but is then attacked and apparently killed by the "Pteranodons". The rest of the group escapes the aviary, leaving the door unlocked. They find a boat and make their way down a river.
Pteranodon sternbergi is the only known species of Pteranodon with an upright crest. The lower jaw of P. sternbergi was 1.25 meters (4.1 feet) long.[24] It was collected by George F. Sternberg in 1952 and described by John Christian Harksen in 1966, from the lower portion of the Niobrara Formation. It was older than P. longiceps and is considered by Bennett to be the direct ancestor of the later species.[6]
Two genre of pterosaurs are present in the formation, both within Pterodactyloidea: the pteranodontid "Pteranodon" (sometimes broken into several genera like "Geosternbergia" and "Dawndraco", though this is dubious) and the nyctosaurid "Nyctosaurus". They are large pterosaurs with elongated cranial crests. The pterosaurs of Niobrara probably spent most of their time at sea and rarely went on land, with "Nyctosaurus" being a probably fully pelagic animal. "Pteranodon" probably foraged on the ocean surface, while "Nyctosaurus" was a frigatebird-like aerial predator.
Some very fragmentary fossils belonging to pteranodontian pterosaurs, and possibly Pteranodon itself, have also been found on the Gulf Coast and East Coast of the United States. For example, some bone fragments from the Mooreville Formation of Alabama and the Merchantville Formation of Delaware may have come from Pteranodon, though they are too incomplete to make a definite identification.[6] Some remains from Japan have also been tentatively attributed to Pteranodon, but their distance from its known Western Interior Seaway habitat makes this identification unlikely.[6]
Pteranodon fossils are known primarily from the Niobrara Formation of the central United States. Broadly defined, Pteranodon existed for more than four million years, during the late Coniacian to early Campanian stages of the Cretaceous period.[6] The genus is present in most layers of the Niobrara Formation except for the upper two; in 2003, Kenneth Carpenter surveyed the distribution and dating of fossils in this formation, demonstrating that Pteranodon sternbergi existed there from 88 to 85 million years ago, while P. longiceps existed between 86 and 84.5 million years ago. A possible third species, which Kellner named Geosternbergia maysei in 2010, is known from the Sharon Springs member of the Pierre Shale Formation in Kansas, Wyoming, and South Dakota, dating to between 81.5 and 80.5 million years ago.[16]
It is likely that, as in other polygynous animals (in which males compete for association with harems of females), Pteranodon lived primarily on offshore rookeries, where they could nest away from land-based predators and feed far from shore; most Pteranodon fossils are found in locations which at the time, were hundreds of kilometres from the coastline.[7]
The wing shape of Pteranodon suggests that it would have flown rather like a modern-day albatross. This is based on the fact that Pteranodon had a high aspect ratio (wingspan to chord length) similar to that of the albatross — 9:1 for Pteranodon, compared to 8:1 for an albatross. Albatrosses spend long stretches of time at sea fishing, and use a flight pattern called "dynamic soaring" which exploits the vertical gradient of wind speed near the ocean surface to travel long distances without flapping, and without the aid of thermals (which do not occur over the open ocean the same way they do over land).[8] While most of a Pteranodon flight would have depended on soaring, like long-winged seabirds, it probably required an occasional active, rapid burst of flapping, and studies of Pteranodon wing loading (the strength of the wings vs. the weight of the body) indicate that they were capable of substantial flapping flight, contrary to some earlier suggestions that they were so big they could only glide.[3]
Two species of Pteranodon are traditionally recognized as valid: Pteranodon longiceps, the type species, and Pteranodon sternbergi. The species differ only in the shape of the crest in adult males (described above), and possibly in the angle of certain skull bones.[6] Because well-preserved Pteranodon skull fossils are extremely rare, researchers use stratigraphy (i.e. which rock layer of the geologic formation a fossil is found in) to determine species identity in most cases.
