Pteranodon sp. (S/F) / (S/F-T/G) / (S/F-S)

Note that this page details both Pteranodon longiceps, which has three known variants, as well as Geosternbergia (which was formerly classified as a species of Pteranodon).

Pteranodon and Geosternbergia are large pteranodontid pterosaurs native to much of the North American Midwest during the late Cretaceous period, 88 to 80.5 million years ago. At the time, this area was a large shallow ocean called the Western Interior Seaway. They are among the largest known pterosaurs, and are notable for their extremely distinct sexual dimorphism.

The first remains were found in the Smoky Hills Chalk of western Kansas by Othniel Charles Marsh in 1870, and named by Marsh in 1871. It was the first pterosaur to be discovered outside of Europe by paleontologists, and was originally named Pterodactylus oweni. However, that species name was already occupied (though it has since been reclassified), so Marsh renamed his discovery Pterodactylus occidentalis. He also named the largest specimen Pterodactylus ingens, and the smallest specimen Pterodactylus velox. The remains consisted mostly of wing bones.

Around the same time, Marsh’s rival Edward Drinker Cope discovered wing bone fossils which he assigned to the genus Ornithocheirus (though he misspelled the name as Ornithochirus). He named two species, Ornithochirus umbrosus and Ornithochirus harpyia. The paper naming these species was published in 1872, only five days after Marsh’s. As the rivals recognized that the fossils they had named belonged to the same species, debate occurred over which name should be given priority. In 1875, Cope relented and allowed Marsh’s classification to take priority, but maintained that the umbrosus species was distinct from Marsh’s other species of Pterodactylus. Later paleontologists would disagree with Cope’s conclusion, determining that P. umbrosus was not a distinct species.

On May 2, 1876, Samuel Wendell Williston discovered the first-known skull of the animal near the Smoky Hill River in Kansas. Shortly after, a smaller skull was also discovered. Williston alerted Marsh, his employer, to the discovery; Marsh noted that the skull lacked teeth and possessed a bony crest on the head. These features were distinctly different from other known pterosaurs at the time, prompting Marsh to reclassify the animal. No longer a member of Pterodactylus, the genus was christened Pteranodon, meaning “wing without teeth.” The species recognized at the time included Pteranodon occidentalis and Pteranodon ingens, along with the newly-named Pteranodon longiceps; the specific epithet of this new species refers to the length of its skull.

Three other species Marsh named at the time included Pteranodon comptusPteranodon nanus, and Pteranodon gracilis. The final of those three was named based on a wing bone which he had mistaken for a pelvic bone. He recognized his mistake, however, and reclassified it as Nyctosaurus gracilis.

Samuel Williston would return to question the classification of all Pteranodon genera beginning in 1892. He had learned of toothless pterosaur jaws discovered in England’s Cambridge Greensand which were assigned to the genus Ornithostoma, and suggested that Pteranodon should also be given this name as it was discovered later. This decision was never formally accepted, however. In the end, the two names were recognized as separate genera, and neither incorporated the other. At one point, Williston believed that there could be seven Pteranodon species; as of 1903, he narrowed this down to just three species. These were the small P. velox, medium-sized P. occidentalis, and large P. ingens. Both P. comptus and P. nanus were reclassified as Nyctosaurus, and Williston considered P. longiceps to potentially be the same species as either P. velox or P. occidentalis.

In 1910, the classification of Pteranodon species would be challenged yet again. George Francis Eaton was the first scientist to describe the whole skeleton of the animal, using this information to determine that Pteranodon longiceps was a valid species and Pteranodon velox was not. This three-species conclusion was upheld for decades.

The discovery of a new species by George F. Sternberg in 1952, named Pteranodon sternbergi later in 1966 complicated the situation, as it was found to have a prominent upright crest. As many of the skeletons were headless, paleontologist Halsey Wilkinson Miller argued in 1972 that only the fossils that had known skulls (Pteranodon longiceps and Pteranodon sternbergi) should be considered valid species. Eaton and Miller would go on to name several other species, assigning them into three subgenera. First, the long-crested species P. longiceps and P. ingens, which was renamed as P. marshi, were classified in the Pteranodon subgenus. Second, the tall-crested species P. sternbergi and the newly-named P. walkeri were classified in the subgenus Geosternbergia. Third, P. occidentalis was renamed as P. eatoni and classified in the subgenus Occidentalia.

Today, the only recognized genus is Pteranodon longiceps. In 2010, Alexander W.A. Kellner reclassified P. sternbergi as Geosternbergia sternbergi, and also named a new species Geosternbergia maiseyi. Paleontologist S. Christopher Bennett has suggested that Geosternbergia, which existed slightly earlier in time, evolved directly into Pteranodon.

Throughout the history of paleontology, around 1,200 Pteranodon fossils have been discovered in Kansas, Alabama, Nebraska, Wyoming, and South Dakota. This makes it more common than any other known pterosaur in the fossil record. Some possible Pteranodon remains are known from Skåne, Sweden and from the East Coast of North America.

International Genetic Technologies cloned the first Pteranodon sometime between 1986 and 1993, along with Geosternbergia, which at the time was classified as Pteranodon. At least one variant of Geosternbergia and three variants of Pteranodon have been created, which Jurassic-Pedia has classified as three subspecies for the sake of clarity. The subspecies are Pteranodon longiceps longiceps, which is known only from Isla Sorna; Pteranodon longiceps hippocratesi, an aberrant subspecies with teeth, which was released from Isla Sorna in 2001; and finally Pteranodon longiceps masranii, which was first hatched in 2004 or 2005 and is the only known surviving subspecies. It is not known if any Geosternbergia are still alive.

Description
Adult P. l. longiceps raises its wings while perching, making a display posture to nearby pterosaurs.

All four of the pterosaurs described here have similar features. They are all members of the family Pteranodontidae, which itself is a part of the pterosaur suborder Pterodactyloidea. This means that they are among the short-tailed pterosaurs. In all four, the beak is quite long and has a pointed tip; the sharpest and widest belongs to P. l. masranii, while the beak of P. l. longiceps has a slightly hooked end rather like a fish-eating bird’s. The head and neck differ between the pterosaurs in a few other notable ways, though the overall anatomy is similar. The most obvious difference are the small, triangular teeth of P. l. hippocratesi; these are an aberrant feature, and are presumed to be the result of genetic engineering (intentional or accidental). The head crests differ between the two genera and among the three P. longiceps subspecies. In Pteranodon, the head crest is roughly triangular and points backward from the skull; in fossil specimens, the female’s crest is much smaller, but at least P. l. hippocratesi, they appear to have been engineered to be more or less equal in size. In P. l. masranii, the females’ crests are much smaller than those of the males, more like fossil Pteranodons. The head crest of P. l. longiceps is narrower and slightly bent, while P. l. hippocratesi has a mostly straight crest. Of course, the crest of the male Geosternbergia is tall, erect, and has a forward-facing protrusion which makes it immediately distinguishable from Pteranodon. In all four, the tongue is narrow and pointed, and does not appear able to extend out of the beak. The nostrils are located near the base of the beak, on the upper side.

Adult male Geosternbergia

Size varies among these four pterosaurs. Below is a list of each of their physical dimensions, including all three subspecies of Pteranodon longiceps and its fossil ancestors, as well as both species of Geosternbergia as it is not yet known which species InGen succeeded in cloning. Dimensions for Geosternbergia are based on the fossil record.

  • Fossil Pteranodon longiceps
    • Male
      • Wingspan: 5.6m (18ft) to 6.25m (20.5ft)
      • Length: 2.4m (8ft)
      • Weight: Debated; estimates range from 20kg (44lbs) to 93kg (205lbs)
    • Female
      • Wingspan: 3.8m (12ft)
      • Length: 1.6m (5.3ft)
      • Weight: Unknown, but lighter than the male
  • Geosternbergia sternbergi
    • Male
      • Wingspan: 6m (19.7ft)
      • Length: 2.5m (8.2ft)
      • Weight: Unknown
    • Female
      • Wingspan: 3m (9.8ft) to 3.5m (11.5ft)
      • Length: 1.5m (4.9ft)
      • Weight: Unknown, but lighter than the male
  • Geosternbergia maiseyi
    • Male
      • Wingspan: 7.25m (23.8ft)
      • Length: Probably slightly longer than male G. sternbergi
      • Weight: Unknown, but probably slightly more than male G. sternbergi
    • Female
      • Wingspan: Unknown, but smaller than the male
      • Length: Unknown, but shorter than the male
      • Weight: Unknown, but lighter than the male
  • Pteranodon longiceps longiceps
    • Male (?)
      • Wingspan: 7.6m (25ft)
      • Length: 3m (10ft)
      • Weight: Unknown
    • Female
      • Wingspan: Unknown
      • Length: Unknown
      • Weight: Unknown
  • Pteranodon longiceps hippocratesi
    • Male
      • Wingspan: At least 9.8m (32ft); 14m (45.9ft) in some 2001 sources
      • Length: At least 3m (10.2ft)
      • Weight: 81.6kg (180lbs) in some 2001 sources; 226.8kg (500lbs) to 272.2kg (600lbs), according to Flyers
    • Female
      • Wingspan: 9.1m (30ft) to 9.8m (32ft)
      • Length: 3m (10.2ft)
      • Weight: 54.4kg (120lbs) in some 2001 sources
  • Pteranodon longiceps masranii
    • Male
      • Wingspan: 7.9m (26ft) to 10.1m (33ft), according to Jurassic World: The Game
      • Length: Presumably about 3.8m (12.3ft) based on female’s proportions
      • Weight: Presumably about 48.9kg (107.8lbs) based on female’s proportions
    • Female
      • Wingspan: 5.5m (18ft) to 6m (19.8ft); may reach 7.5m (24.6ft) according to Jurassic World social media
      • Length: 2.4m (8ft) to 3.1m (10.2ft)
      • Weight: 30kg (66.1lbs) to 31.8kg (70lbs)
Pteranodon longiceps longiceps. This is presumed to be a male, though some have suggested that Geosternbergia instead represents the male of this subspecies rather than being a separate genus or species.

