Ctenosauriscus
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''Ctenosauriscus'' 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
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 ...
-backed
poposauroid Poposauroidea is a clade of advanced pseudosuchians (archosaurs closer to crocodilians than to dinosaurs). It includes poposaurids, shuvosaurids, ctenosauriscids, and other unusual pseudosuchians such as ''Qianosuchus'' and ''Lotosaurus''. Howeve ...
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 ...
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 Ma and Ma (million years ago). Rocks from this epoch are collectively known as the Lower Triassic Series, which is a un ...
deposits of
Lower Saxony Lower Saxony (german: Niedersachsen ; nds, Neddersassen; stq, Läichsaksen) is a German state (') in northwestern Germany. It is the second-largest state by land area, with , and fourth-largest in population (8 million in 2021) among the 16 ...
in northern
Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwe ...
. It gives its name to the family
Ctenosauriscidae Ctenosauriscidae is an extinct family (biology), 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 ...
, which includes other sail-backed poposauroids such as ''
Arizonasaurus ''Arizonasaurus'' was a ctenosauriscid archosaur from the Middle Triassic (243 million years ago). ''Arizonasaurus'' is found in the Middle Triassic Moenkopi Formation of northern Arizona. A fairly complete skeleton was found in 2002 by Sterling ...
''. Fossils have been found in latest
Olenekian In the geologic timescale, the Olenekian is an age in the Early Triassic epoch; in chronostratigraphy, it is a stage in the Lower Triassic series. It spans the time between Ma and Ma (million years ago). The Olenekian is sometimes divided into ...
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 is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...
, GZG.V.4191, a single partial preserved
postcrania Postcrania (postcranium, adjective: postcranial) in zoology and vertebrate paleontology is all or part of the skeleton apart from the skull. Frequently, fossil remains, e.g. of dinosaurs or other extinct tetrapods, consist of partial or isolated sk ...
l
skeleton A skeleton is the structural frame that supports the body of an animal. There are several types of skeletons, including the exoskeleton, which is the stable outer shell of an organism, the endoskeleton, which forms the support structure inside ...
including 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, ...
,
rib In vertebrate anatomy, ribs ( la, costae) are the long curved bones which form the rib cage, part of the axial skeleton. In most tetrapods, ribs surround the chest, enabling the lungs to expand and thus facilitate breathing by expanding the ches ...
s and
girdle A belt, especially if a cord or rope, is called a girdle if it is worn as part of Christian liturgical vestments, or in certain historical, literary or sports contexts. Girdles are used to close a cassock in Christian denominations, including th ...
. 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 Counterpart or Counterparts may refer to: Entertainment and literature * "Counterparts" (short story), by James Joyce * Counterparts, former name for the Reel Pride LGBT film festival * ''Counterparts'' (film), a 2007 German drama * ''Counterp ...
. The holotype was discovered in early 1871 in the Bremke dell locality, near the county of
Göttingen Göttingen (, , ; nds, Chöttingen) 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. At the end of 2019, 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 ; English: ''Oak-field'') is a historical region in the southeast of the state of Lower Saxony (which is called "Untereichsfeld" = lower Eichsfeld) and northwest of the state of Thuringia ("Obereichsfeld" = upper Eichsfeld) in th ...
. After the holotype was uncovered, it was housed in the
University of Göttingen The University of Göttingen, officially the Georg August University of Göttingen, (german: Georg-August-Universität Göttingen, known informally as Georgia Augusta) is a public research university in the city of Göttingen, Germany. Founded ...
. It remained undescribed until 1902, when German paleontologist
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 ...
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 (phylogenetics), basal or primitive Late Paleozoic synapsids, excluding the therapsids and their descendants. Previously, the term ''mammal-like reptile'' had been used, and pelycosaur was considered an ...
, a group of distant mammal relatives otherwise known only from the
Late Carboniferous Late may refer to: * LATE, an acronym which could stand for: ** Limbic-predominant age-related TDP-43 encephalopathy, a proposed form of dementia ** Local-authority trading enterprise, a New Zealand business law ** Local average treatment effect, ...
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 (Son of Dave album), ''01'' (Son of Dave album), 2000 * 01 (Urban ...
. He based this classification on its similarity with sail-backed pelycosaurs like ''
Dimetrodon ''Dimetrodon'' ( or ,) meaning "two measures of teeth,” is an extinct genus of non-mammalian synapsid that lived during the Cisuralian (Early Permian), around 295–272 million years ago (Mya). It is a member of the family Sphenacodontid ...
