Spinosuchus Caseanus
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''Spinosuchus'' (meaning "spined
crocodile Crocodiles (family (biology), family Crocodylidae) or true crocodiles are large semiaquatic reptiles that live throughout the tropics in Africa, Asia, the Americas and Australia. The term crocodile is sometimes used even more loosely to inclu ...
") is an
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of
trilophosaurid Trilophosaurs are lizard-like Triassic allokotosaur reptiles related to the archosaurs. The best known genus is ''Trilophosaurus'', a herbivore up to long. It had a short, unusually heavily built skull, equipped with massive, broad flattened che ...
allokotosaur Allokotosauria is a clade of early archosauromorph reptiles from the Middle Triassic, Middle to Late Triassic known from Asia, Africa, North America and Europe. Allokotosauria was first described and named when a new monophyletic grouping of spec ...
from the
Late Triassic The Late Triassic is the third and final epoch (geology), epoch of the Triassic geologic time scale, Period in the geologic time scale, spanning the time between annum, Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch ...
of
Texas Texas (, ; Spanish language, Spanish: ''Texas'', ''Tejas'') is a state in the South Central United States, South Central region of the United States. At 268,596 square miles (695,662 km2), and with more than 29.1 million residents in 2 ...
, southern United States. It has been assigned to a variety of groups over its history, from
coelophysid Coelophysidae is a family of primitive carnivorous theropod dinosaurs. Most species were relatively small in size. The family flourished in the Late Triassic and Early Jurassic periods, and has been found on numerous continents. Many members of C ...
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
to
pseudosuchian Pseudosuchia is one of two major divisions of Archosauria, including living crocodilians and all archosaurs more closely related to crocodilians than to birds. Pseudosuchians are also informally known as "crocodilian-line archosaurs". Prior to ...
to uncertain
theropod Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally c ...
dinosaur and to
Proterosuchidae Proterosuchidae is an early family of basal archosauriforms whose fossils are known from the Latest Permian and the Early Triassic of Europe, Asia, Africa, Australia and possibly South America. The name comes from Greek πρότερο- ("first") ...
. This uncertainty is not unusual, given that it was only known from a poorly preserved, wall-mounted, partial
vertebral column The vertebral column, also known as the backbone or spine, is part of the axial skeleton. The vertebral column is the defining characteristic of a vertebrate in which the notochord (a flexible rod of uniform composition) found in all chordata, ...
of an animal that lived in a time of diverse, poorly known reptile groups. However, newly collected material and recent phylogenetic studies of early
archosauromorph Archosauromorpha (Greek for "ruling lizard forms") is a clade of diapsid reptiles containing all reptiles more closely related to archosaurs (such as crocodilians and dinosaurs, including birds) rather than lepidosaurs (such as tuataras, liza ...
s suggest that it represents an advanced trilophosaurid very closely related to ''
Trilophosaurus ''Trilophosaurus'' (Greek for "lizard with three ridges") is a lizard-like trilophosaurid allokotosaur known from the Late Triassic of North America. It was a herbivore up to 2.5 m long. It had a short, unusually heavily built skull, equipped wi ...
''.


