Megalosaurus

Megalosaurus is an extinct genus of large carnivorous theropod dinosaurs of the Middle Jurassic Epoch of southern England. Although fossils from other areas have been assigned to the genus, the only certain remains of Megalosaurus come from Oxfordshire and date to the late Middle Jurassic.
The earliest remains of Megalosaurus were described in the 17th century, and were initially interpreted as the remains of elephants or giants. Megalosaurus was named in 1824 by William Buckland, becoming the first genus of dinosaur to be validly named. The type species is M. bucklandii, named in 1827 by Gideon Mantell, after Buckland. In 1842, Megalosaurus was one of three genera on which Richard Owen based his Dinosauria, along with Iguanodon and Hylaeosaurus. On Owen's directions a model was made as one of the Crystal Palace Dinosaurs, which greatly increased the public interest for prehistoric reptiles. Over 50 other species would eventually be classified under the genus; at first, this was because so few types of dinosaur had been identified, but the practice continued even into the 20th century after many other dinosaurs had been discovered. Today it is understood that none of these additional species was directly related to M. bucklandii, which is the only true Megalosaurus species. Because a complete skeleton of it has never been found, much is still unclear about its build.
The first naturalists who investigated Megalosaurus mistook it for a gigantic quadrupedal lizard in length. In 1842, Owen concluded that it was no longer than. Modern scientists were able to obtain a more accurate picture, by comparing Megalosaurus with its direct relatives in the Megalosauridae. Megalosaurus was about long, weighing about. It was bipedal, walking on stout hindlimbs, its horizontal torso balanced by a horizontal tail. Its forelimbs were short, though very robust. Megalosaurus had a rather large head, equipped with long curved teeth. It was generally a robust and heavily muscled animal.
At the time Megalosaurus lived, Europe formed an island archipelago bounded by then narrow Atlantic Ocean and Tethys Ocean, with Megalosaurus inhabiting an island formed by the London–Brabant Massif, where it likely served as the apex predator of its ecosystem, coexisting with other dinosaurs like the large sauropod Cetiosaurus, stegosaurs, ankylosaurs, and heterodontosaurids.

Discovery and naming

Edward Lhuyd's tooth (specimen OU 1328)

In 1699, Edward Lhuyd described what he believed to have been a fish tooth, later believed to be part of a belemnite, that was illustrated alongside the holotype tooth of the sauropod "Rutellum impicatum" and another tooth, from a theropod, in 1699. Later studies found that the theropod tooth, known as specimen 1328 almost certainly was a tooth crown that belonged to an unknown species of Megalosaurus. OU 1328 has since been lost and it was not confidently assigned to Megalosaurus until the tooth was re-described by Delair & Sarjeant.
OU 1328 was collected near Caswell, near Witney, Oxfordshire sometime during the 17th century and became the third dinosaur fossil to ever be illustrated, after "Scrotum humanum" in 1677 and "Rutellum impicatum" in 1699.

"''Scrotum humanum''"

Megalosaurus may have been the first non avian dinosaur to be described in the scientific literature. The earliest possible fossil of the genus, from the Taynton Limestone Formation, was the lower part of a femur, discovered in the 17th century. Part of a bone was recovered from the Taynton Limestone Formation of Stonesfield limestone quarry, Oxfordshire in 1676. Sir Thomas Pennyson gave the fragment to naturalist Robert Plot, who published a description and illustration in his Natural History of Oxfordshire in 1676. It was the first illustration of a dinosaur bone published. Plot correctly identified the bone as the lower extremity of the thighbone or femur of a large animal and he recognized that it was too large to belong to any species known to be living in England. He therefore at first concluded it to be the thigh bone of a Roman war elephant and later that of a giant human, such as those mentioned in the Bible. The bone has since been lost, but the illustration is detailed enough that some have since identified it as that of Megalosaurus.
It has also been argued that this possible Megalosaurus bone was given the very first species name ever applied to an extinct dinosaur. Plot's engraving of the Cornwell bone was again used in a book by Richard Brookes in 1763. Brookes, in a caption, called it "Scrotum humanum", apparently comparing its appearance to a pair of "human testicles". However, it is possible that the attribution of this name stemmed from illustrator error, not Richard Brookes. In 1970, paleontologist Lambert Beverly Halstead pointed out that the similarity of Scrotum humanum to a modern species name, a so-called Linnaean "binomen" that has two parts, was not a coincidence. Linnaeus, the founder of modern taxonomy, had in the eighteenth century not merely devised a system for naming living creatures, but also for classifying geological objects. The book by Brookes was all about applying this latter system to curious stones found in England. According to Halstead, Brookes thus had deliberately used binomial nomenclature, and had in fact indicated the possible type specimen of a new biological genus. According to the rules of the International Code of Zoological Nomenclature, the name Scrotum humanum in principle had priority over Megalosaurus because it was published first. That Brookes understood that the stone did not actually represent a pair of petrified testicles was irrelevant. Merely the fact that the name had not been used in subsequent literature meant that it could be removed from competition for priority, because the ICZN states that if a name has never been considered valid after 1899, it can be made a nomen oblitum, an invalid "forgotten name".
In 1993, after the death of Halstead, his friend William A.S. Sarjeant submitted a petition to the International Commission on Zoological Nomenclature to formally suppress the name Scrotum in favour of Megalosaurus. He wrote that the supposed junior synonym Megalosaurus bucklandii should be made a conserved name to ensure its priority. However, the Executive Secretary of the ICZN at the time, Philip K. Tubbs, did not consider the petition to be admissible, concluding that the term "Scrotum humanum", published merely as a label for an illustration, did not constitute the valid creation of a new name, and stated that there was no evidence it was ever intended as such. Furthermore, the partial femur was too incomplete to definitely be referred to Megalosaurus and not a different, contemporary theropod.

