Siamosaurus
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''Siamosaurus'' (meaning "Siam lizard") is a potentially dubious
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
spinosaurid Spinosauridae (or spinosaurids) is a clade or Family (taxonomy), family of tetanuran theropod dinosaurs comprising ten to seventeen known genera. Spinosaurid fossils have been recovered worldwide, including Africa, Europe, South America, and Asia. ...
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
that lived in what is now
Thailand Thailand, officially the Kingdom of Thailand and historically known as Siam (the official name until 1939), is a country in Southeast Asia on the Mainland Southeast Asia, Indochinese Peninsula. With a population of almost 66 million, it spa ...
and possibly
China China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...
during the
Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name) is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 143.1 ...
period Period may refer to: Common uses * Period (punctuation) * Era, a length or span of time *Menstruation, commonly referred to as a "period" Arts, entertainment, and media * Period (music), a concept in musical composition * Periodic sentence (o ...
(
Barremian The Barremian is an age in the geologic timescale (or a chronostratigraphic stage) between 125.77 Ma (million years ago) and 121.4 ± 1.0 Ma (Historically, this stage was placed at 129.4 million to approximately 125 million years ago) It is a ...
to
Aptian The Aptian is an age (geology), age in the geologic timescale or a stage (stratigraphy), stage in the stratigraphic column. It is a subdivision of the Early Cretaceous, Early or Lower Cretaceous epoch (geology), Epoch or series (stratigraphy), S ...
) and is the first reported spinosaurid from Asia. It is confidently known only from tooth
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserve ...
s; the first were found in the
Sao Khua Formation The Sao Khua Formation () is a middle member of the Khorat Group. It consists of an alteration of pale red to yellowish-gray, fine to medium-grained sandstone and grayish-reddish brown siltstone and clay. Rare pale red to light gray conglomerates ...
, with more teeth later recovered from the younger
Khok Kruat Formation The Khok Kruat Formation () is a geological formation, rock formation found in northeastern Thailand. It is the uppermost formation of the Khorat Group. It is dated to the Aptian Geological stage, stage of the Early Cretaceous Geological period, ...
. The only species ''Siamosaurus suteethorni'', whose name honours Thai
palaeontologist Paleontology, also spelled as palaeontology or palæontology, is the scientific study of the life of the past, mainly but not exclusively through the study of fossils. Paleontologists use fossils as a means to classify organisms, measure geolo ...
Varavudh Suteethorn Varavudh Suteethorn, or Warawut Suteethorn (Thai language, Thai:วราวุธ สุธีธร; born 10 October 1948) is a Thai palaeontologist and geologist. He is the current director of the Palaeontological Research and Education Centre ...
, was formally described in 1986. In 2009, four teeth from China previously attributed to a
pliosaur Pliosauroidea is an extinct clade of plesiosaurs, known from the earliest Jurassic to early Late Cretaceous. They are best known for the subclade Thalassophonea, which contained crocodile-like short-necked forms with large heads and massive toot ...
—under the species "''Sinopliosaurus''" ''fusuiensis''—were identified as those of a spinosaurid, possibly ''Siamosaurus''. It is yet to be determined if two partial spinosaurid skeletons from Thailand and an isolated tooth from Japan also belong to ''Siamosaurus''. Since it is based only on teeth, ''Siamosaurus'' body size is uncertain, though it has been estimated at between in length. The
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
tooth is long. ''Siamosaurus''s teeth were straight, oval to circular in cross-section, and lined with distinct lengthwise grooves. Its teeth had wrinkled enamel, similar to teeth from the related genus ''
Baryonyx ''Baryonyx'' () is a genus of theropod dinosaur which lived in the Barremian stage of the Early Cretaceous period, about 130–125 million years ago. The first skeleton was discovered in 1983 in the Smokejack Clay Pit, of Surrey, England, in ...
''. As a spinosaur it would have had a long, low snout and robust forelimbs, and one possible skeleton indicates the presence of a tall
sail A sail is a tensile structure, which is made from fabric or other membrane materials, that uses wind power to propel sailing craft, including sailing ships, sailboats, windsurfers, ice boats, and even sail-powered land vehicles. Sails may b ...
running down its back, another typical trait of this theropod
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
. ''Siamosaurus'' is considered by some palaeontologists to be a
dubious name In binomial nomenclature, a ''nomen dubium'' (Latin for "doubtful name", plural ''nomina dubia'') is a scientific name that is of unknown or doubtful application. Zoology In case of a ''nomen dubium,'' it may be impossible to determine whether a ...
, with some arguing that its teeth are hard to differentiate from those of other Early Cretaceous spinosaurids, and others that it may not be a dinosaur at all. Based on dental traits, ''Siamosaurus'' and "''S.''" ''fusuiensis'' have been placed in the
subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end botanical subfamily names with "-oideae", and zo ...
Spinosaurinae Spinosauridae (or spinosaurids) is a clade or family of tetanuran theropod dinosaurs comprising ten to seventeen known genera. Spinosaurid fossils have been recovered worldwide, including Africa, Europe, South America, and Asia. Their remains have ...
. Like in all spinosaurids, ''Siamosaurus'' teeth were conical, with reduced or absent serrations. This made them suitable for impaling rather than tearing flesh, a trait typically seen in largely
piscivorous A piscivore () is a carnivorous animal that primarily eats fish. Fish were the diet of early tetrapod evolution (via water-bound amphibians during the Devonian period); insectivory came next; then in time, the more terrestrially adapted rept ...
(fish-eating) animals. Spinosaurids are also known to have consumed
pterosaur Pterosaurs are an extinct clade of flying reptiles in the order Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 million to 66 million years ago). Pterosaurs are the earli ...
s and small dinosaurs, and there is fossil evidence of ''Siamosaurus'' itself feeding on
sauropod Sauropoda (), whose members are known as sauropods (; from '' sauro-'' + '' -pod'', 'lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their b ...
dinosaurs, either via
scavenging Scavengers are animals that consume dead organisms that have died from causes other than predation or have been killed by other predators. While scavenging generally refers to carnivores feeding on carrion, it is also a herbivorous feeding be ...
or active hunting. ''Siamosaurus'' role as a partially piscivorous predator may have reduced the prominence of some contemporaneous crocodilians competing for the same food sources.
Isotope analysis Isotope analysis is the identification of isotopic signature, abundance of certain stable isotopes of chemical elements within organic and inorganic compounds. Isotopic analysis can be used to understand the flow of energy through a food we ...
of the teeth of ''Siamosaurus'' and other spinosaurids indicates
semiaquatic In biology, being semi-aquatic refers to various macroorganisms that live regularly in both aquatic and terrestrial environments. When referring to animals, the term describes those that actively spend part of their daily time in water (in ...
habits. ''Siamosaurus'' lived in a
semi-arid A semi-arid climate, semi-desert climate, or steppe climate is a aridity, dry climate sub-type. It is located on regions that receive precipitation below Evapotranspiration#Potential evapotranspiration, potential evapotranspiration, but not as l ...
habitat of
floodplain A floodplain or flood plain or bottomlands is an area of land adjacent to a river. Floodplains stretch from the banks of a river channel to the base of the enclosing valley, and experience flooding during periods of high Discharge (hydrolog ...
s and meandering rivers, where it coexisted with other dinosaurs, as well as pterosaurs, fishes, turtles,
crocodyliforms Crocodyliformes is a clade of crurotarsan archosaurs, the group often traditionally referred to as "crocodilians". They are the first members of Crocodylomorpha to possess many of the features that define later relatives. They are the only pseudo ...
, and other aquatic animals.


