Moradisaurinae
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Captorhinidae is an extinct family of
tetrapods A tetrapod (; from Ancient Greek τετρα- ''(tetra-)'' 'four' and πούς ''(poús)'' 'foot') is any four- limbed vertebrate animal of the clade Tetrapoda (). Tetrapods include all extant and extinct amphibians and amniotes, with the lat ...
, traditionally considered primitive
reptiles Reptiles, as commonly defined, are a group of tetrapods with an ectothermic metabolism and Amniotic egg, amniotic development. Living traditional reptiles comprise four Order (biology), orders: Testudines, Crocodilia, Squamata, and Rhynchocepha ...
, known from the late
Carboniferous The Carboniferous ( ) is a Geologic time scale, geologic period and System (stratigraphy), system of the Paleozoic era (geology), era that spans 60 million years, from the end of the Devonian Period Ma (million years ago) to the beginning of the ...
to the Late
Permian The Permian ( ) is a geologic period and System (stratigraphy), 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 251.902 Mya. It is the s ...
. 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 (continent), Siberia during the Carboniferous period ...
.


Description

Captorhinids are a clade of small to very large lizard-like animals that date 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 ...
through the
Permian The Permian ( ) is a geologic period and System (stratigraphy), 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 251.902 Mya. It is the s ...
. Their
skull The skull, or cranium, is typically a bony enclosure around the brain of a vertebrate. In some fish, and amphibians, the skull is of cartilage. The skull is at the head end of the vertebrate. In the human, the skull comprises two prominent ...
s were much stronger than those of their relatives, the protorothyridids, and had teeth that were better able to deal with tough plant material. The
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 is similar to those of seymouriamorphs and diadectomorphs; these animals were grouped together with the captorhinids in the order
Cotylosauria image:Labidosaurus hamatus.JPG, Fossil of ''Labidosaurus, Labidosaurus hamatus'' Captorhinida (older name: Cotylosauria) is a doubly paraphyletic grouping of early reptiles. Robert L. Carroll (1988) ranked it as an order in the subclass Anapsida, c ...
as the first
reptiles Reptiles, as commonly defined, are a group of tetrapods with an ectothermic metabolism and Amniotic egg, amniotic development. Living traditional reptiles comprise four Order (biology), orders: Testudines, Crocodilia, Squamata, and Rhynchocepha ...
in the early 20th century, but are now usually regarded as
stem Stem or STEM most commonly refers to: * Plant stem, a structural axis of a vascular plant * Stem group * Science, technology, engineering, and mathematics Stem or STEM can also refer to: Language and writing * Word stem, part of a word respon ...
-
amniote Amniotes are tetrapod vertebrate animals belonging to the clade Amniota, a large group that comprises the vast majority of living terrestrial animal, terrestrial and semiaquatic vertebrates. Amniotes evolution, evolved from amphibious Stem tet ...
s no closer to reptiles than to
mammal A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...
s. 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 mammals h ...
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 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 an appendage, usually as a self-defense mechanism to elude a predator's grasp ...
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.


