Siamosaurus
Siamosaurus | |
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Illustration of the holotype tooth, with British penny for scale | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Dinosauria |
Clade: | Saurischia |
Clade: | Theropoda |
Family: | †Spinosauridae |
Genus: | †Siamosaurus Buffetaut & Ingavat, 1986
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Species: | †S. suteethorni
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Binomial name | |
†Siamosaurus suteethorni Buffetaut & Ingavat, 1986
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Synonyms | |
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Siamosaurus (meaning "Siam lizard") is a
Since it is based only on teeth, Siamosaurus's body size is uncertain, though it has been estimated at between 5.1 to 9.1 metres (17 to 30 feet) in length. The
Like in all spinosaurids, Siamosaurus's teeth were conical, with reduced or absent
History of discovery
The
The best-preserved specimen from the teeth described, designated DMR TF 2043a, was chosen as the holotype of Siamosaurus.[1] The paratypes[4] comprise eight other well-preserved teeth catalogued as DMR TF 2043b to i. The original fossils are currently housed in the palaeontological collection of the Department of Mineral Resources, Bangkok.[1] Siamosaurus teeth are common in the Sao Khua Formation, and further isolated specimens were found later throughout the late 20th and early 21st centuries.[5][6] Sculptures of the animal have been erected in various places across northeastern Thailand, including Si Wiang Dinosaur Park,[7] the Siam Paragon shopping mall in Bangkok,[8][9] the Phu Wiang Dinosaur Museum,[10] and the Sirindhorn Museum.[11] S. suteethorni was also illustrated on Thai postage stamps released in 1997, along with fellow Thai dinosaurs Phuwiangosaurus, Siamotyrannus, and Psittacosaurus.[12]
Thailand's
Siamosaurus is the first reported spinosaurid dinosaur from Asia,[1][18] and subsequently to its naming, material resembling or possibly belonging to the genus has been found across the continent. In 1975, Chinese palaeontologist Hou Lian-Hai and colleagues described five teeth as a new species of the pliosauroid Sinopliosaurus, which they named S. fusuiensis, the specific name is in reference to Fusui County in Guangxi, China, from which the fossils were collected.[19] Four of these teeth—one was not found in the museum collection—were reassigned by Buffetaut and colleagues in 2008 to a spinosaurid theropod and referred to as "Sinopliosaurus" fusuiensis. The researchers deemed it as "closely related to, if not identical with", S. suteethorni.[20] In 2019, "S." fusuiensis was referred to by Thai palaeontologist Wongko Kamonrak and colleagues as Siamosaurus sp. (of uncertain species).[6] Later in 2019, Thai palaeontologist Adun Samathi and colleagues considered the teeth as belonging to an indeterminate spinosaurid.[21] The specimens were retrieved from China's Early Cretaceous Xinlong Formation, in which spinosaurid teeth are frequently reported, though most of them are hard to differentiate from those of Japan or Thailand without more complete fossil material, such as a skull.[22] Buffetaut and Suteethorn suggested that the Xinlong Formation could be geologically related to the Sao Khua or Khok Kruat Formation, since similar types of fossils have been recovered in all three regions.[5]
In 1994, an isolated tooth (specimen GMNH-PV-999) was found by a fossil prospector in the Sebayashi Formation, Japan. The tooth was believed, until 2003, to belong to a marine reptile, when Japanese palaeontologist Yoshikazu Hasegawa and colleagues assigned it to ?Siamosaurus sp.[23] The tooth came from rocks dated to the Barremian, similar in age to sediments that Siamosaurus teeth have been recovered from in Thailand.[23][24] In 2015, a more incomplete tooth was recovered from the same formation by two local children. Kept under the specimen number KDC-PV-0003, the tooth was assigned to an indeterminate spinosaurid in 2017 by Japanese palaeontologist Kubota Katsuhiro and colleagues.[24] Further spinosaurid teeth from unnamed and indeterminate forms have been discovered in central China[25] and Malaysia.[26]
In 2004, excavation began on a partial skeleton from an outcrop of the Khok Kruat Formation near the city of
Description
