Tyrannosaurus (meaning "tyrant lizard", from Greek tyrannos (τυράννος) meaning "tyrant," and sauros (σαύρος) meaning "lizard") is a genus of coelurosaurian theropod dinosaur. The species Tyrannosaurus rex (rex meaning "king" in Latin), commonly abbreviated to T-Rex, is a fixture in popular culture. It lived throughout what is now western North America, with a much wider range than other tyrannosaurids. Fossils are found in a variety of rock formations dating to the Maastrichtian age of the upper Cretaceous Period, 67 to 65.5 million years ago. It was among the last non-avian dinosaurs to exist before the Cretaceous–Paleogene extinction event.
Like other tyrannosaurids, Tyrannosaurus was a bipedal carnivore with a massive skull balanced by a long, heavy tail. Relative to the large and powerful hindlimbs, Tyrannosaurus forelimbs were small, though unusually powerful for their size, and bore two clawed digits. Although other theropods rivaled or exceeded Tyrannosaurus rex in size, it was the largest known tyrannosaurid and one of the largest known land predators, measuring up to 12.8 m (42 ft) in length, up to 4 metres (13 ft) tall at the hips, and up to 6.8 metric tons (7.5 short tons) in weight. By far the largest carnivore in its environment, Tyrannosaurus rex may have been an apex predator, preying upon hadrosaurs and ceratopsians, although some experts have suggested it was primarily a scavenger. The debate over Tyrannosaurus as apex predator or scavenger is among the longest running in paleontology.
More than 30 specimens of Tyrannosaurus rex have been identified, some of which are nearly complete skeletons. Soft tissue and proteins have been reported in at least one of these specimens. The abundance of fossil material has allowed significant research into many aspects of its biology, including life history and biomechanics. The feeding habits, physiology and potential speed of Tyrannosaurus rex are a few subjects of debate. Its taxonomy is also controversial, with some scientists considering Tarbosaurus bataar from Asia to represent a second species of Tyrannosaurus and others maintaining Tarbosaurus as a separate genus. Several other genera of North American tyrannosaurids have also been synonymized with Tyrannosaurus.
Description
Tyrannosaurus rex was one of the largest land carnivores of all time; the largest complete specimen, FMNH PR2081 ("Sue"), measured 12.8 metres (42 ft) long, and was 4.0 metres (13.1 ft) tall at the hips. Mass estimates have varied widely over the years, from more than 7.2 metric tons (7.9 short tons), to less than 4.5 metric tons (5.0 short tons), with most modern estimates ranging between 5.4 and 6.8 metric tons (6.0 and 7.5 short tons). Packard et al. (2009) tested dinosaur mass estimation procedures on elephants and concluded that dinosaur estimations are flawed and produce over-estimations; thus, the weight of Tyrannosaurus could be much less than usually estimated. Other estimations have concluded that the largest known Tyrannosaurus specimens had a weight exceeding 9 tonnes.
Although Tyrannosaurus rex was larger than the well known Jurassic theropod Allosaurus, it was slightly smaller than two other Cretaceous carnivores, Spinosaurus and Giganotosaurus.
The neck of Tyrannosaurus rex formed a natural S-shaped curve like that of other theropods, but was short and muscular to support the massive head. The forelimbs had only two clawed fingers, along with an additional small metacarpal representing the remnant of a third digit. In contrast the hind limbs were among the longest in proportion to body size of any theropod. The tail was heavy and long, sometimes containing over forty vertebrae, in order to balance the massive head and torso. To compensate for the immense bulk of the animal, many bones throughout the skeleton were hollow, reducing its weight without significant loss of strength.
Skulls
The largest known Tyrannosaurus rex skulls measure up to 5 feet (1.5 m) in length. Large fenestrae (openings) in the skull reduced weight and provided areas for muscle attachment, as in all carnivorous theropods. But in other respects Tyrannosaurus's skull was significantly different from those of large non-tyrannosauroid theropods. It was extremely wide at the rear but had a narrow snout, allowing unusually good binocular vision. The skull bones were massive and the nasals and some other bones were fused, preventing movement between them; but many were pneumatized (contained a "honeycomb" of tiny air spaces) which may have made the bones more flexible as well as lighter. These and other skull-strengthening features are part of the tyrannosaurid trend towards an increasingly powerful bite, which easily surpassed that of all non-tyrannosaurids. The tip of the upper jaw was U-shaped (most non-tyrannosauroid carnivores had V-shaped upper jaws), which increased the amount of tissue and bone a tyrannosaur could rip out with one bite, although it also increased the stresses on the front teeth.
Teeth
The teeth of Tyrannosaurus rex displayed marked heterodonty (differences in shape). The premaxillary teeth at the front of the upper jaw were closely packed, D-shaped in cross-section, had reinforcing ridges on the rear surface, were incisiform (their tips were chisel-like blades) and curved backwards. The D-shaped cross-section, reinforcing ridges and backwards curve reduced the risk that the teeth would snap when Tyrannosaurus bit and pulled. The remaining teeth were robust, like "lethal bananas" rather than daggers; more widely spaced and also had reinforcing ridges. Those in the upper jaw were larger than those in all but the rear of the lower jaw. The largest found so far is estimated to have been 30 centimetres (12 in) long including the root when the animal was alive, making it the largest tooth of any carnivorous dinosaur yet found.
Arms
When Tyrannosaurus rex was first discovered, the humerus was the only element of the forelimb known. For the initial mounted skeleton as seen by the public in 1915, Osborn substituted longer, three-fingered forelimbs like those of Allosaurus. However, a year earlier, Lawrence Lambe described the short, two-fingered forelimbs of the closely related Gorgosaurus. This strongly suggested that Tyrannosaurus rex had similar forelimbs, but this hypothesis was not confirmed until the first complete Tyrannosaurus rex forelimbs were identified in 1989, belonging to MOR 555 (the "Wankel rex"). The remains of "Sue" also include complete forelimbs. Tyrannosaurus rex arms are very small relative to overall body size, measuring only 1 metre (3.3 ft) long. However, they are not vestigial but instead show large areas for muscle attachment, indicating considerable strength. This was recognized as early as 1906 by Osborn, who speculated that the forelimbs may have been used to grasp a mate during copulation. It has also been suggested that the forelimbs were used to assist the animal in rising from a prone position.[54] Another possibility is that the forelimbs held struggling prey while it was killed by the tyrannosaur's enormous jaws. This hypothesis may be supported by biomechanical analysis.
Tyrannosaurus rex forelimb bones exhibit extremely thick cortical bone, indicating that they were developed to withstand heavy loads. The biceps brachii muscle of a full-grown Tyrannosaurus rex was capable of lifting 199 kilograms (439 lb) by itself; other muscles such as the brachialis would work along with the biceps to make elbow flexion even more powerful. The M. biceps muscle of T. rex was 3.5 times as powerful as the human equivalent. A Tyrannosaurus rex forearm also had a reduced range of motion, with the shoulder and elbow joints allowing only 40 and 45 degrees of motion, respectively. In contrast, the same two joints in Deinonychus allow up to 88 and 130 degrees of motion, respectively, while a human arm can rotate 360 degrees at the shoulder and move through 165 degrees at the elbow. The heavy build of the arm bones, extreme strength of the muscles, and limited range of motion may indicate a system evolved to hold fast despite the stresses of a struggling prey animal. Carpenter and Smith dismissed notions that the forelimbs were useless or that Tyrannosaurus rex was an obligate scavenger.
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