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Spinosaurus

Spinosaurus is a genus of large spinosaurid theropod dinosaurs that lived in what is now North Africa during the Cenomanian stage of the Late Cretaceous period, about 100 to 94 million years ago. The genus was known first from Egyptian remains discovered in 1912 and described by German palaeontologist Ernst Stromer in 1915. The original remains were destroyed in World War II, but additional material came to light in the early 21st century. It is unclear whether one or two species are represented in the fossils reported in the scientific literature. The type species, S. aegyptiacus, is mainly known from the Bahariya Formation in Egypt and the Kem Kem beds in Morocco. The other species, S. mirabilis, is known from the Farak Formation in Niger. Although another potential species, S. maroccanus, has been recovered from the Kem Kem beds, this dubious species is likely a junior synonym of S. aegyptiacus. Other possible junior synonyms include Sigilmassasaurus from the Kem Kem beds and Oxalaia from the Alcântara Formation in Brazil, though other researchers propose both genera to be distinct taxa.

Discovery and naming
Initial finds (name-bearing) specimen from Stromer (1915) The first discovery of Spinosaurus may have been in October 1898, when French geologist Fernand Foureau unearthed two unusual teeth in Cenomanian-aged sediments of the Djoua Escarpment in what is now southern Algeria, then French Algeria. Foreau found the specimens while on the Foureau-Lamy Expedition of 1898-1900, which traveled across the Sahara from Algeria to Lake Chad. In 1905, French geologist Emile Haug described the fossils unearthed while on the expedition, including the two teeth which he assigned to the fish Saurocephalus. The first confidently referable remains of Spinosaurus were discovered in the autumn of 1912 by Austro-Hungarian paleontologist Richard Markgraf, a fossil collector who was financed by Stromer and the Bavarian Academy of Sciences, in deposits of the Bahariya Formation in the Bahariya Oasis of western Egypt. The sediments of the Bahariya Formation come from the Cenomanian stage of the Late Cretaceous period, one of many Cretaceous-aged formations of North Africa. In 1915, Stromer described these remains as belonging to a new genus and species of dinosaur, Spinosaurus aegyptiacus. The generic name Spinosaurus comes from the Latin spina, meaning "spine", and the Greek sauros, meaning "lizard", and thus "spine lizard". The specific name aegyptiacus derives from Egypt, where the fossils were unearthed. Spinosaurus aegyptiacus type specimen (BSP 1912 VIII 19) consists of elements of the mandible (lower jaw), a maxilla (upper jaw) fragment, cervical (neck), caudal (tail), and dorsal (back) vertebrae, including much of the , ribs, teeth, and some gastralia (belly ribs). the skeleton arrived in Germany in 1922 where it was described by Stromer in 1934. Stromer assigned the skeleton to Spinosaurus, but considered it different enough from S. aegyptiacus that it would belong to its own Spinosaurus species, "Spinosaurus B". In 1936, Stromer worked with Dr. Erhardt (the first name is unknown) to create a skeletal reconstruction of Spinosaurus based on fossils of the holotype, "Spinosaurus B", and related taxa. Although Stromer was hesistant about the lack of known remains, he and Erhardt produced a restoration that filled in missing elements of Spinosaurus' skeleton with the tyrannosaurid theropods Tyrannosaurus and Gorgosaurus.'''' Unfortunately, during the night of April 24/25, 1944, the building of the Paläontologisches Museum München (Bavarian State Collection of Paleontology) of the Bavarian Academy of Sciences was severely damaged during the British Bombing of Munich. Due to personal and political tensions between Stromer and the museum's curator, who was a fervent Nazi, the Spinosaurus fossils held there were not rehoused and subsequently destroyed as a result of the bombing. Stromer's finds, including Spinosaurus, received little academic or public attention. In 1995, Stromer's son donated his father's archives to the Paläontologische Staatssammlung München, leading to a 2006 study by American researcher Joshua Smith and colleagues of the photographs, two of which were of Spinosaurus holotype. Based on these photographs of the mandible and the entire specimen as mounted, Smith concluded that Stromer's original 1915 drawings were slightly inaccurate. In 1986, a paper referred several teeth found Cretaceous deposits in Tunisia to Spinosaurus, being the first evidence of the taxon's existence in the country. In 1989, French paleontologist Eric Buffetaut assigned two jaw fragments and a tooth that had been unearthed in a deposit of the Kem Kem Beds in Taouz, Morocco to Spinosaurus sp., expanding the known distribution of the genus to Morocco. megaraptorans, carcharodontosaurids, or