The largest pterosaur was Hatzegopteryx and close contender is Quetzalcoatlus northropi
Hatzegopteryx ("Hațeg basin wing") is a genus of azhdarchidpterosaur, known from incomplete remains found in Hunedoara County, Transylvania, Romania. The skull fragments, left humerus, and other fossilized remains indicate it was among the largest pterosaurs. The skeleton of Hatzegopteryx has been considered identical to the known remains of Quetzalcoatlus northropi. Q. northropi has not yet been properly described, and if it is not a nomen dubium, Hatzegopteryx is possibly its junior synonym.
The genus was named in 2002 by French paleontologistEric Buffetaut, and Romanian paleontologists Dan Grigorescu and Zoltan Csiki. It is known from only the type species, Hatzegopteryx thambema. The generic name is derived from the Hatzeg (or Hațeg) basin of Transylvania, the so-called Hațeg Island, where the bones were found, and from Greek pteryx, or 'wing'. The specific name thambema is derived from the Greek for 'monster', in reference to its huge size.
Hatzegopteryx hails from the upper part of the Middle Densus Ciula Formation of Valioara, northwestern Hateg Basin, Transylvania, western Romania, which has been dated to the late Maastrichtian stage of the late CretaceousPeriod, around 66 million years ago.
The holotype, FGGUB R 1083A, consists of the back part of the skull and the damaged proximal part of a left humerus.A 38.5 centimeter long middle section of a femur found nearby, FGGUB R1625, may also have belonged to Hatzegopteryx. FGGUB R1625 belonged to an individual with a five to six metres wingspan. The later reported fragments represent medium-sized animals.
In 2011, additional finds were reported: the front of a mandibula, the possibly third phalanx of the wing finger, an unfused scapulocoracoid, a scapula, cervical vertebrae and a piece of a humerus.
Hatzegopteryx apparently had a robust skull broadened in the rear, and a massive jaw. Its lower jaw featured a unique groove in its point of articulation, also seen in some other pterosaurs, that would have allowed the animal to achieve a very wide gape. Many of the fossilized bones of Hatzegopteryx resemble those of the closely related Quetzalcoatlus sp., though in Hatzegopteryx the skull was much more heavily built, and had a markedly different jaw articulation, similar to that seen in Pteranodon. Based on comparisons with other pterosaurs, Nyctosaurus and Anhanguera, Buffetaut and colleagues when initially describing the specimens estimated that the skull of Hatzegopteryx was probably almost three meters (ten feet) in length, which would have made it larger than that of the largest Quetzalcoatlus species and among the largest skulls of any known non-marine animals.
The skull of Hatzegopteryx was also unique in its heavy, robust construction. Most pterosaur skulls are made up of very lightweight plates and struts. In Hatzegopteryx, the skull bones are stout and robust, with large-ridged muscle insertion areas. In their 2002 description, Buffetaut and colleagues suggested that in order to fly, the skull weight of this pterosaur must have been reduced in some unconventional way (while they allowed that it could have been flightless, they found this unlikely due to the similarity of its wing bones to flying pterosaurs). The authors theorized that the necessary weight reduction was accomplished by the internal structure of the skull bones, which were full of small pits and hollows (alveoli) up to 10 mm long, separated by a matrix of incredibly thin bony struts (trabeculae), a feature also found in some parts of Hatzegopteryx wing bones. The authors pointed out that this unusual construction, which differed significantly from the irregular internal structure of other pterosaur skulls, resembles the structure of expanded polystyrene, the substance used to make Styrofoam. They noted that this would allow a sturdy, stress-resistant construction while remaining lightweight, and would have allowed the huge-headed animal to fly.
The authors estimated the size of Hatzegopteryx by comparing the humerus fragment, 236 mm (9.3 in) long, with that of Quetzalcoatlus, of which specimen TMM 41450-3 has a 544 mm (1 ft 9.4 in) long humerus. Observing that the Hatzegopteryx fragment presented less than half of the original bone, they established that it could possibly have been "slightly longer" than that of Quetzalcoatlus. They noted that the wing span of the latter had in 1981 been estimated at 11 to 12 metres (36–39 ft), while earlier estimates had strongly exceeded this at 15 to 20 metres (49-65.6 ft). From this they concluded that an estimate of a 12-metre (39 ft) wing span for Hatzegopteryx was conservative "if its humerus was indeed somewhat longer than that of Q. northropi".In 2003 they moderated the estimates to a close to 12 metres (39 ft) wing span and an over 2.5 m (8 ft 2 in) skull length.In 2010 Mark Witton e.a. stated that any appearance that the Hatzegopteryx humerus was bigger than TMM 41450-3 had been caused by a distortion of the bone after deposition and that the species thus likely had no larger wingspan than Quetzalcoatlus, today generally estimated at 10 to 11 metres (33–36 ft).