In 2004, American dinosaur researcher Don Lessem estimated Siamosaurus at 9.1 metres (30 feet) long.[34] In 2005, British author Sussana Davidson and colleagues gave a lower estimate of 8 m (26 ft) in length and weighing 1 tonne (1.1 short tons; 0.98 long tons).[35] In a 2016 popular book, authors Rubén Molina-Pérez and Asier Larramendi estimated it at approximately 5.1 m (17 ft) long, 1.45 m (4.8 ft) tall at the hips, and weighing 255 kilograms (562 pounds).[36] However, reliable calculations on the weight and body size of fragmentary dinosaurs like Siamosaurus are hindered by the lack of good material, such as a skull or postcranial skeleton, and thus estimates are only tentative.[37] "Phuwiang spinosaurid B" was calculated as approximately 5 m (16 ft) long by Samathi in 2019.[33] As a spinosaurid, Siamosaurus would have had low, narrow, and elongated jaws; well-built forelimbs; relatively short hindlimbs; and elongated neural spines on the vertebrae forming a sail on its back.[37][38]
Type specimens
Fossil theropod teeth are typically identified by attributes such as the proportions, size, and curvature of the crown, as well as the presence and/or shape of the denticles (serrations).[39] The holotype of S. suteethorni (specimen DMR TF 2043a) is 62.5 millimetres (2.46 in) in total length, with the crown being 47.7 mm (1.88 in) long, and 16.6 by 12.5 mm (0.65 by 0.49 in) wide at its base. It is among the larger teeth discovered by Buffetaut and Ingavat. One much smaller specimen (DMR TF 2043b) measures 24.3 mm (0.96 in) in length. According to the authors, this dramatic size range suggests the teeth are from individuals of different ages.[1] Among theropods, this may also indicate size variation along the tooth row in the jaws, which is observed to have been prevalent in spinosaurids.[37][40]
The holotype tooth is relatively straight, with only minor front to back curvature. It is oval in cross-section while other specimens are nearly circular in this aspect. Unlike in most theropods, the carinae (cutting edges) of Siamosaurus teeth lack well-defined serrations,[1] though unworn teeth do exhibit very fine denticles.[21] Some teeth (including the holotype) have a wave-like double recurvature when viewed from the front or back, which Buffetaut and Ingavat compared to that seen in carnosaur teeth from the same formation and one Deinonychus tooth described by John Ostrom in 1969. The S. suteethorni holotype is symmetrically concave front to back, and bears 15 flutes (lengthwise grooves) on its lingual (inward facing) and labial (outward facing) surfaces. These flutes run from the base of the crown before stopping 5 mm (0.20 in) from the rounded tooth tip. A region of the holotype where the enamel (outer layer of the teeth) has weathered away reveals that these flutes extend down to the dentin (second layer of the teeth).[1] The enamel also has a granular (finely wrinkled) texture, as seen in teeth from the spinosaurid Baryonyx.[41] Some of the root is preserved in the holotype and, as in all theropods, there is a large pocket for the tooth pulp, which would have housed blood vessels and nerves.[1]
Khok Kruat teeth
Of the two Khok Kruat Formation tooth morphotypes assigned by Kamonrak and colleagues in 2019, morphotype I, the Khok Kruat morphotype, is on average 46.35 mm (1.825 in) in total length, of which the crown takes up 43.77 mm (1.723 in), with a 18.40 by 13.5 mm (0.724 by 0.531 in) wide base. They are oval in cross-section, have well-defined carinae, and a smooth enamel surface, which becomes wrinkled at the base of the crown. They bear fine, sharply defined flutes, of which there are about 21 to 32 on each side. Morphotype II, the Siamosaurus morphotype, is on average 51.25 mm (2.018 in) long, with a 48.30 mm (1.902 in) tall crown that is 17.30 by 14.65 mm (0.681 by 0.577 in) wide at the base. They are also oval in cross-section and have distinct carinae, but unlike the Khok Kruat morphotype, the entire length of the crown has wrinkled enamel, and the flutes are coarser and fewer in number, with 11 to 16 on each side. The Siamosaurus morphotype also shares with S. suteethorni, GMNH-PV-999, and IVPP V 4793 a wrinkled enamel surface and between 12 and 15 flutes on each side.[6]
Possible material
Teeth
The first Sebayashi Formation specimen (GMNH-PV-999) is an isolated tooth crown with a partially intact root. It is not known in which jaw the tooth was positioned or which surface faced the inside or outside of the mouth. The tooth's front and back carinae are well-defined, though the former is not well-preserved. Besides having a broader, 20 by 14 mm (0.79 by 0.55 in) wide base and being slightly smaller at 51 mm (2.0 in) in length, GMNH-PV-999 has a very similar morphology to the S. suteethorni holotype. Features shared between the two specimens include: a straight and only slightly compressed shape; a somewhat oval cross-section; no serrations on the carinae (possibly due to bad preservation); and flutes on the crown surface, the Japanese specimen having 12 on each side. The teeth also share a crown surface with numerous small granular structures oriented parallel to their lengths. Because of these resemblances, Hasegawa and colleagues regarded GMNH-PV-999 as nearly identical to the S. suteethorni holotype tooth.
