Captorhinidae (also known as cotylosaurs) is an extinct family of
tetrapods
Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids (reptiles, including dinosaurs and therefore birds) and synapsids (pelycosaurs, extinct therapsids ...
, traditionally considered primitive
reptiles
Reptiles, as most commonly defined are the animals in the Class (biology), class Reptilia ( ), a paraphyletic grouping comprising all sauropsid, sauropsids except birds. Living reptiles comprise turtles, crocodilians, Squamata, squamates (lizar ...
, known from the late
Carboniferous
The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carbonifero ...
to the Late
Permian
The Permian ( ) is a geologic period and stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last period of the Paleoz ...
. They had a cosmopolitan distribution across
Pangea
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 approximately 335 million y ...
.
Description
![Captorhinus kierani](https://upload.wikimedia.org/wikipedia/commons/f/f7/Captorhinus_kierani.jpg)
Captorhinids are a clade of small to very large lizard-like reptiles that date 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, ...
through the
Permian
The Permian ( ) is a geologic period and stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last period of the Paleoz ...
. Their
skull
The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, the ...
s were much stronger than those of their relatives, the
Protorothyrididae
Protorothyrididae is an extinct family of small, lizard-like reptiles belonging to Eureptilia. Their skulls did not have fenestrae, like the more derived diapsids. Protorothyridids lived from the Late Carboniferous to Early Permian periods, in wh ...
, and had teeth that were better able to deal with tough plant material. The
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 is very similar to that of advanced
reptiliomorph
Reptiliomorpha (meaning reptile-shaped; in PhyloCode known as ''Pan-Amniota'') is a clade containing the amniotes and those tetrapods that share a more recent common ancestor with amniotes than with living amphibians ( lissamphibians). It was de ...
amphibians, so much in fact that the amphibian
Seymouriamorpha
Seymouriamorpha were a small but widespread group of limbed vertebrates (tetrapods). They have long been considered reptiliomorphs, and most paleontologists may still accept this point of view, but some analyses suggest that seymouriamorphs are ...
and
Diadectomorpha
Diadectomorpha is a clade of large tetrapods that lived in Euramerica during the Carboniferous and Early Permian periods and in Asia during Late Permian (Wuchiapingian), They have typically been classified as advanced reptiliomorphs (transitional ...
were thought to be reptiles and grouped together in "Cotylosauria" as the first
reptiles
Reptiles, as most commonly defined are the animals in the Class (biology), class Reptilia ( ), a paraphyletic grouping comprising all sauropsid, sauropsids except birds. Living reptiles comprise turtles, crocodilians, Squamata, squamates (lizar ...
in the early 20th century.
Captorhinids have broad, robust skulls that are generally triangular in shape when seen in dorsal view. The
premaxilla
The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammal has b ...
e are characteristically downturned. The largest captorhinid, the herbivorous ''
Moradisaurus'', could reach an estimated snout-vent length of 2 meters (6.5 feet). Early, smaller forms possessed single rows of teeth, and were likely carnivorous or omnivorous, while the larger, more
derived
Derive may refer to:
* Derive (computer algebra system), a commercial system made by Texas Instruments
* ''Dérive'' (magazine), an Austrian science magazine on urbanism
*Dérive, a psychogeographical concept
See also
*
*Derivation (disambiguatio ...
captorhinids belonging to the subfamily Moradisaurinae were herbivorous and developed multiple (up to 11) rows of teeth in the jaws alongside propalinal (back and forth) jaw motion, which created an effective apparatus for grinding and shredding plant matter.Histological and SEM analysis of captorhinid tail vertebrae concluded in a 2018 study that captorhinids were the first amniotes to develop
caudal autotomy
Autotomy (from the Greek ''auto-'', "self-" and ''tome'', "severing", αὐτοτομία) or self-amputation, is the behaviour whereby an animal sheds or discards one or more of its own appendages, usually as a self-defense mechanism to elude a ...
as a defensive function. In studied specimens a split line is present in certain caudal vertebrae that is similar to those found in modern reptiles that perform caudal autonomy. This behaviour represented significant evolutionary benefit for the animals, allowing for escape and distracting predators, as well as minimizing blood loss at an injury site.
Discovery and history
Until recently, ''
Concordia cunninghami
''Euconcordia'' is an extinct genus of Late Carboniferous captorhinid known from Greenwood County, Kansas of the United States.