History of discovery

The
Sao Khua Formation The Sao Khua Formation () is a middle member of the Khorat Group. It consists of an alteration of pale red to yellowish-gray, fine to medium-grained sandstone and grayish-reddish brown siltstone and clay. Rare pale red to light gray conglomerates ...
, where the first ''Siamosaurus''
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserve ...
s were discovered, is part of the Khorat Group. The formation is dated to the
Barremian The Barremian is an age in the geologic timescale (or a chronostratigraphic stage) between 125.77 Ma (million years ago) and 121.4 ± 1.0 Ma (Historically, this stage was placed at 129.4 million to approximately 125 million years ago) It is a ...
stage Stage, stages, or staging may refer to: Arts and media Acting * Stage (theatre), a space for the performance of theatrical productions * Theatre, a branch of the performing arts, often referred to as "the stage" * ''The Stage'', a weekly Brit ...
of the
Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name) is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 143.1 ...
period Period may refer to: Common uses * Period (punctuation) * Era, a length or span of time *Menstruation, commonly referred to as a "period" Arts, entertainment, and media * Period (music), a concept in musical composition * Periodic sentence (o ...
, 129.4 to 125 million years ago. In 1983, French
palaeontologist Paleontology, also spelled as palaeontology or palæontology, is the scientific study of the life of the past, mainly but not exclusively through the study of fossils. Paleontologists use fossils as a means to classify organisms, measure geolo ...
Éric Buffetaut Éric Buffetaut (born 19 November 1950) is a French paleontologist, author and researcher at the Centre national de la recherche scientifique since 1976 where he is a Doctor of Science and Director of Research. Buffetaut is a specialist of fossil ...
and his Thai colleague Rucha Ingavat described a set of fossil teeth recovered from the Phu Pratu Teema locality of the Sao Khua Formation, in the Phu Wiang area of Khon Kaen Province. They did not conclude as to what animal they originated from, their opinion being that the specimens belonged "either to an unusual
theropod Theropoda (; from ancient Greek , (''therion'') "wild beast"; , (''pous, podos'') "foot"">wiktionary:ποδός"> (''pous, podos'') "foot" is one of the three major groups (clades) of dinosaurs, alongside Ornithischia and Sauropodom ...
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
or to some unknown
crocodilia Crocodilia () is an order of semiaquatic, predatory reptiles that are known as crocodilians. They first appeared during the Late Cretaceous and are the closest living relatives of birds. Crocodilians are a type of crocodylomorph pseudosuchia ...
n". In
1986 The year 1986 was designated as the International Year of Peace by the United Nations. Events January * January 1 ** Aruba gains increased autonomy from the Netherlands by separating from the Netherlands Antilles. ** Spain and Portugal en ...
, a reassessment of the remains by the same authors attributed them to a new
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
and
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
of
spinosaurid Spinosauridae (or spinosaurids) is a clade or Family (taxonomy), family of tetanuran theropod dinosaurs comprising ten to seventeen known genera. Spinosaurid fossils have been recovered worldwide, including Africa, Europe, South America, and Asia. ...
theropod, which they named ''Siamosaurus suteethorni''. The generic name alludes to the ancient name of Thailand, "Siam", and is combined with the
Ancient Greek Ancient Greek (, ; ) includes the forms of the Greek language used in ancient Greece and the classical antiquity, ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Greek ...
word ('), meaning "lizard" or "reptile". The specific name honours Thai
geologist A geologist is a scientist who studies the structure, composition, and History of Earth, history of Earth. Geologists incorporate techniques from physics, chemistry, biology, mathematics, and geography to perform research in the Field research, ...
and palaeontologist
Varavudh Suteethorn Varavudh Suteethorn, or Warawut Suteethorn (Thai language, Thai:วราวุธ สุธีธร; born 10 October 1948) is a Thai palaeontologist and geologist. He is the current director of the Palaeontological Research and Education Centre ...
, and his contributions to
vertebrate Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...
palaeontology discoveries in Thailand. The best-preserved specimen from the teeth described, designated DMR TF 2043a, was chosen as the
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
of ''Siamosaurus.'' The
paratype In zoology and botany, a paratype is a specimen of an organism that helps define what the scientific name of a species and other taxon actually represents, but it is not the holotype (and in botany is also neither an isotype (biology), isotype ...
s 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 Department of Mineral Resources may refer to: * Department of Mineral Resources (South Africa) * Department of Mineral Resources (Thailand) {{Disambiguation ...
,
Bangkok Bangkok, officially known in Thai language, Thai as Krung Thep Maha Nakhon and colloquially as Krung Thep, is the capital and most populous city of Thailand. The city occupies in the Chao Phraya River delta in central Thailand and has an estim ...
. ''Siamosaurus'' teeth are common in the Sao Khua Formation, and further isolated specimens were found later throughout the late 20th and early 21st centuries. Sculptures of the animal have been erected in various places across northeastern Thailand, including Si Wiang Dinosaur Park, the
Siam Paragon Siam Paragon () is a shopping mall in Bangkok, Thailand. It is one of the largest malls in Thailand, along with Iconsiam, CentralWorld and Central WestGate. It is located right in front of Siam BTS station, an interchange station on both Sukhumvi ...
shopping mall in Bangkok, the
Phu Wiang Dinosaur Museum Phu Wiang Dinosaur Museum () is a geological museum mainly exhibiting fossils. It is under the administration of the Department of Mineral Resources, Ministry of Natural Resources and Environment, of the Royal Thai Government, and situated in th ...
, and the
Sirindhorn Museum Sirindhorn Museum (aka Sirindhorn Dinosaur Museum) () is a geology museum in northeast Thailand. It is located at Non Buri, Sahatsakhan district, Kalasin province. About Dinosaur fossils were found by Phra Kru Vijitsahassakun, the abbot of Wa ...
. ''S''. ''suteethorni'' was also illustrated on Thai postage stamps released in 1997, along with fellow Thai dinosaurs ''
Phuwiangosaurus ''Phuwiangosaurus'' (meaning "Phu Wiang lizard") is a genus of titanosaur dinosaur from the Early Cretaceous (Valanginian-Hauterivian) Sao Khua Formation of Thailand. The type species, ''P. sirindhornae'', was described by Martin, Buffetaut, and ...
,'' ''
Siamotyrannus ''Siamotyrannus'' (meaning "Siamese tyrant") is a genus of theropod dinosaur from the early Cretaceous of Thailand. Discovery and naming In 1993, Somchai Traimwichanon found a partial skeleton of a large theropod at the Phu Wiang 9 site in Khon K ...
'', and ''
Psittacosaurus ''Psittacosaurus'' ( ; "parrot lizard") is a genus of extinct ceratopsian dinosaur from the Early Cretaceous of what is now Asia, existing between 125 and 105 million years ago. It is notable for being the most species-rich non-avian dinosaur ...
''. Thailand's
Khok Kruat Formation The Khok Kruat Formation () is a geological formation, rock formation found in northeastern Thailand. It is the uppermost formation of the Khorat Group. It is dated to the Aptian Geological stage, stage of the Early Cretaceous Geological period, ...
is dated to the
Aptian The Aptian is an age (geology), age in the geologic timescale or a stage (stratigraphy), stage in the stratigraphic column. It is a subdivision of the Early Cretaceous, Early or Lower Cretaceous epoch (geology), Epoch or series (stratigraphy), S ...
age (between 125 and 113 million years ago), younger than the Sao Khua Formation. The Khok Kruat Formation has provided many spinosaurid teeth, including ones from ''Siamosaurus'' and closely allied forms. Given the varied size and
morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines *Morphology (archaeology), study of the shapes or forms of artifacts *Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies, ...
of the teeth found, the presence of multiple spinosaur
taxa In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
in the region is likely.'''' Nearly 60 fossil teeth were recovered from the formation during fieldwork by Thai-French palaeontological teams between 2003 and 2008, including specimens from the Sam Ran, Khok Pha Suam, and Lam Pao Dam localities. Eight of these teeth were described in detail by Kamonrak and colleagues in 2019, and classified into two main
morphotype In biology, polymorphism is the occurrence of two or more clearly different morphs or forms, also referred to as alternative ''phenotypes'', in the population of a species. To be classified as such, morphs must occupy the same habitat at the s ...
s: the Khok Kruat morphotype, which is found only in the Khok Kruat Formation, and the ''Siamosaurus'' morphotype, which includes forms widely recovered from both the Sao Khua and Khok Kruat Formations.'''' ''Siamosaurus'' is the first reported spinosaurid dinosaur from Asia, and subsequently to its naming, material resembling or possibly belonging to the genus has been found across the continent. In
1975 It was also declared the ''International Women's Year'' by the United Nations and the European Architectural Heritage Year by the Council of Europe. Events January * January 1 – Watergate scandal (United States): John N. Mitchell, H. R. ...
, 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 Fusui County is a county (China), county in the southwest of Guangxi, China. It is the easternmost county-level division of the prefecture-level city of Chongzuo. Geography Fusui is located in southwestern Guangxi and in eastern Chongzuo City. I ...
in
Guangxi Guangxi,; officially the Guangxi Zhuang Autonomous Region, is an Autonomous regions of China, autonomous region of the China, People's Republic of China, located in South China and bordering Vietnam (Hà Giang Province, Hà Giang, Cao Bằn ...
, China, from which the fossils were collected. 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.'' In 2019, "''S''." ''fusuiensis'' was referred to by Thai palaeontologist Wongko Kamonrak and colleagues as ''Siamosaurus'' sp. (of uncertain species).'''' Later in 2019, Thai palaeontologist Adun Samathi and colleagues considered the teeth as belonging to an indeterminate spinosaurid. The specimens were retrieved from China's Early Cretaceous
Xinlong Formation The Xinlong Formation (sometimes called the "Napai Formation", or misspelt as "Napan Formation") is an Early Cretaceous geologic Formation (geology), formation in Guangxi, southern China. 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. In 1994, an isolated tooth (specimen GMNH-PV-999) was found by a fossil prospector in the
Sebayashi Formation The Sebayashi Formation is a Barremian to Albian geologic formation in Japan.
, Japan. The tooth was believed, until 2003, to belong to a
marine reptile Marine reptiles are reptiles which have become secondarily adapted for an aquatic or semiaquatic life in a marine environment. Only about 100 of the 12,000 extant reptile species and subspecies are classed as marine reptiles, including mari ...
, when Japanese palaeontologist Yoshikazu Hasegawa and colleagues assigned it to ?''Siamosaurus ''sp. The tooth came from rocks dated to the Barremian, similar in age to
sediment Sediment is a solid material that is transported to a new location where it is deposited. It occurs naturally and, through the processes of weathering and erosion, is broken down and subsequently sediment transport, transported by the action of ...
s that ''Siamosaurus'' teeth have been recovered from in Thailand. 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. Further spinosaurid teeth from unnamed and indeterminate forms have been discovered in central China and
Malaysia Malaysia is a country in Southeast Asia. Featuring the Tanjung Piai, southernmost point of continental Eurasia, it is a federation, federal constitutional monarchy consisting of States and federal territories of Malaysia, 13 states and thre ...
. In 2004, excavation began on a partial skeleton from an outcrop of the Khok Kruat Formation near the city of
Khon Kaen Khon Kaen (, ) is the capital of Khon Kaen province and the List of municipalities in Thailand#Largest cities by urban population, fourth-largest city in Thailand. It is one of the four major cities of Isan, Northeast Thailand, also known as the ...
. The specimen (SM-KK 14) consists of cervical (neck) and
dorsal Dorsal (from Latin ''dorsum'' ‘back’) may refer to: * Dorsal (anatomy), an anatomical term of location referring to the back or upper side of an organism or parts of an organism * Dorsal, positioned on top of an aircraft's fuselage The fus ...
(back)
vertebrae Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spinal ...
, a high
neural spine Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spinal ...
(upwards-extending
process A process is a series or set of activities that interact to produce a result; it may occur once-only or be recurrent or periodic. Things called a process include: Business and management * Business process, activities that produce a specific s ...
from top of vertebra),
pelvis The pelvis (: pelves or pelvises) is the lower part of an Anatomy, anatomical Trunk (anatomy), trunk, between the human abdomen, abdomen and the thighs (sometimes also called pelvic region), together with its embedded skeleton (sometimes also c ...
(hip) fragments, a possible
metacarpal In human anatomy, the metacarpal bones or metacarpus, also known as the "palm bones", are the appendicular bones that form the intermediate part of the hand between the phalanges (fingers) and the carpal bones ( wrist bones), which articulate ...
(long bone of the hand), and a
chevron Chevron (often relating to V-shaped patterns) may refer to: Science and technology * Chevron (aerospace), sawtooth patterns on some jet engines * Chevron (anatomy), a bone * '' Eulithis testata'', a moth * Chevron (geology), a fold in rock la ...
from the tail. The cervical vertebrae and pelvic region resemble those of the European spinosaur ''
Baryonyx walkeri ''Baryonyx'' () is a genus of theropod dinosaur which lived in the Barremian Geological stage, stage of the Early Cretaceous Period (geology), period, about 130–125 million years ago. The first skeleton was discovered in 1983 in the Smokejack ...
'', and the neural spine is elongated, similarly to those of other spinosaurids. A ''Siamosaurus'' tooth found nearby indicates the skeleton may belong to this genus, though this could also represent evidence of
scavenging Scavengers are animals that consume dead organisms that have died from causes other than predation or have been killed by other predators. While scavenging generally refers to carnivores feeding on carrion, it is also a herbivorous feeding be ...
. The skeleton, as well as two well-preserved teeth—SM2016-1-147 and SM2016-1-165, also attributed to ''Siamosaurus—''are currently stored in the vertebrate fossil collection of the Sirindhorn Museum, Kalasin Province. In 2019, a series of spinosaurid
caudal Caudal may refer to: Anatomy * Caudal (anatomical term) (from Latin ''cauda''; tail), used to describe how close something is to the trailing end of an organism * Caudal artery, the portion of the dorsal aorta of a vertebrate that passes into th ...
(tail) vertebrae possibly belonging to ''S''. ''suteethorni'' were recovered from the Sao Khua Formation, and described in a dissertation by Samathi. The fossils (SM-PW9B-11 to 17, SM-PW9B, SM-PW9A-unnumbered, SM-PW9-unnumbered, and SM 2017-1-176) were designated by Samathi as "Phuwiang spinosaurid B", and bear similarities to the possible spinosaurid ''
Camarillasaurus ''Camarillasaurus'' (meaning " Camarillas lizard") is a genus of theropod dinosaur from the Early Cretaceous period (Barremian) of Camarillas, Teruel Province, in what is now northeastern Spain. Described in 2014, it was originally identified as ...
'' and a ''Baryonyx'' specimen discovered in Portugal.