Buckland's research

During the last part of the eighteenth century, the number of fossils in British collections quickly increased. According to a hypothesis published by science historian Robert Gunther in 1925, among them was a partial lower jaw of Megalosaurus. It was discovered about underground in a Stonesfield Slate mine during the early 1790s and was acquired in October 1797 by Christopher Pegge for 10s.6d. and added to the collection of the Anatomy School of Christ Church, Oxford.
In the early nineteenth century, more discoveries were made. In 1815, John Kidd reported the find of bones of giant tetrapods, again at the Stonesfield quarry. The layers there are currently considered part of the Taynton Limestone Formation, dating to the mid-Bathonian stage of the Jurassic Period. The bones were apparently acquired by William Buckland, Professor of Geology at the University of Oxford and fellow of Corpus Christi. Buckland also studied a lower jaw, according to Gunther the one bought by Pegge. Buckland did not know to what animal the bones belonged but, in 1818, after the Napoleonic Wars, the French comparative anatomist Georges Cuvier visited Buckland in Oxford and realized that they were those of a giant lizard-like creature. Buckland further studied the remains with Mary Morland, and his friend William Conybeare who in 1821 referred to them as the "Huge Lizard". In 1822 Buckland and Conybeare, in a joint article to be included in Cuvier's Ossemens, intended to provide scientific names for both gigantic lizard-like creatures known at the time: the remains found near Maastricht would be named Mosasaurus – then seen as a land-dwelling animal – while for the British lizard Conybeare had devised the name "Megalosaurus", from the Greek μέγας, megas, "large". That year a publication failed to occur, but the physician James Parkinson already in 1822 announced the name "Megalosaurus", illustrating one of the teeth and revealing the creature was 40 feet long and eight feet high. It is generally considered that the name in 1822 was still a nomen nudum. Buckland, urged on by an impatient Cuvier, continued to work on the subject during 1823. Mary provided drawings of the bones, that were to be the basis of illustrating lithographies. Finally, on 20 February 1824, during the same meeting of the Geological Society of London in which Conybeare described a very complete specimen of Plesiosaurus, Buckland formally announced Megalosaurus. The descriptions of the bones in the Transactions of the Geological Society, in 1824, constitute a valid publication of the name. Megalosaurus was the first non-avian dinosaur genus named; the first of which the remains had with certainty been scientifically described was Streptospondylus, in 1808 by Cuvier.
By 1824, the material available to Buckland consisted of specimen OUM J13505, a piece of a right lower jaw with a single erupted tooth; OUM J13577, a posterior dorsal vertebra; OUM J13579, an anterior caudal vertebra; OUM J13576, a sacrum of five sacral vertebrae; OUM J13585, a cervical rib; OUM J13580, a rib; OUM J29881, an ilium of the pelvis, OUM J13563, a piece of the pubic bone; OUM J13565, a part of the ischium; OUM J13561, a thigh bone and OUM J13572, the lower part of a second metatarsal. As he himself was aware, these did not all belong to the same individual; only the sacrum was articulated. Because they represented several individuals, the described fossils formed a syntype series. By modern standards, from these a single specimen has to be selected to serve as the type specimen on which the name is based. In 1990, Ralph Molnar chose the famous dentary, OUM J13505, as such a lectotype. Because he was unaccustomed to the deep dinosaurian pelvis, much taller than with typical reptiles, Buckland misidentified several bones, interpreting the pubic bone as a fibula and mistaking the ischium for a clavicle. Buckland identified the organism as being a giant animal belonging to the Sauria – the Lizards, at the time seen as including the crocodiles - and he placed it in the new genus Megalosaurus, repeating an estimate by Cuvier that the largest pieces he described, indicated an animal 12 meters long in life.