other forms of theropod. Later authors have been split on this topic, with some authors noting that the length of the vertebrae can vary from individual to individual, that the holotype specimen was destroyed and thus cannot be compared directly with the S. maroccanus specimen, and that it is unknown which cervical vertebrae the S. maroccanus specimens represent. Therefore, though some have retained the species as valid without much comment, most researchers regard S. maroccanus as a nomen dubium (dubious name), a junior synonym of S. aegyptiacus, or a junior synonym of Sigilmassasaurus brevicollis. The specimens previously ascribed as paratypes of S. maroccanus (NMC 41768 and NMC 50790) were reidentified as spinosaurid specimens that are currently not identifiable at the generic level. In 1996, a study by American paleontologist Paul Sereno and colleagues assigned cervical vertebrae referred to "Spinosaurus B" and the spinosaurid Sigilmassasaurus (a possible synonym of Spinosaurus) however later analyses have proven that "Spinosaurus B" and Sigilmassasaurus are not synonyms or contain fossils from Carcharodontosaurus. Ibrahim recognized that the MSNM fossils and the ones he had seen in 2008 were from a similar sandstone matrix and that the neural spines of the MSNM fossils were similar to those depicted by Stromer in 1915, leading him to think that they may be from the same individual and belonged to Spinosaurus. Following this, an international collaboration began between Ibrahim and other paleontologists to study the bones and visit the original site, with fossils from the MSNM getting sent to the University of Chicago for study. Subsequently, the dealer took them back to the site where the fossils had been found. Over the course of several years, a skeleton of a subadult individual consisting of skull fragments, some cervical, dorsal, and caudal vertebrae, neural spines, a complete sacrum, much of the hindlimbs, several pedal phalanges, and several dorsal ribs was unearthed. This skeleton comes from a locality of the Douria Formation of the Kem Kem Beds in Zrigat, Morocco, dated to the Cenomanian stage of the mid-Cretaceous period. In 2015, British paleontologist Serjoscha Evers and colleagues rejected the neotype proposal for FSAC-KK-11888 on the basis of the lack of association or locality information for FSAC-KK-11888, the geographical separation between the holotype and FSAC-KK-11888 localities, and anatomical differences between FSAC-KK-11888 and the holotype. Evers and colleagues argue for the validity of both S. marrocanus and S. aegyptiacus, stating that FSAC-KK-11888 likely belongs to the former. Additionally, they state that the geographical separation of FSAC-KK-11888 and the holotype as well as the detailed description of the holotype by Stromer are evidence that FSAC-KK-11888 does not meet the ICZN (International Code of Zoological Nomenclature)'s requirements for a neotype and that the creation of a neotype is unnecessary for S. aegyptiacus. Between 2015 and 2019, several additional expeditions to the site unearthed additional fossils of FSAC-KK-11888 including many caudal vertebrae, composing about 80% of the tail with a preserved length of , elements of the pes and hindlimbs, skull fragments, and more. Around 90% of this new material was found in late 2018, whereas some fragments including cranial material were found in debris of previous expeditions. In 2020, Ibrahim and colleagues described the tail of FSAC-KK-11888, which they stated closely compares to the caudal vertebrae of the holotype and "Spinosaurus B". The material found on these expeditions perfectly matches elements of the FSAC-KK-11888, proving that they come from a single individual. The body length of the proposed neotype excluding the tail portion, from the anterior tip of the skull to the posterior region of the pelvic girdle, is estimated up to ,Table S1 and the total length of the tail is estimated up to . ==Description==
Description
Size Since its discovery, Spinosaurus has been a contender for the largest theropod dinosaur. Both Friedrich von Huene in 1926 and Donald F. Glut in 1982 listed it as among the most massive theropods in their surveys, at in length and upwards of in weight. In 1988, Gregory S. Paul also listed it as the longest theropod at , but gave a lower mass estimate of . In 2020, Nicolás Campione and David Evans suggested to take caution with the approach using skull length for extrapolation, namely that of Spinosaurus, since the preorbital region of some dinosaurs grow allometrically (disproportionately with body size) and the cranial properties were likely under different selection pressures unrelated to body size. In 2014, Ibrahim and his colleagues suggested that an adult Spinosaurus aegyptiacus could reach over in length. overestimated the presacral column length by 10%, ribcage depth by 25%, and