Daspletosaurus

Daspletosaurus (/dæsˌpliːtɵˈsɔrəs/ das-PLEET-o-SAWR-əs; meaning "frightful lizard") is a genus of tyrannosaurid theropod dinosaur that lived in western North America between 77 and 74 million years ago, during the Late Cretaceous Period. Fossils of the only named species (D. torosus) were found in Alberta, although other possible species from Alberta and Montana await description. Including these undescribed species makes Daspletosaurus the most species-rich genus of tyrannosaur.

Daspletosaurus is closely related to the much larger and more recent Tyrannosaurus. Like most known tyrannosaurids, it was a multi-tonne bipedal predator equipped with dozens of large, sharp teeth. Daspletosaurus had the small forelimbs typical of tyrannosaurids, although they were proportionately longer than in other genera.

As an apex predator, Daspletosaurus was at the top of the food chain, probably preying on large dinosaurs like the ceratopsid Centrosaurus and the hadrosaur Hypacrosaurus. In some areas, Daspletosaurus coexisted with another tyrannosaurid, Gorgosaurus, though there is some evidence of niche differentiation between the two. While Daspletosaurus fossils are rarer than other tyrannosaurids, the available specimens allow some analysis of the biology of these animals, including social behavior, diet, and life history.

Description
While very large by the standard of modern predators, Daspletosaurus was not the largest tyrannosaurid. Adults could reach a length of 8–9 meters (26–30 ft) from snout to tail.[1] Mass estimates have centered around 2.5 tonnes (2.75 short tons)[1][2][3] but have ranged between 1.8[4] and 3.8 tonnes (2.0 and 4.1 short tons).[5]

Daspletosaurus had a massive skull that could reach more than 1 meter (3.3 ft) in length.[1] The bones were heavily constructed and some, including the nasal bones on top of the snout, were fused for strength. Large fenestrae (openings) in the skull reduced its weight. An adult Daspletosaurus was armed with about six dozen teeth that were very long but oval in cross section rather than blade-like. Unlike its other teeth, those in the premaxilla at the end of the upper jaw had a D-shaped cross section, an example of heterodonty always seen in tyrannosaurids. Unique skull features included the rough outer surface of the maxilla (upper jaw bone) and the pronounced crests around the eyes on the lacrimal, postorbital, and jugal bones. The orbit (eye socket) was a tall oval, somewhere in between the circular shape seen in Gorgosaurus and the 'keyhole' shape of Tyrannosaurus.[6][7][8] Split carinae[9] have been found on Daspletosaurus teeth.[10]

Daspletosaurus shared the same body form as other tyrannosaurids, with a short, S-shaped neck supporting the massive skull. It walked on its two thick hindlimbs, which ended in four-toed feet, although the first digit (the hallux) did not contact the ground. In contrast, the forelimbs were extremely small and bore only two digits, although Daspletosaurus had the longest forelimbs in proportion to body size of any tyrannosaurid. A long, heavy tail served as a counterweight to the head and torso, with the center of gravity over the hips

Classification and Systematics
Daspletosaurus belongs in the subfamily Tyrannosaurinae within the family Tyrannosauridae, along with Tarbosaurus, Tyrannosaurus and Alioramus. Animals in this subfamily are more closely related to Tyrannosaurus than to Albertosaurus and are known – with the exception of Alioramus – for their robust build with proportionally larger skulls and longer femora than in the other subfamily, the Albertosaurinae.[8][11]

Daspletosaurus is usually considered to be closely related to Tyrannosaurus rex, or even a direct ancestor through anagenesis.[12] Gregory Paul reassigned D. torosus to the genus Tyrannosaurus, creating the new combination Tyrannosaurus torosus,[2] but this has not been generally accepted.[6][8] Many researchers believe Tarbosaurus and Tyrannosaurus to be sister taxa or even to be the same genus, with Daspletosaurus a more basal relative.[8][13] On the other hand, Phil Currie and colleagues find Daspletosaurus to be more closely related to Tarbosaurus and other Asian tyrannosaurids like Alioramus than to the North American Tyrannosaurus.[11] The systematics (evolutionary relationships) of Daspletosaurus may become clearer once all the species have been described.

Below is the cladogram of Tyrannosauridae based on the phylogenetic analysis conducted by Loewen et al. in 2013.