Wednesday, June 29, 2022

The "flightless pterodactyl" that never was: Ornithopsis hulkei

Prior to and during the Victoria era, British fossil hunters came upon huge or peculiar bones of reptiles from Middle Jurassic to Early Cretaceous deposits in England that they interpreted as belonging to huge crocodile-like archosaurs, namely those from the Oolite Group of the Midlands, the Kimmeridge Clay of southern and eastern England, and the Wealden Supergroup of Sussex and the Isle of Wight. For instance, the type specimens of the basal eusauropod Cetiosaurus and the basal titanosauriform  Pelorosaurus were initially thought to have represented gigantic sea-going crocodiles, until more complete finds in the 1870s showed that they were actually dinosaurs and not crocodiles. However, most paleontology gurus overlook the fact that one Early Cretaceous titanosauriform sauropod from the UK, Ornithopsis, was misinterpreted by its describer as belonging not to a huge crocodile-like reptile, but instead as a flightless pterosaur!

Anterior view of the lectotype of Ornithopsis hulkei (NHMUK R28632) (from Owen 1875)

The story of the discovery and naming of Ornithopsis begins in the early 1850s, when an anterior dorsal vertebra was found in the Early Cretaceous (Barremian) Wessex Formation of the Isle of Wight along the English Channel coast of southern England and kept by Gideon Mantell (describer of Pelorosaurus) in his personal fossil collection, before being acquired by the British Museum in 1853 (a year after Mantell's death) and assigned the catalogue number BMNH R28632 (now NHMUK R28632). The dorsal vertebra, however, was not published in the scientific literature until Seeley (1870) erected the name Ornithopsis hulkei for NHMUK R28632 as well as NHMUK R2239, a dorsal vertebra found in the Early Cretaceous (late Valanginian) Tunbridge Wells Sand Formation of West Sussex in the 1820s by Gideon Mantell and misidentified by Owen (1854) as a quadrate of Iguanodon. He noted that NHMUK R28632 and NHMUK R2239 had cavities for air sacs seen in the bones of birds and pterosaurs, and thus surmised that Ornithopsis could be a missing link between pterosaurs and birds, but also possibly allied with dinosaurs, hence the name Ornithopsis meaning "bird face" in Greek.

Sir Richard Owen (1804-1892), who correctly determined that Ornithopsis was a sauropod dinosaur and not a flightless pterosaur

The identification of Ornithopsis as potentially being a flightless pterosaur would not hold water for very long, however. Owen (1875) agreed with Seeley (1870) that NHMUK R2239 was a dorsal vertebra rather than a quadrate, but he rejected Seeley's interpretation of Ornithopsis as a close relative of birds and pterosaurs and instead considered NHMUK R2239 and R28632 to be congeneric with his new sauropod genus Bothriospondylus from the Late Jurassic (Kimmerdgian) Kimmeridge Clay Formation of Wiltshire. The hypodigm for Ornithopsis hulkei was split into two species, with NHMUK R28632 receiving the new name Bothriospondylus magnus and NHMUK R2239 being made the holotype of the new species Bothriospondylus elongatus. For one thing, Richard Owen was a creationist and not a fan of Darwinist thought, so his lack of enthusiasm for Charles Darwin's theory of evolution endeared him to recognize that Ornithopsis belonged to a sauropod and not a pterosaur-like archosaur. Owen also must have been aware that because the Bothriospondylus elongatus holotype is from an older horizon than NHMUK R28632, the two vertebrae were most likely not conspecific. As a matter of fact, a year after he assigned the Ornithopsis hulkei material to Bothriospondylus, Owen (1876) changed his mind about B. magnus being congeneric with the Bothriospondylus type species (B. suffosus) and referred it to the sauropod genus Chondrosteosaurus from the same geologic horizon and location as NHMUK R28632. Forthwith, Ornithopsis would now be recognized not as a flightless pterodactyl, but instead as a member of Dinosauria --- although Richard Owen had been familiar with extinct and extant flightless birds, no one ever found a genuine pterosaur fossil with flightless abilities. 