forelimb length by 30% over dimensions based on CT-scanned fossils; these proportional overestimates shift the center of mass anteriorly when translated to a flesh model, and thus the estimate from Ibrahim and his colleagues cannot be considered a reliable body size estimate. Very tall neural spines growing on the back vertebrae of Spinosaurus formed the basis of what is usually called the animal's "sail". The lengths of the neural spines reached over 10 times the diameters of the centra (vertebral bodies) from which they extended. The neural spines were slightly longer front to back at the base than higher up, and were unlike the thin rods seen in the pelycosaur finbacks such as Edaphosaurus, Ianthasaurus and Dimetrodon, contrasting also with the thicker spines in the iguanodontian Ouranosaurus. In 2014, Ibrahim and colleagues instead posited that the spines were covered tightly by skin, similar to a crested chameleon, given their compactness, sharp edges, and likely poor blood flow. ==Classification==
Classification
that did not live in the same time or space; Spinosaurus, Baryonyx, Irritator and Suchomimus|270x270pxSpinosaurus is the type genus of the family Spinosauridae, subfamily Spinosaurinae, and tribe Spinosaurini. and Camarillasaurus. Spinosaurini is defined as a clade including all spinosaurids closer to Spinosaurus aegyptiacus than Irritator challengeri and Oxalaia quilombensis, which are distinguished from spinosaurins due to their less elongate premaxillae and premaxillae which are excluded from the external naris. However, some studies by paleontologists like American researcher Alfred Romer included the high-spined theropods Acrocanthosaurus and Altispinax in Spinosauridae based on their tall neural spines. This has not been supported by later studies, which typically classify these taxa as members of Carcharodontosauridae or Allosauroidea instead of Spinosauridae. In 1986, British paleontologists Alan Charig and Angela Milner described Baryonyx, which they considered distinct enough to be classified in its own family of theropod dinosaurs: Baryonychidae. However, in 1988 American paleontologist Gregory S. Paul argued that Baryonyx and Spinosaurus were related based on their notched snouts and stated they may be late-surviving dilophosaurids. In a 1989 paper, Buffetaut supported this conclusion, stating that they were both piscivorous, related theropods. Descriptions of Irritator, Cristatusaurus, and other spinosaurids (as well as additional fossils of Spinosaurus) led to the realization of a transcontinental clade of spinosaurids. and Spinosauridae as a clade composed of spinosauroids closer related to Spinosaurus aegyptiacus than Torvosaurus tanneri. Additionally, Sereno and colleagues recognized the existence of two subfamilies within Spinosauridae: Baryonychinae, which included Suchomimus and Baryonyx, and Spinosaurinae, which included Spinosaurus and Irritator. of Africa, Europe, and South America. }} To test the relationships and affinities of Spinosaurus mirabilis, Sereno and colleagues included it in an updated version of the phylogenetic matrix published by Sereno et al. (2022). }}|style=font-size:90%; line-height:90%;|label1=Spinosauridae}} ==Paleobiology==
Paleobiology
Function of neural spines The function of the dinosaur's sail or hump is uncertain; scientists have proposed several hypotheses including heat regulation and display. In addition, such a prominent feature on its back could make it appear even larger than it was, intimidating other animals. However, Bailey (1997) was of the opinion that a sail could have absorbed more heat than it radiated. Bailey proposed instead that Spinosaurus and other dinosaurs with long neural spines had fatty humps on their backs for energy storage, insulation, and shielding from heat. Gimsa and others point out that more basal, long-legged spinosaurids had otherwise round or crescent-shaped dorsal sails, whereas in Spinosaurus, the dorsal neural spines formed a shape that was roughly rectangular, similar in shape to the dorsal fins of sailfish. They therefore argue that Spinosaurus used its dorsal neural sail in the same manner as sailfish, and that it also employed its long narrow tail to stun prey like a modern thresher shark. Sailfish employ their dorsal fins for herding schools of fish into a "bait ball" where they cooperate to trap the fish into a certain area where the sailfish can snatch the fish with their bills. The sail could have possibly reduced yaw rotation by counteracting the lateral force in the direction opposite to the slash as suggested by Gimsa and colleagues (2015). Large fish are known from the faunas containing other spinosaurids, including the Mawsonia, in the mid-Cretaceous of northern Africa and Brazil. Direct evidence for spinosaur diet comes from related European and South American taxa. Baryonyx was found with fish scales and