The neck is long and somewhat flexible. Much of its overall body size consists of its wings; Geosternbergia maiseyi has a wingspam of up to 7.25 meters (23.8 feet), though the species cloned by InGen may have been the slightly smaller G. sternbergi. Size among the Pteranodon longiceps subspecies is more variable. In the fossil record, the male of this animal reached wingspans of 5.6 meters (18 feet) to 6.25 meters (20.5 feet); female P. l. masranii reach similar sizes, though fossil females have wingspans of only 3.8 meters (12 feet). What are presumed to be male P. l. longiceps are somewhat larger than female P. l. masranii, while female P. l. hippocratesi have broader wingspans yet. They are also the heaviest subspecies, with males weighing five to six hundred pounds according to Flyers. In the Jurassic Park /// arcade game, Jurassic Park ///: Park Builder, and the Jurassic Park /// Field Guide, the male (referred to as “Giant Pteranodon“) is much larger than the females, having a 14-meter (45.9-foot) wingspan according to the latter two.

The “giant” or “alpha” P. l. hippocratesi, generally believed to represent the male of the subspecies.

Fossil Pteranodon longiceps are estimated to weigh between 20 kilograms (44 pounds) to 93 kilograms (205 pounds) in adult males; P. l. masranii females weigh 66 to 70 pounds as adults. In contrast to the junior novels, the Jurassic Park /// Field Guide lists “giant” Pteranodons as weighing 180 pounds, though it still gives them 45-foot wingspans. The standard Pteranodon is said to have a 30-foot wingspan and weighs 120 pounds. According to the mobile application Jurassic World Facts, the body length of a Pteranodon is about eight feet and the wingspan is twenty-four feet (a different entry claims it is 24.6 feet); it is unclear which subspecies this is meant to refer to. The mobile game Jurassic World: The Game, which features what appears to be a male P. l. masranii, describes their wingspan as ranging from twenty-six to thirty-three feet, which would make it larger than any Pteranodon known in the fossil record. Despite this, the game’s fact file claims that P. l. masranii is most similar to fossil specimens out of all three subspecies.

When standing on all fours, an average adult Pteranodon is roughly as tall as a human, though some can easily grow to a larger size than this.

Adult female P. l. masranii displaying its expansive wing membranes. Note that, when not flaring its wings as a form of display, it is a quadruped.

The wings of Pteranodon and Geosternbergia are formed from the arm bones, as with all pterosaurs. The majority of the wing membrane, called the brachiopatagium, stretches from the tip of the fourth finger to the hips. In addition, all four have a smaller membrane called a propatagium which stretches between the shoulder and the elbow. There are three exposed fingers, as well as the fourth one that supports the wing. The membrane is muscular in order to allow the animal to achieve powered flight. Part of the membrane stretches to the tail, though this is not particularly prominent; this membrane is called the uropatagium. The tail itself is short and stiff, and reaches up to 25 centimeters (9.8 inches) long. Wing morphology differs between InGen’s pterosaurs and their fossil counterparts; the wings of fossil pterosaurs have rounded tips, while InGen’s have narrow, pointed wingtips. Overall, InGen’s pterosaurs have less membrane surface area than the extinct versions, which may increase the effort required to remain airborne.

The muscular wings and strong upper body of the Pteranodon allow it to fly at high altitudes, though its piscivorous diet means that it is often seen closer to sea level.

The legs and feet are substantially different than those of fossil pterosaurs, having very bird-like anatomy that permits them to grasp. While Geosternbergia has not been observed using its feet to grasp objects, P. longiceps has been known to do this extensively, whether it is to perch bipedally or grab prey items. The legs are very muscular, and the feet have four toes; one of these resembles a thumb. When walking, they use a quadruped gait, with the wing finger held upward. Flight in fossil pterosaurs was achieved by pushing off with the wings from a quadrupedal position; InGen’s pterosaurs use a variety of other methods to achieve flight, such as standing on two legs and jumping while forcefully flapping the wings, or standing into the wind and spreading the wings wide. Why they opt to use these more energy-consuming methods is unknown, but may be a result of their altered leg and wing anatomy. According to the mobile application Jurassic World Facts, the flight speed of a Pteranodon is twenty-five to thirty miles per hour. This same application describes their flight pattern as similar to the modern albatross, though P. l. masranii flies with near-constant wing strokes rather than gliding with occasional flapping. It may fly with rapid flaps at times, while using thermal updrafts from the ocean to fly slowly at other times.

This model depicts the animal with a coat of pycnofibers, but these are absent in specimens that have been directly observed.

Fossil pterosaurs are believed to have had coatings of pycnofibers, which are similar to feathers or hair. However, among InGen’s pteranodontids, only Geosternbergia possesses pycnofibers. The others have bare, leathery skin. Some models of P. l. longiceps do depict it with a full coating of pycnofibers; this variant has never been observed directly.

Coloration among the pteranodontids is quite variable. Geosternbergia has a blue body, white countershading, and a yellow beak and crest. The wing membranes are yellow-green. Somewhat similar coloring is seen in Pteranodon longiceps longiceps, but its body has a greener tone, and the beak and crest are darker in color than Geosternbergia. Some artwork has depicted them with a red crest, blue body, white countershading, and yellowish-tan wing membranes. The junior novel Survivor describes one with red and gold wing markings.

Mature female P. l. hippocratesi

Female Pteranodon longiceps hippocratesi are generally tan or pink in color, though the dorsal side is patterned black or gray. The face has elaborate patterning with yellows, tans, pinks, and grays decorating the head and beak, while the crest is colored similarly; the eyes have dark orange sclerae and round black pupils. The area around its eyes, as well as its throat, are red or pink. Its wings are generally lighter in color than the dorsal side of its body, and often have dark mottling toward the tips. A darker-colored individual, presumed to be a male, has been depicted in a number of sources; it is mostly gray, with white or very light gray markings near the wingtips. The neck has whitish speckling and the crest becomes bright red toward its tip, but it lacks other significant coloring. The junior novel Flyers describes a senescent male P. l. hippocratesi with plain brown colors and a sexually-mature adult with crimson markings, a sexually mature female with blue and gray markings, and four subadult males; one of these has gold-tinged wings, another has silver streaks on the wings, a third has red wings, and the final one has blue and red spots on its wings.

Color variants of female P. l. masranii, illustrated by Ian Joyner

Pteranodon longiceps masranii has a more subdued, grayish body color in most individuals, though some have reddish, bluish, purplish, or brown shading. The undersides of the wing membranes are orange, with some black or gray streaking, and the face and crest have a tough mask-like covering which may be light to dark red, cyan, or violet. The red color appears to be most common, appearing on all individuals (about 35 animals) witnessed in 2018. Some individuals have a similar color down the spine, while others have white or pale yellow stripes on the wings.

Growth
Flapling P. l. hippocratesi

Juveniles of P. l. hippocratesi have been observed. They have a color pattern similar to the adult females, but less elaborate and lighter-toned. Young juveniles, often called “flaplings,” are capable of flight already despite not being as powerful as adults. Upon hatching, the wings are actually fully developed; while they are within the egg, the wings are tightly folded. The crests are very small at this stage, and grow out as the animals mature. The head is proportionally larger, and the eyes are considerably larger in proportion to the head. At the stage at which the flaplings were observed in 2001, they had wingspans of approximately nine feet and body lengths of three feet and two inches.

Flapling P. l. masranii

Flapling P. l. longiceps can be hatched in the mobile game Jurassic Park: Builder, which similarly depicts them as having smaller crests and larger heads than the adults. Finally, the flapling stage of P. l. masranii is a part of Jurassic World: The Movie Exhibition, also demonstrating a smaller crest but overall larger skull size in relation to the body. The beak is proportionally shorter than that of the adult, a feature not seen in the other subspecies.

The natural growth rate is unknown, but a female P. l. masranii hatched in 2011 or 2012 was fully mature by 2018. This would mean it matures in six or seven years. In the junior novel Flyers, four juvenile male P. l. hippocratesi were strong enough to fly long distances and hunt for themselves and were large enough to be a threat to humans despite being, at most, around a year old.