Another close contender is Quetzalcoatlus northropi, at 127 kg (280 lb) and with a wingspan of 12 m (39 ft).
When it was first discovered, scientists estimated that the largest Quetzalcoatlus fossils came from an individual with a wingspan as large as 15.9 meters (52 feet), choosing the middle of three extrapolations from the proportions of other pterosaurs that gave an estimate of 11, 15.5 and 21 meters respectively (36 feet, 50.85 feet, 68.9 feet). In 1981, further study lowered these estimates to 11–12 meters (36–39 ft).More recent estimates based on greater knowledge of azhdarchid proportions place its wingspan at 10–11 meters (33–36 ft).
Mass estimates for giant azhdarchids are extremely problematic because no existing species share a similar size or body plan, and in consequence published results vary widely.While some studies have historically found extremely low weight estimates for Quetzalcoatlus, as low as 70 kilograms (150 lb) for a 10-meter (32-foot-10-inch) individual, a majority of estimates published since the 2000s have been higher, around 200–250 kilograms (440–550 lb).
Skull material (from smaller specimens, possibly a related species) shows that Quetzalcoatlus had a very sharp and pointed beak. That is contrary to some earlier reconstructions that showed a blunter snout, based on the inadvertent inclusion of jaw material from another pterosaur species, possibly a tapejarid or a form related to Tupuxuara. A skull crest was also present but its exact form and size are still unknown.
The first Quetzalcoatlus fossils were discovered in Texas, United States, from the MaastrichtianJavelina Formation at Big Bend National Park (dated to around 68 million years ago) in 1971 by a geology graduate student from the University of Texas at Austin's Jackson School of Geosciences, Douglas A. Lawson. The specimen consisted of a partial wing (in pterosaurs composed of the forearms and elongated fourth finger), from an individual later estimated at over 10 m (33 ft) in wingspan. Lawson discovered a second site of the same age, about forty kilometers from the first, where between 1972 and 1974 he and Professor Wann Langston Jr. of the Texas Memorial Museum unearthed three fragmentary skeletons of much smaller individuals. Lawson in 1975 announced the find in an article in Science.That same year, in a subsequent letter to the same journal, he made the original large specimen, TMM 41450-3, the holotype of a new genus and species, Quetzalcoatlus northropi. The genus name refers to the Aztec feathered serpent god Quetzalcoatl. The specific name honors John Knudsen Northrop, the founder of Northrop, who was interested in large tailless flying wing aircraft designs resembling Quetzalcoatlus.At first it was assumed that the smaller specimens were juvenile or subadult forms of the larger type. Later, when more remains were found, it was realized they could have been a separate species. This possible second species from Texas was provisionally referred to as a Quetzalcoatlus sp. by Alexander Kellner and Langston in 1996, indicating that its status was too uncertain to give it a full new species name.The smaller specimens are more complete than the Q. northropi holotype, and include four partial skulls, though they are much less massive, with an estimated wingspan of 5.5 meters (18 feet).
The holotype specimen of Q. northropi has yet to be properly described and diagnosed. Where the known remains overlap, it has been considered by Mark Witton and colleagues (2010) to be indistinguishable from its Romanian contemporary Hatzegopteryx. If Q. northropi is complete enough to be distinguished from other pterosaurs (i.e., if it is not a nomen dubium), Hatzegopteryx may represent the same animal. It is likely that huge pterosaurs such as Q. northropi would have had very large, transcontinental ranges, making its presence in both North America and Europe unsurprising.Mark Witton et al. argued that the skull material of Hatzegopteryx and Q. sp. differ enough that they cannot be regarded as the same animal, making it likely that Q. sp., if not identical to Quetzalcoatlus northropi, represents a distinct genus.
An azhdarchid neck vertebra, discovered in 2002 from the Maastrichtian age Hell Creek Formation, may also belong to Quetzalcoatlus. The specimen (BMR P2002.2) was recovered accidentally when it was included in a field jacket prepared to transport part of a tyrannosaur specimen. Despite this association with the remains of a large carnivorous dinosaur, the vertebra shows no evidence that it was chewed on by the dinosaur. The bone came from an individual azhdarchid pterosaur estimated to have had a wingspan of 5–5.5 m (16–18 ft).