Out of the four teeth attributed to "S." fusuiensis, specimen
Postcrania
Though no skeletal elements were associated with the original Siamosaurus teeth, the Khok Kruat skeleton, SM-KK 14, may be attributable to the genus.[27][28][32] The cervical vertebrae of SM-KK 14 had elongated centra (vertebral bodies) with articulating surfaces that were not offset, as well as prominent epipophyses (processes to which neck muscles attached) and strong ligament scars. All of these characteristics were also present in Baryonyx.[16][27] The cervicals also became longer towards the front of the neck and—based on comparison with Baryonyx—may represent the fourth, sixth, seventh, and tenth vertebrae. The dorsal vertebrae had enlarged infraprezygapophyseal fossae—depressions under the prezygapophyses, which connect adjacent vertebrae[21]—and their neural spines were elongated similarly to those of other spinosaurids, indicating the presence of a sail on the animal's back; like in the Asian spinosaurid Ichthyovenator.[21][28][37] One of the neural spines of SM-KK 14 measured at least 60 centimetres (24 inches) in height.[29] The chevron lacked a process on its front end, as in other spinosaurids. Viewed distally (towards the centre of attachment), the lower end of the pubis had an L-shape, similar to that of Ichthyovenator and the African Suchomimus.[21] Also as in Ichthyovenator, the hind rim of the pubis had a notch-like obturator foramen.[21][42] However, in SM-KK 14 the front of the pubis was concave and the chevrons were curved backwards, in contrast to the straight condition these bones had in Ichthyovenator.[21]
Classification
In 1986, Buffetaut and Ingavat classified Siamosaurus as a theropod because of the straight, tall crown and double sideways recurvature of its teeth. At the time, Siamosaurus's particular combination of dental characteristics, especially the longitudinal fluting and lack of serrations, had not been observed in other theropods. The authors noted similarities in Siamosaurus's teeth to those of ceratosaurian tooth crowns, some of which also have longitudinal flutes. However, this identification was ruled out, since ceratosaur teeth are more narrow and blade-like in cross-section, bear far fewer dental flutes, and have distinct serrations. Buffetaut and Suteethorn concluded that the closest taxon in dentition to Siamosaurus was Spinosaurus aegyptiacus from Egypt, whose fragmentary fossils had been destroyed during World War II. Like Siamosaurus, this African taxon had straight and unserrated conical teeth. Though Spinosaurus lacked the developed flutes seen in Siamosaurus, Buffetaut and Ingavat noted that both smooth and fluted spinosaur teeth had been reported from Africa. Therefore, they tentatively placed Siamosaurus in the family Spinosauridae, based on the close similarities in dentition to S. aegyptiacus.[1][29]
Many palaeontologists later questioned Buffetaut and Ingavat's identification of Siamosaurus, given that spinosaurid teeth, including many from Asia, have often been mistaken for those of aquatic reptiles like crocodilians,
Authors such as Buffetaut and Ingavat in 1986, and Hasegawa and colleagues in 2003, have noted that since crocodilian teeth are usually more strongly recurved than spinosaur teeth, they can be distinguished from each other.[1][23] Crocodilians also lack the lateral double recurvature of Siamosaurus's tooth crowns, which, based on their shape, were vertically inserted into the jaw, whereas long-snouted crocodilian teeth are usually angled outwards from the mouth. Though Siamosaurus and plesiosaur teeth are similar in overall shape, Buffetaut and Ingavat pointed out that plesiosaur teeth were significantly more recurved.[1] Other researchers also noted that compared to plesiosaurs, Asian spinosaurid teeth also have coarser and more numerous flutes that extend almost the whole length of the crown.[1][20][23] In 2008, Buffetaut and colleagues stated that the "S." fusuiensis teeth bear carinae on the plane of the crown's curvature, a condition not observed in plesiosaur teeth. The discovery of the Khok Kruat skeleton and of baryonychine teeth with dental flutes similar to those of Siamosaurus, were also brought up by the researchers as further evidence of Siamosaurus's spinosaurid classification.[20] Later discoveries revealed that largely straight tooth crowns with flutes and a lack or reduction of serrations were unique characteristics of spinosaurid teeth.[37][48]