Description

In 2004, American dinosaur researcher
Don Lessem "Dino" Don Lessem (born 1951) is a writer of more than 50 popular science books, specializing in dinosaurs. He was the founder of the Dinosaur Society and the Jurassic Foundation, which collectively have raised millions of dollars for dinosaur rese ...
estimated ''Siamosaurus'' at long. In 2005, British author Sussana Davidson and colleagues gave a lower estimate of in length and weighing . In a 2016 popular book, authors Rubén Molina-Pérez and Asier Larramendi estimated it at approximately long, tall at the hips, and weighing . 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 The postcranium ("behind the cranium"; plural: postcrania) or postcranial skeleton in zoology and vertebrate paleontology is the skeleton apart from the skull. The postcranium encompasses the axial skeleton, which includes the entirety of the verte ...
skeleton, and thus estimates are only tentative. "Phuwiang spinosaurid B" was calculated as approximately long by Samathi in 2019. 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 A sail is a tensile structure, which is made from fabric or other membrane materials, that uses wind power to propel sailing craft, including sailing ships, sailboats, windsurfers, ice boats, and even sail-powered land vehicles. Sails may b ...
on its back.


Type specimens

Fossil theropod teeth are typically identified by attributes such as the proportions, size, and curvature of the
crown A crown is a traditional form of head adornment, or hat, worn by monarchs as a symbol of their power and dignity. A crown is often, by extension, a symbol of the monarch's government or items endorsed by it. The word itself is used, parti ...
, as well as the presence and/or shape of the denticles (serrations). The holotype of ''S''. ''suteethorni'' (specimen DMR TF 2043a) is in total length, with the crown being long, and wide at its base. It is among the larger teeth discovered by Buffetaut and Ingavat. One much smaller specimen (DMR TF 2043b) measures in length. According to the authors, this dramatic size range suggests the teeth are from individuals of different ages. Among theropods, this may also indicate size variation along the tooth row in the jaws, which is observed to have been prevalent in spinosaurids. 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 ''Carinae'' was an area of ancient Rome. It was one of its most exclusive neighborhoods, where many of the senatorial class lived. Florus described the ''Carinae'' as the "''most celebrated part of the city''" (''celeberrima pars urbis''). Desc ...
(cutting edges) of ''Siamosaurus'' teeth lack well-defined serrations, though unworn teeth do exhibit very fine denticles. 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 Carnosauria is an extinct group of carnivorous theropod dinosaurs that lived during the Jurassic and Cretaceous periods. While Carnosauria was historically considered largely synonymous with Allosauroidea, some recent studies have revived Carn ...
teeth from the same formation and one ''
Deinonychus ''Deinonychus'' ( ; ) is a genus of Dromaeosauridae, dromaeosaurid Theropoda, theropod dinosaur with one described species, ''Deinonychus antirrhopus''. This species, which could grow up to long, lived during the early Cretaceous Period (ge ...
'' tooth described by
John Ostrom John Harold Ostrom (February 18, 1928 – July 16, 2005) was an American paleontologist who revolutionized the modern understanding of dinosaurs. Ostrom's work inspired what his pupil Robert T. Bakker has termed a " dinosaur renaissance". Begin ...
in 1969. The ''S''. ''suteethorni'' holotype is symmetrically concave front to back, and bears 15 flutes (lengthwise grooves) on its
lingual Lingual may refer to: * Tongue, a muscular hydrostat on the floors of the mouths of most vertebrates which manipulates food for mastication * Lingual, in palaeontology, the side of the teeth that faces the tongue * Lingual artery arises from the e ...
(inward facing) and labial (outward facing) surfaces. These flutes run from the base of the crown before stopping 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 Dentin ( ) (American English) or dentine ( or ) (British English) () is a calcified tissue (biology), tissue of the body and, along with tooth enamel, enamel, cementum, and pulp (tooth), pulp, is one of the four major components of teeth. It i ...
(second layer of the teeth). The enamel also has a granular (finely wrinkled) texture, as seen in teeth from the spinosaurid ''Baryonyx''. Some of the
root In vascular plants, the roots are the plant organ, organs of a plant that are modified to provide anchorage for the plant and take in water and nutrients into the plant body, which allows plants to grow taller and faster. They are most often bel ...
is preserved in the holotype and, as in all theropods, there is a large pocket for the tooth
pulp Pulp may refer to: * Pulp (fruit), the inner flesh of fruit * Pulp (band), an English rock band Engineering * Pulp (paper), the fibrous material used to make paper * Dissolving pulp, highly purified cellulose used in fibre and film manufacture ...
, which would have housed blood vessels and nerves.


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 in total length, of which the crown takes up , with a 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 long, with a tall crown that is 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.''''


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, wide base and being slightly smaller at 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. The blood grooves (tiny furrows in the gaps between each denticle) of GMNH-PV-999 have an
oblique Oblique may refer to: * an alternative name for the character usually called a slash (punctuation) ( / ) *Oblique angle, in geometry * Oblique triangle, in geometry * Oblique lattice, in geometry * Oblique leaf base, a characteristic shape of the ...
orientation of 45 degrees, as in ''Baryonyx'' and KDC-PV-0003, the second Sebayashi formation tooth, which consists of a slightly recurved crown fragment with an almost circular cross-section. It has better preservation of small details than the former specimens, such as visible, though poorly defined serrations, with two to three denticles per mm (0.039 in). Like GMNH-PV-999, it has a granular texture and at least 12 flutes on its surface, not all of which stretch to the crown's full length. Out of the four teeth attributed to "''S''." ''fusuiensis'', specimen
IVPP The Institute of Vertebrate Paleontology and Paleoanthropology (IVPP; ) of China is a research institution and collections repository for fossils, including many dinosaur and pterosaur specimens (many from the Yixian Formation). As its name sugges ...
 V 4793 is the most intact, although still somewhat deformed. The crown, which is missing its tip, is long and wide at the base. The tooth is straight, only slightly recurved, and has an oval cross-section. The front and rear carinae are distinct, though their serrations have been heavily eroded, similar to those of KDC-PV-0003. Like the Thai and Japanese teeth, the "''S''." ''fusuiensis'' specimens bear developed flutes and a granular surface. As in both Sebayashi Formation teeth, there are 12 flutes on each face of the "''S''." ''fusuiensis'' teeth. Like in KDC-PV-0003, these flutes vary in length. Buffetaut and colleagues found the "''S''." ''fusuiensis'' teeth most similar to those of ''Siamosaurus'', given their identical crown shape, fluting, and granular enamel.


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. The cervical vertebrae of SM-KK 14 had elongated
centra Centra is a convenience shop chain that operates throughout Ireland. The chain operates as a symbol group owned by Musgrave Group, the food wholesaler, meaning the individual shops are all owned by individual franchisees. The chain has three ...
(vertebral bodies) with articulating surfaces that were not offset, as well as prominent
epipophyses Epipophyses are bony projections of the cervical vertebrae found in archosauromorphs, particularly dinosaurs (including some basal birds). These paired processes sit above the postzygapophyses on the rear of the vertebral neural arch. Their m ...
( processes to which neck muscles attached) and strong
ligament A ligament is a type of fibrous connective tissue in the body that connects bones to other bones. It also connects flight feathers to bones, in dinosaurs and birds. All 30,000 species of amniotes (land animals with internal bones) have liga ...
scars. All of these characteristics were also present in ''Baryonyx''. 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 The articular process or zygapophysis ( + apophysis) of a vertebra is a projection of the vertebra that serves the purpose of fitting with an adjacent vertebra. The actual region of contact is called the ''articular facet''.Moore, Keith L. et al. ...
, which connect adjacent vertebrae—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 ''Ichthyovenator'' is a genus of spinosaurid dinosaur that lived in what is now Laos, sometime between 120 and 113 million years ago, during the Aptian Stage (geology), stage of the Early Cretaceous Period (geologic time), period. It is k ...
''. One of the neural spines of SM-KK 14 measured at least in height. 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 ''Suchomimus'', from Ancient Greek ''σούχος'' (''soúkhos''), meaning "crocodile", and Latin ''mimus'', meaning "actor", is a genus of spinosaurid dinosaur that lived between 125 and 112 million years ago in what is now Niger, North A ...
''. Also as in ''Ichthyovenator'', the hind rim of the pubis had a notch-like
obturator foramen The obturator foramen is the large, Bilateral symmetry, bilaterally paired opening of the bony pelvis. It is formed by the pubis and ischium. It is mostly closed by the obturator membrane except for a small opening, the obturator canal, through wh ...
. 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''.