In the 1860s and 1870s additional sauropod material was uncovered from the Wessex Formation of the Isle of Wight by Reverend William Fox and John Whitaker Hulke, including some vertebrae that would become the type specimens of the titanosauriforms Chondrosteosaurus magnus and Eucamerotus foxi. In his description of the new titanosauriform remains from the Isle of Wight, Hulke (1879) disputed Owen's opinion about the generic name Ornithopsis being misleading by pointing out that the syntypes of O. hulkei were lightly constructed regardless of the reclassification of Ornithopsis as a sauropod. He designated NHMUK R28632 as the lectotype of O. hulkei, making Bothriospondylus magnus a junior objective synonym of Ornithopsis, and the genera Chondrosteosaurus and Eucamerotus (the latter also described from the Isle of Wight) were synonymized with Ornithopsis. Although Hulke (1882, p. 375) treated the holotype of Bothriospondylus elongatus as the O. hulkei lectotype and referred NHMUK R28632 and the type material of Eucamerotus foxi the Wessex Formation of the Isle of Wight to his new species Ornithopsis eucamerotus, the earlier lectotype designation for O. hulkei by Hulke (1879) stands, as pointed out by Lydekker (1888). 

Although Ornithopsis had been wrongly interpreted as a flightless pterosaur when first named in 1870, it was nonetheless one of the first sauropod taxa to be described from the Isle of Wight and the fourth sauropod taxon described from the Early Cretaceous of Europe (after Pelorosaurus, Haestasaurus, and Oplosaurus). On a few occasions, Ornithopsis was synonymized with Pelorosaurus by von Huene (1909), Romer (1956), and Steel (1970) but Blows (1995) noted that the O. hulkei lectotype does not overlap with the holotype of Pelorosaurus conybeari and found Ornihopsis to be a distinct and valid genus of basal titanosauriform (followed by Upchurch et al. 2011).

References:

Blows, W.T., 1995. The Early Cretaceous brachiosaurid dinosaurs Ornithopsis and Eucamerotus from the Isle of Wight, England. Palaeontology 38 (1): 187–197.

Huene, F. v., 1909. Skizze zu einer Systematik und Stammesgeschichte der Dinosaurier. Centralblatt für Mineralogie, Geologie und Paläontologie 1909:12-22.

Hulke, J.W., 1879. Note (3rd) on (Eucamerotus, Hulke) Ornithopsis, H. G. Seeley, = Bothrospondylus magnus, Owen, = Chondrosteous magnus, Owen. Quarterly Journal of the Geological Society 35 (1–4): 752–762.

Hulke, J.W., 1882. Note on the Os Pubis and Ischium of Ornithopsis eucamerotusQuarterly Journal of the Geological Society 38 (1–4): 372–376.

Lydekker, R. (1888). Catalogue of the Fossil Reptilia and Amphibia in the British Museum (Natural History). Part I. Containing the Orders Ornithosauria, Crocodilia, Dinosauria, Squamata, Rhynchocephalia, and Proterosauria. British Museum (Natural History). Department of Geology. 309 pp.

Owen, R., 1854. Monograph on the Fossil Reptilia of the Wealden Formations. Part II. Dinosauria (Iguanodon).  Monographs of the Palaeontographical Society 8 (27): 1–54.

Owen, R., 1875. Monographs on the British Fossil Reptilia of the Mesozoic Formations. Part II. (Genera BothriospondylusCetiosaurusOmosaurus)Monographs of the Palaeontographical Society 29 (133): 15–93.

Owen, R., 1876. Monograph on the Fossil Reptilia of the Wealden and Purbeck Formations. Supplement No. VII. Crocodilia (Poikilopleuron) and Dinosauria? (Chondrosteosaurus). Monographs of the Palaeontographical Society 30 (136): 1–7.

Romer, A.S., 1956. Osteology of the Reptiles. University of Chicago Press: Chicago: IL 772 pp. 

Seeley, H.G., 1870. Ornithopsis, a gigantic animal of the Pterodacyle kind from the Wealden. Annals and Magazine of Natural History 5 (4): 305–318.

Steel, R., 1970. Part 14. Saurischia. Handbuch der Paläoherpetologie. Gustav Fischer Verlag: Stuttgart, 87 pp.

Upchurch, P., Mannion, P.D., and Barrett, P.M., 2011. Sauropod dinosaurs. pp. 476–525. In: Batten, D.J. (ed.). English Wealden Fossils. The Palaeontological Association.

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