Sexual Dimorphism
Pteranodon longiceps hippocratesi. The upper individual is believed to be male, while the lower is believed to be female.

Considerable confusion exists as to the sexual dimorphism in all four of these animals. In fossil specimens of Pteranodon longiceps, females are considerably smaller than males, with an average wingspan of twelve feet (compared to the average male’s eighteen-foot wingspan, while exceptional individuals reached around twenty feet). Male Geosternbergia typically had wingspans between ten and nineteen feet, while particularly large ones could have wingspans of around twenty-three feet; females averaged half that size. In addition, the crests of females in both genera were short, small, and rounded, while males had longer crests. Males can be identified by their extremely narrow hips, while females have wide hips with large pubic openings in order to lay eggs. InGen’s pteranodontids do not appear to follow these patterns.

For a time, there was some speculation that InGen’s Geosternbergia was the male form of Pteranodon longiceps longiceps, while all of the P. l. longiceps were the females of this single species. This has not been confirmed as fact. All individuals of these two taxa that have been observed so far resemble males of their respective genera, but their actual sexes have not been confirmed. Canon consultants have since treated Geosternbergia as separate from Pteranodon, suggesting that InGen did create both genera separately and that they are not intended to represent sexual dimorphism within a single species. However, some concept art does indicate that the females of both would have been duller in coloration.

Pteranodon longiceps hippocratesi females can achieve thirty-two-foot wingspans and possess the long crests of males, though adult males have never been observed directly. A model of a dark-colored, red-crested individual of this subspecies exists, and is generally assumed to represent the male. In the Jurassic Park /// arcade game and in Jurassic Park /// Builder, the dark-colored presumed male is depicted as being significantly larger than the females with a 14-meter (49.5-foot) wingspan, but the junior novel Flyers depicts them as being roughly the same size. Aside from coloration and maximum hypothetical size, there do not appear to be any differences between the sexes.

Only females of Pteranodon longiceps masranii have ever been observed up close. They have very short and small, but triangular, crests; however, they have 18-foot wingspans, making them as large as the average adult male in the fossil record. What appears to be a male version of P. l. masranii appears in the mobile game Jurassic World: The Game, though the Code 19 dialogue still refers to it as a female; it possesses the longer crest of the male. In-game factoids describe it as having a 26- to 33-foot wingspan, while the Jurassic World website gives 18 feet as its wingspan. This would mean that males are between 31% and 45% larger than females, compared to the fossil record’s range of males being 33% to 41% larger than females. If correct, this would mean that Pteranodon longiceps masranii is more or less comparable to fossil P. longiceps in terms of sexual dimorphism, the main exception being its significantly larger size.

Male Pteranodons, assumed to be P. l. masranii, appear in the background in the motion comic episode “Dinosaur Crossing,” but not in significant detail. The stylization of the comic means that its depictions may be unreliable.

Habitat
Preferred Habitat

Pteranodon and Geosternbergia are both marine species, living on coastlines. In prehistory, they inhabited the Western Interior Seaway, a shallow warm-water ocean which separated North America into two subcontinents during the Cretaceous period.

Three female Pteranodon longiceps masranii fly northward along the Pacific coast of North America (date unknown; sometime after 6/25/2018)

Their preferred habitats are rocky islands and cliffs near the ocean, where they can remain safe from terrestrial predators and search for fish and other prey. Pteranodon, at least, appears to be highly migratory; Jurassic World researchers estimate that they would range over territories spanning 120 miles or more. Their habitat must consist of more than 430,000 square feet and contain a large, deep body of water. However, they may venture into desert and grassland environments where larger water sources are rare; when they do so, they are likely to gravitate toward freshwater lakes or artificial sources of water.

When released from captivity, Pteranodons have almost exclusively been seen to migrate northward away from the equator. Following their escape from Isla Sorna, P. l. hippocratesi has been known to venture as far north as British Columbia, farther north than any other subspecies; P. l. masranii has been confirmed reaching Nevada. This makes Pteranodon a highly adaptable creature, capable of surviving in the humid tropics as well as arid deserts and the cooler climates of the Pacific Northwest. In the Nevada desert, temperatures during the daytime in July can reach up to 110 degrees Fahrenheit (43.3 degrees Celsius) and during the night drop as low as 40 degrees Fahrenheit (4.4 degrees Celsius). Such a rapid and extreme temperature change is difficult for many species to adapt to, but at least three P. l. masranii were sighted in Las Vegas, Nevada during July and therefore are known to be able to tolerate these drastically changing temperatures without noticeable difficulty.

Thus far, P. l. longiceps and Geosternbergia have not been known to migrate outside of the Gulf of Fernandez, but this is probably due to their relative scarcity.

Isla Nublar

An aviary was constructed in Jurassic Park on Isla Nublar to contain Pteranodons. However, no DNA from this species was maintained on-site, and the animal itself would likely not be put on display until Phase II of the Park’s operation. This aviary was situated on the Jungle River and was bordered to the east by the Stegosaurus paddock. The main tour road, as well as the Jungle River Cruise and one service road, would have crossed through the aviary. To the north and south, the aviary would have bordered undeveloped land, while to the west it would have bordered some undeveloped land and the southeasternmost corner of the Proceratosaurus paddock.

Known range (red) of P. l. hippocratesi on Isla Nublar, as of June 11, 1993

It is unknown how many Pteranodons were present on Isla Nublar as of June 11, 1993. However, based on Jurassic Park: The Game, there were at least three or four Pteranodon longiceps hippocratesi on the island at the time of the incident, which escaped confinement and traveled northward and eastward across Isla Nublar before disappearing. Three were sighted near the eastern mountain range on June 12, and one was sighted north of the Visitors’ Centre near the Western Ridge later that evening. Their location after the incident is unknown.

Known (red) and hypothetical (purple) range of P. l. hippocratesi on Isla Nublar, following June 12, 1993

In 2004, a new aviary was constructed on Isla Nublar near the northern end of the Jungle River to house the Pteranodons. Originally the structure was going to be made of glass, but the pterosaurs were able to break the glass using their beaks and escape. This necessitated the development of a new transparent polymer to use instead. As of August 2004, there were no pterosaurs on Isla Nublar; however, InGen geneticist Dr. Henry Wu had created the first embryos of Pteranodon longiceps masranii by that time, and twelve eggs were incubating. These included one individual (Egg B) exhibiting polydactyly on one wing, meaning it bore an extra finger. According to Dr. Wu, the incubation period lasts for several months; it is not currently known when the first P. l. masranii hatched.

Between late 2004 and Jurassic World‘s opening date of May 30, 2005, the remaining Pteranodons were removed from Isla Sorna and transported to the Jurassic World Aviary. These would have included any remaining P. l. longiceps and P. l. hippocratesi, though it is not known how many of each remained by that time. Additionally, P. l. masranii would have been introduced to the Aviary as they matured. There do not appear to have been any further attempts to breed the other subspecies, and no Geosternbergia are believed to ever have been on Isla Nublar at all. By the end of 2015, the sole remaining pteranodontid on Isla Nublar appears to have been P. l. masranii.

A female Pteranodon longiceps masranii is confirmed by the Dinosaur Protection Group to have hatched in Jurassic World sometime between May 15, 2011 and May 17, 2012. This individual was said to still be alive as of May 15, 2018 and was a six-year-old adult at that time.

On December 22, 2015, an incident involving the hybrid Indominus rex caused a breach in the Aviary; this permitted the pterosaurs to escape. Upon being released, they initially flocked westward over the forests, but turned south somewhere near the ruins of the old Visitors’ Centre. From here, they flocked toward the largest nearby body of water, the Jurassic World Lagoon. At least 44 adult females were involved in this. The ensuing incident left several Pteranodons deceased; some were shot in defense of JW001 and Simon Masrani, and many were tranquilized in midair by the Asset Containment Unit; some of these either fell from such a height as to sustain fatal injuries or fell into the Lagoon where they likely drowned. One was consumed by the Mosasaurus which inhabited the Lagoon. At least eleven of these animals died during the incident. During the night, by the time Sector 3 had been largely abandoned, the animals began to return; some of them followed the monorails as they continued along their automated paths. At least two monorail trams were destroyed by the Pteranodons; during the second such incident, one animal was struck by a monorail, but it is not confirmed whether it died. It is similarly uncertain whether any died in explosions that occurred when the monorails were destroyed.

Once the island was largely abandoned, the Pteranodons settled and began finding new homes. At least five were confirmed in Gyrosphere Valley as early as December 23, while a few others stayed near the Lagoon; however, since this was not a safe fishing hole due to the mosasaur, not many seem to have fed here. According to the Jurassic World: Fallen Kingdom LEGO website, some of the Pteranodons remained near the Lagoon, roosting in the abandoned hotel complex. Others nested on the island’s coastal cliffs, as the ocean was a far safer place to feed than the Lagoon. Their most prominent nesting site was the Eastern Ridge, where they were confirmed roosting by December 24.

In June 2016, the mosasaur was accidentally released into the ocean and migrated westward, but the pterosaurs do not appear to have returned to the Lagoon in large numbers, since none were sighted there in 2018. As of June 2016 their roosting grounds appear to have been concentrated in the areas bordering the central valley.