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
ceratosauria Ceratosaurs are members of the clade Ceratosauria, a group of dinosaurs defined as all theropods sharing a more recent common ancestor with '' Ceratosaurus'' than with birds. The oldest known ceratosaur, '' Saltriovenator'', dates to the earlies ...
n 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 Dentition pertains to the development of teeth and their arrangement in the mouth. In particular, it is the characteristic arrangement, kind, and number of teeth in a given species at a given age. That is, the number, type, and morpho-physiology ...
to ''Siamosaurus'' was ''Spinosaurus aegyptiacus'' from Egypt, whose fragmentary fossils had been destroyed during
World War II World War II or the Second World War (1 September 1939 – 2 September 1945) was a World war, global conflict between two coalitions: the Allies of World War II, Allies and the Axis powers. World War II by country, Nearly all of the wo ...
. 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 Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
Spinosauridae, based on the close similarities in dentition to ''S''. ''aegyptiacus.'' 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,
plesiosaurs The Plesiosauria or plesiosaurs are an Order (biology), order or clade of extinct Mesozoic marine reptiles, belonging to the Sauropterygia. Plesiosaurs first appeared in the latest Triassic Period (geology), Period, possibly in the Rhaetian st ...
, and
ichthyosaur Ichthyosauria is an order of large extinct marine reptiles sometimes referred to as "ichthyosaurs", although the term is also used for wider clades in which the order resides. Ichthyosaurians thrived during much of the Mesozoic era; based on fo ...
s. In view of this, the German palaeontologist
Hans-Dieter Sues Hans-Dieter Sues (born 1956) is a German-born American palaeontologist who is a Senior Research Geologist and Curator of Vertebrate Paleontology at the National Museum of Natural History of the Smithsonian Institution in Washington, DC. Career ...
and colleagues in 2002 asserted that there is not enough material to confidently identify ''Siamosaurus'' as a dinosaur. In 2004, American palaeontologist
Thomas Holtz Thomas Richard Holtz Jr. (born September 13, 1965) is an American vertebrate palaeontologist, author, and principal lecturer at the University of Maryland's Department of Geology. He has published extensively on the phylogeny, morphology, ecomorp ...
and colleagues considered it a
dubious name In binomial nomenclature, a ''nomen dubium'' (Latin for "doubtful name", plural ''nomina dubia'') is a scientific name that is of unknown or doubtful application. Zoology In case of a ''nomen dubium,'' it may be impossible to determine whether a ...
, stating that the teeth might instead belong to a contemporaneous fish such as a
saurodontid Ichthyodectiformes is an extinct order of marine stem-teleost ray-finned fish. The order is named after the genus ''Ichthyodectes'', established by Edward Drinker Cope in 1870. Ichthyodectiforms are usually considered to be some of the closest re ...
or an
ichthyodectid Ichthyodectiformes is an extinct order of marine stem-teleost ray-finned fish. The order is named after the genus '' Ichthyodectes'', established by Edward Drinker Cope in 1870. Ichthyodectiforms are usually considered to be some of the closest r ...
teleost Teleostei (; Ancient Greek, Greek ''teleios'' "complete" + ''osteon'' "bone"), members of which are known as teleosts (), is, by far, the largest group of ray-finned fishes (class Actinopterygii), with 96% of all neontology, extant species of f ...
. The same year, American palaeontologist
David Weishampel Professor David Bruce Weishampel (born November 16, 1952) is an American palaeontologist in the Center for Functional Anatomy and Evolution at Johns Hopkins University School of Medicine. Weishampel received his Ph.D. in Geology from the Univers ...
and colleagues considered ''Siamosaurus'' an indeterminate theropod. In 2012, an analysis by American palaeontologist Matthew Carrano and colleagues agreed with the possibility of confusion with other reptiles, and regarded the genus as a possible indeterminate spinosaurid. They noted that oftentimes, isolated teeth are an unstable foundation for naming new theropod taxa, and most species based on them turn out to be invalid. This problem is especially common with spinosaurids, given that skull and skeletal fossils from the group are rare. 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. 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. 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.'''' 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.'''' Later discoveries revealed that largely straight tooth crowns with flutes and a lack or reduction of serrations were unique characteristics of spinosaurid teeth. 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''. 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. Carrano and colleagues noted that the Khok Kruat skeleton may provide answers to their identification. Authors such as Milner and colleagues in 2007, Bertin Tor in 2010, Holtz in 2011, and Kamonrak and colleagues in 2019 regarded the Khok Kruat skeleton as first definitive evidence of spinosaurs in Asia. In 2012, French palaeontologist Ronan Allain and colleagues described a partial skeleton from the
Grès supérieurs Formation The Grès supérieurs Formation (French for "Upper sandstone") is a geological formation in Laos whose strata date back to the Aptian to Albian stages of the Early Cretaceous. Dinosaur remains are among the fossils that have been recovered from t ...
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.'' 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. The
taxonomic 280px, Generalized scheme of taxonomy Taxonomy is a practice and science concerned with classification or categorization. Typically, there are two parts to it: the development of an underlying scheme of classes (a taxonomy) and the allocation ...
and
phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...
affinities of the Spinosauridae are subject to active research and debate, given that in comparison to other theropod groups, many of the family's taxa are based on poor fossil material. Traditionally, the group is split into the
subfamilies In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end botanical subfamily names with "-oideae", and zool ...
Spinosaurinae (unserrated, straight teeth with well marked flutes and more circular cross-sections) and Baryonychinae (finely serrated, somewhat recurved teeth with weaker flutes and a more oval cross-section). Since spinosaurines were, on average, larger animals than baryonychines, their teeth were also generally larger. The morphological variation seen in spinosaurid teeth, however, has shown that the aforementioned characteristics are not always consistent within the subfamilies. Likewise, the Khok Kruat skeleton shares mixed characteristics between ''Baryonyx'' and ''Spinosaurus'', and its precise phylogenetic placement is uncertain pending a description of the material. The possibility that Baryonychinae is a
paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
(unnatural) grouping has been suggested by researchers such as Sales and Schultz, on the basis that genera such as ''
Irritator ''Irritator'' is a genus of Spinosauridae, spinosaurid dinosaur that lived in what is now Brazil during the Albian Geological stage, stage of the Early Cretaceous Geological period, Period, about 113 to 110 million years ago. It is known from ...
'' and ''Angaturama'' (the two are possible synonyms) may represent intermediate forms between baryonychines and spinosaurines. As it is definitively known only from teeth, ''Siamosaurus''s exact position within the Spinosauridae is difficult to determine. In 2004, Brazilian palaeontologists Elaine Machado and
Alexander Kellner Alexander Wilhelm Armin Kellner (born September 26, 1961) is a Brazilian geologist and paleontologist who is a leading expert in the field of studying pterosaurs. His research has focused mainly on fossil reptiles from the Cretaceous Period, i ...
suggested it as a possible spinosaurine, given its lack of dental serrations. Likewise in 2010, British palaeontologist David Hone and colleagues placed ''Siamosaurus'' and "''S''." ''fusuiensis'' in the Spinosaurinae. British palaeontologist Thomas Arden and colleagues identified ''Siamosaurus'' as a basal (early diverging or "primitive") member of this subfamily in 2019; their
cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...
can be seen below: 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.'''' 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
heterodont In anatomy, a heterodont (from Greek, meaning 'different teeth') is an animal which possesses more than a single tooth morphology. Human dentition is heterodont and diphyodont as an example. In vertebrates, heterodont pertains to animals wher ...
y (variation within the tooth row) that spinosaurines apparently exhibited. 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.


