''Ctenosauriscus'' is an
extinct genus
Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial n ...
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 ma ...
-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''. ...
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-avi ...
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 ...
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 (FRG),, is a country in Central Europe. It is the most populous member state of the European Union. Germany lies between the Baltic and North Sea to the north and the Alps to the sou ...
. 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 Ani ...
, 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 Sterlin ...
''. 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 divid ...
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 seve ...
,
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 s ...
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 chordate ...
,
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 ch ...
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 ...
. 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 lithostratigraphic and allostratigraphic unit (a sequence of rock strata) in the subsurface of large parts of west and central Europe. The Buntsandst ...
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 i ...
. 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 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 order, but this is now ...
, 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 effe ...
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), 2001
* ''01'' (Son of Dave album), 2000
* ''01'' (Urban Zakapa album), 2011
* ''O1'' (Hiroyuki Sawano ...
. 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 Sphenacodo ...
''. 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 τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') is a diverse order of small to giant tetrapods—often considered primitive amphibians—that flourished worldwide during the Carb ...
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 d ...
'' (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 Crotaphyt ...
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 Be ...
'' 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 ...
Manda Beds 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 ...
.[
]
''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 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 ...
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 Trias ...
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 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 Ani ...
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-avi ...
s to date, along with '' Vytshegdosuchus'' from Russia
Russia (, , ), or the Russian Federation, is a transcontinental country spanning Eastern Europe and Northern Asia. It is the largest country in the world, with its internationally recognised territory covering , and encompassing one-eigh ...
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 ...
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'' 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 Be ...
'' 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 Sterlin ...
'' 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 ...
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 U.S. state, states, a Washington, D.C., federal district, five ma ...
. 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 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 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