At least four adults were seen near Isla Nublar’s southern coast on February 17, 2018; they had longer crests than an average female, but their sex is unconfirmed. They possessed a combination of features of different subspecies; their long and sharp beaks resemble those of P. l. masranii, but they possess teeth like P. l. hippocratesi. This implies that they may be crossbreeds of these two subspecies. If true, they can be differentiated from full-blooded P. l. hippocratesi by the larger size, but smaller number, of their teeth as well as the sharp beak tip.

A photograph procured by the Dinosaur Protection Group and featured in a June 5, 2018 article by Klayton S. confirms the continued presence of P. l. masranii on the island. At least twenty-eight adults or subadults can be seen in the photograph. All of these appear to be female. The location at which the photograph was taken is unknown, but appears to be on one of Isla Nublar’s mountains, probably within the Eastern Ridge as they were known to nest here.

They appear to gradually have migrated north, and on June 23, 2018 they were sighted almost exclusively within the vicinity of Mount Sibo. A flock of at least thirty-five females was seen fleeing from a volcanic disturbance at around midday, and a similarly-sized flock of females could be seen fleeing the island during the most violent part of the volcano’s eruption. At least one was killed during the eruption due to being struck by volcanic debris, while three others were removed from the island via the S.S. Arcadia by the mercenary team led by Ken Wheatley at the behest of Eli Mills.

The 2018 population size can be used to estimate the 2015 population. At least 35 individuals were seen west of Mount Sibo, and at least 35 were later seen east of Mount Sibo. These two flocks may have been the same group of animals, but if they were two different flocks, there would have been at least 70 animals on the island at that time. Additionally, three were already caged on the Arcadia, plus at least 11 which died during the 2015 incident. This means that there were at least between 49 and 84 Pteranodon longiceps masranii in the Jurassic World Aviary before the incident.

After the eruption of Mount Sibo, it is likely that many of the Pteranodons fled the island. Some, however, were seen perching on coastal rocks while the eruption continued, and as their diet of fish would have been uninterrupted by the volcano’s activity, it is not impossible for a decent number to have remained on Isla Nublar’s coastal cliffs even after the eruption occurred.

Isla Sorna

Pteranodon longiceps longiceps, Pteranodon longiceps hippocratesi, and Geosternbergia were all cloned by InGen between 1986 and 1993. At least three P. l. hippocratesi were fully mature by June 12, 1993 and were present on Isla Nublar, so it is likely that they were cloned some years prior to this. An aviary was constructed on Isla Sorna to contain the P. l. hippocratesi, but the containment measures for the other pterosaurs is currently unknown. The existence of other aviary structures on the island is currently unconfirmed. As of InGen’s last count in 1993, there were ten Pteranodons living on the island; how many were actually Geosternbergia is not known, or if any Geosternbergia remained by 1993. Including both P. l. longiceps and P. l. hippocratesi, there were precisely ten adult Pteranodons encountered between 1997 and 2001, though the P. l. hippocratesi are said to have been a covert project according to film director Joe Johnston and thus may not have factored into InGen’s overall headcount.

By 1997, the pterosaurs with the exception of P. l. hippocratesi had broken free of whatever containment measures were in place on Isla Sorna. Six adult P. l. longiceps were sighted in the island’s northern grasslands on May 25; they are presumed to be males due to their crest morphology, but this is not confirmed for certain. Older concepts for The Lost World: Jurassic Park would have depicted either Pteranodon or Geosternbergia living within and near the Workers’ Village. Several may have lived off of the island’s southwestern shore according to older Jurassic Park /// scripts, which indicate that these pterosaurs were responsible for the deaths of Enrique Cardoso and his associate. The junior novel Survivor depicts at least two Pteranodons, one living on the sandy western coast of Isla Sorna (which may have been killed by a Tyrannosaurus) and another circling near the Embryonics, Administration, and Laboratories Compound. The first sighting was on May 23, while the second sighting was on May 27. Several days later, a Pteranodon was sighted in the southwestern part of the island over a dry and deforested valley; Eric speculated that it might have been the same individual he saw on May 23, suggesting that it was colored similarly. The next junior novel in the series, Prey, depicts one adult Pteranodon roosting on Mount Hood on the night of December 30 and one flying near the Jurassic Park Ranger Station on the morning of December 31.

Known (red) and hypothetical (purple) range of P. l. longiceps on Isla Sorna, between 1997 and December 31, 2001. Note that the range of Geosternbergia may overlap with the central waterway range of Pteranodon.

The range of Pteranodon longiceps hippocratesi was much more restricted. Unlike the other pterosaurs, they failed to escape containment on their own. However, they did successfully breed, which implies that at least one male existed in the aviary at some fairly recent point. On July 19, 2001, there were four adult females and six flaplings in the aviary. An incident occurring that day resulted in the death of one of the adults and possibly one of the flaplings; the surviving three adults were unintentionally released and left Isla Sorna.

According to the junior novel Flyers, the third in the series, a family of escaped Pteranodons were eventually returned to the aviary on Isla Sorna in July of 2002. The novelization implies that they are the individuals released on July 19, but the descriptions do not match up, and later material confirmed that those particular animals were neutralized in Victoria, British Columbia. Additionally, the novelization makes mention of several other adult males in the aviary which do not appear. It has been speculated since the release of Jurassic Park: The Game that these inconsistencies could be cleared up by explaining the pterosaurs from Flyers as the escapees from the Jurassic Park Aviary, but as Universal Studios has not confirmed the second two Jurassic Park Adventures junior novels or the existence of the game’s Pteranodons as part of the official film canon, there is no evidence for the hypothesis. Nonetheless, The Evolution of Claire officially confirms that escaped Pteranodons were, in general, recaptured and held on Isla Sorna.

Location (red) of the Site B Aviary which contained P. l. hippocratesi

Between late 2004 and May 2005, the remaining Pteranodons from Isla Sorna were rounded up and transported to Isla Nublar by Masrani Global Corporation, where they eventually died out while being replaced by Pteranodon longiceps masranii. Supposedly, none remain on Isla Sorna. The fate of the Geosternbergia is unknown; official sources have described them as “missing in action” when inquired about them, implying that InGen lost track of them at some point.

Other

Because of their strong flight capabilities, Pteranodons have left the islands in the Gulf of Fernandez where they were bred on numerous occasions. Three adult female P. l. hippocratesi which were released from the Site B Aviary on July 19, 2001 were eventually sighted over Victoria, British Columbia later that year. American security contractor Vic Hoskins apprehended and neutralized them.

Adult female P. l. masranii attacks one of its captors at the Lockwood Estate near Orick, CA, 6/24/2018

A different fate is given for the escapees in the junior novel series Jurassic Park Adventures; they are mentioned in Prey and physically appear in Flyers, by which point the flock consists of an old male, a mated pair including the elder’s daughter, and four sons belonging to the mated pair. The pterosaurs travel the subtropics around the world, with confirmed sightings and attacks in South America, Mexico, Texas, and the Florida Keys. Unconfirmed rumors had placed the pterosaurs in farther parts of the world, such as Britain and Japan. In July of 2002, the family had reached Florida and flocked to Universal Studios at Orlando due to the large artificial pool in the center. A tense day-long standoff between the pterosaurs and parkgoers ends with the animals being led to Isla Sorna and housed in the aviary. As discussed above, the idea that these are the Isla Sorna Pteranodons contradicts several aspects of the film canon, and it has been hypothesized that they could actually be the Isla Nublar Pteranodons from Jurassic Park: The Game, whose existence is also questionable.

Jurassic World’s official website states that the natural range of their pterosaurs would cover over 120 miles, which would allow them to comfortably fly between Isla Nublar and western Costa Rica.

Mercenaries led by Ken Wheatley retrieved at least three adult female Pteranodon longiceps masranii from Isla Nublar shortly before the eruption of Mount Sibo on June 23, 2018. They were removed from the island via the S.S. Arcadia and transported to the Lockwood Estate near Orick, California where they remained during the night of June 24. They were released from captivity by Maisie Lockwood to save them from hydrogen cyanide poisoning, upon which they fled into the surrounding pine forest. Shortly after this date, three adult female Pteranodons were seen flying north along the American West Coast; these were presumably the same three that were released previously. At least two adult male and two adult female Pteranodons were seen flanking a passenger aircraft in June or July 2018.

Also in July 2018, three adult female P. l. masranii were sighted at the Vegas Strip in Las Vegas, Nevada. This would place them roughly 150 miles from the nearest ocean, making this the farthest that Pteranodons have been confirmed to travel from the coast. The animals most likely followed the Colorado River north from its estuary in the Gulf of California, searching for sources of water. Pteranodons were sighted diving into Lake Mead earlier in the same week as the Vegas incident, suggesting that they had migrated northeast in search of food and water in the desert.

In 2019, a single female P. l. masranii was filmed by wedding guests preying on a released dove at the ceremony. The location of the sighting has not been released. The pterosaur involved in the sighting was smaller than typical, suggesting that it may have been a subadult.