Palaeobiology


Diet and feeding

Buffetaut and Ingavat suggested in 1986 that ''Siamosaurus'' probably led a heavily
piscivorous A piscivore () is a carnivorous animal that primarily eats fish. Fish were the diet of early tetrapod evolution (via water-bound amphibians during the Devonian period); insectivory came next; then in time, the more terrestrially adapted rept ...
(fish-eating) lifestyle, since its dentition—like those of other spinosaurids—had a highly specialised morphology better suited for piercing rather than tearing flesh, due to the long, straight conical tooth crowns with reduced or absent serrations. The authors noted that this dental morphology is also seen in other piscivorous predators like plesiosaurs and long-snouted crocodilians. Such a dietary preference had been suggested for ''Baryonyx'' the same year by British palaeontologists
Angela Milner Angela Cheryl Milner (3 October 1947 – 13 August 2021) was a British paleontologist who, in 1986 alongside Alan Charig, described the dinosaur ''Baryonyx''. Early life Milner was born Angela Girven in Gosforth, daughter of Cyril and Luci ...
and Alan Charig, and was later confirmed in 1997 with the discovery of acid-etched fish scales inside the body cavity of its holotype skeleton. The elongate, interlocking jaws of spinosaurids also had snout tips that fanned out into a
rosette Rosette is the French diminutive of ''rose''. It may refer to: Flower shaped designs * Rosette (award), a mark awarded by an organisation * Rosette (design), a small flower design *hence, various flower-shaped or rotational symmetric forms: ** R ...
-like shape—a trait also observed in highly piscivorous crocodilians such as
gharials Gavialidae is a family of large semiaquatic crocodilians with elongated, narrow snouts. Gavialidae consists of two living species, the gharial (''Gavialis gangeticus'') and the false gharial (''Tomistoma schlegelii''), both occurring in Asia. Man ...
—which made them well-adapted to catching and feeding on fish. Fossil evidence has shown that besides aquatic prey, spinosaurids also consumed other dinosaurs and
pterosaur Pterosaurs are an extinct clade of flying reptiles in the order Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 million to 66 million years ago). Pterosaurs are the earli ...
s. In the Sao Khua Formation, localities such as Wat Sakawan have yielded
sauropod Sauropoda (), whose members are known as sauropods (; from '' sauro-'' + '' -pod'', 'lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their b ...
remains in association with tooth crowns from ''Siamosaurus'', documenting either predation or scavenging on part of the latter. 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
scanning electron microscopy A scanning electron microscope (SEM) is a type of electron microscope that produces images of a sample by scanning the surface with a focused beam of electrons. The electrons interact with atoms in the sample, producing various signals that ...
. Lauprasert found that spinosaurids and crocodilians may have employed similar feeding tactics and been under comparable mechanical constraints, based on resemblances in the
microstructure Microstructure is the very small scale structure of a material, defined as the structure of a prepared surface of material as revealed by an optical microscope above 25× magnification. The microstructure of a material (such as metals, polymer ...
of their tooth enamel. Therefore, Lauprasert suggested that ''Siamosaurus''—as a piscivorous predator—could have replaced the
ecological niche In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of Resource (biology), resources an ...
of contemporaneous long-snouted crocodilians. He noted that this likely occurred in correlation with the rising aridity of the Sao Khua and Khok Kruat Formations during the Early Cretaceous, since ''Siamosaurus'' had better mobility in a dry environment than crocodilians did. This might explain the absence of long-snouted crocodilian fossils from that time and place.
Goniopholidid Goniopholididae is an extinct family of moderate-sized semi-aquatic neosuchian crocodyliformes. Their bodyplan and morphology are convergent on living crocodilians. They lived across Laurasia (Asia, Europe and North America) between the Middle J ...
crocodilians were prevalent, however, and since this group had broader, shorter snouts and thus more varied diets, Lauprasert suggested that this would have kept them from
competing Competition is a rivalry where two or more parties strive for a common goal which cannot be shared: where one's gain is the other's loss (an example of which is a zero-sum game). Competition can arise between entities such as organisms, individ ...
with ''Siamosaurus''. A similar scenario was proposed for spinosaurids by Hone and colleagues in 2010, who also noted that compared to large crocodilians and obligate aquatic predators, they could more easily travel from one body of water to another in search of prey.


Aquatic habits

In 2008, French palaeontologist Romain Amiot and colleagues compared the
oxygen isotope There are three known stable isotopes of oxygen (8O): , , and . Radioactive isotopes ranging from to have also been characterized, all short-lived. The longest-lived radioisotope is with a half-life of , while the shortest-lived isotope is ...
ratios of remains from theropod and sauropod dinosaurs, crocodilians, turtles, and freshwater fish recovered from eight localities in northeastern Thailand. The study revealed that ''Siamosaurus'' teeth had isotope ratios closer to those of crocodilians and freshwater turtles than other theropods, and so it may have had
semiaquatic In biology, being semi-aquatic refers to various macroorganisms that live regularly in both aquatic and terrestrial environments. When referring to animals, the term describes those that actively spend part of their daily time in water (in ...
habits similar to these animals, spending much of its daily life near or in water. Discrepancies between the ratios of sauropods, ''Siamosaurus'', and other theropods also indicate these dinosaurs drank from different sources, whether river, pond, or plant water. In 2010, Amiot and colleagues published another oxygen isotope study on turtle, crocodilian, spinosaurid, other theropod remains, this time including fossils from Thailand, China, England, Brazil, Tunisia, and Morocco. The analysis showed that Thai spinosaurid teeth tended to have the largest difference from the ratios of other, more terrestrial theropods, while those of ''Spinosaurus'' from Tunisia and Morocco tended to have the least difference, despite the advanced piscivorous adaptations in the skull observed for this genus. The authors suggested that piscivory and semiaquatic habits may explain how spinosaurids coexisted with other large theropods. By feeding on different prey items and occupying a distinct ecological role, a phenomenon that is known as
niche partitioning In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of resources and competitors (for e ...
, the different types of theropods would have been out of direct competition. Further lines of evidence have since demonstrated that spinosaurids, especially those within the Spinosaurinae, developed strong adaptations for aquatic environments, such as dense limb bones for
buoyancy Buoyancy (), or upthrust, is the force exerted by a fluid opposing the weight of a partially or fully immersed object (which may be also be a parcel of fluid). In a column of fluid, pressure increases with depth as a result of the weight of t ...
control; reduction of the pelvic girdle; and elongated neural spines on the tail, likely used for underwater propulsion.


