Ctenosauriscus
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''Ctenosauriscus'' is an
extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
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
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 ...
-backed
poposauroid Poposauroidea is a clade of advanced pseudosuchians. It includes poposaurids, shuvosaurids, ctenosauriscids, and other unusual pseudosuchians such as '' Qianosuchus'' and ''Lotosaurus''. It excludes most large predatory quadrupedal "rauisuchian ...
archosaur Archosauria () or archosaurs () is a clade of diapsid sauropsid tetrapods, with birds and crocodilians being the only extant taxon, extant representatives. Although broadly classified as reptiles, which traditionally exclude birds, the cladistics ...
from
Early Triassic The Early Triassic is the first of three epochs of the Triassic Period of the geologic timescale. It spans the time between 251.9 Ma and Ma (million years ago). Rocks from this epoch are collectively known as the Lower Triassic Series, which ...
deposits of
Lower Saxony Lower Saxony is a States of Germany, German state (') in Northern Germany, northwestern Germany. It is the second-largest state by land area, with , and fourth-largest in population (8 million in 2021) among the 16 ' of the Germany, Federal Re ...
in northern
Germany Germany, officially the Federal Republic of Germany, is a country in Central Europe. It lies between the Baltic Sea and the North Sea to the north and the Alps to the south. Its sixteen States of Germany, constituent states have a total popu ...
. It gives its name to the family
Ctenosauriscidae Ctenosauriscidae is an extinct family of pseudosuchian archosaurs within the clade Poposauroidea. Ctenosauriscids existed in Africa, Asia, Europe and North America during the Early Triassic to the Middle Triassic period (latest Olenekian to Anisi ...
, which includes other sail-backed poposauroids such as ''
Arizonasaurus ''Arizonasaurus'' (meaning "Arizona reptile") was a ctenosauriscid archosaur from the Middle Triassic (243 million years ago) Moenkopi Formation of what is now Arizona. The taxon is known for having a large back sail formed by elongated neural sp ...
''. Fossils have been found in latest
Olenekian In the geologic timescale, the Olenekian is an age (geology), age in the Early Triassic epoch (geology), epoch; in chronostratigraphy, it is a stage (stratigraphy), stage in the Lower Triassic series (stratigraphy), series. It spans the time betw ...
deposits around 247.5-247.2 million years old, making it one of the first known archosaurs.


