Tarbosaurus

Tarbosaurus (/ˌtɑrbɵˈsɔrəs/ TAR-bo-SAWR-əs; meaning "alarming lizard") is a genus of tyrannosaurid theropod dinosaur that flourished in Asia about 70 million years ago, at the end of the Late Cretaceous Period. Fossils have been recovered in Mongolia, with more fragmentary remains found further afield in parts of China.

Although many species have been named, modern paleontologists recognize only one, T. bataar, as valid. Some experts see this species as an Asian representative of the North American genus Tyrannosaurus; this would make the genus Tarbosaurus redundant. Tarbosaurus and Tyrannosaurus, if not synonymous, are considered to be at least closely related genera. Alioramus, also from Mongolia, is thought by some authorities to be the closest relative of Tarbosaurus.

Like most known tyrannosaurids, Tarbosaurus was a large bipedal predator, weighing up to six tonnes and equipped with about sixty large teeth. It had a unique locking mechanism in its lower jaw and the smallest forelimbs relative to body size of all tyrannosaurids, renowned for their disproportionately tiny, two-fingered forelimbs.

Tarbosaurus lived in a humid floodplain criss-crossed by river channels. In this environment, it was an apex predator at the top of the food chain, probably preying on other large dinosaurs like the hadrosaur Saurolophus or the sauropod Nemegtosaurus. Tarbosaurus is very well represented in the fossil record, known from dozens of specimens, including several complete skulls and skeletons. These remains have allowed scientific studies focusing on its phylogeny, skull mechanics, and brain structure.

Description
Although smaller than Tyrannosaurus, Tarbosaurus was one of the largest tyrannosaurids. The largest known individuals were between 10 and 12 meters (33 to 40 ft) long.[1] The mass of a fully grown individual is considered comparable to or slightly smaller than Tyrannosaurus, often estimated to be around 4–5 metric tons.[2][3]

The largest known Tarbosaurus skull is more than 1.3 meters (4 ft) long, larger than all other tyrannosaurids except Tyrannosaurus.[4] The skull was tall, like that of Tyrannosaurus, but not as wide, especially towards the rear. The unexpanded rear of the skull meant that Tarbosaurus eyes did not face directly forwards, suggesting that it lacked the binocular vision of Tyrannosaurus. Large fenestrae (openings) in the skull reduced its weight. Between 58 and 64 teeth lined its jaws, slightly more than in Tyrannosaurus but fewer than in smaller tyrannosaurids like Gorgosaurus and Alioramus. Most of its teeth were oval in cross section, although the teeth of the premaxilla at the tip of the upper jaw had a D-shaped cross section. This heterodonty is characteristic of the family. The longest teeth were in the maxilla (upper jaw bone), with crowns up to 85 millimeters (3.3 in) long. In the lower jaw, a ridge on the outer surface of the angular bone articulated with the rear of the dentary bone, creating a locking mechanism unique to Tarbosaurus and Alioramus. Other tyrannosaurids lacked this ridge and had more flexibility in the lower jaw.

Tyrannosaurids varied little in body form, and Tarbosaurus was no exception. The head was supported by an S-shaped neck, while the rest of the vertebral column, including the long tail, was held horizontally. Tarbosaurus had tiny forelimbs, proportionably to body size the smallest of all members of the family. The hands had two clawed digits each, with an additional unclawed third metacarpal found in some specimens, similar to closely related genera. Holtz has suggested that Tarbosaurus also has a theropod reduction of fingers IV-I "developed further" than in other tyrannosaurids,[6] as the second metacarpal in the Tarbosaurus specimens he studied is less than twice the length of the first metacarpal (other tyrannosaurids have a second metacarpal about twice the length of the first metacarpal). Also, the third metacarpal in Tarbosaurus is proportionally shorter than in other tyrannosaurids; in other tyrannosaurids (like Albertosaurus and Daspletosaurus), the third metacarpal is often longer than the first metacarpal, while in the Tarbosaurus specimens studied by Holtz, the third metacarpal is shorter than the first.[4]

In contrast to the forelimbs, the three-toed hindlimbs were long and thick, supporting the body in a bipedal posture. The long, heavy tail served as a counterweight to the head and torso and placed the center of gravity over the hips.

Classification and systematics
Tarbosaurus is classified as a theropod in the subfamily Tyrannosaurinae within the family Tyrannosauridae. Other members include Tyrannosaurus and the earlier Daspletosaurus, both from North America,[7] and possibly the Mongolian genus Alioramus.[5][8] Animals in this subfamily are more closely related to Tyrannosaurus than to Albertosaurus and are known for their robust build with proportionally larger skulls and longer femurs than in the other subfamily, the Albertosaurinae.[4]

Tarbosaurus bataar was originally described as a species of Tyrannosaurus,[9] an arrangement that has been supported by some more recent studies.[7][10] Others prefer to keep the genera separate, while still recognizing them as sister taxa.[4] A 2003 cladistic analysis based on skull features instead identified Alioramus as the closest known relative of Tarbosaurus, as the two genera share skull characteristics that are related to stress distribution and that are not found in other tyrannosaurines. If proven, this relationship would argue against Tarbosaurus becoming a synonym for Tyrannosaurus and would suggest that separate tyrannosaurine lineages evolved in Asia and North America.[5][8] The two known specimens of Alioramus, which show juvenile characteristics, are not likely juvenile individuals of Tarbosaurus because of their much higher tooth count (76 to 78 teeth) and their unique row of bony bumps along the top of their snouts.[11]

The discovery of Lythronax argestes, a much earlier tyrannosaurine further reveals the close relationship between Tyrannosaurus and Tarbosaurus, and it was discovered that Lythronax is a sister taxon to a clade consisting of Campanian genus Zhuchengtyrannus, and Maastrichtian genera Tyrannosaurus and Tarbosaurus. Further studies of Lythronax also suggest that the Asian tyrannosauroids were part of one evolutionary radiation.

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