Description
''Euconcordia'' is known from the holotype KUVP 8702a&b, well preserved skull in dorsal view along ...
'' was thought to be the basalmost known member of Captorhinidae. A novel phylogenic study of primitive reptile relationships by Muller & Reisz in 2006 recovered ''
Thuringothyris
''Thuringothyris'' is an extinct genus of Early Permian eureptiles known from the Thuringian Forest in central Germany.
Description
''Thuringothyris'' is known from the holotype MNG 7729, articulated well-preserved skull and partial postcra ...
'' as a sister taxon of the Captorhinidae.
[Muller, J. and Reisz, R.R. (2006). "The phylogeny of early eureptiles: Comparing parsimony and Bayesian approaches in the investigation of a basal fossil clade." ''Systematic Biology'', 55(3):503-511. ] The same results were obtained in later phylogenic analyses.
[ ''Concordia'' is still the earliest known captorhinid as all other captorhinid taxa are known only from ]Permian
The Permian ( ) is a geologic period and stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last period of the Paleoz ...
deposits.
Captorhinidae contains a single subfamily, the Moradisaurinae. Moradisaurinae was named and assigned to the family
Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
Captorhinidae by A. D. Ricqlès and P. Taquet in 1982. Moradisaurinae was defined as "all captorhinids more closely related to ''Moradisaurus'' than to ''Captorhinus
''Captorhinus'' (from el, καπτō , 'to gulp down' and el, ῥῑνός , 'nose') is an extinct genus of captorhinid reptiles that lived during the Permian period. Its remains are known from Oklahoma, Texas, Europe, India, the Pedra de Fog ...
''". The moradisaurines inhabited what is now 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 ...
, Morocco
Morocco (),, ) officially the Kingdom of Morocco, is the westernmost country in the Maghreb region of North Africa. It overlooks the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria to ...
, Niger
)
, official_languages =
, languages_type = National languages[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 ...](_blank)
, 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 ...
and Oklahoma
Oklahoma (; Choctaw language, Choctaw: ; chr, ᎣᎧᎳᎰᎹ, ''Okalahoma'' ) is a U.S. state, state in the South Central United States, South Central region of the United States, bordered by Texas on the south and west, Kansas on the nor ...
.
Captorhinids were once thought to be the ancestors of turtle
Turtles are an order of reptiles known as Testudines, characterized by a special shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Cryptodira (hidden necked tu ...
s. The Middle Permian
The Guadalupian is the second and middle series/epoch of the Permian. The Guadalupian was preceded by the Cisuralian and followed by the Lopingian. It is named after the Guadalupe Mountains of New Mexico and Texas, and dates between 272.95 ± 0. ...
reptile ''Eunotosaurus
''Eunotosaurus'' (''Latin'': Stout-backed lizard) is an extinct genus of amniote, possibly a close relative of turtles. ''Eunotosaurus'' lived in the late Middle Permian (Capitanian stage) and fossils can be found in the Karoo Supergroup of Sout ...
'' from South Africa
South Africa, officially the Republic of South Africa (RSA), is the southernmost country in Africa. It is bounded to the south by of coastline that stretch along the South Atlantic and Indian Oceans; to the north by the neighbouring countri ...
was seen as the " missing link" between cotylosaurs and chelonians throughout much of the early 20th century. However, more recent fossil finds have shown that ''Eunotosaurus'' is a parareptile
Parareptilia ("at the side of reptiles") is a subclass or clade of basal sauropsids (reptiles), typically considered the sister taxon to Eureptilia (the group that likely contains all living reptiles and birds). Parareptiles first arose near the ...
unrelated to either turtles or captorhinids.
Classification
Taxonomy
The following taxonomy follows Reisz ''et al.'', 2011 and Sumida ''et al.'', 2010 unless otherwise noted.
*Family Captorhinidae
** '' Captorhinoides''?
** '' Acrodenta''
** '' Baeotherates''
** ''Captorhinus
''Captorhinus'' (from el, καπτō , 'to gulp down' and el, ῥῑνός , 'nose') is an extinct genus of captorhinid reptiles that lived during the Permian period. Its remains are known from Oklahoma, Texas, Europe, India, the Pedra de Fog ...
''
** ''Euconcordia
''Euconcordia'' is an extinct genus of Late Carboniferous captorhinid known from Greenwood County, Kansas of the United States.
