Pteranodon longiceps model.
| Pteranodon Fossil range: Mid-Late Cretaceous |
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Pteranodon (pronounced /tɨˈrænədɒn/; from Greek πτερ- "wing" and αν-οδων "toothless"), from the Late Cretaceous (Coniacian-Campanian, 89.3-70.6 million years ago) of North America (Kansas, Alabama, Nebraska, Wyoming, and South Dakota), was one of the largest pterosaur genera, with a wingspan of up to 9 m (30 ft).
Description
Unlike earlier pterosaurs such as Rhamphorhynchus and Pterodactylus, Pteranodon had toothless beaks, similar to those of modern birds.
Pteranodon fossils were found first (1870) in the Late Cretaceous chalk Smoky Hill Chalk of western Kansas. These chalk beds were deposited at the bottom of what was once a large epicontinental sea over what is now midsection of the North American continent. While the first Pteranodon wing bones were collected by Marsh (1870) and Cope (1871), the first Pteranodon skull was found on May 2, 1876, along the Smoky Hill River in Wallace County (now Logan County), Kansas, USA, by S. W. Williston, a fossil collector working for Othniel C. Marsh.
The Smoky Hill Chalk is the upper part of the Niobrara Formation and is famous for the fossils collected there since 1869. Other fossils found in this formation include those of sea turtles, mosasaurs, and early birds with teeth.[1]
Pteranodon were reptiles, but not dinosaurs. By definition, all dinosaurs were diapsid reptiles with an upright stance, and consist of the group containing saurischians and ornithischians. While the advanced pterodactyloid pterosaurs (like Pteranodon) had a semi-upright stance, it evolved independently of the upright stance in dinosaurs, and pterosaurs lacked the distinctive adaptations in the hip associated with the dinosaurian posture. However, dinosaurs and pterosaurs may have been closely related, and most paleontologists place them together in the group Ornithodira, or "bird necks".
Discovery and species
A number of species of Pteranodon have been named since 1870, with the most well-supported of them being the type species, P. longiceps. This species was named by Othniel Charles Marsh in 1876 from a fairly complete specimen collected by S.W. Williston. This individual had a wingspan of 7 m (23 ft).[2][3] Other valid species include the possibly larger P. sternbergi, with a wingspan of 9 m (30 ft),[2]. P. occidentalis, P. velox, P. umbrosus, P. harpyia, and P. comptus are considered to be nomen dubium by Bennett (1994) and others. All are probably synonymous with the more well-known species.
Pteranodon sternbergi is the first known species of Pteranodon with an upright-crest. The lower jaw of P. sternbergi was 1.25 meters (4 ft) long.[4] It was collected by G.F. Sternberg in 1952 and described by Harksen in 1966, and is currently believed to be ancestral of the later species Pteranodon longiceps.[5] See also paper by H.W. Miller (1971) [6]
Although the first description of Pteranodon (initially called Pterodactylus Oweni) by Marsh erroneously noted it had teeth (1982, p. 244), he subsequently described Pteranodon as "distinguished from all previously known genera of the order {pterosauria} by the entire absence of teeth." This meant that any toothless pterosaur jaw fragment, wherever it was found in the world, tended to be attributed to Pteranodon. Hence there came to be a plethora of species and a great deal of confusion. The name became a wastebasket taxon, rather like the dinosaur Megalosaurus, to label any pterosaur remains that could not be distinguished other than by the absence of teeth. Notable authors who have discussed the various aspects of Pteranodon include Bennett, Padian, Unwin, Kellner, and Wellnhofer. One species, P. orogensis is not actually a pteranodontid and has been renamed Bennettazhia oregonensis. Likewise, P. orientalis has been renamed Bogolubovia orientale (Nessov & Yarkov, 1989) and transferred to the Azhdarchidae.
Paleobiology
The diet of Pteranodon is known to have included fish; fossilized fish bones have been found in the stomach area of one Pteranodon, and a fossilized fish bolus has been found between the rami of another Pteranodon. Pteranodon's wing shape suggests that it would have flown rather like a modern-day albatross. This is a suggestion based on the fact that the Pteranodon had a high aspect ratio (wingspan to chord length) similar to that of the albatross — 9:1 for Pteranodon, compared to 8:1 for an albatross. Albatrosses spend long stretches of time at sea fishing, and utilize a flight pattern called "dynamic soaring" which exploits the vertical gradient of wind speed near the ocean surface to travel long distances without flapping,[7] and without the aid of thermals (which do not occur over the open ocean the same way they do over land). However, other scientists have suggested that Pteranodon could flap their wings and fly with power. These two flight styles need not have been mutually exclusive in Pteranodon, or in pterosaurs in general. Recent wind tunnel tests on model pterosaur wings with the pteroid bone in an extended antero-ventral orientation supporting a large, highly cambered propatagium show that such a configuration enables the wing to develop up to 30% more lift, even at very high angles of attack. This anatomical feature, based on the pteroid bone - the bone unique to the pterosaur clade - may have enabled pterosaurs to be active, powered flyers in spite of the lack of other features associated with strong fliers. For example, pterosaurs usually had a small (relative to modern birds) sternum keel as an anchor point for the pectoralis muscle.