Behavior and Ecology
Activity Patterns

Pteranodons are cathemeral, active at some times during the day but resting at other times. They appear to sleep at night, but are shown to become active at night if disturbed in the junior novel Prey as well as deleted concepts for The Lost World: Jurassic Park (which also considered featuring Geosternbergia, implying a similar behavior pattern). In both cases, the animals were portrayed becoming aggressive toward the people who had disturbed them. Sometimes, P. l. masranii has been observed active at night, typically in areas where human activity alters their behavior patterns; they are particularly easily disturbed by noise, light, and visible moving objects.

The junior novel Flyers, on the other hand, portrays a family of P. l. hippocratesi adopting a nocturnal lifestyle in order to avoid human activity, a behavior that has been observed in numerous mammals in real life. However, they still hunt during the day, though they prefer to actually feed on their captured prey in the evening or at night.

Diet and Feeding Behavior

Pteranodon and Geosternbergia are considered to be piscivores, meaning that their diet consists largely of fish. While the feeding behaviors of Geosternbergia have never been witnessed and can only be inferred, these behaviors have been observed extensively in Pteranodon.

Nesting family of P. l. hippocratesi. Note the juvenile Tyrannosaurus rex carcass in the nest.

Despite being considered a piscivore, this animal does actually feed on animals other than fish. It has been observed targeting humans and small dinosaurs for food, and likely preys on many types of small animal. When permitted to live in the wild, it will usually hunt fish in the ocean; however, it has done well in captivity on a diet of brackish-water or freshwater fish and other small animals. In the junior novel Survivor, a Pteranodon longiceps longiceps was observed flying low over the ocean’s surface and grabbing fish as they jumped; the species of fish is unidentified. The same animal pursued a human across the beach shortly thereafter, but was unsuccessful. When hunting small prey, the sharp and pointed beak is used as the primary weapon, stabbing or scratching at the prey item. Biting at prey is most effective in Pteranodon longiceps hippocratesi due to its teeth, which are absent in all other forms of pteranodontid. Often, these pterosaurs will soar high above the ground riding warm updrafts while scanning the area below for food. Both P. l. hippocratesi and P. l. masranii are known to feed on birds by diving at them while in flight and rapidly grabbing them with their beaks.

Due to genetic modification, InGen’s Pteranodons (and presumably Geosternbergia) have grasping feet similar to those of birds of prey. While fossil specimens would only be able to use their feet to walk, InGen’s have adapted to their altered anatomy in several ways, most prominently by using their feet as hunting implements. A Pteranodon will grasp the upper body of a potential prey item and lift it into the air, dropping it into an ideal feeding location. However, this is not always a viable way to capture prey. An adult female Pteranodon longiceps hippocratesi struggles to hold an adult human aloft, though it can carry a child without much difficulty. On the other hand, an adult female Pteranodon longiceps masranii can easily lift an adult human into the air with no trouble at all; they have even been observed trying to pick up juvenile Triceratops, which can weigh around five hundred pounds, but can only hold them above the ground for a few seconds before dropping them. If a Pteranodon drops its prey from a great enough height, or onto dangerous terrain, it may easily injure its prey.

Additionally, Pteranodons can take advantage of water to collect prey. In the junior novel Flyers, as well as during the 2015 Isla Nublar incident, Pteranodons were observed picking up humans and dropping them into large bodies of water where they would feed on them later. This was shown to be highly organized in Flyers, due to the dramatically increased intelligence of the P. l. hippocratesi and their strong familial relationships. Collected food would be eaten later, when the strongest male of the group permitted the family to feed. This cacheing method was less organized with the P. l. masranii on Isla Nublar, which only successfully captured one human this way, and continuously squabbled over their prey. However, unlike P. l. hippocratesi, which does not appear able to swim, P. l. masranii is an adept diver and can swim under the surface of the water. It may plunge into water from a great height by folding up its wings and essentially turning its body into a giant dart. Fossil evidence suggests that they could also float on water similarly to seabirds, plucking prey from beneath.

Escaped P. l. masranii engage in a feeding frenzy at Main Street, Isla Nublar (12/22/2015)

A feeding behavior only observed in Pteranodon longiceps masranii involves a “feeding frenzy” method in which a large number of pterosaurs generates panic in their prey by swooping and vocalizing loudly. This initiates a fight-or-flight response in prey items, which may then act illogically while attempting to escape. While the prey items are panicked, the Pteranodons may easily pick off stragglers or injured animals. They have been observed associating with the smaller Dimorphodon in such feeding frenzies. Despite the apparent group effort of these hunts, there is no actual cooperation among the animals; they are known to fight over food and do not assist one another in any intentional way.

Other methods of subduing prey used by Pteranodon include plowing into prey feet-first with wings outspread to knock over multiple victims, landing on top of a prey item and repeatedly bashing it into a hard surface, and making hit-and-run strikes with their talons while airborne. They may also scavenge carrion; a model produced for Jurassic Park /// portrays P. longiceps hippocratesi perching on a rocky structure with the carcass of an infant male Tyrannosaurus rex nearby.

Pteranodon dung is distinctively birdlike, being whitish and caking onto surfaces. When dry, it crumbles easily. Its appearance and texture are similar to that of other piscivorous animals.

Social Behavior

Pteranodons are highly gregarious animals, nearly always seen among members of their own kind. It is possible that they may flock with Geosternbergia, which is presumed to be similarly social, but this is not known. Pteranodons are frequently seen in groups of three; however, they have also been sighted solitarily from time to time.

Pteranodon rookery on Isla Nublar, 2017/2018. Image from the Dinosaur Protection Group.

Due to its population size, the largest flocks of Pteranodon were those of the P. l. masranii on Isla Nublar. Flocks of these pterosaurs may contain thirty to forty animals. Since the eruption of Mount Sibo disturbed their nesting ground on the volcano’s slopes, it is not known if their flock sizes are still similarly large. Groups of one to four have been sighted commonly since 2018.

When socializing with one another, Pteranodons tend to be quite loud. However, not all of their social displays involve vocalization. The head crests are presumed to be display structures, and the variability of facial and crest color in P. l. masranii in particular implies that they engage in visual displays. The bright red crest of the black P. l. hippocratesi also suggests a display purpose, and if this is the male of the subspecies, it is likely used for courtship or to intimidate rivals.

Large flock of Pteranodons, accompanied by smaller Dimorphodons. Both of these pterosaurs are highly social, gregarious animals.

According to the account in Flyers, familial bonds among Pteranodon longiceps hippocratesi can be extremely strong. In this account, a family group consisting of a senescent male, his daughter, her mate, and their four sons settled in Orlando, Florida after being attracted to a large artificial pool in the Universal Studios theme park during a drought. The eldest male, grandfather to the four young males, acted as a kind of mentor figure to the younger animals. In animals with strong family ties, the role of mentor is often taken up by animals that have passed reproductive age. The sexually-mature male was in charge of coordinating the hunting efforts and was high in the family’s pecking order; however, he would not start feeding until all his family was present and safe, indicating a level of empathy that likely resulted from the unusually increased intelligence of P. l. hippocratesi demonstrated throughout this account. The entire family group delayed their migration through the area due to their second-youngest flock member becoming injured; they refused to abandon one of their offspring even when there was no chance of his recovery without human intervention.

A deleted concept for Jurassic World also demonstrates a similar empathic tendency in Pteranodon longiceps masranii, despite that subspecies’s lesser intelligence and tendency toward squabbling. In this scene, one of the Pteranodons would have been seized by the Mosasaurus while attempting to extract a human from a damaged monorail over the Jurassic World Lagoon. Several other Pteranodons would have rushed to its aid, pecking at the mosasaur and risking their own safety in an attempt to rescue one of their flock.

Reproduction

Pteranodon and Geosternbergia both lay eggs in the manner common to pterosaurs. Pteranodon in particular roosts in safe areas where predators cannot reach, such as cliffs, rocky pinnacles, and tall buildings; it nearly always nests near a source of food and water. At least P. l. masranii may decorate its nests with attractive objects it finds, since some have been seen collecting small metal objects. Even non-reproductive Pteranodons will build nests to live in; the eggs are probably laid in preexisting nests.

According to Flyers, the males of at least Pteranodon longiceps hippocratesi will engage in physical combat in order to win the rights to mate; their efforts may be judged not only by the females, but by the females’ male relatives. In this subspecies, pair-bonds are very strong and may persist for years, if not a lifetime.

The eggs are bright white, roughly three times the size of a chicken‘s egg; twelve eggs from surrogate parents were incubated simultaneously to hatch the first-ever Pteranodon longiceps masranii. Which subspecies was used to obtain the surrogate eggs is currently unknown. Similarly, a nesting colony of either P. l. longiceps or Geosternbergia was planned to appear in The Lost World: Jurassic Park and would have contained twelve eggs (some of which would be unintentionally destroyed by a helicopter, prompting the parents to attack). The Pteranodon longiceps hippocratesi living in the Site B Aviary as of the summer of 2001 were observed to have six living offspring, while the breeding pair of the family group from Flyers had four living male offspring as of July 2002; however, the original number of eggs laid in each case is not known. Nests consist of sticks, driftwood, foliage, and other debris that the parents collect from the environment, and are typically placed on top of natural or artificial structures where predators cannot easily reach.

