Spinosauridae
Spinosaurids | |
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Montage of four spinosaurids, clockwise from top left: Baryonyx, Irritator, Spinosaurus and Suchomimus | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Dinosauria |
Clade: | Saurischia |
Clade: | Theropoda |
Clade: | Avetheropoda |
Clade: | †Carnosauria (?) |
Family: | †Spinosauridae Stromer, 1915 |
Type species | |
†Spinosaurus aegyptiacus Stromer, 1915
| |
Subgroups | |
*disputed validity; =Spinosaurus?[5][6][7] | |
Synonyms | |
Spinosauridae (or spinosaurids) is a clade or
Spinosaurids were large bipedal carnivores. Their crocodilian-like skulls were long, low and narrow, bearing conical teeth with reduced or absent serrations. The tips of their upper and lower jaws fanned out into a spoon-shaped structure similar to a rosette, behind which there was a notch in the upper jaw that the expanded tip of the lower jaw fit into. The nostrils of spinosaurids were retracted to a position further back on the head than in most other theropods, and they had bony crests on their heads along the midline of their skulls. Their robust shoulders wielded stocky forelimbs, with three-fingered hands that bore an enlarged claw on the first digit. In many species, the upwards-projecting neural spines of the vertebrae (backbones) were significantly elongated and formed a sail on the animal's back (hence the family's etymology), which supported either a layer of skin or a fatty hump.
The genus
History of discovery
The first spinosaurid
The first fossils referred to a spinosaurid were discovered in 1912 at the
In 1983, a relatively complete skeleton was excavated from the Smokejacks pit in
On 2021 a recent discovery in Isle of Wight an island off the south coast of England, remains of a spinosaurid which is said to be of a new species is found. As per the findings, it is about 10 meters in length and weighed several tons. The prehistoric bones of the spinosaurid were found in a geological layer of rock known as the Vectis Formation in Compton Chine, it is the first identifiable theropod from the Vectis Formation. The study was led by Christopher Barker, a PhD doctoral student in vertebrate paleontology at the University of Southampton.[21]
Description
Although reliable size and weight estimates for most known spinosaurids are hindered by the lack of good material, all known spinosaurids were large animals.[18] The smallest genus known from good material is Irritator, which was between 6 and 8 meters (20 and 26 feet) long and around 1 metric ton (1.1 short tons; 0.98 long tons) in weight.[22][23] Ichthyovenator, Baryonyx, and Suchomimus ranged from 7.5 to 11 m (25 to 36 ft) long, and weighed between 1 and 5.2 t (1.1 and 5.7 short tons; 0.98 and 5.12 long tons).[24][23][25] Oxalaia may have reached a length of between 12 and 14 m (39 and 46 ft) and a weight of 5 to 7 t (5.5 to 7.7 short tons; 4.9 to 6.9 long tons).[26] The largest known genus is Spinosaurus, which was capable of reaching lengths of 14 m (46 ft) and weighed around 7.4 t (8.2 short tons; 7.3 long tons), making it the longest known theropod dinosaur and terrestrial predator.[27] The closely allied Sigilmassasaurus may have grown to a similar or greater length, though its taxonomic relationship with Spinosaurus is uncertain.[28] This consistency in large body size among spinosaurids could have evolved as a byproduct of their preference for semiaquatic lifestyles, as without the need to compete with other large theropod dinosaurs for food, they would have been able to grow to massive lengths.[29]
Skull
Spinosaurid
Despite their highly modified skulls, analysis of the endocasts of Baryonyx walkeri and Ceratosuchops inferodios reveals spinosaurid brains shared a high degree of similarity with those of other non-maniraptoriform theropods.[33]
Lengthwise atop their skulls ran a thin and shallow sagittal crest that was usually tallest near or above the eyes, either becoming shorter or disappearing entirely towards the front of the head.[18][34][35] Spinosaurus's head crest was comb-shaped and bore distinct vertical grooves,[34] while those of Baryonyx and Suchomimus looked like small triangular bumps.[36][7] Irritator's median crest stopped above and behind the eyes in a bulbous, flattened shape. However, given that no fully preserved skulls are known for the genus, the complete shape of Irritator's crest is unknown.[31] Cristatusaurus and Suchomimus (a possible synonym of the former) both had narrow premaxillary crests.[37] Angaturama (a possible synonym of Irritator) had an unusually tall crest on its premaxillae that nearly overhung the tip of the snout with a small forward protrusion.[35]
Spinosaurid nostrils were set far back on the skull, at least behind the teeth of the premaxillae, instead of at the front of the snout as in most theropods.[18] Those of Baryonyx and Suchomimus were large and started between the first and fourth maxillary teeth, while Spinosaurus's nostrils were far smaller and more retracted. Irritator's nostrils were positioned similarly to those of Baryonyx and Suchomimus, and were between those of Spinosaurus and Suchomimus in size.[35] Spinosaurids had long secondary palates, bony and rugose structures on the roof of their mouths that are also found in extant crocodilians, but not in most theropod dinosaurs.[30] Oxalaia had a particularly elaborate secondary palate, while most spinosaurs had smoother ones.[26] The teeth of spinosaurids were conical, with an oval to circular cross section and either absent or very fine serrations. Their teeth ranged from slightly recurved, such as those of Baryonyx and Suchomimus, to straight, such as those of Spinosaurus and Siamosaurus, and the crown was often ornamented with longitudinal grooves or ridges.[35][38]
