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''Captorhinus'' (from el, καπτō , 'to gulp down' and el, ῥῑνός , 'nose') 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 nom ...
of captorhinid reptiles that lived during 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 251.9 Mya. It is the last ...
period. Its remains are known from Oklahoma,
Texas Texas (, ; Spanish: ''Texas'', ''Tejas'') is a state in the South Central region of the United States. At 268,596 square miles (695,662 km2), and with more than 29.1 million residents in 2020, it is the second-largest U.S. state by ...
,
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,
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, the Pedra de Fogo Formation, Parnaíba Basin,
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and the Madumabisa Mudstone,
Zambia Zambia (), officially the Republic of Zambia, is a landlocked country at the crossroads of Central, Southern and East Africa, although it is typically referred to as being in Southern Africa at its most central point. Its neighbours are t ...
.


Description

While there are several forms of ''Captorhinus'', there are three main species that are the best known. The previously mentioned ''Captorhinus aguti'' is the
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specime ...
of ''Captorhinus'', but there is also a fair amount of material collected on ''Captorhinus magnus'' and ''Captorhinus laticeps''. The most distinguishable trait of ''Captorhinus'' is its namesake: the hooking of the snout from prominent ventral angulation of the premaxillary process. Other notable characteristics include the dorsally positioned alary process of the jugal on the medial surface and flushed with the orbital margin, the retroarticular process longer anteroposteriorly than broad, and the anteriormost dentary tooth strongly procumbent. The posterior teeth are either chisel-shaped or ogival. Until the late 1990s, ''Captorhinus'' was diagnosed by the presence of multiple rows of
marginal teeth Marginal may refer to: * ''Marginal'' (album), the third album of the Belgian rock band Dead Man Ray, released in 2001 * ''Marginal'' (manga) * '' El Marginal'', Argentine TV series * Marginal seat or marginal constituency or marginal, in polit ...
on the maxillary and dentary bones. However, single-rowed captorhinid elements have been discovered, proving this hypothesis incorrect.


''Captorhinus aguti''

''C. aguti'' was a small captorhinid reptile that lived during the Permian Period 286 MYA to 245 MYA when the continents were still connected as Pangea. An abundance of fossils have been found in Oklahoma, Brazil, and Texas, including a skull, hindlimb bones, spinal vertebra, ribs, and forelimb bones. The entire skeleton of ''C. aguti'' resembles a modern lizard. Although a subfamily of Captorhinidae,
Moradisaurinae Captorhinidae (also known as cotylosaurs) is an extinct family of tetrapods, traditionally considered primitive reptiles, known from the late Carboniferous to the Late Permian. They had a cosmopolitan distribution across Pangea. Description C ...
, also possessed multiple-tooth-rows, the best-known type species ''Captorhinus aguti'' clearly acquired multiple-rowed-teeth independently. In contrast to the teeth rows in moradisaurines, the ''C. aguti'' rows are oriented obliquely to the margins, where each row is followed posterolaterally by the next one. The toothed areas of the maxilla and dentary are broader in ''C. aguti'' than in single-tooth-rowed captorhinids. ''C. aguti'' likely practiced lateralized feeding, as enamel on the teeth of the upper and lower right jaws was more worn down than on the left side. The vertebral structure in ''C. aguti'' is that commonly possessed by primitive reptiles. The centra are amphicoelous and notochordal, with swollen, relatively massive
neural arches 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 ...
. The
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 ...
is differentiated into presacral, sacral, and postsacral or caudal vertebrae. Similarly to extant reptiles, ''C. aguti'' has a functional “mesotarsal” joint. It divides the tarsus into a proximal and distal unit, where the centrale is linked mechanically to the proximal (astragalus-calcaneum) unit. The increased flexibility due to this multiple-jointed arrangement allowed for the articulations between the centrale and first three distal tarsals on the medial side of the mesotarsal joint to have mechanical independence from the lateral articulation between the astragalus-calcaneum and the fourth and fifth distal tarsals. This is almost certainly closely correlated with the primitive sprawling step cycle. Due to the shape of the distal femoral articulation, ''Captorhinus aguti'' would have had little capacity to compensate for lateral movement of the femur.


