Giganotosaurus

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Not to be confused with Gigantosaurus.

Giganotosaurus
Temporal range:
Ma
↓
Reconstructed skeleton, possibly with a proportionally too long skull, Fernbank Museum of Natural History
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Clade: Saurischia
Clade: Theropoda
Family: †Carcharodontosauridae
Tribe: †
Giganotosaurini
Genus: †Giganotosaurus
Coria & Salgado, 1995[1]
Species:
†G. carolinii
Binomial name
†Giganotosaurus carolinii
Coria & Salgado, 1995

Giganotosaurus (

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

pectoral girdle
proportionally small.

Part of the family

sauropod
dinosaurs.

Discovery

Holotype skeleton with reconstructed skull, arm, and feet, on the floor in Ernesto Bachmann Paleontological Museum

In 1993, the amateur Argentine

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.[6][1]

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/γίγας (meaning "giant"), notos/νότος (meaning "austral/southern", in reference to its provenance) and -sauros/-σαύρος (meaning "lizard"). The specific name honors Carolini, the discoverer.[1][7] 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.[4][8]

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

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 1.43 m (4.7 ft), 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 6 to 8 t (6.6 to 8.8 short tons).[1]

Reconstructed skeleton, EBPM

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 (much of its fossils were destroyed in World War II). They estimated the skull to have been 1.60 m (5 ft) 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.[9] 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".[10] 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.[11]

In a 1997 interview, Coria estimated Giganotosaurus to have been 13.7 (45 ft) to 14.3 (47 ft) m long and weighing 8 to 10 t (8.8 to 11.0 short tons) 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".

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 1.80 m (6 ft) long, and estimated the skull of the larger specimen to have been 1.95 m (6.4 ft) long, the longest skull of any theropod.[13][14][15]

In 1999, Calvo referred 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

sauropod dinosaurs that are also known from the same strata as Giganotosaurus.[14]

Continued size estimations

Skull reconstructed in Japan from the side and front; it is possibly reconstructed as too long

In 2001, the

regression equations, these authors also suggested an alternative weight of 6.5 t (7.2 short tons) for the holotype and 8.2 t (9.0 short tons) for the larger specimen, and that the latter was therefore the largest known terrestrial carnivore.[17]

Full size model nicknamed "Lady Giga", Frankfurt Hauptbahnhof.

In 2005, the paleontologist

bipedal animal.[20] In 2010, the paleontologist Gregory S. Paul suggested that the skulls of carcharodontosaurs had been reconstructed as too long in general.[21]

In 2012, the paleontologist Matthew T. Carrano and colleagues noted that though Giganotosaurus had received much attention due to its enormous size, and in spite of the holotype being relatively complete, it had not yet been described in detail, apart from the braincase. They pointed out that many contacts between skull bones were not preserved, which lead to the total length of the skull being ambiguous. They found instead that the skulls of Giganotosaurus and Carcharodontosaurus were exactly the same size as that of Tyrannosaurus. They also measured the femur of the Giganotosaurus holotype to be 1.365 m (4 ft) long, in contrast to the original measurement, and proposed that the body mass would have been smaller overall.[22] In 2013, the paleontologist Scott Hartman published a Graphic Double Integration mass estimate (based on drawn skeletal reconstructions) on his blog, wherein he found Tyrannosaurus ("Sue") to have been larger than Giganotosaurus overall. He estimated the Giganotosaurus holotype to have weighed 6.8 t (7.5 short tons), and the larger specimen 8.2 t (9.0 short tons). Tyrannosaurus was estimated to have weighed 8.4 t (9.3 short tons), and Hartman noted that it had a wider torso, though the two seemed similar in side view. He also pointed out that the Giganotosaurus dentary that was supposedly 8% larger than that of the holotype specimen would rather have been 6.5% larger, or could simply have belonged to a similarly sized animal with a more robust dentary. He conceded that with only one good Giganotosaurus specimen known, it is possible that larger individuals will be found, as it took most of a century to find "Sue" after Tyrannosaurus was discovered.[23]

Comparison between the estimated sizes of the holotype (light green) and assigned specimen (dark green)

In 2014, the paleontologist Nizar Ibrahim and colleagues estimated the length of Spinosaurus to have been over 15 m (49 ft), by extrapolating from a new specimen scaled up to match the snout described by Dal Sasso and colleagues.[24] This would make Spinosaurus the largest known carnivorous dinosaur.[25] In 2019, the paleontologist W. Scott Persons and colleagues described a Tyrannosaurus specimen (nicknamed "Scotty"), and estimated it to be more massive than other giant theropods, but cautioned that the femoral proportions of the carcharodontosaurids Giganotosaurus and Tyrannotitan indicated a body mass larger than other adult Tyrannosaurus. They noted that these theropods were known by far fewer specimens than Tyrannosaurus, and that future finds may reveal specimens larger than "Scotty", as indicated by the large Giganotosaurus dentary. While "Scotty" had the greatest femoral circumference, the femoral length of Giganotosaurus was about 10% longer, but the authors stated it was difficult to compare proportions between large theropod clades.[26][27]