Postcranial skeleton
The coracoid bones of the shoulders in spinosaurids were robust and hook shaped.[7] The arms were relatively large and well-built; the radius (long bone of the forearm) was stout and usually only half as long as the humerus (upper arm bone). Spinosaurid hands had three fingers, typical of tetanurans, and wielded an enlarged ungual on the first finger (or "thumb"), which formed the bony core of a keratin claw. In genera like Baryonyx and Suchomimus, the phalanges (finger bones) were of conventional length for large theropods, and bore hook-shaped, strongly curved hand claws.[18][24] Based on fragmentary material from the forelimbs of Spinosaurus, it appears to have had longer, more gracile hands and straighter claws than other spinosaurids.[39]
The hindlimbs of Suchomimus and Baryonyx were somewhat short and mostly conventional of other
The upward-projecting
Classification
The family Spinosauridae was named by Stromer in 1915 to include the single
Traditionally, Spinosauridae is divided into two subfamilies: Spinosaurinae, which contains the genera Icthyovenator, Irritator, Oxalaia, Sigilmassasaurus and Spinosaurus, is marked by unserrated, straight teeth, and external nares which are further back on the skull than in baryonychines,[7][47] and Baryonychinae, which contains the genera Baryonyx, Cristatusaurus, Suchosaurus, Suchomimus, Ceratosuchops, and Riparovenator,[48] which is marked by serrated, slightly curved teeth, smaller size, and more teeth in the lower jaw behind the terminal rosette than in spinosaurines.[7][47] Others, such as Siamosaurus, may belong to either Baryonychinae or Spinosaurinae, but are too incompletely known to be assigned with confidence.[48] Siamosaurus was classified as a spinosaurine in 2018, but the results are provisional and not entirely conclusive.[28]
The subfamily Spinosaurinae was named by Sereno in 1998, and defined by
The 2017 study mentioned above indicates that Baryonychinae may in fact be non-monophyletic. Their cladogram can be seen below.[35]
Spinosauridae | |
The next cladogram displays an analysis of Tetanurae simplified to show only Spinosauridae from Allain colleagues in 2012:[40]
Spinosauridae |
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The 2018 phylogenetic analysis by Arden and colleagues, which included many unnamed taxa, resolved Baryonychinae as monophyletic, and also coined the new term Spinosaurini for the clade of Sigilmassasaurus and Spinosaurus.[28]
Spinosauridae |
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In 2021, Chris Barker, Hone,
Spinosauridae |
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See also the phylogenetic results in the 2022 article describing Iberospinus.[50]
Evolution
Spinosaurids appear to have been widespread from the
Several theories have been proposed about the
In 2016, the Spanish palaeontologist Alejandro Serrano-Martínez and colleagues reported the oldest known spinosaurid fossil, a tooth from the Middle Jurassic of Niger, which they found to suggest that spinosaurids originated in Gondwana, since other known Jurassic spinosaurid teeth are also from Africa, but they found the subsequent dispersal routes unclear.
Paleobiology
Diet and feeding
Spinosaurid teeth resemble those of crocodiles, which are used for piercing and holding prey. Therefore, teeth with small or no serrations, such as in spinosaurids, were not good for cutting or ripping into flesh but instead helped to ensure a strong grip on a struggling prey animal.
In the past, spinosaurids have often been considered
Further study by Andrew R. Cuff and Rayfield in 2013 on the skulls of Spinosaurus and Baryonyx did not recover similarities in the skulls of Baryonyx and the gharial that the previous study did. Baryonyx had, in models where the size difference of the skulls was corrected for, greater resistance to torsion and dorsoventral bending than both Spinosaurus and the gharial, while both spinosaurids were inferior to the gharial, alligator, and
Direct fossil evidence shows that spinosaurids fed on fish as well as a variety of other small to medium-sized animals, including dinosaurs. Baryonyx was found with scales of the prehistoric fish
A 2018 study by Auguste Hassler and colleagues of
Forelimb function
The use of the robust forelimbs and giant recurved claws of spinosaurs remains a debated topic. Charig and Milner speculated in 1986 that Baryonyx may have crouched by the riverbank and used its claws to
In 1986, Charig and Milner suggested that the robust forelimbs and giant thumb claws would have been Baryonyx's primary method of capturing, killing, and tearing apart large prey; whereas its long snout would have been used mostly for fishing.[36] A 2005 study by Canadian paleontologist the François Therrien and colleagues agreed that spinosaur forelimbs were probably used for hunting larger prey items, given that their snouts could not resist the bending stress.[73] In a 2017 review of the family, David Hone and Holtz considered possible functions in digging for water sources or hard to reach prey, as well as burrowing into soil to construct nests.[18]
Cranial crests and neural spines
Theropod heads are often decorated with some form of crest, horn, or rugose structure, which in life, would have been extended by keratin.