Quetzalcoatlus was abundant in Texas during the Lancian in a fauna dominated by Alamosaurus.The Alamosaurus-Quetzalcoatlus association probably represents semi-arid inland plains.Quetzalcoatlus had precursors in North America and its apparent rise to widespreadness may represent the expansion of its preferred habitat rather than an immigration event, as some experts have suggested.
There have been a number of different ideas proposed about the lifestyle of Quetzalcoatlus. Because the area of the fossil site was four hundred kilometers removed from the coastline and there were no indications of large rivers or deep lakes nearby at the end of the Cretaceous, Lawson in 1975 rejected a fish-eating lifestyle, instead suggesting that Quetzalcoatlus scavenged like the Marabou Stork, but then on the carcasses of titanosaursauropods such as Alamosaurus. Lawson had found the remains of the giant pterosaur while searching for the bones of this dinosaur, which formed an important part of its ecosystem.
In 1996, Thomas Lehman and Langston rejected the scavenging hypothesis, pointing out that the lower jaw bent so strongly downwards that even when it closed completely a gap of over five centimeters remained between it and the upper jaw, very different from the hooked beaks of specialized scavenging birds. They suggested that with its long neck vertebrae and long toothless jaws Quetzalcoatlus fed like modern-day skimmers, catching fish during flight while cleaving the waves with its beak.While this skim-feeding view became widely accepted, it was not subjected to scientific research until 2007 when a study showed that for such large pterosaurs it was not a viable method because the energy costs would be too high due to excessive drag.In 2008 pterosaur workers Mark Paul Witton and Darren Naish published an examination of possible feeding habits and ecology of azhdarchids. Witton and Naish noted that most azhdarchid remains are found in inland deposits far from seas or other large bodies of water required for skimming. Additionally, the beak, jaw, and neck anatomy are unlike those of any known skimming animal. Rather, they concluded that azhdarchids were more likely terrestrial stalkers, similar to modern storks, and probably hunted small vertebrates on land or in small streams. Though Quetzalcoatlus, like other pterosaurs, was a quadruped when on the ground, Quetzalcoatlus and other azhdarchids have fore and hind limb proportions more similar to modern running ungulate mammals than to their smaller cousins, implying that they were uniquely suited to a terrestrial lifestyle.
The nature of flight in Quetzalcoatlus and other giant azhdarchids was poorly understood until serious biomechanical studies were conducted in the 21st century. One early (1984) experiment by Paul MacCready used practical aerodynamics to test the flight of Quetzalcoatlus. MacCready constructed a model flying machine or ornithopter with a simple computer functioning as an autopilot. The model successfully flew with a combination of soaring and wing flapping;however, the model was half scale based on a then-current weight estimate of around 80 kg, far lower than more modern estimates of over 200 kg.The method of flight in these pterosaurs depends largely on weight, which has been controversial, and widely differing masses have been favored by different scientists. Some researchers have suggested that these animals employed slow, soaring flight, while others have concluded that their flight was fast and dynamic.In 2010, Donald Henderson argued that the mass of Q. northropi had been underestimated, even the highest estimates, and that it was too massive to have achieved powered flight. He estimated it in his 2010 paper as 540 kg. Henderson argued that it may have been flightless.
However, most other flight capability estimates have disagreed with Henderson's research, suggesting instead an animal superbly adapted to long-range, extended flight. In 2010, Mike Habib, a professor of biomechanics at Chatham University, and Mark Witton, a British paleontologist, undertook a further investigation into the claims of flightlessness in large pterosaurs.After factoring wingspan, body weight, and aerodynamics, a sophisticated computer program led the two researchers to conclude that Q. northropi was capable of flight "up to 80 miles an hour for 7 to 10 days at altitudes of 15,000 feet".Mike Habib further suggested a maximum flight range of 8,000 to 12,000 miles for Q. northropi.Henderson's work was further criticized by Habib, who pointed out that although Henderson used excellent mass estimations, they were based on outdated pterosaur models, and that anatomical study of Q. northropi and other large pterosaur forelimbs show a higher degree of robustness than would be expected if they were purely quadrupedal.Habib believes that large pterosaurs most likely utilized a short burst of powered flight in order to then transition to thermal soaring.