Palaeoenvironment and palaeobiogeography

Of all the
Mesozoic The Mesozoic Era is the Era (geology), era of Earth's Geologic time scale, geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceous Period (geology), Periods. It is characterized by the dominance of archosaurian r ...
formations in northeastern Thailand, the Sao Khua is the most abundant and diverse in vertebrate fossil discoveries. The Khorat Group yields fossil taxa only of continental origin, with no definitive evidence for marine fossils or
sedimentary Sedimentary rocks are types of rock formed by the cementation of sediments—i.e. particles made of minerals (geological detritus) or organic matter (biological detritus)—that have been accumulated or deposited at Earth's surface. Sedime ...
structures found so far. In 1963, Yoshitsugu Kobayashi of
Hokkaido University , or , is a public research university in Sapporo, Hokkaido, Japan. Founded in 1918, it is the fifth-oldest government-authorised university in Japan and one of the former Imperial Universities. The university finds its roots in Sapporo A ...
reported ichthyosaur and plesiosaur teeth from the Sao Khua Formation, but these have now been identified as belonging to ''Siamosaurus'' and a crocodilian respectively. The sediments of the Sao Khua Formation, which comprise
red clay Ultisol, commonly known as red clay soil, is one of twelve soil orders in the United States Department of Agriculture soil taxonomy. The word "Ultisol" is derived from "ultimate", because Ultisols were seen as the ultimate product of continuous ...
s,
mudstone Mudstone, a type of mudrock, is a fine-grained sedimentary rock whose original constituents were clays or muds. Mudstone is distinguished from ''shale'' by its lack of fissility.Blatt, H., and R.J. Tracy, 1996, ''Petrology.'' New York, New York, ...
s,
sandstone Sandstone is a Clastic rock#Sedimentary clastic rocks, clastic sedimentary rock composed mainly of grain size, sand-sized (0.0625 to 2 mm) silicate mineral, silicate grains, Cementation (geology), cemented together by another mineral. Sand ...
s,
siltstone Siltstone, also known as aleurolite, is a clastic sedimentary rock that is composed mostly of silt. It is a form of mudrock with a low clay mineral content, which can be distinguished from shale by its lack of fissility. Although its permeabil ...
s, and conglomerate rocks, record a
fluvial A river is a natural stream of fresh water that flows on land or inside caves towards another body of water at a lower elevation, such as an ocean, lake, or another river. A river may run dry before reaching the end of its course if it ru ...
environment Environment most often refers to: __NOTOC__ * Natural environment, referring respectively to all living and non-living things occurring naturally and the physical and biological factors along with their chemical interactions that affect an organism ...
dominated by lakes,
floodplain A floodplain or flood plain or bottomlands is an area of land adjacent to a river. Floodplains stretch from the banks of a river channel to the base of the enclosing valley, and experience flooding during periods of high Discharge (hydrolog ...
s, and meandering low-energy rivers. This is consistent with the types of vertebrate fauna present in the formation, which comprise only terrestrial or freshwater animals.'''' Besides ''Siamosaurus'', there were theropod dinosaurs like the
metriacanthosaurid Metriacanthosauridae (Greek for "moderately-spined lizards") is an extinct family of allosauroid theropod dinosaurs that lived in Europe and Asia from the Middle Jurassic to the Early Cretaceous. The family is split into two subgroups: Metriacant ...
''Siamotyrannus'', the ornithomimosaur ''
Kinnareemimus ''Kinnareemimus'' (, meaning "Kinnara#Thailand, Kinnaree mimic", after a figure from Thai folklore) is an extinct genus of ornithomimosaurian theropod dinosaur that was discovered in the Early Cretaceous Sao Khua Formation in what is now Thailand. ...
'', the
megaraptora Megaraptora is a clade of carnivorous theropod dinosaurs. Its derived members, the Megaraptoridae are noted for their large hand claws and powerfully-built forelimbs, which are usually reduced in size in other large theropods. Although undoubt ...
n ''
Phuwiangvenator ''Phuwiangvenator'' () is an extinct genus of megaraptoran Theropoda, theropod that lived during the Early Cretaceous period in what is now Thailand. It contains only the type species, ''P. yaemniyomi''. The genus, generic name of ''Phuwiangven ...
'', the basal
coelurosaur Coelurosauria (; from Greek, meaning "hollow-tailed lizards") is the clade containing all theropod dinosaurs more closely related to birds than to carnosaurs. Coelurosauria is a subgroup of theropod dinosaurs that includes compsognathids, ty ...
''
Vayuraptor ''Vayuraptor'' (meaning "wind thief") is a genus of basal coelurosaurian (possibly megaraptoran) theropod dinosaur that lived in the Early Cretaceous (Barremian) Sao Khua Formation of Thailand. The genus contains a single species, ''V. nongbual ...
'', a
compsognathid Compsognathidae is a family of coelurosaurian theropod dinosaurs. Compsognathids were small carnivores, generally conservative in form, hailing from the Jurassic and Cretaceous Periods. The bird-like features of these species, along with other d ...
theropod, and indeterminate birds. Theropod
eggs An egg is an organic vessel in which an embryo begins to develop. Egg, EGG or eggs may also refer to: Biology * Egg cell, the female reproductive cell (gamete) in oogamous organisms Food * Eggs as food Places * Egg, Austria * Egg, Switzerland ...
with
embryo An embryo ( ) is the initial stage of development for a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male sp ...
s have also been recovered from the formation.'''' There were also sauropods like the
titanosauriform Macronaria is a clade of sauropod dinosaurs. Macronarians are named after the large diameter of the nasal opening of their skull, known as the external naris, which exceeded the size of the orbit, the skull opening where the eye is located (hence ...
''Phuwiangosaurus'',
mamenchisaurids Mamenchisauridae is a family of sauropod dinosaurs belonging to the Eusauropoda known from the Jurassic and Early Cretaceous of Asia and Africa. Mamenchisaurids are characterized by their proportionately extremely long necks. Some members of the ...
, and indeterminate forms.'''' Sauropod remains are some of the most abundant in the Sao Khua and Khok Kruat Formations. No
ornithischia Ornithischia () is an extinct clade of mainly herbivorous dinosaurs characterized by a pelvic structure superficially similar to that of birds. The name ''Ornithischia'', or "bird-hipped", reflects this similarity and is derived from the Greek ...
n (or "bird-hipped") dinosaur fossils have been found in the Sao Khua Formation, possibly suggesting that they were uncommon compared to
saurischia Saurischia ( , meaning "reptile-hipped" from the Greek ' () meaning 'lizard' and ' () meaning 'hip joint') is one of the two basic divisions of dinosaurs (the other being Ornithischia), classified by their hip structure. Saurischia and Ornithi ...
n (or "lizard-hipped") dinosaurs. The faunal assemblage also included indeterminate pterosaurs; carettochelyid,
adocid The Adocidae are an extinct family of aquatic and omnivorous turtles. They are freshwater cryptodiran turtles and are mainly known from Cretaceous and Paleogene Asia and North America. Taxonomy Phylogeny modified from Danilov ''et al.'' (2013) ...
, and softshell turtles;
hybodont Hybodontiformes, commonly called hybodonts, are an extinct group of shark-like cartilaginous fish (chondrichthyans) which existed from the late Devonian to the Late Cretaceous. Hybodonts share a close common ancestry with modern sharks and rays ( ...
sharks like hybodontids, ptychodontids, and lonchidiids;
pycnodontiform Pycnodontiformes is an extinct order of primarily marine bony fish. The group first appeared during the Late Triassic and disappeared during the Eocene. The group has been found in rock formations in Africa, Asia, Europe, North and South America. ...