''Captorhinus magnus''

''Captorhinus magnus'' has thus far been identified from only the
Richards Spur Richards Spur is a Permian fossil locality located at the Dolese Brothers Limestone Quarry north of Lawton, Oklahoma. The locality preserves clay and mudstone fissure fills of a karst system eroded out of Ordovician limestone and dolomite (rock), d ...
locality in Oklahoma, a site that also produced the remains of ''C. aguti''. Fossils of ''C. magnus'' are found predominately in the deeper regions of the fissure complex, whereas in the upper, younger sediments of the fillings, ''C. magnus'' remains are extremely rare. This suggests the ecological replacement of ''C. magnus'' by the smaller, multiple-rowed ''C. aguti'' during the Early Permian. The skeletal elements of ''Captorhinus magnus'' possess a nearly identical morphology to those elements of ''Captorhinus aguti'', with the lone exception of the size difference. ''Captorhinus magnus'' averaged about twice as large as ''C. aguti.'' Another important difference between the two species is that the teeth are arranged in a single row. ''Captorhinus magnus'' possesses ogival check teeth, where the distal tips of the teeth exhibit a triangular shape when viewed in anterior view. Unlike ''C. aguti'', the femur of ''C. magnus'' possesses a concave proximal articular surface in both immature and mature individuals. This allows for distinction between similar-sized femora of ''C. aguti'' and ''C. magnus'', regardless of the
ontogenetic Ontogeny (also ontogenesis) is the origination and development of an organism (both physical and psychological, e.g., moral development), usually from the time of fertilization of the egg to adult. The term can also be used to refer to the st ...
stage of the individuals preserved. Without the presence of the femur or a tooth-bearing element, it would prove difficult to distinguish between ''C. magnus'' and ''C. aguti'', although the two can likely be differentiated with size as the sole criterion.


''Captorhinus laticeps''

''C. laticeps'' is distinguished from ''C. aguti'' by the lack of multiple tooth rows, but is very similar in all other aspects. ''Captorhinus laticeps'' is considerably smaller than ''C. magnus'', and possesses chisel-shaped teeth instead of ogival teeth on the posterior portion of the maxilla and dentary.


History of discovery

In 1882, Edward Cope described a fragmentary skull from the Lower Permian of Texas collected by
W. F. Cummins W. may refer to: * SoHo (Australian TV channel) (previously W.), an Australian pay television channel * ''W.'' (film), a 2008 American biographical drama film based on the life of George W. Bush * "W.", the fifth track from Codeine's 1992 EP ''Bar ...
at Coffee Creek as ''Ectocynodon aguti''. The name had then been revised several times by different paleontologists as more genera were discovered. In 1911, paleontologist
Ermine Cowles Case Ermine Cowles Case (1871–1953), invariably known as E.C. Case, was a prominent American paleontologist in the second generation that succeeded Othniel Charles Marsh and Edward Drinker Cope. A graduate of the University of Kansas, with a PhD ...
revised the recently discovered species. He decided to refer ''P.'' (''Ectodynodon'') ''aguti'', ''P. aduncus'', and ''P. isolomus'' to ''Captorhinus'', and established a new family, Captorhinidae. Named by Cope from the Latin word, “captor,” meaning “one who catches something,” and the Greek word, “rhino,” meaning, “of the nose.” This is based on the theory that the characteristically curved
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 ...
of ''Captorhinus'' may have been used to catch prey.


