Hypsilophodon

Hypsilophodon (/ˌhɪpsɪˈlɒfoʊdɒn/; meaning "high-crested tooth") is a neornithischian dinosaur genus from the Early Cretaceous period of England. It has traditionally been considered an early member of the group Ornithopoda, but recent research has put this into question.

The first remains of Hypsilophodon were found in 1849; the type species, Hypsilophodon foxii, was named in 1869. Abundant fossil discoveries were made on the Isle of Wight, giving a good impression of the build of the species. It was a small, agile bipedal herbivore, measuring 1.5–2 m (4.9–6.6 ft) long and weighing 20 kg (44 lb). It had a pointed head equipped with a sharp beak used to bite off plant material, much like modern-day parrots.

Some outdated studies have given rise to a number of misconceptions about Hypsilophodon, including that it was an armoured, arboreal animal, and that it could be found in areas outside of the Isle of Wight. However, research from the following years has shown these ideas to be incorrect.

The first specimen of Hypsilophodon was recovered in 1849, when workers dug up the soon-called Mantell-Bowerbank block from an outcrop of the Wessex Formation, part of the Wealden Group, about one hundred yards west of Cowleaze Chine, on the south-west coast of Isle of Wight. The larger half of the block (including seventeen vertebrae, parts of ribs and a coracoid, some of the pelvis, and assorted hindleg remains) was given to naturalist James Scott Bowerbank, and the remainder (including eleven caudal vertebrae and most of the rest of hindlegs) to Gideon Mantell. After his death, Mantell's portion was acquired by the British Museum; Bowerbank's was acquired later, bringing both halves back together. Richard Owen studied both halves and, in 1855, published a short article on the specimen, considering it to be a young Iguanodon rather than a new taxon. This was unquestioned until 1867, when Thomas Henry Huxley compared the vertebrae and metatarsals of the specimen more closely to those of known Iguanodon, and concluded that it must be a different animal entirely. The next year, he saw a fossil skull discovered by William Fox on exhibition at the Norwich Meeting of the British Associations. Fox, who had also found his fossil in the Cowleaze Chine area, along with several other specimens, considered it to belong to a juvenile Iguanodon, or to represent a new, small species in the genus. Huxley noticed its unique dentition and edentulous premaxilla, reminiscent of but obviously distinct from that of Iguanodon. He concluded this specimen, too, represented a distinct animal from Iguanodon. After losing track of the specimen for some months, Huxley requested Fox grant him permission to study the specimen to a more extensive degree. The request was granted, and Huxley began work on his new species.

Huxley first announced the new species in 1869 in a lecture; the text of this, published the same year, forms the official naming article, because it contained a sufficient description. The species was named Hypsilophodon foxii, and the holotype was the Fox skull (which today has the inventory number NHM NHMUK PV R 197). The next year, Huxley published the expanded full description article. Within the same block of stone as the Fox skull, the centrum of a dorsal vertebra had been preserved. This allowed comparison with the Mantell-Bowerbank block, confirming it to belong to the same species. Further supporting this, Fox had confirmed that the block was found in the same geological bed as his material. As such, Huxley described this specimen in addition to the skull and centrum. It would become the paratype; its two pieces are now registered in the Natural History Museum as specimen NHMUK PV OR 28707, NHMUK PV OR 39560–1. Later in the same year, Huxley classified Hypsilophodon taxonomically, considering it to belong to the family Iguanodontidae, related to Iguanodon and Hadrosaurus. There would later be a persistent misunderstanding as to the meaning of the generic name, which is often translated directly from the Greek as "high-ridged tooth". In reality Huxley, analogous to the way the name of the related genus Iguanodon ("iguana-tooth") had been formed, intended to name the animal after an extant herbivorous lizard, choosing for this role Hypsilophus and combining its name with Greek ὀδών, odon, "tooth". Hypsilophodon thus means "Hypsilophus-tooth". The Greek ὑψίλοφος, hypsilophos, means "high-crested" and refers to the back frill of the lizard, not to the teeth of Hypsilophodon itself, which are not high-ridged in any case. The specific name foxii honours Fox.