Pteranodon was notable for its skull crest. These may have been used as mating displays, or it might have acted as a rudder, or perhaps both. It is also believed that the crest kept it stable when flying.[4] It has been suggested that males of the species bore larger crests, but with fossil animals it is often difficult to tell whether differences in crest shape reflect different sexes or different species.
Consensus regarding the terrestrial locomotion of Pteranodon (whether it was bipedal or quadrupedal) has historically been the subject of debate. Today, most pterosaur researchers agree that pterosaurs were quadrupedal, thanks largely to the discovery of several pterosaur trackways; however, some new research suggest it was bipedal [1]. The possibility of swimming has been discussed briefly in two papers (Bennett 2001 and Bramwell & Whitfield),[8] and has been studied in detail at Michigan State University (Smith , 2007) through the use of quantitative morphometrics and an extant phylogenetic bracket (a morphologically comparative technique invented by Larry Witmer).
Pteranodon sternbergi skeleton at the Royal Ontario Museum, Toronto. (*)
In popular culture
See also: Biological issues in Jurassic Park
This Painting of Pteranodon was made to illustrate one card of a set of 30 collector cards from "Tiere der Urwelt" (Animals of the Prehistoric World). Painting by Heinrich Harder, 1916.
In colloquial language, Pteranodon is often erroneously called a "pterodactyl." However, "pterodactyl" is not actually the name of a specific species; rather, it is a term for all short-tailed pterosaurs (the suborder Pterodactyloidea), which includes Pteranodon, Pterodactylus, and Quetzalcoatlus. This misuse is most likely due to Pteranodon's high profile in popular culture as the quintessential pterodactyloid. Also, Pteranodon is not the name of a specific species, but of the genus as a whole.
Pteranodon was seen briefly at the end of the 1997 film The Lost World: Jurassic Park, and also appeared in its sequel, Jurassic Park III. The depiction in Jurassic Park III featured many inaccuracies, including toothed jaws, exaggerated strength and, presumably, aggression. (Pteranodon is thought to have eaten fish, and was incapable of grasping with its feet). In the two novels that inspired the first two films, some of these inaccuracies were attributed to the genetic engineering process used to create the animals. More scientifically accurate Pteranodon appeared in the television programs Chased by Dinosaurs, Sea Monsters, and Primeval. A trained Pteranodon named Turu appears in the Jonny Quest series. Pteranodon appear in two Ray Harryhausen movies, The Valley of Gwangi and One Million Years B.C., as well as the pre-Harryhausen classic King Kong.
In the British television series Torchwood, the Torchwood team keep a semi-tame Pteranodon named Myfanwy at the Torchwood headquarters in Cardiff, Wales. In one episode, it is used to destroy a partially constructed Cyberman.
Petrie from The Land Before Time (series) is a Pteranodon.
In the 1981 comedy Caveman, one of the animals the cavemen encounter is a large Pteranodon.
The latest and most accurate portrayal of a Pteranodon (actually a whole flock of them) was in the National Geographic IMAX movie, Sea Monsters, released in October 2007.
References
1. ^ Bennett SC. (2000) Inferring stratigraphic position of fossil vertebrates from the Niobrara Chalk of western Kansas. Current Research in Earth Sciences, Kansas Geological Survey Bulletin 244, Part 1, 26 p
2. ^ a b Wellnhofer, Peter (1996) [1991]. The Illustrated Encyclopedia of Pterosaurs. New York: Barnes and Noble Books. pp. 139. ISBN 0-7607-0154-7.
3. ^ Identified as P. ingens in Wellnhofer, 1991.
4. ^ a b Beyond the Dinosaurs! by Howard Zimmerman, ISBN 0-689-84113-2.
5. ^ www.dinodata.org Retrieved on May 7th, 2008.
6. ^ Transactions of the Kansas Academy of Science (1903-), Vol. 74, No. 1 (Spring, 1971), pp. 1-19 (article consists of 19 pages)
7. ^ Padian K. (1983)A functional analysis of flying and walking in pterosaurs. Paleobiology 9(3):218-239
8. ^ Bramwell CD & Whitfield GR (1974) "Biomechanics of Pteranodon", Phil. Trans. Roy. Soc. B. 267
Smith, Amy C. 2007. Pteranodont claw morphology and its implications for aquatic locomotion. Master's Thesis, Michigan State University.
