Acrocanthosaurus

Acrocanthosaurus (meaning 'high-spined lizard') is a genus of theropod dinosaur that existed in what is now North America during the Aptian and early Albian stages of the Early Cretaceous. Like most dinosaur genera, Acrocanthosaurus contains only a single species, A. atokensis. Its fossil remains are found mainly in the U.S. states of Oklahoma and Texas, although teeth attributed to Acrocanthosaurus have been found as far east as Maryland. Acrocanthosaurus was a bipedal predator. As the name suggests, it is best known for the high neural spines on many of its vertebrae, which most likely supported a ridge of muscle over the animal's neck, back and hips. Acrocanthosaurus was one of the largest theropods, approaching 12 meters (40 ft) in length, and weighing up to about 2.40 metric tons (2.65 short tons). Large theropod footprints discovered in Texas may have been made by Acrocanthosaurus, although there is no direct association with skeletal remains.

Recent discoveries have elucidated many details of its anatomy, allowing for specialized studies focusing on its brain structure and forelimb function. However, there is still debate over its evolutionary relationships, with some scientists classifying it as an allosaurid, and others as a carcharodontosaurid. Acrocanthosaurus was the largest theropod in its ecosystem and likely an apex predator which possibly preyed on large sauropods and ornithopods.

Description
Although slightly smaller than gigantic relatives like Giganotosaurus, Acrocanthosaurus was still among the largest theropods ever to exist. The longest known individual measured 11.5 meters (38 ft) from snout to tail tip and weighed an estimated 2400 kilograms (5300 lb). Its skull alone was nearly 1.3 meters (4.3 ft) in length.[1]

The skull of Acrocanthosaurus, like most other allosauroids, was long, low and narrow. The weight-reducing opening in front of the eye socket (antorbital fenestra) was quite large, more than a quarter of the length of the skull and two-thirds of its height. The outside surface of the maxilla (upper jaw bone) and the upper surface of the nasal bone on the roof of the snout were not nearly as rough-textured as those of Giganotosaurus or Carcharodontosaurus. Long, low ridges arose from the nasal bones, running along each side of the snout from the nostril back to the eye, where they continued onto the lacrimal bones.[1] This is a characteristic feature of all allosauroids.[2] Unlike Allosaurus, there was no prominent crest on the lacrimal bone in front of the eye. The lacrimal and postorbital bones met to form a thick brow over the eye, as seen in carcharodontosaurids and the unrelated abelisaurids. Nineteen curved, serrated teeth lined each side of the upper jaw, but a tooth count for the lower jaw has not been published. Acrocanthosaurus teeth were wider than those of Carcharodontosaurus and did not have the wrinkled texture that characterized the carcharodontosaurids. The dentary (tooth-bearing lower jaw bone) was squared off at the front edge, as in Giganotosaurus, and shallow, while the rest of the jaw behind it became very deep. Acrocanthosaurus and Giganotosaurus shared a thick horizontal ridge on the outside surface of the surangular bone of the lower jaw, underneath the articulation with the skull.

The most notable feature of Acrocanthosaurus was its row of tall neural spines, located on the vertebrae of the neck, back, hips and upper tail, which could be more than 2.5 times the height of the vertebrae from which they extended.[3] Other dinosaurs also had high spines on the back, sometimes much higher than those of Acrocanthosaurus. For instance, the unrelated Spinosaurus had spines nearly 2 meters (6.5 ft) tall, about 11 times taller than the bodies of its vertebrae.[4] Rather than supporting a skin 'sail' as seen in Spinosaurus, the lower spines of Acrocanthosaurus had attachments for powerful muscles like those of modern bison, probably forming a tall, thick ridge down its back.[3] The function of the spines remains unknown, although they may have been involved in communication, fat storage, or temperature control. All of its cervical (neck) and dorsal (back) vertebrae had prominent depressions (pleurocoels) on the sides, while the caudal (tail) vertebrae bore smaller ones. This is more similar to carcharodontosaurids than to Allosaurus.[5]

Aside from its vertebrae, Acrocanthosaurus had a typical allosauroid skeleton. Acrocanthosaurus was bipedal, with a long, heavy tail counterbalancing the head and body, maintaining its center of gravity over its hips. Its forelimbs were relatively shorter and more robust than those of Allosaurus but were otherwise similar: each hand bore three clawed digits. Unlike many smaller fast-running dinosaurs, its femur was longer than its tibia and metatarsals,[1][5] suggesting that Acrocanthosaurus was not a fast runner.[6] Unsurprisingly, the hind leg bones of Acrocanthosaurus were proportionally more robust than its smaller relative Allosaurus. Its feet had four digits each, although as is typical for theropods, the first was much smaller than the rest and did not make contact with the ground.

Classification and systematics
Acrocanthosaurus is classified in the superfamily Allosauroidea within the infraorder Tetanurae. This superfamily is characterized by paired ridges on the nasal and lacrimal bones on top of the snout and tall neural spines on the neck vertebrae, among other features.[2] It was originally placed in the family Allosauridae with Allosaurus,[3] an arrangement also supported by some more recent studies.[1] Other scientists classify it as a member of the related family Carcharodontosauridae.[2][5][7]

At the time of its discovery, Acrocanthosaurus and most other large theropods were known from only fragmentary remains, leading to highly variable classifications for this genus. J. Willis Stovall and Wann Langston, Jr. first assigned it to the "Antrodemidae," the equivalent of Allosauridae, but it was transferred to the taxonomic wastebasket Megalosauridae by Alfred Sherwood Romer in 1956.[8] To other authors, the long spines on its vertebrae suggested a relationship with Spinosaurus.[9][10] This interpretation of Acrocanthosaurus as a spinosaurid persisted into the 1980s,[11] and was repeated in the semi-technical dinosaur books of the time.[12][13]

Tall spined vertebrae from the Early Cretaceous of England were once considered to be very similar to those of Acrocanthosaurus,[14] and in 1988 Gregory S. Paul named them as a second species of the genus, A. altispinax.[15] These bones were originally assigned to Altispinax, an English theropod otherwise known only from teeth, and this assignment led to at least one author proposing that Altispinax itself was a synonym of Acrocanthosaurus.[14] These vertebrae were later assigned to the new genus Becklespinax, separate from both Acrocanthosaurus and Altispinax.[16]

If Acrocanthosaurus was an allosaurid, it would have been most closely related to Jurassic genera like Allosaurus and Saurophaganax.[1] Those cladistic analyses which find it to be a carcharodontosaurid usually place it in a basal position relative to the African Carcharodontosaurus and Giganotosaurus from South America.[2][5]Neovenator, discovered in England, is often considered an even more basal carcharodontosaurid.[2][5][6] This suggests that the family originated in Europe and then dispersed into the southern continents (at the time united as the supercontinent Gondwana). If Acrocanthosaurus was a carcharodontosaurid, then dispersal would also have occurred into North America.[5] All known carcharodontosaurids lived during the early to middle Cretaceous Period.