Wednesday, November 30, 2016

Artist’s impression of how Victorian paleontologists thought Megalosaurus bucklandii looked (right), compared with how we now understand it to have looked (left). Image credit: Mark Garlick / University of Warwick.

Megalosaurus (meaning "Great Lizard") is a genus of large meat-eating theropod dinosaurs of the Middle Jurassic period (Bathonian stage, 166 million years ago) 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.

Megalosaurus was, in 1824, the first genus of non-avian dinosaur to be validly named. The type species is Megalosaurus bucklandii, named in 1827. In 1842, Megalosaurus was one of three genera on which Richard Owen based his Dinosauria. On Owen's directions a model was made as one of the Crystal Palace Dinosaurs, which greatly increased the public interest for prehistoric reptiles. Subsequently, over fifty other species would be classified under the genus, originally because dinosaurs were not well known, but even during the 20th century after many dinosaurs had been discovered. Today it is understood these additional species were not 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.

Size comparison (human in blue, lectotype in pink, largest specimen in red)

Buckland had not provided a specific name, as was not uncommon in the early nineteenth century, when the genus was still seen as the more essential concept. In 1826, Ferdinand von Ritgen gave this dinosaur a complete binomial, Megalosaurus conybeari, which however was not much used by later authors and is now considered a nomen oblitum. A year later, in 1827, Gideon Mantell included Megalosaurus in his geological survey of southeastern England, and assigned the species its current valid binomial name, Megalosaurus bucklandii. Until recently, the form Megalosaurus bucklandi was often used, a variant first published in 1832 by Christian Erich Hermann von Meyer – and sometimes erroneously ascribed to von Ritgen – but the more original M. bucklandii has priority.

During the last part of the eighteenth century, the number of fossils in British collections quickly increased. According to an hypothesis published by science historian Robert William Theodor Gunther in 1925, among them was a partial lower jaw of Megalosaurus, acquired in October 1797 by Christopher Pegge for 10s.6d. and added to the collection of the Anatomy School of Christ Church college.

Lithography from William Buckland's "Notice on the Megalosaurus or great Fossil Lizard of Stonesfield", 1824. Caption reads "anterior extremity of the right lower jaw of the Megalosaurus from Stonesfield near 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 dean of Christ Church. 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 realised that they were those of a giant lizard-like creature. Buckland further studied the remains with 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 forty feet long and eight feet high. It is generally considered the name in 1822 was still a nomen nudum ("naked name"). Buckland, urged on by an impatient Cuvier, continued to work on the subject during 1823, letting his later wife Mary Morland provide 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.

Fossil specimens referred to M. bucklandii, Oxford University Museum of Natural History. The display shows most of the original syntype series, including the lectotype dentary, identified by Buckland in 1824

Early reconstructions

The first reconstruction was given by Buckland himself. He considered Megalosaurus to be a quadruped. He thought it was an "amphibian", i.e. an animal capable of both swimming in the sea and walking on land. Generally, in his mind Megalosaurus resembled a gigantic lizard, but Buckland already understood from the form of the thighbone head that the legs were not so much sprawling as held rather upright. In the original description of 1824, Buckland repeated Cuvier's size estimate that Megalosaurus would have been forty feet long with the weight of a seven foot tall elephant. However, this had been based on the remains present at Oxford. Buckland had also been hurried into naming his new reptile by a visit he had made to the fossil collection of Mantell, who during the lecture announced to have acquired a fossil thighbone of enormous magnitude, twice as long as that just described. Today, this is known to have belonged to Iguanodon, or at least some iguanodontid, but at the time both men assumed this bone belonged to Megalosaurus also. Even taking into account the effects of allometry, heavier animals having relatively stouter bones, Buckland was forced in the printed version of his lecture to estimate the maximum length of Megalosaurus at sixty to seventy feet. The existence of Megalosaurus posed some problems for Christian orthodoxy, which typically held that suffering and death had only come into the world through Original Sin, which seemed irreconcilable with the presence of a gigantic devouring reptile during a pre-Adamitic phase of history. Buckland rejected the usual solution, that such carnivores would originally have been peaceful vegetarians, as infantile and claimed in one of the Bridgewater Treatises that Megalosaurus had played a beneficial rôle in creation by ending the lives of old and ill animals, "to diminish the aggregate amount of animal suffering".

1854 reconstruction in Crystal Palace Park guided by Richard Owen presents Megalosaurus as a quadruped; modern reconstructions make it bipedal, like most theropods

In 1824, Buckland assigned Megalosaurus to the Sauria, assuming within the Sauria a close affinity with modern lizards, more than with crocodiles. In 1842, Owen made Megalosaurus one of the first three genera placed in the Dinosauria. In 1850, Prince Charles Lucien Bonaparte coined a separate family Megalosauridae with Megalosaurus as the type genus. For a long time, the precise relationships of Megalosaurus remained vague. It was seen as a "primitive" member of the Carnosauria, the group in which most large theropods were united.

In the late 20th century the new method of cladistics allowed for the first time to exactly calculate how closely various taxa were related to each other. In 2012, Matthew Carrano et al. showed that Megalosaurus was the sister species of Torvosaurus within the Megalosaurinae

Megalosaurus By Manuelsaurus

Living in what is now Europe, during the Jurassic Period (~201 to ~145 million years ago), Megalosaurus may have hunted stegosaurs and sauropods. Repeated descriptions during the nineteenth and early twentieth century of Megalosaurus hunting Iguanodon (another of the earliest dinosaurs named) through the forests that then covered the continent are now known to be inaccurate, because Iguanodon skeletons are found in much younger Early Cretaceous formations. The only specimens belonging to Megalosaurus bucklandii are from the Lower/Middle Bathonian of Oxfordshire and Gloucestershire. No material from outside of the Bathonian formations of England can be referred to Megalosaurus. It lived alongside the theropods Cruxicheiros, Iliosuchus and Streptospondylus, and the sauropods Cardiodon, Cetiosaurus, and possibly Cetiosauriscus. The pterosaur Rhamphocephalus, and indeterminate sauropod and ornithopod remains have also been found alongside fossils of Megalosaurus.