Paleobiology


Diet

Possible prey from the known fauna of the early Permian of both Texas and Oklahoma may have included other small amniotes, small dissorophid
temnospondyls 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 Carbo ...
, and microsaurian lepospondyls. The comparatively small body size of basal captorhinids suggests that they were probably in competition for food with only the youngest of
varanopids Varanopidae is an extinct family of amniotes that resembled monitor lizards and may have filled a similar niche, hence the name. Typically, they are considered synapsids that evolved from an '' Archaeothyris''-like synapsid in the Late Carbonife ...
and sphenacodontids. These species most likely compromised the smallest
carnivore A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, fat and other s ...
guild among early Permian predators.


Palaeoecology

''Captorhinus'' is known from the Admiral, Belle Plains,
Clyde Clyde may refer to: People * Clyde (given name) * Clyde (surname) Places For townships see also Clyde Township Australia * Clyde, New South Wales * Clyde, Victoria * Clyde River, New South Wales Canada * Clyde, Alberta * Clyde, Ontario, a tow ...
, Arroyo, Vale, and possibly the
Choza Formation The Choza Formation is a geologic formation in Texas. It preserves fossils dating back to the Permian period Period may refer to: Common uses * Era, a length or span of time * Full stop (or period), a punctuation mark Arts, entertainment, a ...
s, Lower Permian, of Texas. The genus is also known from the Lower Permian fissure deposits at
Richards Spur Richards Spur is a Permian fossil locality located at the Dolese Brothers Limestone Quarry north of Lawton, Oklahoma. The locality preserves clay and mudstone fissure fills of a karst system eroded out of Ordovician limestone and dolomite (rock), d ...
, Oklahoma, and the
Cutler Formation The Cutler Formation or Cutler Group is a rock unit that is exposed across the U.S. states of Arizona, northwest New Mexico, southeast Utah and southwest Colorado. It was laid down in the Early Permian during the Wolfcampian epoch. Descri ...
,
Rio Arriba County Rio Arriba County is a county in the U.S. state of New Mexico. As of the 2010 census, the population was 40,246. Its county seat is Tierra Amarilla. Its northern border is the Colorado state line. Rio Arriba County comprises the Española, N ...
, New Mexico. The
morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines * Morphology (archaeology), study of the shapes or forms of artifacts * Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies ...
of small captorhinids is best known from the vast amount of material collected near
Fort Sill, Oklahoma Fort Sill is a United States Army post north of Lawton, Oklahoma, about 85 miles (136.8 km) southwest of Oklahoma City. It covers almost . The fort was first built during the Indian Wars. It is designated as a National Historic Landmark ...
. The majority of bones found belong to the multiple-tooth-rowed captorhinid, ''Captorhinus aguti''. The fossils found are currently in the
Oklahoma Museum of Natural History Oklahoma (; Choctaw: ; chr, ᎣᎧᎳᎰᎹ, ''Okalahoma'' ) is a state in the South Central region of the United States, bordered by Texas on the south and west, Kansas on the north, Missouri on the northeast, Arkansas on the east, New ...
(OMNH). Two specimens held in the OMNH that have been part of the diagnostic process of these species are OMNH 52366, an almost complete right
maxilla The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. T ...
, and OMNH 52367, a partial right
dentary In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
. It is uncertain if the two elements belong to the same individual. Another commonly mentioned captorhinid, ''Labidosaurus hamatus'', was found in a Lower Permian geologic group in Texas, called the Clear Fork Group. There is a skull and incomplete postcranial skeleton of ''Captorhinus'' in the possession of the American Museum of Natural History. Another significant specimen belongs to the Chicago Museum of Natural History taken from the Richard’s Spur, Oklahoma locality.Gaffney 1979


References


Bibliography

* * * * * * * * * {{Taxonbar, from=Q5036974 Captorhinids Prehistoric reptile genera Permian reptiles of Africa Fossils of Zambia Permian reptiles of North America Fossils of the United States Permian reptiles of South America Permian Brazil Fossils of Brazil Fossil taxa described in 1895 Taxa named by Edward Drinker Cope