In 2021, the paleontologist Matías Reolid and colleagues compiled various mass estimates of theropods (including Giganotosaurus) to calculate the average, but did not include Therrien and Henderson's 2007 estimates of Carnotaurus and Giganotosaurus, considering them outliers. This resulted in a body mass range for Giganotosaurus between 5.5 and 8.5 t (6.1 and 9.4 short tons), with an average of 6.75 t (7.44 short tons). They also applied the skull length and body length ratio proposed by Therrien and Henderson and reconstructed various digital 3D models of theropods to measure body mass distribution and volume, resulting in the mass of a 13 m (43 ft) long Giganotosaurus up to 7.2 t (7.9 short tons). These researchers found the estimates consistent with the values proposed by previous studies.

Meraxes, which has the most completely known Carcharodontosaurine skull, with an estimated length of 1.27 m (4.2 ft). Extrapolating from that skull, they estimated the skull of Giganotosaurus to have been 1.634 m (5.36 ft) long, making it one of the largest known theropod skulls.[29] Henderson suggested in 2023 that there was a close relation between the dimensions of the pelvic area and body size in theropods, allowing size estimates for incomplete specimens. Based on this idea, he found Giganotosaurus to have been 12.5 m (41 ft) long, identical to the estimate proposed in the 1995 description.[30]

Description

Size (green) compared to that of other large theropods

Giganotosaurus is thought to have been one of the largest theropod dinosaurs, but the incompleteness of its remains have made it difficult to estimate its size reliably. It is therefore impossible to determine with certainty whether it was larger than Tyrannosaurus, for example, which has been considered the largest theropod historically. Different size estimates have been reached by several researchers, based on various methods, and depending on how the missing parts of the skeleton have been reconstructed. Length estimates for the holotype specimen have varied between 12 and 13 m (39 and 43 ft), with a skull between 1.53 and 1.80 m (5.0 and 5.9 ft) long, a femur (thigh bone) between 1.365 and 1.43 m (4.48 and 4.69 ft) long, and a weight between 4.2 and 13.8 t (4.6 and 15.2 short tons).[6][1][13][20] Fusion of sutures (joints) in the braincase indicates the holotype specimen was a mature individual.[6] A second specimen, consisting of a dentary bone from a supposedly larger individual, has been used to extrapolate a length of 13.2 m (43 ft), a skull 1.95 m (6.4 ft) long, and a weight of 8.2 t (9.0 short tons). Some writers have considered the largest size estimates for both specimens exaggerated.[13][31][23][22] Giganotosaurus has been compared to an oversized version of the well-known genus Allosaurus.[10]

Skull

Diagram showing the known skull remains of the holotype (left) and the assigned specimen

Though incompletely known, the skull of Giganotosaurus appears to have been low. The

foramina (holes) on the inner side.[1][19]

Partial holotype skull (white parts are reconstructed) and teeth, EBPM

The

tyrannosaurs.[6] A latex endocast of the brain cavity of Giganotosaurus showed that the brain was similar to that of the related genus Carcharodontosaurus, but larger. The endocast was 29 mm (1 in) long, 64 mm (3 in) wide, and had a volume of 275 ml (9.7 imp fl oz).[32]

The dentary of the lower jaw expanded in height towards the front (by the

interdental plates, where each tooth had a foramen. The Meckelian groove ran along the lower border. The curvature of the dentary shows that the mouth of Giganotosaurus would have been wide. It is possible that each dentary had twelve alveoli (tooth sockets). Most of the alveoli were about 3.5 cm (1.3 in) long from front to back. The teeth of the dentary were of similar shape and size, except for the first one, which was smaller. The teeth were compressed sideways, were oval in cross-section, and had serrations at the front and back borders, which is typical of theropods.[13][33] The teeth were sigmoid-shaped when seen in front and back view.[34] One tooth had nine to twelve serrations per mm (0.039 in).[14] The side teeth of Giganotosaurus had curved ridges of enamel, and the largest teeth in the premaxilla (front of the upper jaw) had pronounced wrinkles (with their highest relief near the serrations).[35]

Postcranial skeleton

Life restoration

The neck of Giganotosaurus was strong, and the

triceps muscle. The coracoid was small and hook-shaped.[1]

The

pubic foot was pronounced and shorter at the front than behind. The ischium was straight and expanded hindwards, ending in a lobe-shape. The femur was sigmoid-shaped, and had a very robust, upwards pointing head, with a deep sulcus (groove). The lesser trochanter of the femoral head was wing-like, and placed below the greater trochanter, which was short. The fourth trochanter was large and projected backwards. The tibia of the lower leg was expanded at the upper end, its articular facet (where it articulated with the femur) was wide, and its shaft was compressed from front to back.[1]