In 2014, Italian palaeontologist Federico Fanti and colleagues considered the various spinosaurid teeth from East Asia, including those of S. suteethorni, as identical to those of Spinosaurus.[49] In 2017, Brazilian palaeontologists Marcos Sales and Cesar Schultz suggested that the various Asian teeth might eventually be attributed to Ichthyovenator-like forms. The researchers accepted Siamosaurus as a spinosaurid, but stated that its teeth and those of "S." fusuiensis are too similar to those of other Early Cretaceous spinosaurids to erect new taxa unequivocally; and thus considered both taxa as dubious.[48] Carrano and colleagues noted that the Khok Kruat skeleton may provide answers to their identification.[47] Authors such as Milner and colleagues in 2007,[28] Bertin Tor in 2010,[29] Holtz in 2011,[50] and Kamonrak and colleagues in 2019 regarded the Khok Kruat skeleton as first definitive evidence of spinosaurs in Asia.[17] In 2012, French palaeontologist Ronan Allain and colleagues described a partial skeleton from the Grès supérieurs Formation of Laos, and used it to name the new spinosaurid genus and species Ichthyovenator laosensis. They considered it the first definitive evidence of spinosaurids in Asia, in light of the debated identity of Siamosaurus and "S." fusuiensis.[42] In a 2014 abstract, Allain announced that further Ichthyovenator material, including three teeth, had been excavated. Typically of spinosaurines, Ichthyovenator's teeth bore straight and unserrated crowns, though no comparison was made to the other Asian teeth.[51]
The
Spinosauridae |
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Later in 2019, the Khok Kruat Formation teeth were also referred to the Spinosaurinae by Kamonrak and colleagues, on the basis that both the Khok Kruat and Siamosaurus morphotypes lack characteristics seen in baryonychines, such as long and slender roots, 0–10 flutes on each side, no well defined carinae, a sculptured surface of the crown base, and 45 degree orientation of the blood grooves. But they share with spinosaurines a sub-circular to oval cross-section, fluted tooth crowns, well defined front and rear carinae, distinct striations on the crown, varying denticle size, and a wrinkled surface of the crown base. The authors also noted that unlike spinosaurines such as Irritator and Spinosaurus, Asian spinosaurines usually have more laterally compressed tooth crowns, and wrinkles across more of the enamel surface.[6] In 2020, a paper by British palaeontologist Robert Smyth and colleagues considered S. suteethorni a dubious name and attributed its teeth to an indeterminate spinosaurine, given the uncertainties of classifying spinosaurid teeth at the genus or species level, as well as the degree of heterodonty (variation within the tooth row) that spinosaurines apparently exhibited.[53] Due to new discoveries and research on spinosaurid teeth since Siamosaurus was named in 1986, a reassessment of the genus' validity is currently being prepared by Buffetaut.[21][29]
Palaeobiology
Diet and feeding
Buffetaut and Ingavat suggested in 1986 that Siamosaurus probably led a heavily
In 2006, Thai biologist Komsorn Lauprasert examined fossils collected from the Phu Kradung, Sao Khua, and Khok Kruat Formations. In this study, the teeth of Siamosaurus and a Moroccan spinosaurid were compared to those of crocodilians using
Aquatic habits
In 2008, French palaeontologist Romain Amiot and colleagues compared the
Palaeoenvironment and palaeobiogeography
Of all the
Besides Siamosaurus, there were theropod dinosaurs like the
The Khok Kruat Formation is composed mostly of sandstones, conglomerates, siltstones, and
In 2007, Milner and colleagues suggested that spinosaurids and iguanodontians may have spread from western to eastern
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External links
- Media related to Siamosaurus at Wikimedia Commons
- Data related to Siamosaurus at Wikispecies