Megalosauridae is a monophyletic family of carnivorous theropod dinosaurs within the group Megalosauroidea. Appearing in the Middle Jurassic, megalosaurids were among the first major radiation of large theropod dinosaurs. They were a relatively primitive group of basal tetanurans containing two main subfamilies, Megalosaurinae and Afrovenatorinae, along with the basal genus Eustreptospondylus, an unresolved taxon which differs from both subfamilies. The defining megalosaurid, Megalosaurus bucklandii, was first named and described in 1824 by William Buckland after multiple finds in Stonesfield, Oxfordshire, UK. Megalosaurus was the first formally described dinosaur and was the basis for the establishment of the clade Dinosauria. It is also one of the largest known Middle Jurassic carnivorous dinosaurs, with the best-preserved femur at 805 mm and a proposed body mass of around 943 kg. Megalosauridae has mainly been recognized as a European group of dinosaurs, based on fossils found in France and the UK, but fossils show that the group is also found in North America, Africa, South America and possibly Asia. The family Megalosauridae was first defined by Thomas Huxley in 1869, yet it has been contested throughout history due to its role as a "waste-basket" for many partially described dinosaurs or unidentified remains. In the early years of paleontology, most large theropods were grouped together and up to 48 species were included in the clade Megalosauria, the basal clade of Megalosauridae. Over time, most of these taxa were placed in other clades and the parameters of Megalosauridae were narrowed significantly. However, some controversy remains over whether Megalosauridae should be considered its own distinct group, and dinosaurs in this family remain some of the most problematic taxa in all Dinosauria. Some paleontologists, such as Paul Sereno in 2005, have disregarded the group due to its shaky foundation and lack of clarified phylogeny. However, recent research by Carrano, Benson, and Sampson has systematically analyzed all basal tetanurans and determined that Megalosauridae should exist as its own family. They have been generally closely related to the family Spinosauridae.
Description
Body size
Like other tetanurans, megalosaurids are carnivorous theropods characterized by large size and bipedalism. Specifically, megalosaurids exhibit especially giant size, with some members of the family weighing more than one tonne. Over time, there is evidence of size increase within the family. Basal megalosaurids from the Early Jurassic had smaller body size than those appearing in the late Middle Jurassic. Due to this size increase over time, Megalosauridae appear to follow a size increase pattern similar to that of other giant sized theropods like Spinosauridae. This pattern follows Cope's rule, the postulation by paleontologist Edward Cope about evolutionary increase in body size.
Anatomical characteristics
One unambiguous synapomorphy of Megalosauridae is a lower and longer skull with a length to height ratio of 3:1. In addition, the typical skull roof tends to be much less ornamented than that of other tetanurans, and crests or horns are either very small or absent entirely. Megalosaurids also have femoral heads with an orientation 45 degrees between anteromedial and fully medial. Megalosauridae are also defined by the following unique unambiguous synapomorphies:
A humeral deltopectoral crest that terminates about halfway along the humeral shaft The absence of a fibular anterolateral tubercle. Nares which extend as far as the premaxillary teeth, yet the portion of the premaxilla anterior to the nares being longer than the portion under them; angled snout tip (angle between the anterior and alveolar margins <70 degrees). Medial foramina on the quadrate adjacent to the mandibular condyles. Pleurocoelous fossae on the sacral vertebrae. The oblique ligament groove on the posterior surface of the femoral head is shallow. Megalosaurinae (all megalosaurids more closely related to Megalosaurus than Afrovenator) are characterized by a moderate (0.5–2.0) height/length ratio of the premaxilla below the level of the nares, compared to other megalosaurids which have a lower ratio and thus less tall snout tip. Afrovenatorinae (all megalosaurids more closely related to Afrovenator than Megalosaurus) are characterized by a squared anterior margin of the antorbital fossa and the puboischiadic plate being broadly open along the midline.
Dental morphology
Dental findings are frequently used to differentiate between various theropods and to further inform cladistic phylogeny. Tooth morphology and dental evolutionary markers are prone to homoplasy and disappear or reappear throughout history. However, megalosaurids have several specific denture conditions that differentiate them from other basal theropods. One dental condition present in Megalosauridae is multiple enamel wrinkles near the carinae, the sharp edge or serration row of the tooth. Ornamented teeth and a well-marked enamel surface also characterize basal megalosaurids. The ornamentation and well-marked surface appears in early megalosaurids but disappears in derived megalosaurids, suggesting that the condition was lost over time as megalosaurids grew in size.
Classification
Historical classification
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