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 , 'to gulp down' and , 'nose') is an extinct genus of Captorhinidae, captorhinid reptiles that lived during the Permian period. Its remains are known from North America (Oklahoma, Texas) and possibly South America. Descr ...
'' ** '' Euconcordia'' ** ''
Labidosauriscus ''Labidosauriscus'' is an extinct genus of captorhinid tetrapods from the Permian period of North America. Fossils have been discovered in Oklahoma Oklahoma ( ; Choctaw language, Choctaw: , ) is a landlocked U.S. state, state in the Sout ...
'' ** '' Opisthodontosaurus'' ** '' Protocaptorhinus'' ** '' Reiszorhinus'' ** '' Rhiodenticulatus'' ** '' Romeria'' ** '' Saurorictus'' ** 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 Robe ...
'' *** '' Gecatogomphius''The Paleobiology Database: Moradisaurinae
*** '' Indosauriscus'' *** '' Kahneria'' *** '' Labidosaurikos'' *** '' Labidosaurus'' *** '' Moradisaurus'' *** '' Rothianiscus'' *** '' Sumidadectes'' *** '' Tramuntanasaurus'' * Dubious Captorhinids ** '' Puercosaurus'' ** '' Riabininus'' ** ''Chamasaurus'' '' Euconcordia cunninghami'' is thought to be the basalmost known member of Captorhinidae. A phylogenic study of primitive reptile relationships by Muller & Reisz in 2006 recovered '' Thuringothyris'' 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. Captorhinidae contains a single subfamily, the Moradisaurinae. Moradisaurinae was named and assigned to the
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
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 , 'to gulp down' and , 'nose') is an extinct genus of Captorhinidae, captorhinid reptiles that lived during the Permian period. Its remains are known from North America (Oklahoma, Texas) and possibly South America. Descr ...
''". The moradisaurines inhabited what is now
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 ...
,
Morocco Morocco, officially the Kingdom of Morocco, is a country in the Maghreb region of North Africa. It has coastlines on the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria to Algeria–Morocc ...
,
Niger Niger, officially the Republic of the Niger, is a landlocked country in West Africa. It is a unitary state Geography of Niger#Political geography, bordered by Libya to the Libya–Niger border, north-east, Chad to the Chad–Niger border, east ...
,
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 ...
,
Texas Texas ( , ; or ) is the most populous U.S. state, state in the South Central United States, South Central region of the United States. It borders Louisiana to the east, Arkansas to the northeast, Oklahoma to the north, New Mexico to the we ...
and
Oklahoma Oklahoma ( ; Choctaw language, Choctaw: , ) is a landlocked U.S. state, state in the South Central United States, South Central region of the United States. It borders Texas to the south and west, Kansas to the north, Missouri to the northea ...
. Captorhinids were once thought to be the ancestors of
turtle Turtles are reptiles of the order (biology), order Testudines, characterized by a special turtle shell, shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Crypt ...
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 (language), 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 ...
'' from
South Africa South Africa, officially the Republic of South Africa (RSA), is the Southern Africa, southernmost country in Africa. Its Provinces of South Africa, nine provinces are bounded to the south by of coastline that stretches along the Atlantic O ...
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'' was either a
parareptile Parareptilia ("near-reptiles") is an extinct group of Basal (phylogenetics), basal Sauropsida, sauropsids ("Reptile, reptiles"), traditionally considered the sister taxon to Eureptilia (the group that likely contains all living reptiles and birds ...
or a
diapsid Diapsids ("two arches") are a clade of sauropsids, distinguished from more primitive eureptiles by the presence of two holes, known as temporal fenestrae, in each side of their skulls. The earliest traditionally identified diapsids, the araeosc ...
, and therefore unrelated to captorhinids.


Phylogeny

The cladogram below follows the topology from a 2011 analysis by paleontologists Robert R. Reisz, Jun Liu, Jin-Ling Li and Johannes Müller. Traditional classification of Captorhinidae within
Eureptilia Sauropsida (Greek for "lizard faces") is a clade of amniotes, broadly equivalent to the class Reptilia, though typically used in a broader sense to also include extinct stem-group relatives of modern reptiles and birds (which, as theropod dinos ...
, after Muller and Reisz (2006):Simões ''et al.'' (2022) recovered captorhinids as stem-amniotes instead, as the sister group to ''
Protorothyris archeri ''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'' ...
'', while the clade including captorhinids and ''P. archeri'' was recovered as the sister group to
Araeoscelidia Araeoscelidia or Araeoscelida is a clade of extinct tetrapods (traditionally classified as diapsid reptiles) superficially resembling lizards, extending from the Late Carboniferous to the Early Permian. The group contains the genera '' Araeosceli ...
. A cladogram from that study is shown below.Using the same data matrix, Klembara ''et al.'' (2023) found a similar result.


References

{{Taxonbar, from=Q141936 Prehistoric tetrapod families Carboniferous tetrapods Permian tetrapods Pennsylvanian first appearances Lopingian extinctions Taxa named by Ermine Cowles Case