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Megalosaurus
Temporal range: Middle Jurassic
Fossil specimens referred to M. bucklandii, Oxford University Museum of Natural History.
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Sauropsida
Superorder: Dinosauria
Order: Saurischia
Suborder: Theropoda
Family: Megalosauridae
Genus: Megalosaurus
Buckland, 1824

Megalosaurus was a large meat-eating theropod dinosaur of the Middle Jurassic of Europe. It was found in 1824, and was the earliest dinosaur named. In 1827, Gideon Mantell included Megalosaurus in his geological survey of southeastern England. He gave the species its name, Megalosaurus bucklandii.

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 lizard of twenty metres length. In 1842 Richard Owen concluded that it was no longer than nine metres, standing on upright legs as a quadruped.

Later it was realised that all theropods were bipedal. Today is known that the early proto-dinosaurs in the late Middle Triassic were also bipedal. Certainly Eoraptor, from 231.4 million years ago, was bipedal. That means the dinosaurs were bipedal from the start, and only the heavier sauropods and armoured dinosaurs reverted to standing on all four legs.

Description

Since the first finds, many other Megalosaurus bones have been recovered; however, no complete skeleton has yet been found. Therefore, the details of its physical appearance cannot be certain. However, a full osteology of all known material was published in 2010 by Benson.

Megalosaurus size chart
Size comparison

Size

Traditionally, most texts, following Owen's estimate of 1841, give a body length of thirty feet or nine metres for Megalosaurus. The lack of an articulated dorsal vertebral series makes it difficult to determine an exact size. David Bruce Norman in 1984 thought Megalosaurus was seven to eight metres long. Gregory S. Paul in 1988 estimated the weight tentatively at 1.1 tonnes, given a thighbone seventy-six centimetres long. The trend in the early twenty-first century to limit the material to the lectotype inspired even lower estimates, disregarding outliers of uncertain identity. Paul in 2010 stated Megalosaurus was six metres long and seven hundred kilogrammes heavy. However, the same year Benson claimed that Megalosaurus, though medium-sized, was still among the largest of Middle Jurassic theropods. Specimen BMNH 31806, a thighbone 803 millimetres long, would indicate a body weight of 943 kilogrammes, using the extrapolation method of J.F. Anderson — which method, optimised for mammals, tends to underestimate theropod masses by at least a third. Furthermore, thighbone specimen OUM J13561 has a length of about eighty-six centimetres.

General build

Megalosaurus fragments
Hip, femur and sacrum

In general, Megalosaurus had the typical build of a large theropod. It was bipedal, the horizontal torso being balanced by a long horizontal tail. The hindlimbs were long and strong with three forward-facing weight-bearing toes, the forelimbs relatively short but exceptionally robust and probably carrying three digits. Being a carnivore, its large elongated head bore long dagger-like teeth to slice the flesh of its prey. The skeleton of Megalosaurus is highly ossified, indicating a robust and muscular animal, though the lower leg was not as heavily built as that of Torvosaurus, a close relative.

Skull and lower jaws

Megalosaurus bucklandi tooth 2
Referred tooth of M. bucklandii.

The skull of Megalosaurus is poorly known. The discovered skull elements are generally rather large in relation to the rest of the material. This can either be coincidental or indicate that Megalosaurus had an uncommonly large head. The praemaxilla is not known, making it impossible to determine whether the snout profile was curved or rectangular. A rather stubby snout is suggested by the fact that the front branch of the maxilla was short. In the depression around the antorbital fenestra to the front, a smaller non-piercing hollowing can be seen that is probably homologous to the fenestra maxillaris. The maxilla bears thirteen teeth. The teeth are relatively large, with a crown length up to seven centimetres. The teeth are supported from behind by tall, triangular, unfused interdental plates. The cutting edges bear eighteen to twenty denticula per centimetre. The tooth formula is probably 4, 13–14/13–14. The jugal bone is pneumatised, pierced by a large foramen from the direction of the antorbital fenestra. It was probably hollowed out by an outgrowth of an air sac in the nasal bone. Such a level of pneumatisation of the jugal is not known from other megalosaurids and might represent a separate autapomorphy.

Megalosaurus bucklandi
Cast of the lower jaw

The lower jaw is rather robust. It is also straight in top view, without much expansion at the jaw tip, suggesting the lower jaws as a pair, the mandibula, were narrow. Several traits in 2008 identified as autapomorphies, later transpired to have been the result of damage. However, a unique combination of traits is present in the wide longitudinal groove on the outer side (shared with Torvosaurus), the small third dentary tooth and a vascular channel, below the row of interdental plates, that only is closed from the fifth tooth position onwards. The number of dentary teeth was probably thirteen or fourteen, though the preserved damaged specimens show at most eleven tooth sockets. The interdental plates have smooth inner sides, whereas those of the maxilla are vertically grooved; the same combination is shown by Piatnitzkysaurus. The surangular has no bony shelf, or even ridge, on its outer side. There is laterally an oval opening present in front of the jaw joint, a foramen surangulare posterior, but a second foramen surangulare anterior to the front of it is lacking.

Vertebral column

Megalosaurus sacrum (39971421271)
OUM J13576, an articulated sacrum

Although the exact numbers are unknown, the vertebral column of Megalosaurus was probably divided into ten neck vertebrae, thirteen dorsal vertebrae, five sacral vertebrae and fifty to sixty tail vertebrae, as is common for basal Tetanurae.