bones from juvenile Iguanodon in its stomach, while a tooth embedded in a South American pterosaur bone suggests that spinosaurs occasionally preyed on pterosaurs, but Spinosaurus was likely to have been a generalized and opportunistic predator, possibly a Cretaceous equivalent of large grizzly bears, being biased toward fishing, though it undoubtedly scavenged and took many kinds of small or medium-sized prey. In 2009, Dal Sasso and colleagues. reported the results of X-ray computed tomography of the MSNM V4047 snout. As the foramina on the outside all communicated with a space on the inside of the snout, the authors speculated that Spinosaurus had pressure receptors inside the space that allowed it to hold its snout at the surface of the water to detect swimming prey species without seeing them. A 2013 study by Andrew R. Cuff and Emily J. Rayfield concluded that bio-mechanical data suggests that Spinosaurus was not an obligate piscivore and that its diet was more closely associated with each individual's size. The characteristic rostral morphology of Spinosaurus allowed its jaws to resist bending in the vertical direction, but its jaws were poorly adapted with respect to resisting lateral bending compared to other members of this group (Baryonyx) and modern alligators. This suggests that Spinosaurus preyed more regularly on fish than it did on land animals, although considered predators of the former too. In 2022, Sakamoto estimated that Spinosaurus had an anterior bite force of 4,829 newtons and a posterior bite force of 11,936 newtons . Based on this estimate, he asserted that the jaws of Spinosaurus are adapted for generating relatively faster shutting speeds with less muscle input force, indicating that the animal likely killed its prey with fast-snapping jaws rather than slow-crushing bites, a trait commonly observed in animals which have a semi-aquatic feeding habit. A 2024 paper suggests that Spinosaurus and other spinosaurines in addition to fish also preyed upon small to medium-sized terrestrial vertebrates, and had relatively weak bite forces compared to those of other theropods. Aquatic habits Onchopristis A 2010 isotope analysis by Romain Amiot and colleagues found that oxygen isotope ratios of spinosaurid teeth, including teeth of Spinosaurus, indicate semiaquatic lifestyles. Isotope ratios from tooth enamel and from other parts of Spinosaurus (found in Morocco and Tunisia) and of other predators from the same area such as Carcharodontosaurus were compared with isotopic compositions from contemporaneous theropods, turtles, and crocodilians. The study found that Spinosaurus teeth from five of six sampled localities had oxygen isotope ratios closer to those of turtles and crocodilians when compared with other theropod teeth from the same localities. The authors postulated that Spinosaurus switched between terrestrial and aquatic habitats to compete for food with large crocodilians and other large theropods respectively. A 2018 study by Donald Henderson, however, refutes the claim that Spinosaurus was semiaquatic. By studying the buoyancy in lungs of crocodilians and comparing it to the lung placement in Spinosaurus, it was discovered that Spinosaurus could not sink or dive below the water surface. It was also capable of keeping its entire head above the water surface while floating, much like other non-aquatic theropods. Furthermore, the study found that Spinosaurus had to continually paddle its hind legs to prevent itself from tipping over onto its side, something that extant semiaquatic animals do not need to perform. Henderson therefore theorized that Spinosaurus probably did not hunt completely submerged in water as previously hypothesized, but instead would have spent much of its time on land or in shallow water. Later studies of the tail vertebrae of Spinosaurus refute Henderson's proposal that Spinosaurus mainly inhabited areas of land near and in shallow water and was too buoyant to submerge. Studies of the tail, thanks to fossils recovered and analyzed by Ibrahim, Pierce, Lauder, and Sereno and colleagues in 2018 indicate that Spinosaurus had a keeled tail that was well adapted to propelling the animal through water. The elongated neural spines and chevrons, which run to the end of the tail on both dorsal and ventral sides, indicate that Spinosaurus was able to swim in a similar manner to modern crocodilians. Through experimentation by Lauder and Pierce, the tail of Spinosaurus was found to have eight times as much forward thrust as the tails of terrestrial theropods like Coelophysis and Allosaurus, as well as being twice as efficient at achieving forward thrust. The discovery indicates that Spinosaurus may have had a lifestyle comparable to modern alligators and crocodiles, remaining in water for long periods of time while hunting. They highlight the positioning of the nostrils and orbits as one