SUGGESTED ADDITIONAL REFERENCES:
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Bennett, S. C. 1987. New evidence on the tail of the pterosaur Pteranodon (Archosauria: Pterosauria). pp. 18-23 In Currie, P. J. and E. H. Koster (eds.), Fourth Symposium on Mesozoic Terrestrial Ecosystems, Short Papers. Occasional Papers of the Tyrrell Museum of Paleontology, #3.
Bennett, S. C. 1990. Inferring stratigraphic position of fossil vertebrates from the Niobrara Chalk of western Kansas. pp. 43-72, In Bennett, S. C. (ed.), Niobrara Chalk Excursion Guidebook, The University of Kansas Museum of Natural History and the Kansas Geological Survey.
Bennett, S. C. 1992. Sexual dimorphism of Pteranodon and other pterosaurs, with comments on cranial crests. Journal of Vertebrate Paleontology 12 p. 422-434.
Bennett, S. C. 1994. Taxonomy and systematics of the Late Cretaceous pterosaur Pteranodon (Pterosauria, Pterodactyloida). Occasional Papers of the Natural History Museum, University of Kansas. 169:1-70.
Bennett, S. C. 2000. New information on the skeletons of Nyctosaurus. Journal of Vertebrate Paleontology 20(Supplement to Number 3): 29A. (Abstract)
Bennett, S. C. 2001. The osteology and functional morphology of the Late Cretaceous pterosaur Pteranodon. Part I. General description of osteology. Palaeontographica, Abteilung A, 260:1-112.
Bennett, S. C. 2001.The osteology and functional morphology of the Late Cretaceous pterosaur Pteranodon. Part II. Functional morphology. Palaeontographica, Abteilung A, 260:113-153.
Bennett, S. C. 2003. New crested specimens of the Late Cretaceous pterosaur Nyctosaurus. Paläontologische Zeitschrift, 77:61-75.
Bennett, S. C. 2003. A survey of pathologies of large pterydactyloid pterosaurs. Palaeontology 46(1):185-198.
Bennett, S. C. 2007. Articulation and function of the pteroid bone of pterosaurs. Journal of Vertebrate Paleontology, 27:881-891.
Betts, C. W. 1871. The Yale College Expedition of 1870. Harper’s New Monthly Magazine, 43(257):663-671. (Issue of October, 1871)
Bonner, O. W. 1964. An osteological study of Nyctosaurus and Trinacromerum with a description of a new species of Nyctosaurus. Unpub. Masters Thesis, Fort Hays State University, 63 pages.
Brower, J. C. 1983. The aerodynamics of Pteranodon and Nyctosaurus, two large pterosaurs from the Upper Cretaceous of Kansas. Journal of Vertebrate Paleontology 3(2):84-124.
Cope, E. D. 1872. On the geology and paleontology of the Cretaceous strata of Kansas. Annual Report of the U. S. Geological Survey of the Territories 5:318-349 (Report for 1871).
Cope, E. D. 1872. On two new Ornithosaurians from Kansas. Proceedings of the American Philosophical Society 12(88):420-422.
Cope, E. D. 1874. Review of the Vertebrata of the Cretaceous period found west of the Mississippi River. U. S. Geological Survey of the Territories Bulletin 1(2):3-48.
Cope, E. D. 1875. The Vertebrata of the Cretaceous formations of the West. Report, U. S. Geological Survey of the Territories (Hayden). 2:302 p, 57 pls.
Eaton, G. F. 1903. The characters of Pteranodon. American Journal of Science, ser. 4, 16(91):82-86, pl. 6-7.
Eaton, G. F. 1904. The characters of Pteranodon (second paper). American Journal of Science, ser. 4, 17(100):318-320, pl. 19-20.
Eaton, G. F. 1908. The skull of Pteranodon. Science (n. s.) XXVII 254-255.
Eaton, G. F. 1910. Osteology of Pteranodon. Memoirs of the Connecticut Academy of Arts and Sciences, 2:1-38, pls. i-xxxi.
Everhart, M. J. 1999. An early occurrence of Pteranodon sternbergi from the Smoky Hill Member (Late Cretaceous) of the Niobrara Chalk in western Kansas. Kansas Academy of Science, Transactions 18(Abstracts):27.
Everhart, M. J. 2005. Oceans of Kansas - A Natural History of the Western Interior Sea. Indiana University Press, 320 pp.
Harksen, J. C. 1966. Pteranodon sternbergi, a new fossil pterodactyl from the Niobrara Cretaceous of Kansas. Proceedings South Dakota Academy of Science 45:74-77.