The Stonesfield Slate material contains no neck vertebrae; but a single broken anterior cervical vertebra is known from the New Park Quarry, specimen BMNH R9674. The breakage reveals large internal air chambers. The vertebra is also otherwise heavily pneumatised, with large pleurocoels, pneumatic excavations, on its sides. The rear facet of the centrum is strongly concave. The neck ribs are short. The front dorsal vertebrae are slightly opisthocoelous, with convex front centrum facets and concave rear centrum facets. They are also deeply keeled, with the ridge on the underside representing about 50% of the total centrum height. The front dorsals perhaps have a pleurocoel above the diapophysis, the lower rib joint process. The rear dorsal vertebrae, according to Benson, are not pneumatised. They are slightly amphicoelous, with hollow centrum facets. They have secondary joint processes, forming a hyposphene–hypantrum complex, the hyposphene having a triangular transverse cross-section. The height of the dorsal spines of the rear dorsals is unknown, but a high spine on a tail vertebra of the New Park Quarry material, specimen BMNH R9677, suggests the presence of a crest on the hip area. The spines of the five vertebrae of the sacrum form a supraneural plate, fused at the top. The undersides of the sacral vertebrae are rounded but the second sacral is keeled; normally it is the third or fourth sacral having a ridge. The sacral vertebrae seem not to be pneumatised but have excavations at their sides. The tail vertebrae are slightly amphicoelous, with hollow centrum facets on both the front and rear side. They have excavations at their sides and a longitudinal groove on the underside. The neural spines of the tail basis are transversely thin and tall, representing more than half of the total vertebral height.

Appendicular skeleton

Megalosaurus femur
Lithograph of the femur, from a posterior (left) and medial view (right). Work by J. Erxleben in the 1800s

The shoulderblade or scapula is short and wide, its length about 6.8 times the minimum width; this is a rare and basal trait within Tetanurae. Its top curves slightly to the rear in side view. On the lower outer side of the blade a broad ridge is present, running from just below the shoulder joint to about midlength where it gradually merges with the blade surface. The middle front edge over about 30% of its length is thinned forming a slightly protruding crest. The scapula constitutes about half of the shoulder joint, which is oriented obliquely sideways and to below. The coracoid is in all known specimens fused with the scapula into a scapulocoracoid, lacking a visible suture. The coracoid as such is an oval bone plate, with its longest side attached to the scapula. It is pierced by a large oval foramen but the usual boss for the attachment of the upper arm muscles is lacking.

The humerus is very robust with strongly expanded upper and lower ends. Humerus specimen OUMNH J.13575 has a length of 388 millimetres. Its shaft circumference equals about half of the total humerus length. The humerus head continues to the front and the rear into large bosses, together forming a massive bone plate. On the front outer side of the shaft a large triangular deltopectoral crest is present, the attachment for the Musculus pectoralis major and the Musculus deltoideus. It covers about the upper half of the shaft length, its apex positioned rather low. The ulna is extremely robust, for its absolute size more heavily built than with any other known member of the Tetanurae. The only known specimen, BMNH 36585, has a length of 232 millimetres and a minimal shaft circumference of 142 millimetres. The ulna is straight in front view and has a large olecranon, the attachment process for the Musculus triceps brachii. Radius, wrist and hand are unknown.

Megalosaurus pelvis
Lithograph of the pelvis. Work from the 1800s drawn by J. Erxleben

In the pelvis, the ilium is long and low, with a convex upper profile. Its front blade is triangular and rather short; at the front end there is a small drooping point, separated by a notch from the pubic peduncle. The rear blade is roughly rectangular. The outer side of the ilium is concave, serving as an attachment surface for the Musculus iliofemoralis, the main thigh muscle. Above the hip joint, on this surface a low vertical ridge is present with conspicuous vertical grooves. The bottom of the rear blade is excavated by a narrow but deep trough forming a bony shelf for the attachment of the Musculus caudofemoralis brevis. The outer side of the rear blade does not match the inner side, which thus can be seen as a separate "medial blade" that in side view is visible in two places: in the corner between outer side and the ischial peduncle and as a small surface behind the extreme rear tip of the outer side of the rear blade. The pubic bone is straight. The pubic bones of both pelvis halves are connected via narrow bony skirts that originated at a rather high position on the rear side and continued downwards to a point low on the front side of the shaft. The ischium is S-shaped in side view, showing at the transition point between the two curvatures a rough boss on the outer side. On the front edge of the ischial shaft an obturator process is present in the form of a low ridge, at its top separated from the shaft by a notch. To below, this ridge continues into an exceptionally thick bony skirt at the inner rear side of the shaft, covering over half of its length. Towards the end of the shaft, this skirt gradually merges with it. The shaft eventually ends in a sizeable "foot" with a convex lower profile.

Megalosaurus right tibia (& left metatarsals)
Tibia and metatarsals

The thighbone is straight in front view. Seen from the same direction its head is perpendicular to the shaft, seen from above it is orientated 20° to the front. The greater trochanter is relatively wide and separated from the robust lesser trochanter in front of it, by a fissure. At the front base of the lesser trochanter a low accessory trochanter is present. At the lower end of the thighbone a distinct front, extensor, groove separates the condyles. At the upper inner side of this groove a rough area is present continuing inwards into a longitudinal ridge, a typical megalosauroid trait. The shinbone, or tibia, is relatively straight, slightly curving inwards. To below, its shaft progressively flattens from front to rear, resulting in a generally oval cross-section. For about an eighth of its length the front lower end of the shaft is covered by a vertical branch of the astragalus. Of the foot, only the second, third and fourth metatarsals are known, the bone elements that were connected to the three weight-bearing toes. They are straight and robust, showing ligament pits at their lower sides. The third metatarsal has no clear condyles at its lower end, resulting in a more flexible joint, allowing for a modicum of horizontal movement. The top inner side of the third metatarsal carries a unique ridge that fits into a groove along the top outer side of the second metatarsal, causing a tighter connection.

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