History of discovery

''Ctenosauriscus'' is known from 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 ...
, GZG.V.4191, a single partial preserved
postcrania 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 ...
l
skeleton A skeleton is the structural frame that supports the body of most animals. There are several types of skeletons, including the exoskeleton, which is a rigid outer shell that holds up an organism's shape; the endoskeleton, a rigid internal fra ...
including partial
vertebral column The spinal column, also known as the vertebral column, spine or backbone, is the core part of the axial skeleton in vertebrates. The vertebral column is the defining and eponymous characteristic of the vertebrate. The spinal column is a segmente ...
,
rib In vertebrate anatomy, ribs () are the long curved bones which form the rib cage, part of the axial skeleton. In most tetrapods, ribs surround the thoracic cavity, enabling the lungs to expand and thus facilitate breathing by expanding the ...
s and
girdle A belt without a buckle, especially if a cord or rope, is called a girdle in various contexts, especially historical ones, where girdles were a very common part of everyday clothing from antiquity until perhaps the 15th century, especially for w ...
. The holotype is composed of four slabs, which were labeled A1, A2, B1, and B2 in a 2011 study. Slabs A1 and B1 form the part and slabs A2 and B2 form the counterpart. The holotype was discovered in early 1871 in the Bremke dell locality, near the county of
Göttingen Göttingen (, ; ; ) is a college town, university city in Lower Saxony, central Germany, the Capital (political), capital of Göttingen (district), the eponymous district. The River Leine runs through it. According to the 2022 German census, t ...
. The holotype was found in a deposit called the Solling-Bausandstein, which is an outcropping of the upper Middle
Buntsandstein The Buntsandstein (German for ''coloured'' or ''colourful sandstone'') or Bunter sandstone is a lithostratigraphy, lithostratigraphic and allostratigraphy, allostratigraphic unit (a sequence of rock strata) in the Subsurface (geology), subsurface ...
in the region of
Eichsfeld The Eichsfeld ( or ; 'Oak-field') is a historical region in the southeast of the state of Lower Saxony (which is called , 'lower Eichsfeld') and northwest of the state of Thuringia (, 'upper Eichsfeld') in the south of the Harz mountains in Germany ...
. After the holotype was uncovered, it was housed in the
University of Göttingen The University of Göttingen, officially the Georg August University of Göttingen (, commonly referred to as Georgia Augusta), is a Public university, public research university in the city of Göttingen, Lower Saxony, Germany. Founded in 1734 ...
. It remained undescribed until 1902, when German paleontologist
Friedrich von Huene Baron Friedrich Richard von Hoyningen-Huene (22 March 1875 – 4 April 1969) was a German nobleman paleontologist who described a large number of dinosaurs, more than anyone else in 20th-century Europe. He studied a range of Permo-Carbonife ...
designated it to the new species ''Ctenosaurus koeneni''. Von Huene considered ''C. koeneni'' to be a late-surviving species of
Pelycosauria Pelycosaur ( ) is an older term for basal or primitive Late Paleozoic synapsids, excluding the therapsids and their descendants. Previously, the term mammal-like reptile was used, and Pelycosauria was considered an order, but this is now though ...
, a group of distant mammal relatives otherwise known only from the
Late Carboniferous Late or LATE may refer to: Everyday usage * Tardy, or late, not being on time * Late (or the late) may refer to a person who is dead Music * Late (The 77s album), ''Late'' (The 77s album), 2000 * Late (Alvin Batiste album), 1993 * Late!, a pseudo ...
and
Early Permian 01 or 01 may refer to: * The year 2001, or any year ending with 01 * The month of January * 1 (number) Music * '01 (Richard Müller album), ''01'' (Richard Müller album), 2001 * 01 (Urban Zakapa album), ''01'' (Urban Zakapa album), 2011 * ''01011 ...
. He based this classification on its similarity with sail-backed pelycosaurs like ''
Dimetrodon ''Dimetrodon'' ( or ; ) is an extinct genus of sphenacodontid synapsid that lived during the Cisuralian (Early Permian) Epoch (geology), epoch of the Permian period, around 295–272 million years ago. With most species measuring long and ...
''. Austrian paleontologist
Othenio Abel Othenio Lothar Franz Anton Louis Abel (20 June 1875 – 4 July 1946) was an Austrian paleontologist and evolutionary biologist. Together with Louis Dollo, he was the founder of " paleobiology" and studied the life and environment of fossilized ...
placed ''C. koeneni'' as a
temnospondyl Temnospondyli (from Greek language, Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') or temnospondyls is a diverse ancient order (biology), order of small to giant tetrapods—often considered Labyrinth ...
amphibian closely related to the sail-backed ''
Platyhystrix ''Platyhystrix'' (from Greek: πλατύς ''platús'', 'flat' and Greek: ῠ̔́στρῐξ ''hústrix'', 'porcupine') is an extinct temnospondyl amphibian with a distinctive sail along its back, similar to the unrelated synapsids, '' Dimetro ...