''. Austrian paleontologist
Othenio Abel Othenio Lothar Franz Anton Louis Abel (June 20, 1875 – July 4, 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 or ...
placed ''C. koeneni'' as a
temnospondyl Temnospondyli (from Greek language, Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') is a diverse order (biology), order of small to giant tetrapods—often considered Labyrinthodontia, primitive amphi ...
amphibian closely related to the sail-backed ''
Platyhystrix ''Platyhystrix'' (from el, πλατύς , 'flat' and el, ῠ̔́στρῐξ , 'porcupine') was a temnospondyl amphibian with a distinctive sail along its back, similar to the unrelated synapsids, ''Dimetrodon'' and ''Edaphosaurus''. It lived du ...
'' (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: ctenos (κτεν ...
''), so a new generic name, ''Ctenosauriscus'', was erected by paleontologist
Oskar Kuhn Oskar Kuhn (7 March 1908, Munich – 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 Munich, fr ...
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 Bed ...
'' from the
Middle Triassic In the geologic timescale, the Middle Triassic is the second of three epochs of the Triassic period or the middle of three series in which the Triassic system is divided in chronostratigraphy. The Middle Triassic spans the time between Ma and ...
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 ( sw, Jamhuri ya Muungano wa Tanzania), is a country in East Africa within the African Great Lakes region. It borders Uganda to the north; Kenya to the northeast; Comoro Islands and ...
. ''Ctenosauriscus'' was found in the
Solling Formation The Solling Formation is a geologic formation in Germany.
, 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 date materials such as rocks or carbon, in which trace radioactive impurities were selectively incorporated when they were formed. The method compares t ...
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 Serbian geophysicist and astronomer Milutin Milanković. In the 1920s, he hypot ...
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 epochs of the Triassic period or the middle of three series in which the Triassic system is divided in chronostratigraphy. The Middle Triassic spans the time between Ma and ...
after Krebs placed the Middle Buntsandstein within the
Anisian In the geologic timescale, the Anisian is the lower stage or earliest age of the Middle Triassic series or epoch and lasted from million years ago until million years ago. The Anisian Age succeeds the Olenekian Age (part of the Lower Triassic Ep ...
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 t ...
for ''Ctenosauriscus'', but they noted that the holotype can be distinguished from other
ctenosauriscid Ctenosauriscidae is an extinct family (biology), 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 ...
s by a unique combination of characters. ''Ctenosauriscus'' is one of the oldest
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 ...
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, ''Vytsheg ...
'' from
Russia Russia (, , ), or the Russian Federation, is a List of transcontinental countries, transcontinental country spanning Eastern Europe and North Asia, Northern Asia. It is the List of countries and dependencies by area, largest country in the ...
and probably ''Xilousuchus'', a ctenosauriscid from China.


Description

The most prominent feature of ''Ctenosauriscus'' is its sail-like back, formed from elongated
neural spine The spinal column, a defining synapomorphy shared by nearly all vertebrates,Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic i ...
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 Poposauroidea is a clade of advanced pseudosuchians (archosaurs closer to crocodilians than to dinosaurs). It includes poposaurids, shuvosaurids, ctenosauriscids, and other unusual ...
'' 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 Bed ...
'' and ''
Xilousuchus ''Xilousuchus'' is an extinct genus of poposauroid from lower Triassic (Olenekian stage) deposits of Fugu County of northeastern Shanxi Province, China. It is known from the holotype, IVPP V 6026, a single well-preserved partial skele ...
'' 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'' was a ctenosauriscid archosaur from the Middle Triassic (243 million years ago). ''Arizonasaurus'' is found in the Middle Triassic Moenkopi Formation of northern Arizona. A fairly complete skeleton was found in 2002 by Sterling ...
'' from the
Middle Triassic In the geologic timescale, the Middle Triassic is the second of three epochs of the Triassic period or the middle of three series in which the Triassic system is divided in chronostratigraphy. The Middle Triassic spans the time between Ma and ...
of the southwestern
United States The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territorie ...
. 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 stores are all owned by individual franchisees. The chain has three different f ...
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. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...
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 (plural: ''sacra'' or ''sacrums''), in human anatomy, is a large, 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 ...
, 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 vertebrae and they are intermediate in size between the cervical ...
. 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 Ctenosauriscids Triassic archosaurs Prehistoric pseudosuchian genera