Many theories have been proposed over the years for the use of spinosaurid dorsal sails, such as
In 2015, the German biophysicist Jan Gimsa and colleagues suggested that this feature could also have aided aquatic movement by improving manoeuvrability when submerged, and acted as fulcrum for powerful movements of the neck and tail (similar to those of
Ontogeny
Juvenile spinosaurid fossils are somewhat rare. However, an ungual phalanx measuring 21 mm (0.83 in) belonging to a very young Spinosaurus indicates that Spinosaurus, and probably by extent other spinosaurids, may have developed their semiaquatic adaptations at birth or at a very young age and maintained the adaptations throughout their lives. The specimen, found in 1999 and described by Simone Maganuco, Cristiano Dal Sasso and colleagues in 2018, is believed to have come from a very small juvenile measuring 1.78 m (5.8 ft), making said specimen the smallest known example of a spinosaurid currently described.[83][84]
Paleoecology
Habitat preference
A 2010 publication by Romain Amiot and colleagues found that oxygen isotope ratios of spinosaurid bones indicates semiaquatic lifestyles. Isotope ratios from teeth from Baryonyx, Irritator, Siamosaurus, and Spinosaurus were compared with isotopic compositions from contemporaneous theropods, turtles, and crocodilians. The study found that, among theropods, spinosaurid isotope ratios were closer to those of turtles and crocodilians. Siamosaurus specimens tended to have the largest difference from the ratios of other theropods, and Spinosaurus tended to have the least difference. The authors concluded that spinosaurids, like modern crocodilians and hippopotamuses, spent much of their daily lives in water. The authors also suggested that semiaquatic habits and piscivory in spinosaurids can explain how spinosaurids coexisted with other large theropods: by feeding on different prey items and living in different habitats, the different types of theropods would have been out of direct competition.[85] In 2018, an analysis was conducted on the partial tibia of an indeterminate spinosaurine from the early Albian, the bone was from a sub-adult between 7 and 13 m (22 and 42 ft) in length still growing moderately fast before its death. This specimen (LPP-PV-0042) was found in the Araripe Basin of Brazil and taken to the University of San Carlos for a CT Scan, where it revealed osteosclerosis (high bone density).[29] This condition had previously only been observed in Spinosaurus, as a possible way of controlling its buoyancy.[39] The presence of this condition on the leg fragment showed that semi-aquatic adaptations in spinosaurids were already present at least 10 million years before Spinosaurus aegyptiacus appeared. According to the phylogenetic bracketing method, this high bone density might have been present in all spinosaurines.[29] In 2020, a scientific paper by paleontologists published in the scientific journal Cretaceous Research found taphonomic evidence in the Kem Kem group that would support Spinosaurus being a semi-aquatic dinosaur.[86] However, research conducted in 2023 cited the immediate assumption of Spinosaurids being avid divers due to correlations in bone compactness as being subject to errors, such as flawed statistical methods and measurements, as well as sampling bias.[87] A study conducted in 2023 by Stephanie Baumgart also found similar results with the previous studies as; given amount of variation in specimens and in data collection techniques, they concluded that previous evidence isn’t strong enough to put Spinosaurus swimming and diving entirely submerged. Spinosaurus still more likely mostly hung out on shore, akin to wader lifestyle previously interfered.[88]
A 2018 study of buoyancy (through simulation with 3D models) by the Canadian palaeontologist Donald M. Henderson found that distantly related theropods floated as well as the tested spinosaurs, and instead supported they would have stayed by the shorelines or shallow water rather than being semi-aquatic.[89]
Distribution
Confirmed spinosaurids have been found on every continent except for North America, Australia and Antarctica, the first of which was Spinosaurus aegyptiacus, discovered at the
The earliest record of spinosaurines is from Europe, with the Barremian species
Partial skeletons and numerous fossil teeth indicate spinosaurids were widespread in Asia; three taxa—all spinosaurines—have been named: Siamosaurus suteethorni from Thailand,
Timeline of genera
Timeline of genera descriptions
See also
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External links
- Spinosauridae on the Theropod Database