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Daspletosaurus (or das-PLEET-o-SAWR-us; meaning 'frightful lizard') is a genus of tyrannosaurid theropod dinosaur that lived in western North America between 80 and 73 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, Montana and New Mexico await description. Including these undescribed species, Daspletosaurus is 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-ton 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's 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 wasn't the largest tyrannosaurid. Adults could reach a length of 8–9 meters (26–30 ft) from snout to tail. Mass estimates have centered around 2.5 tonnes (2.75 short tons) but have ranged between 1.8 tonnes (2 tons) and 3.8 tonnes (4.1 tons). Daspletosaurus had a massive skull that could reach more than 1 meter (3.3 ft) in length. 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) didn't 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. Daspletosaurus is usually considered to be closely related to Tyrannosaurus rex, or even a direct ancestor through anagenesis. Gregory Paul reassigned D. torosus to the genus Tyrannosaurus, creating the new combination Tyrannosaurus torosus, 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. The type species is D. torosus, which is Latin for 'muscular' or 'brawny.'

Unnamed species

Two or three additional species have been assigned to the genus Daspletosaurus over the years, although as of 2007 none of these species have received a proper description or scientific name. In the meantime, all are assigned to Daspletosaurus sp. although this doesn't imply that they all are the same species. Many later authors have reassigned this specimen, along with a few others from New Mexico, to yet another unnamed species of Daspletosaurus. However, recent unpublished research has suggested that this species, from the Hunter Wash Member of the Kirtland Formation, is actually an early tyrannosaurid, perhaps related to Appalachiosaurus. There is currently disagreement over the age of the Kirtland Formation, with some workers claiming a late Campanian age, while others suggest a younger age in the early Maastrichtian stage.
   In 1992, Jack Horner and colleagues published an extremely preliminary report of a tyrannosaurid from the upper parts of the Campanian Two Medicine Formation in Montana, which was interpreted as a transitional species between Daspletosaurus and the later Tyrannosaurus. The remains of at least three more Daspletosaurus have also been described in a Two Medicine bonebed. These specimens have not been described in detail, but Currie believes all of the Two Medicine material represents an as-yet-unnamed third species of Daspletosaurus. Several studies have attempted to explain niche differentiation in Daspletosaurus and Gorgosaurus.
   Dale Russell hypothesized that the more lightly built and more common Gorgosaurus may have preyed on the abundant hadrosaurs of the time, while the more robust and less common Daspletosaurus may have specialized on the less prevalent but better-defended ceratopsids, which may have been more difficult to hunt.
   Other authors have suggested that competition was limited by geographical separation. Unlike some other groups of dinosaurs, there appears to be no correlation with distance from the sea. Neither Daspletosaurus nor Gorgosaurus was more common at higher or lower elevations than the other.
   Evidence that Daspletosaurus lived in social groups comes from a bonebed found in the Two Medicine Formation of Montana. The bonebed includes the remains of three Daspletosaurus, including a large adult, a small juvenile, and another individual of intermediate size. At least five hadrosaurs are preserved at the same location. Geologic evidence indicates that the remains were not brought together by river currents but that all of the animals were buried simultaneously at the same location. The hadrosaur remains are scattered and bear numerous marks from tyrannosaur teeth, indicating that the Daspletosaurus were feeding on the hadrosaurs at the time of death. The cause of death is unknown. Currie speculates that the daspletosaurs formed a pack, although this can't be stated with certainty. Brian Roach and Daniel Brinkman have suggested that Daspletosaurus social interaction would have more closely resembled the modern Komodo dragon, where non-cooperative individuals mob carcasses, frequently attacking and even cannibalizing each other in the process.

Life history

Paleontologist Gregory Erickson and colleagues have studied the growth and life history of tyrannosaurids. Analysis of bone histology can determine the age of a specimen when it died. Growth rates can be examined when the age of various individuals are plotted against their size on a graph. Erickson has shown that after a long time as juveniles, tyrannosaurs underwent tremendous growth spurts for about four years midway through their lives. After the rapid growth phase ended with sexual maturity, growth slowed down considerably in adult animals. Erickson only examined Daspletosaurus from the Dinosaur Park Formation, but these specimens show the same pattern. Compared to albertosaurines, Daspletosaurus showed a faster growth rate during the rapid growth period due to its higher adult weight. The maximum growth rate in Daspletosaurus was 180 kilograms (400 lb) per year, based on a mass estimate of 1800 kilograms (2 tons) in adults. Other authors have suggested higher adult weights for Daspletosaurus; this would change the magnitude of the growth rate but not the overall pattern.

Paleoecology

All known Daspletosaurus fossils have been found in formations dating to the middle to late Campanian stage of the Late Cretaceous Period, between 80 and 73 million years ago. Since the middle of the Cretaceous, North America had been divided in half by the Western Interior Seaway, with much of Montana and Alberta below the surface. However, the uplift of the Rocky Mountains in the Laramide Orogeny to the west, which began during the time of Daspletosaurus, forced the seaway to retreat eastwards and southwards. Rivers flowed down from the mountains and drained into the seaway, carrying sediment along with them that formed the Two Medicine Formation, the Judith River Group, and other sedimentary formations in the region. About 73 million years ago, the seaway began to advance westwards and northwards again, and the entire region was covered by the Bearpaw Sea, represented throughout the western United States and Canada by the massive Bearpaw Shale. Daspletosaurus lived in a vast floodplain along the western shore of the interior seaway. Large rivers watered the land, occasionally flooding and blanketing the region with new sediment. When water was plentiful, the region could support a great deal of plant and animal life, but periodic droughts also struck the region, resulting in mass mortality as preserved in the many bonebed deposits found in Two Medicine and Judith River sediments, including the Daspletosaurus bonebed. Similar conditions exist today in East Africa. Volcanic eruptions from the west periodically blanketed the region with ash, also resulting in large-scale mortality, while simultaneously enriching the soil for future plant growth. It is these ash beds that allow precise radiometric dating as well. Fluctuating sea levels also resulted in a variety of other environments at different times and places within the Judith River Group, including offshore and nearshore marine habitats, coastal wetlands, deltas and lagoons, in addition to the inland floodplains.

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