History

In 1922,
Ermine Cowles Case Ermine Cowles Case (1871–1953), invariably known as E.C. Case, was a prominent American paleontologist in the second generation that succeeded Othniel Charles Marsh and Edward Drinker Cope. A graduate of the University of Kansas, with a PhD ...
described a partial vertebral column (UMMP 7507) he'd discovered in 1921 from the
Tecovas Member Tecovas may refer to: * Tecovas Creek, a creek in Potter County, Texas, USA *Tecovas Formation, a geological formation in Texas See also *''Tecovasaurus ''Tecovasaurus'' (te-KOH-va-SAWR-us) is an extinct Late Triassic amniote genus of unknown a ...
of the
Carnian The Carnian (less commonly, Karnian) is the lowermost stage of the Upper Triassic Series (or earliest age of the Late Triassic Epoch). It lasted from 237 to 227 million years ago (Ma). The Carnian is preceded by the Ladinian and is followed by t ...
-age
Upper Triassic The Late Triassic is the third and final epoch of the Triassic Period in the geologic time scale, spanning the time between Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch and followed by the Early Jurassic Epoch. Th ...
Dockum Formation The Dockum is a Late Triassic (approximately late Carnian through Rhaetian, or 223–200 Ma) geologic group found primarily on the Llano Estacado of western Texas and eastern New Mexico with minor exposures in southwestern Kansas, eastern Color ...
of
Crosby County Crosby County is a county located in the U.S. state of Texas. As of the 2020 census, its population was 5,133. The county seat is Crosbyton. The county was founded in 1876 and later organized in 1886. Both the county and its seat are named for ...
,
Texas Texas (, ; Spanish language, Spanish: ''Texas'', ''Tejas'') is a state in the South Central United States, South Central region of the United States. At 268,596 square miles (695,662 km2), and with more than 29.1 million residents in 2 ...
, as ''
Coelophysis ''Coelophysis'' ( traditionally; or , as heard more commonly in recent decades) is an extinct genus of coelophysid theropod dinosaur that lived approximately 228 to 201.3 million years ago during the latter part of the Triassic Period from t ...
'' sp. (''Coelophysis'' at that time also being poorly known).Case, E.C. (1922). New reptiles and stegocephalians from the Upper Triassic of western Texas. ''Carnegie Institution of Washington Publication'' 321. 84 p. He considered it to be about 2.5 meters (8.5 ft) long. Additional material was referred to it, including a
femur The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates with ...
(UMMP 3396), an ilium (UMMP 8870), and a
basicranium The base of skull, also known as the cranial base or the cranial floor, is the most inferior area of the skull. It is composed of the endocranium and the lower parts of the calvaria. Structure Structures found at the base of the skull are for ...
(UMMP 7473).Case, E.C. (1927). The vertebral column of ''Coelophysis'' Cope. ''Contributions from the Museum of Paleontology, University of Michigan'' 2:209-222. These additional remains have since been recognized as belonging to a variety of other Triassic animals, all of which were poorly known or unknown at the time: the femur to an
aetosaur Aetosaurs () are heavily armored reptiles belonging to the extinct order Aetosauria (; from Greek, (aetos, "eagle") and (, "lizard")). They were medium- to large-sized omnivorous or herbivorous pseudosuchians, part of the branch of archosaurs ...
, possibly ''
Desmatosuchus ''Desmatosuchus'' (, from Greek δεσμός ''desmos'' 'link' + σοῦχος ''soûkhos'' 'crocodile') is an extinct genus of archosaur belonging to the Order Aetosauria. It lived during the Late Triassic. Description ''Desmatosuchus'' was a ...
'',Hunt, A.P, Lucas, S.G., Heckert, A.B., Sullivan, R.M., and Lockley, M.G. (1998). Late Triassic dinosaurs from the western United States. ''Geobios'' 31(4):511-531. the ilium to a
herrerasaurid Herrerasauridae is a family of carnivorous dinosaurs, possibly basal to either theropods or even all of saurischians, or even their own branching from dracohors, separate from dinosauria altogether. They are among the oldest known dinosaurs, fi ...
, either ''
Chindesaurus ''Chindesaurus'' ( ) is an extinct genus of basal saurischian dinosaur from the Late Triassic (213-210 million years ago) of the southwestern United States. It is known from a single species, ''C. bryansmalli'', based on a partial skeleton recovere ...