Classification

Coria and Salgado originally found Giganotosaurus to group more closely with the theropod

synapomorphies) in the legs, skull, and pelvis. Other features showed that it was outside the more derived (or "advanced") clade Coelurosauria.[1] In 1996, Sereno and colleagues found Giganotosaurus, Carcharodontosaurus, and Acrocanthosaurus to be closely related within the superfamily Allosauroidea, and grouped them in the family Carcharodontosauridae. Features shared between these genera include the lacrimal and postorbital bones forming a broad "shelf" over the orbit, and the squared front end of the lower jaw.[9]

As more carcharodontosaurids were discovered, their interrelationships became clearer. The group was defined as all allosauroids closer to Carcharodontosaurus than Allosaurus or

Giganotosaurini.[37] In 2010, Paul listed Giganotosaurus as "Giganotosaurus (or Carcharodontosaurus) carolinii" without elaboration.[21] Giganotosaurus is one of the most complete and informative members of Carcharodontosauridae.[36]

The following cladogram shows the placement of Giganotosaurus within Carcharodontosauridae according to Sebastián Apesteguía et al., 2016:[38]

Natural History Museum, Helsinki

Allosaurus

Carcharodontosauria

Neovenatoridae

Carcharodontosauridae

Concavenator

Shaochilong

Carcharodontosaurinae

Carcharodontosaurus saharicus

Carcharodontosaurus iguidensis

Giganotosaurini

Tyrannotitan

Mapusaurus

Giganotosaurus

Evolution

Coria and Salgado suggested that the

vicariance, when Gondwana broke up during the Aptian–Albian ages of the Early Cretaceous.[34]

Paleobiology

Restoration of a walking individual

In 1999, the paleontologist Reese E. Barrick and the geologist William J. Showers found that the bones of Giganotosaurus and Tyrannosaurus had very similar

warm-bloodedness"). The metabolism of an 8 t (8.8 short tons) Giganotosaurus would be comparable to that of a 1 t (1.1 short tons) mammalian carnivore, and would have supported rapid growth.[40]

In 2001, the physicist Rudemar Ernesto Blanco and Mazzetta evaluated the

kinematic limit of the running speed to be 14 m/s (50 km/h; 31 mph). They also found comparison between the running capability of Giganotosaurus and birds like the ostrich based on the strength of their leg-bones to be of limited value, since theropods, unlike birds, had heavy tails to counterbalance their weight.[41]

A 2017 biomechanical study of the running ability of Tyrannosaurus by the biologist William I. Sellers and colleagues suggested that skeletal loads were too great to have allowed adult individuals to run. The relatively long limbs, which were long argued to indicate good running ability, would instead have mechanically limited it to walking gaits, and it would therefore not have been a high-speed

pursuit predator. They suggested that these findings would also apply to other long-limbed giant theropods such as Giganotosaurus, Mapusaurus, and Acrocanthosaurus.[42]

Feeding

sauropod Limaysaurus, Hungarian Natural History Museum

In 2002, Coria and Currie found that various features of the rear part of the skull (such as the frontwards slope of the occiput and low and wide occipital condyle) indicate that Giganotosaurus would have had a good capability of moving the skull sideways in relation to the front neck vertebrae. These features may also have been related to the increased mass and length of the jaw muscles; the jaw articulation of Giganotosaurus and other carcharodontosaurids was moved hindwards to increase the length of the jaw musculature, enabling faster closure of the jaws, whereas tyrannosaurs increased the mass of the lower jaw musculature, to increase the power of their bite.[6]

In 2005 Therrien and colleagues estimated the relative

torsional stress and crushing bones. Estimates in absolute values like newtons were impossible. The bite force of Giganotosaurus was weaker than that of Tyrannosaurus, and the force decreased hindwards along the tooth row. The lower jaws were adapted for slicing bites, and it probably captured and manipulated prey with the front part of the jaws. These authors suggested that Giganotosaurus and other allosaurs may have been generalized predators that fed on a wide spectrum of prey smaller than themselves, such as juvenile sauropods. The ventral process (or "chin") of the lower jaw may have been an adaptation for resisting tensile stress when the powerful bite was delivered with the front of the jaws against the prey.[43]

The first known fossils of the closely related Mapusaurus were found in a

National Geographic article, Coria stated that the bonebed was probably the result of a catastrophic event and that the presence of mainly medium-sized individuals, with very few young or old, is normal for animals that form packs. Therefore, Coria said, large theropods may have hunted in groups, which would be advantageous when hunting gigantic sauropods.[44]

Paleoenvironment

dinosaurs

Giganotosaurus was discovered in the

claystones are present, some of which represent swamp conditions.[47]

Giganotosaurus was probably the apex predator in its ecosystem. It shared its environment with herbivorous dinosaurs such as the

ornithopods and pterosaurs as well.[47][36]

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External links
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