Giganotosaurus ( ) is a genus of large theropod dinosaur that lived in what is now Argentina, during the Albian age of the Early Cretaceous period, approximately 111 to 106 million years ago. The holotype specimen was discovered in the Candeleros Formation of Patagonia in 1993 and is almost 70% complete. The animal was named Giganotosaurus carolinii in 1995; the genus name translates to "giant southern lizard", and the specific name honors the discoverer, Ruben Carolini. A dentary bone, a tooth, and some tracks, discovered before the holotype, were later assigned to this animal. The genus attracted much interest and became part of a scientific debate about the maximum sizes of theropod dinosaurs.
Giganotosaurus was one of the largest known terrestrial carnivores, but the exact size has been hard to determine due to the incompleteness of the remains found so far. Estimates for the most complete specimen range from a length of , a skull in length, and a weight of . The dentary bone that belonged to a supposedly larger individual has been used to extrapolate a length of . Some researchers have found the animal to be larger than Tyrannosaurus, which has historically been considered the largest theropod, while others have found them to be roughly equal in size and the largest size estimates for Giganotosaurus exaggerated. The skull was low, with rugose (rough and wrinkled) nasal bones and a ridge-like crest on the lacrimal bone in front of the eye. The front of the lower jaw was flattened and had a downward-projecting process (or "chin") at the tip. The teeth were compressed sideways and had serrations. The neck was strong and the pectoral girdle proportionally small.
Part of the family Carcharodontosauridae, Giganotosaurus is one of the most completely known members of the group, which includes other very large theropods, such as the closely related Mapusaurus, Tyrannotitan and Carcharodontosaurus. Giganotosaurus is thought to have been homeothermic (a type of "warm-bloodedness"), with a metabolism between that of a mammal and a reptile, which would have enabled fast growth. It would have been capable of closing its jaws quickly, capturing and bringing down prey by delivering powerful bites. The "chin" may have helped in resisting stress when a bite was delivered against prey. Giganotosaurus is thought to have been the apex predator of its ecosystem, and it may have fed on juvenile sauropod dinosaurs.
Discovery
In 1993, the amateur Argentine fossil hunter discovered the tibia (lower leg bone) of a theropod dinosaur while driving a dune buggy in the badlands near Villa El Chocón, in the Neuquén province of Patagonia, Argentina. Specialists from the National University of Comahue were sent to excavate the specimen after being notified of the find. The discovery was announced by the paleontologists Rodolfo Coria and Leonardo Salgado at a Society of Vertebrate Paleontology meeting in 1994, where science writer Don Lessem offered to fund the excavation, after having been impressed by a photo of the leg-bone. The partial skull was scattered over an area of about 10 m2 (110 sq ft), and the postcranial skeleton was disarticulated. The specimen preserved almost 70% of the skeleton, and included most of the vertebral column, the pectoral and pelvic girdles, the femora, and the left tibia and fibula.
In 1995, this specimen was preliminarily described by Coria and Salgado, who made it the holotype of the new genus and species Giganotosaurus carolinii (parts of the skeleton were still encased in plaster at this time). The generic name is derived from the Ancient Greek words gigas/ (meaning "giant"), notos/ (meaning "austral/southern", in reference to its provenance) and -sauros/- (meaning "lizard"). The specific name honors Carolini, the discoverer. The holotype skeleton is now housed in the Ernesto Bachmann Paleontological Museum (where it is catalogued as specimen MUCPv-Ch1) in Villa El Chocón, which was inaugurated in 1995 at the request of Carolini. The specimen is the main exhibition at the museum, and is placed on the sandy floor of a room devoted to the animal, along with tools used by paleontologists during the excavation. A mounted reconstruction of the skeleton is exhibited in an adjacent room.
One of the features of theropod dinosaurs that has attracted most scientific interest is the fact that the group includes the largest terrestrial predators of the Mesozoic Era. This interest began with the discovery of one of the first known dinosaurs, Megalosaurus, named in 1824 for its large size. More than half a century later in 1905, Tyrannosaurus was named, and it remained the largest known theropod dinosaur for 90 years, though other large theropods were also known. The discussion of which theropod was the largest was revived in the 1990s by new discoveries in Africa and South America. In their original description, Coria and Salgado considered Giganotosaurus at least the largest theropod dinosaur from the Southern Hemisphere, and perhaps the largest in the world. They conceded that comparison with Tyrannosaurus was difficult due to the disarticulated state of the cranial bones of Giganotosaurus, but noted that at , the femur of Giganotosaurus was 5 cm (2 in) longer than that of "Sue", the largest known Tyrannosaurus specimen, and that the bones of Giganotosaurus appeared to be more robust, indicating a heavier animal. They estimated the skull to have been about 1.53 m (5 ft) long, and the whole animal to have been 12.5 m (41 ft) long, with a weight of about .