'' or ''
Caseosaurus ''Caseosaurus'' ( ) is a dubious genus of saurischian dinosaur that lived approximately 221.5 to 212 million years ago during the latter part of the Triassic Period in what is now Texas, in North America. It was a small, lightly-built, bipedal, ...
'', depending on the
taxonomic Taxonomy is the practice and science of categorization or classification. A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...
authority,Long, R.A., and Murry, P.A. (1995). ''Late Triassic (Carnian and Norian) tetrapods from the southwestern United States.'' ''New Mexico Museum of Natural History and Science Bulletin 4'':1-154. and the basicranium to the
rauisuchia "Rauisuchia" is a paraphyletic group of mostly large and carnivorous Triassic archosaurs. Rauisuchians are a category of archosaurs within a larger group called Pseudosuchia, which encompasses all archosaurs more closely related to crocodilia ...
n ''
Postosuchus ''Postosuchus'', meaning "Crocodile from Post", is an extinct genus of rauisuchid reptiles comprising two species, ''P. kirkpatricki'' and ''P. alisonae'', that lived in what is now North America during the Late Triassic. ''Postosuchus'' is a me ...
''.Chatterjee, S. (1985). ''Postosuchus'', a new thecodontian reptile from the Triassic of Texas and the origin of tyrannosaurs . ''Philosophical Transactions of the Royal Society of London, Series B: Biological Sciences'' 309(1139):395-460.
Friedrich von Huene Friedrich von Huene, born Friedrich Richard von Hoinigen, (March 22, 1875 – April 4, 1969) was a German paleontologist who renamed more dinosaurs in the early 20th century than anyone else in Europe. He also made key contributions about v ...
recognized it as a new genus in 1932, and named it in honor of Case.von Huene, F. (1932). Die fossile Reptil-Ordnung Saurischia, ihte Entwicklung und Geschichte. ''Monographien zur Geologie und Palaeontologie'' 1(4). 361 p.
erman Erman Rašiti may refer to: Given name * Erman Bulucu (born 1989), Turkish footballer * Erman Eltemur (born 1993), Turkish karateka * Erman Güraçar (born 1974), Turkish footballer * Erman Kılıç (born 1983), Turkish footballer * Erman Kunter (b ...
/ref> He considered it to be a " podokesaurid". Since the 1970s, though, it has been considered a nondinosaurian.Zhang, F-K. (1975). A new thecodont ''Lotosaurus'', from Middle Triassic of Hunan. ''Vertebrata PalAsiatica'' 8(3):144-147.Padian, K. (1986). On the type material of ''Coelophysis'' Cope (Saurischia: Theropoda) and a new specimen from the Petrified Forest of Arizona (Late Triassic: Chinle Formation). In: Padian, K. (ed.). ''The Beginning of the Age of Dinosaurs''. Cambridge University Press: Cambridge. 45-60. Murry, P.A., and Long, R.A. (1989). Geology and paleontology of the Chinle Formation, Petrified Forest National Park and vicinity, Arizona and a discussion of vertebrate fossils of the southwestern Upper Triassic. In: Lucas, S.G., and Hunt, A.P. (eds.). ''Dawn of the Age of Dinosaurs in the American Southwest''. New Mexico Museum of Natural History: Albuquerque, New Mexico. 29-64. However, a review by Hunt ''et al.'' in 1998 suggested that it was a theropod, possibly a herrerasaurid, citing its hollow centra as evidence for dinosaurian affinities. In an abstract for the 1999
Society of Vertebrate Paleontology The Society of Vertebrate Paleontology (SVP) is a professional organization that was founded in the United States in 1940 to advance the science of vertebrate paleontology around the world. Mission and Activities SVP has about 2,300 members inter ...
meeting and his unpublished thesis, Richards recognized that it had a variety of characters that are
apomorphic In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to have ...
for various dinosaur subgroups, but that these are also found in different basal
archosaur Archosauria () is a clade of diapsids, with birds and crocodilians as the only living representatives. Archosaurs are broadly classified as reptiles, in the cladistic sense of the term which includes birds. Extinct archosaurs include non-avian d ...
groups, and that the poorly preserved, distorted, and reconstructed vertebrae offer no evidence for assignment to any major archosaur group; it does, though, show some resemblances to the
trilophosaur Trilophosaurs are lizard-like Triassic allokotosaur reptiles related to the archosaurs. The best known genus is ''Trilophosaurus'', a herbivore up to long. It had a short, unusually heavily built skull, equipped with massive, broad flattened che ...