Due to the subtle variations between specimens of pteranodontid from the Niobrara Formation, most researchers have assigned all of them to the single genus Pteranodon, in at least two species (P. longiceps and P. sternbergi) distinguished mainly by the shape of the crest. However, the classification of these two forms has varied from researcher to researcher. In 1972, Halsey Wilkinson Miller published a paper arguing that the various forms of Pteranodon were different enough to be placed in distinct subgenera. He named these Pteranodon (Occidentalia) occidentalis (for the now-disused species P. occidentalis) and Pteranodon (Sternbergia) sternbergi. However, the name Sternbergia was preoccupied, and in 1978 Miller re-named the species Pteranodon (Geosternbergia) sternbergi, and named a third subgenus/species combination for P. longiceps, as Pteranodon (Longicepia) longiceps. Most prominent pterosaur researchers of the late 20th century however, including S. Christopher Bennett and Peter Wellnhofer, did not adopt these subgeneric names, and continued to place all pteranodont species into the single genus Pteranodon.
Pteranodontia Nyctosauridae Muzquizopteryx coahuilensis "Nyctosaurus" lamegoi Nyctosaurus gracilis Alamodactylus byrdiPteranodontoidea Pteranodon longiceps Pteranodon sternbergi Istiodactylidae Longchengpterus zhaoi Nurhachius ignaciobritoi Liaoxipterus brachyognathus Istiodactylus latidens Istiodactylus sinensis Lonchodectes compressirostris Aetodactylus halli Cearadactylus atrox Brasileodactylus araripensis Ludodactylus sibbicki Ornithocheirae
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 (now Logan County), Kansas, USA, by Samuel Wendell Williston, a fossil collector working for Marsh.[1] 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 ("wing without tooth") 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.[20][6] 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.[21][6]
The most distinctive characteristic of Pteranodon is its cranial crest. These crests consisted of skull bones (frontals) projecting upward and backward from the skull. The size and shape of these crests varied due to a number of factors, including age, sex, and species. Male Pteranodon sternbergi, the older species of the two described to date (and sometimes placed in the distinct genus Geosternbergia), had a more vertical crest with a broad forward projection, while their descendants, Pteranodon longiceps, evolved a narrower, more backward-projecting crest.[1] Females of both species were smaller and bore small, rounded crests.[6] The crests were probably mainly display structures, though they may have had other functions as well.[7]
Pteranodon were pterosaurs, not dinosaurs. By definition, all dinosaurs belong to one of the two groups within Dinosauria, i.e. Saurischia or Ornithischia. As such, this excludes pterosaurs. Nonetheless, Pteranodon are frequently featured in dinosaur media and are strongly associated with dinosaurs by the general public.[2] While not dinosaurs, they form a sister clade to dinosaurs within the clade Avemetatarsalia.
Methods used to estimate the mass of large male Pteranodon specimens (those with wingspans of about 7 meters) have been notoriously unreliable, producing a wide range of estimates from as low as 20 kilograms (44lb) to as high as 93 kilograms (205lb). In a review of pterosaur size estimates published in 2010, researchers Mark Witton and Mike Habib demonstrated that the latter, largest estimates are almost certainly incorrect given the total volume of a Pteranodon body, and could only be correct if the animal "was principally comprised of aluminium".[3] Witton and Habib considered the methods used by researchers who obtained smaller mass estimates equally flawed. Most have been produced by scaling modern animals such as bats and birds up to Pteranodon size, despite the fact that pterosaurs have vastly different body proportions and soft tissue anatomy from any living animal.[3]
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 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 (P. longiceps and P. sternbergi) 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 categories or "subgenera" for them. 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 already in use ("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 Occidentalia (with a small crest), which in turn gave rise to Pteranodon with its long backwards crest, finally leading to Geosternbergia with its large, upright crest. 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, though Wellnhofer (1978) followed Miller's species list. and Schoch (1984) somewhat oddly published another revision that essentially returned to Marsh's original classification scheme, most notably sinking P. longiceps as a synonym of P. ingens.[6]
The diet of Pteranodon is known to have included fish; fossilized fish bones have been found in the stomach area of one Pteranodon, and a fossilized fish bolus has been found between the jaws of another Pteranodon, specimen AMNH 5098. Numerous other specimens also preserve fragments of fish scales and vertebrae near the torso, indicating that fish made up a majority of the diet of Pteranodon (though they may also have taken invertebrates).[2]
It is based on the type specimen or holotype BPV-078, an incomplete skeleton and skull. Its skull, including a large crest, was toothless and similar to that of "Pteranodon". The skull lacks the point of the snout but it was in life less than 200 millimeters long (7.9 inches), and the animal had a wingspan of about 1.1 meters (3.6 feet). A second specimen, D2526, described in 2006, had a larger wingspan. Despite its similarities to "Pteranodon", "Eopteranodon" was not placed into a family by its describers, who put it into the clade Pteranodontia as "incertae sedis" (uncertain position).