Lane, H. H. 1946. A survey of the fossil vertebrates of Kansas, Part III, The Reptiles, Kansas Academy Science, Transactions 49(3):289-332, figs. 1-7.
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Marsh, O. C. 1871. Notice of some new fossil reptiles from the Cretaceous and Tertiary formations. American Journal of Science, Series 3, 1(6):447-459.
Marsh, O. C. 1871. Note on a new and gigantic species of Pterodactyle. American Journal of Science, Series 3, 1(6):472.
Marsh, O. C. 1872. Discovery of additional remains of Pterosauria, with descriptions of two new species. American Journal of Science, Series 3, 3(16) :241-248.
Marsh, O. C. 1876. Notice of a new sub-order of Pterosauria. American Journal of Science, Series 3, 11(65):507-509.
Marsh, O. C., 1876. Principal characters of American pterodactyls. American Journal of Science, Series 3, 12(72):479-480.
Marsh, O. C. 1881. Note on American pterodactyls. American Journal of Science, Series 3, 21(124):342-343.
Marsh, O. C. 1882. The wings of Pterodactyles. American Journal of Science, Series 3, 23(136):251-256, pl. III.
Marsh, O. C. 1884. Principal characters of American Cretaceous pterodactyls. Part I. The skull of Pteranodon. American Journal of Science, Series 3, 27(161):422-426, pl. 15.
Miller, H. W. 1971. The taxonomy of the Pteranodon species from Kansas. Kansas Academy of Science, Transactions 74(1):1-19.
Miller, H. W. 1971. A skull of Pteranodon (Longicepia) longiceps Marsh associated with wing and body parts. Kansas Academy of Science, Transactions 74(10):20-33.
Padian, K. 1983. A functional analysis of flying and walking in pterosaurs. Paleobiology 9(3):218-239.
Russell, D. A. 1988. A check list of North American marine cretaceous vertebrates Including fresh water fishes, Occasional Paper of the Tyrrell Museum of Palaeontology, (4):57.
Schultze, H.-P., L. Hunt, J. Chorn and A. M. Neuner, 1985. Type and figured specimens of fossil vertebrates in the collection of the University of Kansas Museum of Natural History, Part II. Fossil Amphibians and Reptiles. Miscellaneous Publications of the University of Kansas Museum of Natural History 77:66 pp.
Seeley, Harry G. 1871. Additional evidence of the structure of the head in ornithosaurs from the Cambridge Upper Greensand; being a supplement to "The Ornithosauria." The Annals and Magazine of Natural History, Series 4, 7:20-36, pls. 2-3. (Discovery of toothless pterosaurs in England)
Shor, E. N. 1971. Fossils and flies; The life of a compleat scientist - Samuel Wendell Williston, 1851-1918, University of Oklahoma Press, 285 pp.
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Sternberg, G. F. and M. V. Walker. 1958. Observation of articulated limb bones of a recently discovered Pteranodon in the Niobrara Cretaceous of Kansas. Kansas Academy of Science, Transactions, 61(1):81-85.
Stewart, J. D. 1990. Niobrara Formation vertebrate stratigraphy. pp. 19-30 in Bennett, S. C. (ed.), Niobrara Chalk Excursion Guidebook, The University of Kansas Museum of Natural History and the Kansas Geological Survey.
Wang, X. and Z. Zhou. 2004. Pterosaur embryo from the Early Cretaceous. Nature 429:621.
Wellnhofer, P. 1991. The illustrated encyclopedia of pterosaurs. Crescent Books, New York, 192 pp.
Williston, S. W. 1891. The skull and hind extremity of Pteranodon. American Naturalist 25(300):1124-1126.
Williston, S. W. 1892. Kansas pterodactyls. Part I. Kansas University Quarterly 1:1-13, pl. i.
Williston, S. W. 1893. Kansas pterodactyls. Part II. Kansas University Quarterly 2:79-81, with 1 fig.
Williston, S. W. 1895. Note on the mandible of Ornithostoma. Kansas University Quarterly 4:61.
Williston, S. W. 1896. On the skull of Ornithostoma. Kansas University Quarterly 4(4):195-197, with pl. i.
Williston, S. W. 1897. Restoration of Ornithostoma (Pteranodon). Kansas University Quarterly 6:35-51, with pl. ii.
Williston, S. W. 1902. On the skeleton of Nyctodactylus, with restoration. American Journal of Anatomy. 1:297-305.
Williston, S. W. 1902. On the skull of Nyctodactylus, an Upper Cretaceous pterodactyl. Journal of Geology, 10:520-531, 2 pls.
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Williston, S. W. 1903. On the osteology of Nyctosaurus (Nyctodactylus), with notes on American pterosaurs. Field Mus. Publ. (Geological Ser.) 2(3):125-163, 2 figs., pls. XL-XLIV.
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