Megaraptora

Megaraptora is a clade of carnivorous theropod dinosaurs with elongated hand claws and controversial relations to other theropods. Megaraptorans are incompletely known, and no complete megaraptoran skeleton has been found. However, they still possessed a number of unique features. Their forelimbs were large and strongly built, and the ulna bone had a unique shape in members of the family Megaraptoridae, a subset of megaraptorans which excludes Fukuiraptor. The first two fingers were elongated, with massive curved claws, while the third finger was small. Megaraptoran skull material is very incomplete, but a juvenile Megaraptor described in 2014 preserved a portion of the snout, which was long and slender. Leg bones referred to megaraptorans were also quite slender and similar to those of coelurosaurs adapted for running. Although megaraptorans were thick-bodied theropods, their bones were heavily pneumatized, or filled with air pockets. The vertebrae, ribs, and the ilium bone of the hip were pneumatized to an extent which was very rare among theropods, only seen elsewhere in taxa such as Neovenator. Other characteristic features include opisthocoelous neck vertebrae and compsognathid-like teeth. The clade was originally named in 2010 as a subset of the family Neovenatoridae, a group of lightly-built allosauroids related to the massive carcharodontosaurids such as Giganotosaurus and Carcharodontosaurus. A 2013 phylogenetic analysis by Fernando Novas and his colleagues disagreed with this classification scheme, and instead argued that the megaraptorans evolved deep within Tyrannosauroidea, a superfamily of basal coelurosaurs including the famous Tyrannosaurus. Subsequent refinements to Novas's data and methodologies have supported a third position for the group, at the base of Coelurosauria among other controversial theropods such as Gualicho, but not within the Tyrannosauroidea. Regardless of their position, it is clear that megaraptorans experienced a large amount of convergent evolution with either Neovenator-like allosauroids or basal coelurosaurs. Megaraptorans were most diverse in the early Late Cretaceous of South America, particularly Patagonia. However, they had a widespread distribution. Fukuiraptor, the most basal ('primitive') known member of the group, lived in Japan. Megaraptoran material is also common in Australia, and the largest known predatory dinosaur from the continent, Australovenator, was a megaraptoran. Megaraptorans were medium to large-sized theropods, ranging from Fukuiraptor, which was about 4.2 meters (14 feet) in length, to the 9 meter (30 feet)-long Aerosteon, and the 42 foot long Bahariasaurus, if it is a member. Most megaraptorans are known from very fragmentary remains, although certain characteristics can be identified in multiple members of the clade. At least some megaraptorans, such as Murusraptor and Aerosteon, had extensively pneumatic bones (most noticeably the ilia and ribs), which likely housed sinuses connected to the lungs, similar to modern birds. The slender leg bones and long metatarsals of several species indicate that members of this group likely had cursorial habits. Most megaraptorans are part of the family Megaraptoridae, which was named by Fernando Novas and his colleagues in 2013. This family is united by several adaptations of the ulna and claws which are not present in the basal megaraptoran Fukuiraptor. No megaraptoran fossil is known to preserve a complete skull, although skull material is known for several taxa. Aerosteon, Orkoraptor, and Murusraptor preserve several bones of the rear part of the skull, lower jaws are known from Australovenator, and a juvenile specimen of Megaraptor described in 2014 preserved much of the snout as well as parietal fragments. Teeth have been found in many genera. Collectively, megaraptorans can be reconstructed as having a long, lightly built skull with many relatively small teeth. Based on Megaraptor, the premaxillary bone at the tip of the snout is small, with a long and rod-like branch of bone which extends above the external nares (nostril holes). The nares themselves were very large and elongated, akin to some early tyrannosauroids (Dilong, Proceratosaurus, etc). The snout also had some similarities to carcharodontosaurids, namely the straight upper edge of the maxilla and rectangular nasal bones. The parietal bones at the top of the skull, behind the eyes, had a strongly developed sagittal crest, as in tyrannosauroids. Otherwise, the rear part of the skull is rather simple, without any pronounced crests or bosses, although the lacrimal and postorbital bones did have rugose patches in some genera. Aerosteon and Murusraptor possessed a pneumatic quadrate, as in a few allosauroids (Sinraptor, Mapusaurus) and tyrannosauroids. The dentary, which is only known in Australovenator, is long and graceful, with the first tooth smaller than the rest (as in tyrannosauroids). The mandible as a whole has only a single meckelian foramen, as in carcharodontosaurians, tyrannosaurids, and ornithomimids. However, the rear part of the mandible (as seen in Murusraptor) was significantly more lightly built than that of tyrannosauroids. Preserved braincase material has similarities to both carcharodontosaurians and tyrannosauroids. The premaxillary teeth of Megaraptor were variably similar to those of tyrannosauroids, being small, incisiform (chisel-like) and D-shaped in cross section. However, Murusraptor's premaxillary teeth were fang-like, as in non-tyrannosauroid theropods. Megaraptoran maxillary teeth show much variety between genera, although they were generally small compared to the snout with minimal enamel ornamentation. Some megaraptorans, such as Orkoraptor, Australovenator, and Megaraptor, had teeth which were 8-shaped in cross section and completely unserrated from the front (similar to dromaeosaurids and compsognathids), while Murusraptor had anterior serrations only at the tip of its teeth. Fukuiraptor had very laterally compressed and blade-like teeth (similar to carcharodontosaurs) with both anterior and posterior serrations.