A mother Pteranodon provides live food to her flaplings. This allows them to practice killing their own prey without having to leave the shelter of their nesting area.

After an incubation period of several months, the eggs will hatch; the young are sometimes called “flaplings” at this stage because they are capable of limited flight fairly quickly. Young flaplings are already tenacious predators, eagerly pursuing prey and taking it down using methods similar to the adults. In P. l. hippocratesi, adults may actively teach their young how to hunt. One individual (presumed to be the mother) will often deliver live food to give the flaplings the opportunity to hunt and kill live prey, helping them develop essential survival skills for later in life. While other adults will cooperatively protect the nesting territory, only the parent appears to provide direct care to its own offspring. The account in Flyers depicts a multigenerational group of P. l. hippocratesi maintaining very strong familial bonds between animals of three successive generations, even after the youngest had become capable of defending themselves. This behavior is unconfirmed in the less intelligent subspecies.

During the 2001 Isla Sorna incident, the pterosaur that was presumed to be the mother of six flaplings died due to the collapse of abandoned aviary infrastructure. Following her death, the other adults abandoned the offspring to fend for themselves, seeking out new nesting grounds on the North American mainland. While it is unusual for adult animals to vacate a territory for juveniles to inherit, rather than evict the juveniles, it may be that the limited size of the aviary and competition for prey with other piscivores made this a less suitable territory. The flaplings, while capable of flight, were too young to journey with the adults and were abandoned. This confirms that, while adults will tolerate the offspring of other Pteranodons, they do not act as surrogate parents. Adoptive behaviors like those seen in many dinosaurs are not known in pterosaurs at all, which may be due to the precocious nature of pterosaur young. Parents provide care for their own offspring, but not to juveniles with no genetic relation.

Communication

The sounds produced by Pteranodon are highly varied. Screeching, squawking, and cawing noises are often used to communicate with one another while flying, with different calls used while traveling or hunting. These sounds are probably used to relay information throughout the flock, and to coordinate their movements as to avoid midair collisions. Even lone Pteranodons can be heard making such vocalizations, suggesting that they serve purposes beyond intraspecies communication. The coloration of the body is variable enough to suggest that it helps the animals identify one another, and the crest is believed to serve as a visual signal as well. The junior novel Flyers details the ways that Pteranodon longiceps hippocratesi visually identify one another using body coloration, as well as their specific social, aggressive, and distress calls.

Cawing sounds are often used to signify aggression, and may be used during hunts to startle prey into moving. Growls and hisses are also used when confronting threats, in addition to loud shrieks which are used to intimidate enemies or alert other Pteranodons to danger.

Ecological Interactions

A primarily coastal-dwelling species, the Pteranodon feeds on fish that venture into shallow water, such as possibly bonitos. As they sometimes hunt on tidal rivers or bodies of freshwater, they could potentially come into conflict with spinosaurs; at least one has been witnessed competing for prey with a Velociraptor antirrhopus masranii. Prey other than fish are confirmed to include humans, which Pteranodons hunt with surprising frequency. They have been witnessed on occasion attempting to prey on juvenile Triceratops and Ankylosaurus, and chasing after Franklin’s gulls (Leucophaeus pipixcan). Since their introduction to North America, they have been observed feeding on domestic pigeons (Columba liva domestica).

Because of their wide-ranging habitat, Pteranodons share territory with numerous other animals. As they are rather aggressive toward other species and highly territorial, they likely come into conflict with rival species fairly often. On Isla Nublar, they have been sighted in areas that were home to carnivorous dinosaurs such as Teratophoneus, CarnotaurusCeratosaurus, Allosaurus, and the fish-eating Baryonyx and Suchomimus, with which they likely competed for prey. Many species of herbivorous dinosaurs also live in their territory, as do the omnivorous Gallimimus and diminutive Compsognathus, which have unknown relationships to Pteranodon; it may have preyed on the smaller dinosaurs in its environment such as compies and Microceratus. Juveniles of many dinosaurs may have been food items; it has been seen near populations of Sinoceratops, Stegosaurus, Peloroplites, Ouranosaurus, Brachiosaurus, Apatosaurus, Stygimoloch, and Pachycephalosaurus, as well as possibly Pachyrhinosaurus.

On Isla Sorna, it was known to live close to animals such as Stegosaurus and Tyrannosaurus, which it does not appear to fear. This is despite two known attempts by Tyrannosaurus throughout the Jurassic Park Adventures junior novels to prey on Pteranodon; in one case, the Pteranodon had been intentionally harassing a tyrannosaur on a beach by diving low over its head. The tyrannosaur retaliated by snatching the pterosaur and thrashing it violently, possibly killing it. The second known interaction between the two species was a simple attempt at predation. Pteranodons on Isla Sorna also lived within the ranges of Brachiosaurus and, while confined to the aviary, Spinosaurus.

When walking, Pteranodon is a quadruped.

A supposed deleted scene from Jurassic Park /// would have had a violent conflict between Velociraptors and Pteranodons, but the nature of the scene is not known. However, a confirmed deleted scene from The Lost World: Jurassic Park would have featured Pteranodons and/or Geosternbergia living comfortably in Velociraptor territory and even laying eggs on top of a building within said territory. This suggests that these two types of animals can coexist without major issue, likely because they mostly hunt different prey. However, as mentioned above, a Pteranodon longiceps masranii and a Velociraptor antirrhopus masranii were in 2018 seen to compete for the same prey item, a juvenile Triceratops which had wandered away from its own kind.

Mosasaurus maximus is known to prey on Pteranodon; the artificial species of theropod Indominus rex has been observed attempting to attack Pteranodon without success. Its smaller relative, Scorpius rex, may have tried to prey on these pterosaurs in 2016; they were observed active during times of night when they would usually be asleep around the same time a Scorpius was seen in the area.

Not all of this pterosaur’s relationships with other animals are confrontational, however. Concept art implies that both Pteranodon and Geosternbergia can cohabit the same area, and Pteranodon longiceps masranii will tolerate the presence of Dimorphodon willingly. The exact nature of the relationship between Pteranodon and Dimorphodon is not fully understood, but it appears to be at least a commensal relationship, as the smaller pterosaurs benefit from the protection of the larger ones. Its aggressive territoriality may even indirectly benefit some animals. For example, the Troodon Jeanie laid her eggs in a hidden nest underneath a Pteranodon hunting ground; the pterosaurs’ aggression would keep away potential scavengers, meaning that the Pteranodons unwittingly acted as the eggs’ protectors.

It shows territorial behavior toward the huge Brachiosaurus, but is helpless to drive this sauropod away; to such a large animal, Pteranodon is a mere annoyance.

Cultural Significance
Symbolism

Pteranodon, and to a lesser extent Geosternbergia, has long been known to the general public as one of the most famous (if not the singly most famous) variety of pterosaur. It is universally common in paleoart, both professional and amateur, with plenty of reference material available thanks to the abundance of this reptile’s fossilized remains. Other forms of media, such as films, comic books, and video games, almost always utilize Pteranodon as the go-to pterosaur. Whether it is depicted as ambient wildlife or as an aggressive killer varies based on the creator.

Many misconceptions about Pteranodon exist due to its ubiquitous presence in media. Laypeople may not know its name specifically, recognizing it only as a “pterodactyl.” While it is indeed a pterodactyloid, it is far from the only one, and the term “pterodactyl” is often incorrectly applied to all pterosaurs regardless of their classification. It should, of course, be used only for the suborder Pterodactyloidea (especially pedantic scientists will insist that it be used only as a common name for the genus Pterodactylus in particular). Another misclassification by the general public is that pterosaurs are some type of bird, or that they are dinosaurs. Although they are neither, they are within the clade Ornithodira as essentially cousins to the dinosaurs; this makes birds their closest living relatives.

This unused Extinction Now! poster would have used imagery of the Victoria incident to sway public opinion in favor of re-extinction. Note that it contains not only grammatical errors, but terminology errors such as using “dinosaur” to mean “de-extinct animal.” Pteranodons are not dinosaurs, while naturally-extant birds are dinosaurs.

In media, Pteranodon is often depicted as the ecological equivalent of a bird of prey, hunting mainly terrestrial animals which it grabs in its feet and carries to a nest. In reality it is more comparable to a gull, a bold and opportunistic creature hunting and feeding mainly in water. While InGen’s pterosaurs have been genetically modified to have birdlike grasping feet, this feature was not present in the original animal, which grabbed food in its beak instead. It usually eats its prey not far form where it catches it, rather than bringing it to a nest to feed its young; in fact the young hatch with their wings nearly fully formed and can fly and hunt at a very young age.

Along with Tylosaurus kansasensis, the U.S. state of Kansas has adopted Pteranodon longiceps as one of its two state fossils as of 2014.

Since being brought back to life, Pteranodon has presented the clearest example of a de-extinct life form that is able to advance into new regions without human assistance. It is a good long-distance flier and has dispersed itself along the North American coastline multiple times. Due to several incidents of it arriving in areas used by humans, and due to its territorial nature, it has become a symbol of the threat de-extinct animals can pose to modern ecosystems and human safety. This has made it one of the most politicized de-extinct species, as discussed below.