Adult Pteranodon specimens may be divided into two distinct size classes, small and large, with the large size class being about one and a half times larger than the small class, and the small class being twice as common as the large class. Both size classes lived alongside each other, and while researchers had previously suggested that they represent different species, Christopher Bennett showed that the differences between them are consistent with the concept that they represent females and males, and that Pteranodon species were sexually dimorphic. Skulls from the larger size class preserve large, upward and backward pointing crests, while the crests of the smaller size class are small and triangular. Some larger skulls also show evidence of a second crest that extended long and low, toward the tip of the beak, which is not seen in smaller specimens.[7]
Adult Pteranodon specimens from the two major species can be divided into two distinct size classes. The smaller class of specimens have small, rounded head crests and very wide pelvic canals, even wider than those of the much larger size class. The size of the pelvic canal probably allowed the laying of eggs, indicating that these smaller adults are females. The larger size class, representing male individuals, have narrow hips and very large crests, which were probably for display.
Specimens assigned to Pteranodon have been found in both the Smoky Hill Chalk deposits of the Niobrara Formation, and the slightly younger Sharon Springs deposits of the Pierre Shale Formation. When Pteranodon was alive, this area was covered by a large inland sea, known as the Western Interior Seaway. Famous for fossils collected since 1870, these formations extend from as far south as Kansas in the United States to Manitoba in Canada. However, Pteranodon specimens (or any pterosaur specimens) have only been found in the southern half of the formation, in Kansas, Wyoming, and South Dakota. Despite the fact that numerous fossils have been found in the contemporary parts of the formation in Canada, no pterosaur specimens have ever been found there. This strongly suggests that the natural geographic range of Pteranodon covered only the southern part of the Niobrara, and that its habitat did not extend farther north than South Dakota.[6]
The fact that females appear to have outnumbered males two to one suggests that, as with modern animals with size-related sexual dimorphism, such as sea lions and other pinnipeds, Pteranodon might have been polygynous, with a few males competing for association with groups consisting of large numbers of females. Similar to modern pinnipeds, Pteranodon may have competed to establish territory on rocky, offshore rookeries, with the largest, and largest-crested, males gaining the most territory and having more success mating with females. The crests of male Pteranodon would not have been used in competition, but rather as "visual dominance-rank symbols", with display rituals taking the place of physical competition with other males. If this hypothesis is correct, it also is likely that male Pteranodon played little to no part in rearing the young; such a behavior is not found in the males of modern polygynous animals who father many offspring at the same time.[7]
Manly then tries to get one of the "Pteranodons" to become famous, but Amanda punches him out cold. The "Pteranodons" are returned to the island, by tricking them to follow an aircraft disguised as a "Pteranodon".
In 1892, Samuel Williston examined the question of Pteranodon classification. He noticed that, in 1871, Seeley had mention 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.[6] 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.[22]