'' (which, like ''Dimetrodon'', was from the Early Permian). The name ''Ctenosaurus'' was preoccupied by a species of
iguanid The Iguanidae is a family of lizards composed of the iguanas, chuckwallas, and their prehistoric relatives, including the widespread green iguana. Taxonomy Iguanidae is thought to be the sister group to the collared lizards (family Crotaphy ...
lizard (now called ''
Ctenosaura ''Ctenosaura'' is a lizard genus commonly known as spinytail iguanas or ctenosaurs. The genus is part of the large lizard family Iguanidae and is native to Mexico and Central America. The name is derived from two Greek words: (), meaning "com ...
''), so a new generic name, ''Ctenosauriscus'', was erected by paleontologist
Oskar Kuhn Oskar Kuhn (7 March 1908, Munich – 1 May 1990) was a German palaeontologist. Life and career Kuhn was educated in Dinkelsbühl and Bamberg and then studied natural science, specialising in geology and paleontology, at the University of Mun ...
in 1964. Paleontologist B. Krebs redescribed the holotype and reclassified ''Ctenosauriscus'' as an archosaur based on similarities with the sail-backed pseudosuchian ''
Hypselorhachis ''Hypselorhachis'' is a genus of extinct reptile, possibly a ctenosauriscid archosaur related to ''Ctenosauriscus''. It lived during the Triassic Period. It is currently known only from a single vertebra found from the Middle Triassic Manda Beds ...
'' from the
Middle Triassic In the geologic timescale, the Middle Triassic is the second of three epoch (geology), epochs of the Triassic period (geology), period or the middle of three series (stratigraphy), series in which the Triassic system (stratigraphy), system is di ...
Manda Beds The Manda Formation (also known as the Manda Beds) is a Middle Triassic (Anisian?) or possibly Late Triassic (Carnian?) geologic formation in Tanzania. It preserves fossils of many terrestrial vertebrates from the Triassic, including some of the e ...
of
Tanzania Tanzania, officially the United Republic of Tanzania, is a country in East Africa within the African Great Lakes region. It is bordered by Uganda to the northwest; Kenya to the northeast; the Indian Ocean to the east; Mozambique and Malawi to t ...
. ''Ctenosauriscus'' was found in the Solling Formation, which was deposited about 247.5 to 247.2 million years ago in the latest Olenekian stage. This age is based on
radiometric dating Radiometric dating, radioactive dating or radioisotope dating is a technique which is used to Chronological dating, date materials such as Rock (geology), rocks or carbon, in which trace radioactive impurity, impurities were selectively incorporat ...
and records of
Milankovitch cycles Milankovitch cycles describe the collective effects of changes in the Earth's movements on its climate over thousands of years. The term was coined and named after the Serbian geophysicist and astronomer Milutin Milanković. In the 1920s, he pr ...
in the formation. ''Ctenosauriscus'' was traditionally thought to have lived during the
Middle Triassic In the geologic timescale, the Middle Triassic is the second of three epoch (geology), epochs of the Triassic period (geology), period or the middle of three series (stratigraphy), series in which the Triassic system (stratigraphy), system is di ...
after Krebs placed the Middle Buntsandstein within the
Anisian In the geologic timescale, the Anisian is the lower stage (stratigraphy), stage or earliest geologic age, age of the Middle Triassic series (stratigraphy), series or geologic epoch, epoch and lasted from million years ago until million years ag ...
stage. A redescription of the holotype was published by Richard J. Butler, Stephen L. Brusatte, Mike Reich, Sterling J. Nesbitt, Rainer R. Schoch and Jahn J. Hornung in 2011. They did not identify any
autapomorphies 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 ...
for ''Ctenosauriscus'', but they noted that the holotype can be distinguished from other
ctenosauriscid Ctenosauriscidae is an extinct family of pseudosuchian archosaurs within the clade Poposauroidea. Ctenosauriscids existed in Africa Africa is the world's second-largest and second-most populous continent after Asia. At about 30.3 mill ...
s by a unique combination of characters. ''Ctenosauriscus'' is one of the oldest
archosaur Archosauria () or archosaurs () is a clade of diapsid sauropsid tetrapods, with birds and crocodilians being the only extant taxon, extant representatives. Although broadly classified as reptiles, which traditionally exclude birds, the cladistics ...
s to date, along with ''
Vytshegdosuchus ''Vytshegdosuchus'' is an extinct genus of paracrocodylomorph archosaur known from the Early Triassic (latest Olenekian stage) Yarenskian Gorizont of the Komi Republic of the European section of Russia. It contains a single species, ''Vytshegdo ...
'' from
Russia Russia, or the Russian Federation, is a country spanning Eastern Europe and North Asia. It is the list of countries and dependencies by area, largest country in the world, and extends across Time in Russia, eleven time zones, sharing Borders ...
and probably ''Xilousuchus'', a ctenosauriscid from China.