In Captivity

The first Pteranodon or Geosternbergia to be brought back from extinction hatched sometime between 1986 and 1993, presumably at least a few years before 1993. There are two schools of thought on which Pteranodon subspecies was created first. One holds that P. l. hippocratesi was a kind of failed first attempt, with phenotypic aberrations that made it less suitable for Park exhibition. This hypothesis is slightly supported by Jurassic World: Evolution, where this subspecies is the default appearance and a phenotype resembling P. l. longiceps requires genetic modification. The other, which is presented in the junior novel Flyers, suggests that P. l. hippocratesi was modified intentionally to be smarter, stronger, and larger than a normal Pteranodon so that it could perform in shows for entertainment purposes. This second hypothesis is supported by Jurassic Park: The Game, which depicts this subspecies as being imported to Isla Nublar by 1993. Whichever the reason, P. l. hippocratesi was clearly held under much more secure conditions than P. l. longiceps.

It was intended to be displayed in Jurassic Park, where it would be visible from the main tour, the Jungle River Cruise, and the lodge within the aviary on Isla Nublar. At the moment little is known about the Jurassic Park Aviary other than it was scheduled to open for Phase II of park operations.

In late 2004, Dr. Henry Wu succeeded in producing Pteranodon longiceps masranii, which is believed to have fewer genetic anomalies than the preceeding subspecies. It was the first new breed of animal created for Jurassic World; twelve individuals were hatched. Eventually, this subspecies completely replaced those that had existed before it. For the full ten years of Jurassic World’s operation, these pterosaurs existed in the Jurassic World Aviary, where visitors could observe them from the secure walkways below as they soared about and fed on fish from the rivers. The Aviary remained a popular attraction throughout Jurassic World’s history and was frequently advertised, with the Pteranodons being prominently featured in marketing.

The needs of this pterosaur in captivity are extreme and hard to satisfy. While not as intelligent as some theropod dinosaurs, it still has very high stimulation needs and will quickly start exhibiting maladaptive behaviors if it becomes bored or stressed. These include increased aggression levels, making it dangerous for its handlers. In the wild, its gets the stimulation it needs from hunting, including diving into lakes or the ocean to chase fish. To keep it satisfied in captivity, it is recommended to give it the chance to hunt down its food in deep water. Unfortunately, artificial rivers are not large enough to provide for this need. Lakes and lagoons, as well as some larger natural rivers, are preferable. Cliffs are another beneficial habitat feature, since they like to perch where predators cannot reach. While the Jurassic World Aviary did not have deep enough water sources, it did provide a dynamic cliff-filled environment in which to roost.

Another challenge is keeping its habitat large enough for its population. This is a gregarious pterosaur and prefers to live in flocks, so having a large number is often favored; Jurassic World failed to make its aviary large enough for the huge flock it housed, however. This led to the pterosaurs becoming cramped, and therefore stressed. During the 2015 incident, the flock behaved very reactively, exhibiting aggression toward humans and violently attacking tourists. Such a tragedy could have been avoided by providing for the pterosaurs’ needs. However, it is also possible to overstimulate them. Too many moving targets, excessive noise, and dramatic lighting contrasts can make them agitated and are equally as bad as understimulation. There is a happy medium which must be met in order to safely and humanely keep this pterosaur in captivity.

During the park’s operation, a minor incident occurred in which a Pteranodon stole a tourist’s hat.

Science

Before Pteranodon, all the known pterosaur species were smaller toothed varieties chiefly known from Europe. This was the first major North American find, the first truly huge pterosaur found by paleontologists, and the first to demonstrate a toothless beak. This is where its name, which means “wing without teeth,” comes from. Since it is one of the most common pterosaurs in the fossil record, it has provided a wealth of information about these reptiles’ biology, including their anatomy, sexual dimorphism, evolution, and ecology. While less common, Geosternbergia has also made significant contributions to our knowledge of pteranodontid pterosaurs, especially its evolutionary relationship to Pteranodon. Fossils of distinct males and females, as well as different growth stages, are well-known. These pterosaurs are presumed to have been a major ecological presence in the Western Interior Seaway, a marine ecosystem which existed in North America during the middle of the Cretaceous period.

While it was the first pterosaur confirmed added to InGen’s genetic database and therefore the first pterosaur to have its genome analyzed by modern science, it has sadly not been quite as useful as a scientific model as it perhaps could have been. The first specimens were engineered in the 1990s, when paleontological knowledge about pterosaurs was not as complete as it is now, and numerous phenotypic errors were produced. Some, such as the presence of teeth in one variant, would have been obvious anomalies; others appear to have gone under the radar and are still present in modern Pteranodons. As a result, some aspects of its behavior are significantly altered. Some may have been altered by InGen on purpose for display and entertainment reasons. So, while some of its behaviors are probably similar to its ancient ancestor, any conclusions drawn from InGen Pteranodons cannot be completely applied to paleontology.

Politics
Footage of Pteranodons attacking tourists during the 2015 Isla Nublar incident has circulated for years and become emblematic of the supposed threat de-extinction poses to human life.

Because of its ability to fly, Pteranodon has never been possible to contain on remote islands with complete confidence and certainty, and this makes it more of a public safety risk than other de-extinct animals. Any escapes from the islands in the Gulf of Fernandez have invariably resulted in the pterosaurs migrating north along the American coastline, often taking them through inhabited regions. These are not limited to Latin America; some have been sighted as far north as British Colombia. Because of their migratory patterns and territorial behavior, Pteranodon has been an international political issue since the summer of 2001, when the first such incident was reported.

To deal with the Victoria incident, the Canadian government hired American security contractor Vic Hoskins to lead a team and neutralize the animals. His success at doing so led to him being hired by Simon Masrani to become the new head of InGen Security. Pteranodon remained out of the news for a little over a decade, but then returned in force during the 2015 Isla Nublar incident, which involved the escape of the entire Jurassic World Aviary population. The flock, already stressed from their insufficient living conditions and further agitated by the incident’s events, pursued a jeep to the Main Street area where they found a sufficient marine water source and abundant food. Unfortunately, their food items were the Jurassic World tourists and staff members. This led to a violent confrontation between the animals and park security, with the whole incident being recorded and publicized almost instantly. Scores of injuries and several deaths, both of humans and pterosaurs, resulted. It was this conflict more than anything else that ensured Jurassic World would close forever; recover from such an incident was impossible.

The attack on Main Street is essentially the centerpoint of the modern de-extinct animal rights debate. It has been cited as proof by organizations such as Extinction Now! that de-extinct animals are too dangerous to exist, an ideology rooted in the concept of prehistoric animals being inherently more vicious and bloodthirsty than their modern counterparts. On the other hand, experts cited by the Dinosaur Protection Group have explained that the pterosaurs were made aggressive by the poor conditions they were kept in, and that their behavior was only the result of mistreatment and neglect. This makes Pteranodon central to two separate but related animal rights debates: first, the issue of animal welfare in captivity, and second, the argument of which species should be prioritized over others.

During the Mount Sibo debates in 2017 and 2018, the issue of whether or not to voluntarily save Pteranodon from extinction was discussed at length by world governments, corporate entities, and special interest groups. The DPG was the leader in lobbying for their survival, while many people had suggested leaving carnivorous species like this one to die out while saving only herbivorous animals. In the end, the U.S. and Costa Rican governments as well as Masrani Global opted to let all of Isla Nublar’s inhabitants die, but the Pteranodons proved to be quite capable of saving themselves. Since the June 2018 eruption, they have been sighted up and down the North American coast, as well as occuring as far inland as Las Vegas, Nevada. Drawn to water sources and stimulated by bright lights, they are among those species most likely to come into dangerous conflict with humans, and authorities are still struggling to determine how best to deal with their presence on the continent. Whether they have established in other parts of the world, and how countries other than the United States are coping, is unknown at this time.

Resources

Since the 1990s these pterosaurs have been used as tourist attractions, being the first non-dinosaurian reptiles confirmed to have been brought back from extinction. Some may even have been genetically modified intentionally to make them more entertaining to park visitors. InGen intended to exhibit Pteranodon in Jurassic Park, and had researched this reptile extensively in the Site B facilities beforehand. While Jurassic Park never opened, its successor Jurassic World had massive success at exhibiting pterosaurs including Pteranodon, going so far as to engineer a new variant. This was the first new creature bred for the park, created in 2004.

Attempts to house Pteranodon also prompted Masrani Global Corporation to develop new technologies. While metal mesh had proven capable of containing Pteranodon, the company’s CEO Simon Masrani had wanted a transparent barrier for the aviary’s dome. Glass was considered, but shattered too easily if struck by the pterosaurs’ beaks. Eventually, one of his companies developed a transparent polymer which was much more resistant, and this was used to construct the aviary. It is believed that the polymer was also utilized for other industrial purposes.