reason why a crocodile-like lifestyle is unlikely: they are ventrally positioned in such a way that the whole head would have to be lifted inefficiently out of the water in order to breathe. Additionally, they argue that the general body shape of Spinosaurus is poorly adapted for this lifestyle, drawing on the amount of water drag and aquatic instability In the same year, contradicting the study by Fabbri and colleagues, Sereno and his colleagues suggested that Spinosaurus was wholly bipedal on land and an unstable, slow moving surface swimmer in deep water. Their results, taken from reconstructing a CT model of the skeleton, and then adding internal air and muscles. Their results, coupled with fossils from Spinosaurus that showed it also lived further inland along rivers and lakes, suggest it was a semi-aquatic, ambush piscivore that preferred waterside environments both along the coasts and further inland along rivers and lakes. Simultaneously, they suggested that the large tail fin was probably utilized more for display than swimming, as tails in living animals have the same function when they possess comparably tall neural spines. A 2024 paper by Myrhvold et al. also disputes that Spinosaurus and Baryonyx were diving pursuit predators. Instead they also argue that Spinosaurus and Baryonyx hunted more like herons instead of diving after prey. Another paper in the same year analyzed the linear measurements of the skull of Spinosaurus, and concluded that the skull morphology and hunting method of Spinosaurus would likely be the most similar to those of wading birds like herons, though the authors noted that they are uncertain how beneficial the skull would have been for the diving pursuit predation method. Locomotion and posture Although traditionally depicted in the scientific community as a biped, Spinosaurus was occasionally depicted in the mid-20th century as an obligate quadruped akin to Dimetrodon. Starting in the mid-1970s, it was hypothesized Spinosaurus was at least an occasional quadruped, Because of the mass of the hypothesized fatty dorsal humps of Spinosaurus, Bailey (1997) was open to the possibility of a quadrupedal posture, The hypothesis that Spinosaurus had a typical quadrupedal gait since fell out of favor, however it was still believed that spinosaurids may have crouched in a quadrupedal posture, due to biological and physiological constraints. The possibility of a quadrupedal Spinosaurus was revived by a 2014 paper by Ibrahim and colleagues that described new material of the animal. The paper found that the hind limbs of Spinosaurus were much shorter than previously believed, and that its center of mass was located in the midpoint of the torso region, as opposed to near the hip as in typical bipedal theropods. It was therefore proposed that Spinosaurus was poorly adapted for bipedal terrestrial locomotion, and must have been an obligate quadruped on land. The reconstruction used in the study was an extrapolation based on different sized individuals, scaled to what were assumed to be the correct proportions. Scott Hartman also expressed criticism because he believed the legs and the pelvis were inaccurately scaled (27% too short) and didn't match the published lengths. However, Mark Witton expressed agreement with the proportions reported in the paper. In their 2015 re-description of Sigilmassasaurus, Evers and colleagues argued that Sigilmassasaurus was in fact a distinct genus from Spinosaurus, and therefore doubted whether the material assigned to Spinosaurus by Ibrahim et al. (2014) should be assigned to Spinosaurus or Sigilmassasaurus. Ontogeny An ungual phalanx measuring belonging to a very young juvenile cf. S. aegyptiacus indicates that the theropod developed its semiaquatic adaptations at a very young age or at birth and maintained them throughout its life. The specimen, discovered in 1999 and described by Simone Maganuco and Cristiano Dal Sasso in 2018, is believed to have come from an animal measuring (assuming it resembled a smaller version of the adult), making it the smallest specimen of Spinosaurus currently known. Palaeopathology A cf. Spinosaurus sp. tooth from the Ifezouane Formation displays enhanced lingual curvature to the tooth's crown, the development of three deep grooves extending from crown root junction in the direction of the crown's apex, an attenuated carina that does not extend apically nor to the base of the tooth, and a wear facet at the tip. == Paleoecology ==
Paleoecology