Description
''Euconcordia'' is known from the holotype KUVP 8702a&b, well preserved skull in dorsal view along ...
''
** '' Labidosauriscus''
** '' Opisthodontosaurus''
** '' Protocaptorhinus''
** '' Reiszorhinus''
** '' Rhiodenticulatus''
** ''Romeria
Christianity has a strong tradition of pilgrimages, both to sites relevant to the New Testament narrative (especially in the Holy Land) and to sites associated with later saints or miracles.
History
Christian pilgrimages were first made to si ...
''
** '' Saurorictus''
** ''Thuringothyris
''Thuringothyris'' is an extinct genus of Early Permian eureptiles known from the Thuringian Forest in central Germany.
Description
''Thuringothyris'' is known from the holotype MNG 7729, articulated well-preserved skull and partial postcra ...
''
** Subfamily Moradisaurinae
*** '' Balearosaurus''
*** '' Captorhinikos''
*** ''Gansurhinus
''Gansurhinus'' is an extinct genus of moradisaurine captorhinid known from the Middle Permian Qingtoushan Formation of the Qilian Mountains and the Late Permian Naobaogou Formation in the Daqing Mountains of China. It was first named by R ...
''
*** ''Gecatogomphius
''Gecatogomphius'' is an extinct genus of Middle Permian captorhinid with multiple tooth rows known from the Kirov Oblast and the Republic of Tatarstan of Russia.
Description
''Gecatogomphius'' is known from the holotype PIN 1156/1, a th ...
''[The Paleobiology Database: Moradisaurinae](_blank)
*** '' Kahneria''[
*** '']Labidosaurikos
''Labidosaurikos'' is a genus of extinct captorhinid anapsid reptile that lived around 279 to 272 million years ago during Kungurian age of the lower Permian. The American paleontologist John Willis Stovall first described ''Labidosaurikos'' in ...
''
*** ''Labidosaurus
''Labidosaurus'' (from el, λᾰβῐ́δος , 'forceps' and el, σαῦρος , 'lizard') is an extinct genus of reptile from the Permian period of North America. Fossils have been discovered in Texas.
It was heavily built, resembling a liza ...
''
*** '' Moradisaurus''
*** '' Rothianiscus''
* Dubious
Doubt is a mental state in which the mind remains suspended between two or more contradictory propositions, unable to be certain of any of them.
Doubt on an emotional level is indecision between belief and disbelief. It may involve uncertainty ...
Captorhinids
** '' Puercosaurus''
** '' Riabininus''
Phylogeny
The cladogram
A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...
below was recovered in a study by Sumida ''et al.'', 2010.[
The cladogram below follows the topology from a 2011 analysis by paleontologists Robert R. Reisz, Jun Liu, Jin-Ling Li and Johannes Müller.][
The majority of phylogenetic studies recover captorhinids as basal members of Eureptilia; however, Simões ''et al.'' (2022) recover them as ]stem
Stem or STEM may refer to:
Plant structures
* Plant stem, a plant's aboveground axis, made of vascular tissue, off which leaves and flowers hang
* Stipe (botany), a stalk to support some other structure
* Stipe (mycology), the stem of a mushro ...
-amniote
Amniotes are a clade of tetrapod vertebrates that comprises sauropsids (including all reptiles and birds, and extinct parareptiles and non-avian dinosaurs) and synapsids (including pelycosaurs and therapsids such as mammals). They are disti ...
s instead, as the sister group to ''Protorothyris
''Protorothyris'' is an extinct genus of Early Permian protorothyridid known from Texas and West Virginia of the United States. It was first named by Llewellyn Ivor Price in 1937 and the type species is ''Protorothyris archeri''. ''P. archeri ...
archeri'', while the clade including captorhinids and ''P. archeri'' is recovered as the sister group to Araeoscelidia
Araeoscelidia or Araeoscelida is a clade of extinct diapsid reptiles superficially resembling lizards, extending from the Late Carboniferous to the Early Permian.
The group contains the genera ''Araeoscelis'', ''Petrolacosaurus'', the possi ...
.
References
{{Taxonbar, from=Q141936
Prehistoric reptile families
Carboniferous reptiles
Permian reptiles
Pennsylvanian first appearances
Lopingian extinctions
Taxa named by Ermine Cowles Case