In 1996, the paleontologist Paul Sereno and colleagues described a new skull of the related genus Carcharodontosaurus from Morocco, a theropod described in 1927 but previously known only from fragmentary remains. They estimated the skull to have been long, similar to Giganotosaurus, but perhaps exceeding that of the Tyrannosaurus "Sue", with a 1.53 m (5 ft) long skull. They also pointed out that carcharodontosaurs appear to have had the proportionally largest skulls, but that Tyrannosaurus appears to have had longer hind limbs. In an interview for a 1995 article entitled "new beast usurps T. rex as king carnivore", Sereno noted that these newly discovered theropods from South America and Africa competed with Tyrannosaurus as the largest predators, and would help in the understanding of Late Cretaceous dinosaur faunas, which had otherwise been very "North America-centric". In the same issue of the journal in which Carcharodontosaurus was described, the paleontologist Philip J. Currie cautioned that it was yet to be determined which of the two animals were larger, and that the size of an animal is less interesting to paleontologists than, for example, adaptations, relationships, and distribution. He also found it remarkable that the two animals were found within a year of each other, and were closely related, in spite of being found on different continents.
In a 1997 interview, Coria estimated Giganotosaurus to have been 13.7 (45 ft) to 14.3 (47 ft) m long and weighing based on new material, larger than Carcharodontosaurus. Sereno countered that it would be difficult to determine a size range for a species based on few, incomplete specimens, and both paleontologists agreed that other aspects of these dinosaurs were more important than settling the "size contest". In 1998, the paleontologist Jorge O. Calvo and Coria assigned a partial left dentary bone (part of the lower jaw) containing some teeth (MUCPv-95) to Giganotosaurus. It had been collected by Calvo near Los Candeleros in 1988 (found in 1987), who described it briefly in 1989, while noting it may have belonged to a new theropod taxon. Calvo and Coria found the dentary to be identical to that of the holotype, though 8% larger at 62 cm (24 in). Though the rear part of it is incomplete, they proposed that the skull of the holotype specimen would have been long, and estimated the skull of the larger specimen to have been long, the longest skull of any theropod.
In 1999, Calvo assigned an incomplete tooth, (MUCPv-52), to Giganotosaurus; this specimen was discovered near Lake Ezequiel Ramos Mexia in 1987 by A. Delgado, and is therefore the first known fossil of the genus. Calvo further suggested that some theropod trackways and isolated tracks (which he made the basis of the ichnotaxon Abelichnus astigarrae in 1991) belonged to Giganotosaurus, based on their large size. The largest tracks are long with a pace of , and the smallest is long with a pace of . The tracks are tridactyl (three-toed) and have large and coarse digits, with prominent claw impressions. Impressions of the digits occupy most of the track-length, and one track has a thin heel. Though the tracks were found in a higher stratigraphic level than the main fossils of Giganotosaurus, they were from the same strata as the single tooth and some sauropod dinosaurs that are also known from the same strata as Giganotosaurus.
Continued size estimations
In 2001, the physician-scientist Frank Seebacher proposed a new polynomial method of calculating body-mass estimates for dinosaurs (using body-length, depth, and width), and found Giganotosaurus to have weighed (based on the original length estimate). In their 2002 description of the braincase of Giganotosaurus, Coria and Currie gave a length estimate of for the holotype skull, and calculated a weight of by extrapolating from the circumference of the femur-shaft. This resulted in an encephalization quotient (a measure of relative brain size) of 1.9. In 2004, the paleontologist Gerardo V. Mazzetta and colleagues pointed out that though the femur of the Giganotosaurus holotype was larger than that of "Sue", the tibia was shorter at . They found the holotype specimen to have been equal to Tyrannosaurus in size at (marginally smaller than "Sue"), but that the larger dentary might have represented an animal of , if geometrically similar to the holotype specimen. By using multivariate regression equations, these authors also suggested an alternative weight of for the holotype and for the larger specimen, and that the latter was therefore the largest known terrestrial carnivore.
In 2005, the paleontologist Cristiano Dal Sasso and colleagues described new skull material (a snout) of Spinosaurus (the original fossils of which were also destroyed during World War II), and concluded this dinosaur would have been long with a weight , exceeding the maximum size of all other theropods. In 2006, Coria and Currie described the large theropod Mapusaurus from Patagonia; it was closely related to Giganotosaurus and of approximately the same size. In 2007, the paleontologists François Therrien and Donald M. Henderson found that Giganotosaurus would have approached in length and in weight, while Carcharodontosaurus would have approached in length and in weight (surpassing Tyrannosaurus), and estimated the Giganotosaurus holotype skull to have been long. They cautioned that these measurements depended on whether the incomplete skulls of these animals had been reconstructed correctly, and that more complete specimens were needed for more accurate estimates. They also found that Dal Sasso and colleagues' reconstruction of Spinosaurus was too large, and instead estimated it to have been long, weighing , and possibly as low as in length and in weight. They concluded that these dinosaurs had reached the upper biomechanical size limit attainable by a strictly bipedal animal. In 2010, the paleontologist Gregory S. Paul suggested that the skulls of carcharodontosaurs had been reconstructed as too long in general.