The holotype specimen of Spinosaurus is known from the Bahariya Formation, one of many dinosaur-bearing Cretaceous-aged fossil formations in North Africa. However, fossils of Spinosaurus have been described from several sites including the Kem Kem Beds of Morocco, the Farak, and Echkar Formations of Niger, the Gara Samani Formation of Algeria, and the Chenini Sandstones. though Deltadromeus taxonomy and distribution too is debated. North Africa during this period bordered the Tethys Sea, which transformed the region into a mangrove-dominated coastal environment filled with vast tidal flats and waterways. The composition of the dinosaur fauna of North Africa at this time is an anomaly, as there are fewer herbivorous dinosaur species relative to carnivorous dinosaur species than in most fossil sites. This abundance of theropods compared to that of non-theropods was dubbed "Stromer's Riddle", which despite suggestions that this is due to ecological, preservation, or other biases, is supported by the fossil record. Isotopic evidence supports this, which found greater quantities of sizable, terrestrial animals in the diets of carcharodontosaurids and ceratosaurs from both the Kem Kem Beds and Elrhaz Formation. North Africa was dominated by a triumvirate of Abelisauroidea, Spinosauridae, and Carcharodontosauridae during the mid-Cretaceous, with all of these groups present in the Kem Kem Beds, Echkar, Elrhaz, and Bahariya Formations. Alongside the coeval theropods Tameryraptor and Bahariasaurus, Spinosaurus is one of several theropod dinosaurs known from the Bahariya Formation. Fossils of sauropods include those of the titanosaurs Paralititan and Aegyptosaurus, the former reaching great sizes. The faunal composition of both the Bahariya Formation and the Kem Kem Beds were thought to be similar in the past, but the describers of Tameryraptor suggested that such superficial comparisons require further examination. An isolated wing phalanx of a pterosaur, potentially Anhanguera, was unearthed from the Bahariya Formation. A diverse fauna of aquatic animals is known from the Bahariya Formation. Underwater life diversity exploded during this period in the mangroves of North Africa, with turtles represented by the pleurodian Apertotemporalis, large bony fish like Mawsonia and Paranogmius, sawskates Onchopristis and Schizorhiza, sharks like Squalicorax and Cretolamna, and a broad selection of invertebrates. Additionally, several crocodylomorphs like the stomatosuchid Stomatosuchus and the eunotosuchian Libycosuchus are known from the formation. Kem Kem Beds The Kem Kem Beds is composed of three geologic formations: the Gara Sbaa Formation, the Douria Formation, and the Izefouane Formation. This river system was freshwater based on the presence of lungfishes and other typically freshwater vertebrates. This indicates that the Kem Kem Beds had a wide variety of features, including river channels, river banks, sandbars, and more. an indeterminate somphospondylian titanosauriform, and an indeterminate titanosaur, one comparable in size to the giant Paralititan. Ornithischian fossils are extremely rare, only being represented from an isolated thyreophoran tooth As for theropods, many are known, including Spinosaurus, Sigilmassasaurus, if it is valid, However, many of these dinosaurs are known from isolated or incomplete remains, have complicated taxonomies, or are under study. == In popular culture ==
In popular culture
, Barcelona Spinosaurus appeared in the 2001 film Jurassic Park III, replacing Tyrannosaurus as the main antagonist. The film's consulting paleontologist John R. Horner was quoted as saying, "If we base the ferocious factor on the length of the animal, there was nothing that ever lived on this planet that could match this creature [Spinosaurus]. Also my hypothesis is that T-rex was actually a scavenger rather than a killer. Spinosaurus was really the predatory animal." He has since retracted the statement about T. rex being a scavenger. In the film, Spinosaurus was portrayed as larger and more powerful than Tyrannosaurus: in a scene depicting a battle between the two resurrected predators, Spinosaurus emerges victorious by snapping the Tyrannosaurus neck. In the fourth film, Jurassic World, there is a nod to this fight where the T. rex smashes through the skeleton of a Spinosaurus in the climactic fight near the end of the film. The Spinosaurus would appear in many Jurassic Park games, most notably Jurassic World Evolution and its sequels. The same Spinosaurus from the third film returns in the fourth, and fifth season of Jurassic World Camp Cretaceous, this time battling two T. rex. Spinosaurus return in Jurassic World Rebirth in a more realistic depiction, and having a coexisting relationship with a Mosasaurus. Spinosaurus has long been depicted in popular books about dinosaurs, although only recently has there been enough information about spinosaurids for an accurate depiction. After an influential 1955 skeletal reconstruction by Lapparent and Lavocat based on a 1936 diagram by Stromer, it has been treated as a generalized upright theropod, with a skull similar to that of other large theropods and a sail on its back, even having four-fingered hands. ==See also==
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