Pteranodon has no exceptionally close living relatives, aside from other de-extinct pterosaurs. Its closest naturally-living relatives are the birds, since both pterosaurs and dinosaurs are in the clade Ornithodira. This would mean that Pteranodon has biochemistry unlike most modern life, which could make it a source of very unique biopharmaceuticals. As the first pterosaur added to InGen’s genetic library, it had more time to be studied than others such as Dimorphodon, and is probably better understood by genetic biologists. For this and other reasons, several Pteranodons were captured by Ken Wheatley in 2018 at the orders of Eli Mills, the manager of the Lockwood Foundation; they were meant to be sold on the black market to finance Henry Wu’s research and add to the Lockwood fortune. None are confirmed to have been sold, instead being released during the night to save them from a toxic gas leak.

In the modern day, Pteranodon has become an air traffic hazard as well as a transient threat to people living near large sources of water. It often mistakes aircraft, especially helicopters and smaller airplanes, for territorial threats and attacks them; this can frequently be lethal for the pterosaurs. Several crashes have been attributed to Pteranodon. It has also been known to view parasails or paragliders as threats, and may on rare occasion attack boats.

Pteranodon presents an economic threat as it can hinder air traffic, cause vehicular accidents, damage infrastructure, and fatally attack people. Since it prefers marine environments it is more of a pressing issue in coastal cities, but has been known to wander inland; it may follow rivers upstream and find cities and towns this way. At least three in mid-2018 were sighted near Lake Mead, entering the city of Las Vegas later that week and attacking at least one citizen at the Vegas Strip.

Safety

Humans are well within the size range of animals that Pteranodon feeds on, making this one of the few de-extinct animals that truly does routinely hunt humans as prey. As a flying animal, it attacks from above, picking up prey items and dropping them from sufficient height to kill them on impact. Then it can eat at its leisure, not having to worry about its prey struggling. A fast-moving predator that relies on surprise to catch its prey, Pteranodon is difficult to dodge, and fatal attacks on humans have occurred. Now that it has spread out from the Gulf of Fernandez to other parts of the world, keeping yourself safe from it is more important than ever.

It is not impossible to survive an attack. The first and easiest way to do this is to remain perpetually aware of whether or not Pteranodons have been sighted in your local area, which you should do as a community. The more eyes watching the sky, the more likely someone in your community will see these pterosaurs. Warn others if you see any, especially if your area has geographic features they prefer such as large lakes, ocean coastline, or mountains. Tall enough buildings may simulate their natural habitat of coastal cliffs, so bigger cities might attract them by accident. Bright lights draw their attention, and too many lights can make them confused and on edge. Attacks will be less common if you reduce nighttime light pollution. Deter their presence by keeping food sources secure; do not leave food waste (especially fish and other meat) out where they can reach it, and keep all pets indoors. Small children are especially at risk, and should be supervised at all times if you believe Pteranodons may be in the area. If you have outdoor water sources, especially with fish stocked in them, consider putting up covers to discourage unwanted use by pterosaurs. Under no circumstances should you feed a Pteranodon.

Due to its aggressive and territorial behavior, Pteranodon is likely to feel threatened by aircraft such as helicopters and may attack in defense of its territory or its flock.

Driving problematic Pteranodons away from an area is not simple, since they cannot be easily intimidated and harassing them will only make them retaliate violently. We also do not recommend trying to chase them away using aircraft, especially not helicopters; they frequently attack airborne vehicles as territorial threats rather than flee. If you happen to be in an aircraft during an attack, it will help to have previously reviewed the safety procedures for surviving a crash (if you are a pilot, you should already know what to do in case of a bird strike; now simply apply this principle to a bird which is actively trying to be involved in a strike). To date, there are no extremely reliable ways to make Pteranodons disperse other than to make your area inhospitable to them. Since this is not possible for some communities, the only way to deal with them may be to have experts capture and relocate the problem animals. If they keep returning, extermination may unfortunately be necessary.

Get inside as soon as you see a Pteranodon around, and if no buildings are available, get underneath tree cover or in other tight spaces where this large animal cannot maneuver. It is capable of walking on all fours, but most humans can outpace a Pteranodon on the ground, especially if undergrowth and other obstacles exist. Roofed vehicles also provide decent shelter from a Pteranodon; it is not strong enough to topple most large vehicles such as cars and trucks. However, it can shatter glass with its sharp beak, so stay away from windows. If you think it has spotted you and is likely to attack, lie flat on the vehicle floor where it is harder to reach. Do not attempt to drive away if you are under attack; loud noises agitate it, and moving objects are sure to draw its attention. Remain sheltered until it goes away. If you cannot get into shelter, use a natural-looking cover to hide yourself.

The amount of weight a Pteranodon can heft into the air depends on its subspecies, and is further compounded by its age and health, but assume it can lift you at least a short way off the ground. Most people are grabbed by the shoulders when attacked, since this is the easiest part of a standing human to grasp from above. Try to exercise some control over where you are dropped: if it deposits you on land, it will try to have you killed on impact. Grab hold of the pterosaur if you can, for example by locking an arm around its leg, and only allow yourself to be dropped if you can survive the fall. Observe your surroundings; if you can grab a tree branch or land on a soft surface, you may survive. Avoid letting go if there are hazards underneath you. Try to increase air resistance while falling. If you can, land on your rear to absorb as much of the impact as possible, not your front and absolutely not your head.

Alternatively to just dropping you to your death, a Pteranodon may try to drop you into any available source of water where it can feed more easily. If you are high enough in the air, landing in water can be just as dangerous as landing on a solid surface. To minimize the threat, try to enter the water smoothly, feet-first. An impact may still break your legs (especially if you are dropped in shallow water), but this is better than breaking your skull. The Pteranodon will pursue you into the water to feed. It is an excellent swimmer, but you are now in a position to defend yourself. Attack its vulnerable spots, such as its eyes or throat. You might be able to fight it off by attempting to drown it or threatening to damage its wing membranes. If you can put up enough of a struggle, it may have to back down and recollect itself for a second attempt, which will give you time to escape to shelter.

Defending yourself against a Pteranodon attack is more effective in groups, so that same community spirit you use to keep watch for pterosaur activity should be employed in case of conflict. An adult Pteranodon can lift a human into the air, but in groups you can try to weigh it down and fight it off with whatever weapons you have available. It is possible that a feeding frenzy might occur if the conditions are right, and this is far harder to defend against other than simply taking shelter. These happen when large groups of Pteranodons become aggressive and are presented with ample hunting grounds, so reducing such circumstances are key to preventing feeding frenzies. Immediately seek safety if you see large flocks moving about.

While no direct observations of Geosternbergia have been made (and it is unclear if any still exist), they are believed to be behaviorally similar to Pteranodon, so these same guidelines apply.

Behind the Scenes

Geosternbergia is known to have been cloned in the film universe due to physically appearing in an official InGen species guide, the mobile laboratory’s screensaver, and in concept art and models made for the film. It has been suggested that the intended use was for this species to be the male of P. longiceps, though this has never been confirmed and would be a gross scientific inaccuracy if true. Those in favor of considering these two genera as sexually dimorphic forms of the same species have cited these quotes in The Making of The Lost World: Jurassic Park, page 25.

“‘We had to design new paint schemes not only for the new dinosaurs, but for some of the already-designed dinosaurs from the last movie,’ Winston explained, ‘because now there were male dinosaurs, as well as females; and typically in nature the males of any species are far more brightly colored. We also wanted to make sure that the audience would be able to tell the males and females apart. It was a great of fun to run the gamut of color and come up with interesting designs. The colors on the females for the first film had been fairly subdued; but with male animals, there were may more possible colorations.'”
[…]
“As they had for the first film, the designers at Winston studio adhered closely to paleontological fact-or, in the absence of fact, paleontological theory- in their dinosaur renderings and sculptures, ‘We were very concerned that whatever we came up with made sense scientifically,’ Winston said, ‘so we drew from the science of paleontology. But since that science is limited in detail and has to rely on bone structure and hypothesis, we also drew from nature-what we see around us today.'”
[…]
“‘We looked at the color schemes and hide textures of living animals, especially those that live in a similar environment and have a similar lifestyle to the dinosaurs. The third thing we pulled from, of course, was imagination and instinct. No one really knows what these animals looked like; but our instincts told us if what we’d come up with made sense.'” (Duncan 25)

Despite Geosternbergia not featuring in any films, it does appear in the arcade game and PS1 video game based on The Lost World: Jurassic Park. In the former, it attacks the player near a laboratory facility, while in the latter it appears as a non-threatening background character.

Notable Individuals

Pteranodon longiceps “hippocratesi” – Isla Sorna – adults and offspring housed on Isla Sorna

Pteranodon longiceps “hippocratesi” – Isla Nublar – multiple animals possibly bred for Jurassic Park

Eggs A-F – first specimens of Pteranodon longiceps masranii bred in 2004

Jurassic World Pteranodons – predominantly-female flock bred for Jurassic World

Disambiguation Links

Pteranodon longiceps “hippocratesi” (JN)

Pteranodon longiceps (L/M)

Pteranodon longiceps “hippocratesi” (IDW-DitD)

Pteranodon longiceps “hippocratesi” (IDW-JPR)

Pteranodon (sp.) (CB-Topps)