Immediate reception to Huxley's proposal of a new genus, distinct from Iguanodon, was mixed. The issue of distinctiveness was seen as important as more information on the form of Iguanodon was in demand, and the cranial anatomy in particular was of importance. If the Cowleaze Chine material was a distinct genus, it ceased being useful in this respect. Whilst some palaeontologists such as William Boyd Dawkins and Harry Seeley supported distinction, Fox rejected Huxley's proposal of a distinct genus and subsequently took back his skull and gave it to Owen to study. In attempt to clarify the situation, John Whitaker Hulke returned to the Hypsilophodon fossil bed on the Isle of Wight to obtain more material. He remarked that the whole of the skeleton seemed to be present, but that fragility limited excavation. He published a description of his new specimens in 1873, and based on his examination of the new teeth fossils echoed Fox's sentiments of doubt. Owen followed this with a study comparising at length the teeth of known Iguanodon and those from Fox's specimens. He agreed there were differences, but found them lacking in sufficient distinctiveness to be considered a distinct genus. As such, he renamed the species Iguanodon foxii. But Hulke had by then shifted his opinion, having obtained two new more informative specimens. Building on Huxley's comments on the Mantell-Bowerbank block, he gave focus to vertebral characters. As a result of his study, he concluded Hypsilophodon was a distinct genus related to Iguanodon. He published these findings in a supplementary note, also in 1874. Finally, in 1882 he published a full osteology of the species, considering it of great importance to properly document the taxon as such a wealth of specimens had been discovered and comparison with American dinosaurs was necessary. Fox had by this point died, and no further argument against generic distinctiveness had occurred in the intervening time.

Later, the number of specimens was increased by Reginald Walter Hooley.^{[citation needed][clarification needed]} In 1905, Baron Franz Nopcsa dedicated a study to Hypsilophodon, and in 1936 William Elgin Swinton did the same, on the occasion of the mounting of two restored skeletons in the British Museum of Natural History.^{[clarification needed]} Most known Hypsilophodon specimens were discovered between 1849 and 1921 and are in the possession of the Natural History Museum that acquired the collections of Mantell, Fox, Hulke and Hooley. These represent about twenty individual animals. Apart from the holotype and paratype, the most significant specimens are: NHM R5829, the skeleton of a large animal; NHMUK PV R 5830 and NHMUK PV R 196/196a, both skeletons of juvenile animals; and NHMUK PV R R2477, a block with a skull together with two separate vertebral columns. Although this was the largest find, new ones continue to be made.

Modern research of Hypsilophodon began with^{[citation needed]} the studies of Peter Malcolm Galton, starting with his thesis of 1967. He and James Jensen briefly described a left femur, AMNH 2585, in 1975, and in 1979 formally coined a second species, Hypsilophodon wielandi, for the specimen. The femur was diagnosed with two supposed minor differences from that of H. foxii. The specimen was found in 1900 in the Black Hills of South Dakota, United States, by George Reber Wieland, who the species was named after. Geologically, it comes from the Lakota Sandstone. This species was seen at the time as indicative of a probable late land bridge between North America and Europe, and of the dinosaur fauna of both continents being similar. Spanish Palaeontologist José Ignacio Ruiz-Omeñaca proposed that H. wielandi was not a species of Hypsilophodon but instead related to or synonymous with "Camptosaurus" valdensis from England, both species being dryosaurids. Galton refuted this in his contribution to a 2012 book, noting the femurs of the two species to be quite different, and that of H. wielandi to be unlike those of dryosaurs. He, as well as other studies before and after Ruiz-Omeñaca's proposal, considered H. wielandi a dubious basal ornithopod, with H. foxii the only species in the genus. Galton elaborated on the invalidity of the species in 2009, noting that the two supposed diagnostic characters were variable in both H. foxii and Orodromeus makelai, making the species dubious. He speculated that it may belong to Zephyrosaurus, from a similar time and place, as no femur was known from that taxon.