Description

The most prominent feature of ''Ctenosauriscus'' is its sail-like back, formed from elongated
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 ...
s of the dorsal and cervical vertebrae. These spines curve slightly forward at the front of the sail and slightly backward at the back of the sail. Although other poposauroids like ''
Lotosaurus ''Lotosaurus'' is an extinct genus of sail-backed poposauroid known from Hunan Province of central China. Discovery ''Lotosaurus'' is known from the holotype IVPP V 4881 (or possibly V 4880), an articulated and well-preserved s ...
'' and the ctenosauriscids ''
Hypselorhachis ''Hypselorhachis'' is a genus of extinct reptile, possibly a ctenosauriscid archosaur related to ''Ctenosauriscus''. It lived during the Triassic Period. It is currently known only from a single vertebra found from the Middle Triassic Manda Beds ...
'' and ''
Xilousuchus ''Xilousuchus'' is an extinct genus of poposauroid from lower Triassic (Olenekian stage) deposits of Fugu County of northeastern Shanxi Province, China. Xilousuchus is one of the oldest archosaurs known to date. Discovery and naming It is ...
'' also have elongated spines, the sail of ''Ctenosauriscus'' is one of the largest in the group. Among ctenosauriscids, ''Ctenosauriscus'' is most similar to ''
Arizonasaurus ''Arizonasaurus'' (meaning "Arizona reptile") was a ctenosauriscid archosaur from the Middle Triassic (243 million years ago) Moenkopi Formation of what is now Arizona. The taxon is known for having a large back sail formed by elongated neural sp ...
'' from the
Middle Triassic In the geologic timescale, the Middle Triassic is the second of three epoch (geology), epochs of the Triassic period (geology), period or the middle of three series (stratigraphy), series in which the Triassic system (stratigraphy), system is di ...
of the southwestern
United States The United States of America (USA), also known as the United States (U.S.) or America, is a country primarily located in North America. It is a federal republic of 50 U.S. state, states and a federal capital district, Washington, D.C. The 48 ...
. Both of these ctenosauriscids have elongated vertebrae spines up to 12 times the height of the bodies of the vertebrae. The ends of the spines are wider in ''Ctenosauriscus'', and ''Ctenosauriscus'' also has larger projections on the
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 ...
of the dorsal vertebrae. ''Hypselorhachis'' also has neural spines that are widened at the end, but they are shorter than those of ''Ctenosauriscus''. ''Lotosaurus'' from the Middle Triassic of
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 ...
also has elongate spines, but they are straighter, broader, and much shorter than those of ''Ctenosauriscus''.


Paleobiology

A 1998 study proposed that ''Ctenosauriscus'' was bipedal and that its elongated neural spines served to absorb the forces exerted from walking on two legs. Although limb bones are unknown, the study found that forces on the tips of the spines were focused on a point below the spinal column, hypothesized to be the knee joint. In the 2011 redescription of ''Ctenosauriscus'', the authors rejected this idea because for forces on the spine to meet at the knee joint, muscles would have to form a direct connection between the knee and the back. Forces exerted from movement travel from the hind legs to the hip and
sacral vertebrae The sacrum (: sacra or sacrums), in human anatomy, is a triangular bone at the base of the spine that forms by the fusing of the sacral vertebrae (S1S5) between ages 18 and 30. The sacrum situates at the upper, back part of the pelvic cavity, ...
, not the
dorsal vertebrae In vertebrates, thoracic vertebrae compose the middle segment of the vertebral column, between the cervical vertebrae and the lumbar vertebrae. In humans, there are twelve thoracic vertebra (anatomy), vertebrae of intermediate size between the ce ...
. The sail of ''Ctenosauriscus'' would also have shifted its center of weight toward the front of the body, making bipedal locomotion difficult or impossible.


References

{{Taxonbar, from=Q143972 Fossil taxa described in 1964 Prehistoric reptiles of Europe Poposauroidea Triassic archosaurs Prehistoric pseudosuchian genera