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Alioramus skull steveoc

Alioramus

Alioramus (/ˌælioʊˈreɪməs/; meaning 'different branch') is a genus of tyrannosaurid theropod dinosaurs from the Late Cretaceous period of Asia. It currently contains two species. The type species, A. remotus is known from a partial skull and three foot bones recovered from the Mongolian Nemegt Formation, which was deposited in a humid floodplain about 70 million years ago. These remains were named and described by Soviet paleontologist Sergei Kurzanov in 1976. A second species, A. altai, known from a much more complete skeleton, was named and described by Stephen L. Brusatte and colleagues in 2009. Its relationships to other tyrannosaurid genera are unclear, with some evidence supporting a hypothesis that Alioramus is closely related to the contemporary species Tarbosaurus bataar.

Alioramus were bipedal like all known theropods, and their sharp teeth indicate that they were carnivores. Known specimens were smaller than other tyrannosaurids like Tarbosaurus bataar and Tyrannosaurus rex, but their adult size is difficult to estimate since both Alioramus species are known only from juvenile or sub-adult remains. The recent discovery of Qianzhousaurus indicates that it belongs to a distinct branch of tyrannosaur. The genus Alioramus is characterized by a row of five bony crests along the top of the snout, a greater number of teeth than any other genus of tyrannosaurid, and a lower skull than most other tyrannosaurids.

History of discovery[]

The holotype (PIN 3141/1) of Alioramus is a partial skull associated with three metatarsals. A joint Soviet-Mongolian expedition to the Gobi Desert in the early 1970s found these remains at a locality known as Nogon-Tsav in the Mongolian province of Bayankhongor, Nemegt Formation. Alioramus was named and described by Russian paleontologist Sergei Kurzanov in 1976. Its crests and low skull profile looked so different from other tyrannosaurids that Kurzanov believed his find was far removed from other members of the family. Accordingly, he gave it the generic name Alioramus, derived from the Latin alius ('other') and ramus ('branch'), and the specific name A. remotus, which means 'removed' in Latin. A second species, A. altai, was discovered back in 2009 at the Tsagan Khushu locality also from the Nemegt Formation. However, several faunal differences may suggest that the respective locations of A. remotus and A. altai differ in age. The holotype IGM 100/1844 is a partial skeleton that includes a very complete skull—more so than A. remotus—with partial vertebrae, pelvic girdle and hindlimbs. The name for this species, altai, is in reference to the Altai Mountains.

Description[]

Alioramus remotus was estimated at 5 to 6 m (16 to 20 ft) in length when originally described by Sergei Kurzanov in 1976. In 1988 Paul gave a similar length of 6 m (20 ft) and a weight of 700 kg (1,500 lb). In 2016 Molina-Pérez and Larramendi estimated A. remotus at 5.5 m (18 ft) and 500 kg (1,100 lb), and A. altai at 5 m (16 ft) and 385 kg (849 lb). Kurzanov, however, did not correct for lengthening of the skull by deformation during fossilization, which may indicate a shorter overall body length for this individual. If this specimen is a juvenile, then adult Alioramus would have reached greater lengths, but no confirmed adult specimens are known.

The skull of A. remotus was approximately 45 cm (1.48 ft) long. In general, it is long and low, a shape typical of more basal tyrannosauroids and juveniles of larger tyrannosaurids. The premaxillary bones at the tip of the snout in Alioramus remotus have not been found, but are taller than wide in all tyrannosauroids for which they are known. The nasal bones are fused and ornamented with a row of five irregular bony crests that protrude upwards from the midline, where the nasal bones are sutured together. These crests all measure more than 1 cm (0.39 in) tall.

At the back of the skull there is a protrusion, called the nuchal crest, arising from the fused parietal bones, a feature shared with all tyrannosaurids. In Alioramus, the nuchal crest is greatly thickened, similarly to Tarbosaurus and Tyrannosaurus. Like the rest of the skull, the lower jaw of Alioramus was long and slender, another possible juvenile characteristic. As in Tarbosaurus, a ridge on the outer surface of the angular bone of the lower jaw articulated with the rear of the dentary bone, locking the two bones together and removing much of the flexibility seen in other tyrannosaurids. Other tyrannosaurids had four premaxillary teeth, D-shaped in cross section, on each side. Including 16 or 17 in each maxilla, and 18 in each dentary, Alioramus had 76 or 78 teeth, more than any other tyrannosaurid. The braincase of A. altai was intermediate between the basal theropod and avialan conditions.

The rest of the skeleton of Alioramus remotus is completely unknown except for three metatarsals (bones of the upper foot), but the discovery of A. altai, which is known from substantially more complete remains, has shed light on the anatomy of the genus.

A examination of Qianzhousaurus and its comparisons with both species of Alioramus published in 2022 suggests that both Alioramus species are known from juvenile specimens in different growth stages, and that Qianzhousaurus represents an adult example of the alioramini. The examinations also suggest that the variation seen between the various species is consistent with the growth trends seen in other tyrannosaurid genera, though specimens that could constitute a full growth series from infant to adult for each species have not been recovered for any of these tyrannosaurs. One part of the growth series across all specimens in this study was discovered to remain unique to alioramin tyrannosaurs; the rugose process of the jugal starts small and conical in early life, but becomes massive and indistinct as the animals grow. This same study also suggests Alioramins did not undergo a secondary metamorphosis from slender juveniles to robust adults like other tyrannosaurs such as Tarbosaurus and Tyrannosaurus did, but maintained a unique physiology better suited to pursuit of fast, small prey.

Classification and systematics[]

Paleontologists have long classified Alioramus within the superfamily Tyrannosauroidea, but because its remains were for many years poorly known, a more precise classification had remained elusive until the discovery of A. altai. A cladistic analysis published in 2003 found Alioramus could be further classified into the family Tyrannosauridae and the subfamily Tyrannosaurinae, alongside Tyrannosaurus, Tarbosaurus and Daspletosaurus. A 2004 study supported this result but suggested it was equally probable that Alioramus belonged outside the family Tyrannosauridae entirely, with its supposed juvenile characters actually reflecting a more basal position within Tyrannosauroidea. Another study omitted Alioramus altogether due to the only specimen's fragmentary nature. The description of A. altai in 2009 confirmed the placement of the genus within the Tyrannosaurinae.

Tarbosaurus and Alioramus shared several skull features, including a locking mechanism in the lower jaw between the dentary and angular bones, and both lacked the prong of the nasal bones which connected to the lacrimal bones in all other tyrannosaurids except adult Daspletosaurus. The two genera may be closely related, representing an Asian branch of the Tyrannosauridae. Some specimens of Tarbosaurus have a row of bumps on the nasal bones like those of Alioramus, although much lower. The long and low shape of the only known Alioramus remotus skull indicated that it was immature when it died and might even have been a juvenile Tarbosaurus, which lived in the same time and place. The more prominent nasal crests and much higher tooth count of Alioramus, however, suggested it was a separate taxon, even if it is known only from juvenile remains, confirmed by the discovery of A. altai. Specimens identified as immature Tarbosaurus have the same tooth count as adults.

Recently a cladogram has been published finding Alioramus just outside Tyrannosauridae. Below is the cladogram by Loewen (2013).



Dilong paradoxus




Eotyrannus lengi




Bagaraatan ostromi




Raptorex kriegsteini




Dryptosaurus aquilunguis





Alectrosaurus olseni



Xiongguanlong baimoensis





Appalachiosaurus montgomeriensis





Alioramus altai



Alioramus remotus




Tyrannosauridae










Paleoecology[]

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