s.Richards, H.R. (1999a). Is ''Spinosuchus'' a dinosaur? ''Journal of Vertebrate Paleontology'' 19(Supplement to 3), Abstracts of Papers, Fifty-Ninth Annual Meeting, 70A.Richards, H.R. M.S. (1999b). Osteology and relationships of ''Spinosuchus caseanus'' Huene, 1932 from Texas (Dockum Group, Upper Triassic): a new interpretation. December 17, 1999. Richard J. Zakrzewski, Thesis Advisor. Fort Hays University, Department of Geosciences. Further review, as part of a larger series of papers on the evolution of dinosaurs in the Late Triassic by Sterling Nesbitt, Randall Irmis, and William Parker, found ''Spinosuchus'' to be a valid genus. However, the authors could not classify it beyond
Archosauriformes Archosauriformes (Greek for 'ruling lizards', and Latin for 'form') is a clade of diapsid reptiles that developed from archosauromorph ancestors some time in the Latest Permian (roughly 252 million years ago). It was defined by Jacques Gauthi ...
, and disagreed with Richards' trilophosaur
hypothesis A hypothesis (plural hypotheses) is a proposed explanation for a phenomenon. For a hypothesis to be a scientific hypothesis, the scientific method requires that one can test it. Scientists generally base scientific hypotheses on previous obse ...
. Justin Spielmann and colleagues published a redescription in 2009 that concluded ''Spinosuchus'' was a trilophosaurid closely related to ''
Trilophosaurus ''Trilophosaurus'' (Greek for "lizard with three ridges") is a lizard-like trilophosaurid allokotosaur known from the Late Triassic of North America. It was a herbivore up to 2.5 m long. It had a short, unusually heavily built skull, equipped wi ...
''.Spielmann, J.A., Lucas, S.G., Heckert, A.B., Rinehart, L.F., and Richards, H.R., III. 2009
Redescription of ''Spinosuchus caseanus'' (Archosauromorpha: Trilophosauridae) from the Upper Triassic of North America
''Palaeodiversity'' 2:283-313.
Nesbitt ''et al.'' (2015) performed a
phylogenetic analysis In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
focusing on relations within
Allokotosauria Allokotosauria is a clade of early archosauromorph reptiles from the Middle to Late Triassic known from Asia, Africa, North America and Europe. Allokotosauria was first described and named when a new monophyletic grouping of specialized herbivor ...
and recovered '' Trilophosaurus jacobsi'' to be more closely related to ''S. caseanus'' than to the type species of ''Trilophosaurus'', ''
Trilophosaurus buettneri ''Trilophosaurus'' (Greek language, Greek for "lizard with three ridges") is a lizard-like trilophosaurid allokotosaur known from the Late Triassic of North America. It was a herbivore up to 2.5 m long. It had a short, unusually heavily built sku ...
''. To further test this possibility, the types of ''S. caseanus'' and ''T. jacobsi'' were scored separately from the referred Kahle ''Trilophosaurus'' Quarry elements (referred to ''T. jacobsi'' by Spielmann ''et al.'' (2008) or to ''S. caseanus'' by Spielmann ''et al.'' (2009)). A phylogenetic analysis recovered the three in a
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
clade to the exclusion of ''T. buettneri'' based on a single
autapomorphy In phylogenetics, an autapomorphy is a distinctive feature, known as a derived trait, that is unique to a given taxon. That is, it is found only in one taxon, but not found in any others or outgroup taxa, not even those most closely related to t ...
. Furthermore, the types of ''S. caseanus'' and ''T. jacobsi'' as well as the Kahle Quarry material all scored identically, suggesting that ''T. jacobsi'' not only should be reassigned to ''Spinosuchus'', but in fact represents the junior synonym of its type and only species (''S. caseanus''). Nesbitt ''et al.'' (2015) refrained from officially synonymizing the two taxa pending further study of other advanced trilophosaurids.


References


Sources

*Glut, D.F. (2002). Is ''Spinosuchus'' a theropod? In: Glut, D.F. ''Dinosaurs: The Encyclopedia.'' 2nd Supplement. McFarland & Company, Inc.:Jefferson, North Carolina, 46-48.


External links


Case, 1927
{{Taxonbar, from=Q3080413 Allokotosaurs Prehistoric reptile genera Late Triassic reptiles of North America Triassic geology of Texas Paleontology in Texas Fossil taxa described in 1932 Taxa named by Friedrich von Huene