fish;
ray-finned fishes Actinopterygii (; ), members of which are known as ray-finned fish or actinopterygians, is a class (biology), class of Osteichthyes, bony fish that comprise over 50% of living vertebrate species. They are so called because of their lightly built ...
such as sinamiids and semionotids;'''' and the goniopholidid
crocodyliforms Crocodyliformes is a clade of crurotarsan archosaurs, the group often traditionally referred to as "crocodilians". They are the first members of Crocodylomorpha to possess many of the features that define later relatives. They are the only pseudo ...
''
Sunosuchus ''Sunosuchus'' is an extinct genus of goniopholidid mesoeucrocodylian. Fossils are known from China, Kyrgyzstan, and Thailand and are Jurassic in age, although some may be Early Cretaceous. Four species are currently assigned to the genus: the t ...
'','' Siamosuchus'','''' and ''
Theriosuchus ''Theriosuchus'' is an extinct genus of atoposaurid neosuchian from Late Jurassic to Early Cretaceous of Europe (Hungary & southern England), Southeast Asia (Thailand) and western North America (Wyoming), with fragmentary records from Middle Jur ...
''. The Sao Khua and Khok Kruat Formations had a more
semi-arid A semi-arid climate, semi-desert climate, or steppe climate is a aridity, dry climate sub-type. It is located on regions that receive precipitation below Evapotranspiration#Potential evapotranspiration, potential evapotranspiration, but not as l ...
climate than the older, more humid
Phu Kradung Formation The Early Cretaceous Phu Kradung Formation is the lowest member of the Mesozoic Khorat Group which outcrops on the Khorat Plateau in Isan, Thailand. This geological formation consists of micaceous, brown to reddish-brown siltstone beds with mino ...
, dated to the
Berriasian In the geological timescale, the Berriasian is an age/ stage of the Early/Lower Cretaceous. It is the oldest subdivision in the entire Cretaceous. It has been taken to span the time between 143.1 ±0.6 Ma and 137.05 ± 0.2 (million years ago) ...
.'''' The Khok Kruat Formation is composed mostly of sandstones, conglomerates, siltstones, and
shale Shale is a fine-grained, clastic sedimentary rock formed from mud that is a mix of flakes of Clay mineral, clay minerals (hydrous aluminium phyllosilicates, e.g., Kaolinite, kaolin, aluminium, Al2Silicon, Si2Oxygen, O5(hydroxide, OH)4) and tiny f ...
s. Similar to the Sao Khua Formation, the deposition of these sediments occurred in an arid to semi-arid floodplain environment of slow-moving, meandering rivers. This ecosystem included pterosaurs, sinamiid fish; carettochelyid and acocid turtles; ptychodontid, hybodontid, and thaiodontid sharks; and the crocodyliform '' Khoratosuchus'', as well as goniopholidids. Besides ''Siamosaurus'', the dinosaur fauna of the Khok Kruat Formation included the
carcharodontosaurid Carcharodontosauridae (carcharodontosaurids; from the Greek καρχαροδοντόσαυρος, ''carcharodontósauros'': "shark-toothed lizards") is a group of carnivorous theropod dinosaurs. In 1931, Ernst Stromer named Carcharodontosauridae ...
''
Siamraptor ''Siamraptor'' is an extinct genus of carcharodontosaurian dinosaur, containing the single species ''S. suwati'', known from the Khok Kruat Formation of Thailand. It is possibly the first definitive named carcharodontosaurian species known from ...
'';
iguanodontia Ornithopoda () is a clade of ornithischian dinosaurs, called ornithopods (). They represent one of the most successful groups of herbivorous dinosaurs during the Cretaceous. The most primitive members of the group were bipedal and relatively sm ...
ns like ''
Sirindhorna ''Sirindhorna'' is a genus of hadrosauroid ornithopod dinosaur from Early Cretaceous deposits of northeastern Thailand. Discovery and naming In 2007, the Fukui Prefectural Dinosaur Museum (FPDM) approached the Northeastern Research Institute ...
'', '' Ratchasimasaurus'', and '' Siamodon''; a titanosauriform sauropod similar to ''Phuwiangosaurus''; an indeterminate
ceratopsia Ceratopsia or Ceratopia ( or ; Ancient Greek, Greek: "horned faces") is a group of herbivore, herbivorous, beaked dinosaurs that thrived in what are now North America, Asia and Europe, during the Cretaceous Period (geology), Period, although ance ...
n; and various indeterminate theropods. The formation is probably equivalent to the Grès supérieurs Formation of Laos, since animals like spinosaurids, sauropods, and derived ("advanced") iguanodontians have also been found there. In 2007, Milner and colleagues suggested that spinosaurids and iguanodontians may have spread from western to eastern
Laurasia Laurasia () was the more northern of two large landmasses that formed part of the Pangaea supercontinent from around ( Mya), the other being Gondwana. It separated from Gondwana (beginning in the late Triassic period) during the breakup of Pa ...
—the northern
supercontinent In geology, a supercontinent is the assembly of most or all of Earth's continent, continental blocks or cratons to form a single large landmass. However, some geologists use a different definition, "a grouping of formerly dispersed continents", ...
at the time—during the Aptian, based on their distribution and presence in the Khok Kruat Formation. American palaeontologist
Stephen Brusatte Stephen Louis "Steve" Brusatte FRSE (born April 24, 1984) is an American author and evolutionary biologist who specializes in the anatomy and evolution of dinosaurs. He was educated at the University of Chicago for his Bachelor's degree, at the ...
and colleagues noted in 2010 that the discovery of spinosaurids in Asia, a family previously known only from Europe, Africa, and South America, suggests a faunal interchange between Laurasia and
Gondwana Gondwana ( ; ) was a large landmass, sometimes referred to as a supercontinent. The remnants of Gondwana make up around two-thirds of today's continental area, including South America, Africa, Antarctica, Australia (continent), Australia, Zea ...
(in the south) during the early Late Cretaceous. Though it may also be possible that spinosaurids already had a
cosmopolitan distribution In biogeography, a cosmopolitan distribution is the range of a taxon that extends across most or all of the surface of the Earth, in appropriate habitats; most cosmopolitan species are known to be highly adaptable to a range of climatic and en ...
before the Middle Cretaceous, preceding the breakup of Laurasia from Gondwana. However, the authors noted that more evidence is needed to test this hypothesis. In 2012, Allain and colleagues suggested such a global distribution may have occurred earlier across
Pangaea Pangaea or Pangea ( ) was a supercontinent that existed during the late Paleozoic and early Mesozoic eras. It assembled from the earlier continental units of Gondwana, Euramerica and Siberia during the Carboniferous period approximately 335 mi ...
before the Late Jurassic, even if Asia became separated from the supercontinent first. In 2019, Spanish palaeontologist Elisabete Malafaia and colleagues also indicated a complex
biogeographical Biogeography is the study of the distribution of species and ecosystems in geographic space and through geological time. Organisms and biological communities often vary in a regular fashion along geographic gradients of latitude, elevation, ...
pattern for spinosaurs during the Early Cretaceous, based on anatomical similarities between ''Ichthyovenator'' and the European genus ''
Vallibonavenatrix ''Vallibonavenatrix'' (meaning "Vallibona huntress" after the town near where its remains were found) is a genus of spinosaurid dinosaur from the Early Cretaceous (Barremian) Arcillas de Morella Formation of Castellón, Spain. The type and onl ...
''.


References


External links

* * {{Taxonbar, from1=Q134636, from2=Q95951117 Spinosauridae Dinosaur genera Early Cretaceous dinosaurs Taxa named by Éric Buffetaut Fossil taxa described in 1996 Dinosaurs of Thailand Tooth taxa