Caseasauria

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Caseasaurs
Temporal range: Late Carboniferous-Late Permian, 306–254  Ma
Cotylorhynchus romeri.jpg
Fossil skeleton of Cotylorhynchus romeri
Scientific classification Red Pencil Icon.png
Kingdom: Animalia
Phylum: Chordata
Clade: Synapsida
Clade: Caseasauria
Williston, 1912
Families

Caseidae
Eothyrididae

Caseasauria is one of the two main clades of early synapsids, the other being the Eupelycosauria. Caseasaurs are currently known only from the Late Carboniferous and the Permian, and include two superficially different families, the small insectivorous or carnivorous Eothyrididae, and the large, herbivorous Caseidae. These two groups share a number of specialised features associated with the morphology of the snout and external naris.

Contents

The ancestor of caseasaurs can be traced back to an insect eating or an omnivorous reptile-like synapsid from the Pennsylvanian time of the Carboniferous, possibly resembling Archaeothyris , the earliest known synapsid. The caseasaurs were abundant during the later part of the Early Permian, but by the Middle Permian caseasaur diversity declined because the group was outcompeted by the more successful therapsids. The last caseasaurs became extinct at the end of the Guadelupian (Middle Permian).

Description

Among the most conspicuous characteristics uniting caseasaurs are an enlarged nostril and a snout tip that overhangs the tooth row. [1]

Early caseasaurs, including all eothyridids, were relatively small animals. However, most caseids reached larger sizes, and some caseids, such as Cotylorhynchus and Alierasaurus, were among the largest terrestrial animals of the early Permian. These large herbivorous taxa reached a length of 4 to 6 meters (13 to 20 ft) and a mass of 330 to 500 kilograms (730 to 1,100 lb). [2]

Evolution

Caseasaurs first appear in the fossil record in the late Carboniferous, alongside many other early amniote groups. The earliest known synapsid, Asaphestera from the Bashkirian age, may be an eothyridid caseasaur. [3] The earliest definitive caseasaur is Eocasea . [4]

Caseids thrived during the Kungurian age, and numerous large herbivorous caseids are known from the Kungurian of the United States.

Caseasaurs are one of the two synapsid clades known to have survived into the Guadalupian epoch, along with varanopids. Two caseasaur taxa are known from the Guadalupian of Russia, representing the geologically youngest known caseasaurs: the small, possibly omnivorous or insectivorous Phreatophasma , and the large, herbivorous Ennatosaurus . [5]

Classification

Seymouriamorpha

Diadectomorpha?

Sauropsida

Synapsida

Diadectomorpha?

Caseasauria

Varanopidae?

Ophiacodontidae

Edaphosauridae

Sphenacodontidae

Therapsida

Phylogenetic position of Caseasauria within Reptiliomorpha, showing the possible alternate positions of Diadectomorpha. Varanopidae may belong to Sauropsida.

Caseasauria is generally regarded as the most basal clade of synapsids, with all other synapsids being grouped in the clade Eupelycosauria. However, not all studies have supported this position. In 2012, Roger Benson argued that most of the characters supporting a basal position for caseasaurs pertained to the skull, and presented a phylogenetic analysis incorporating more postcranial data that resolved a clade comprising ophiacodontids and varanopids as the basalmost synapsid clade. [6] However, new postcranial data from eothyridids and basal caseids established that caseasaurs were more basal than ophiacodontids and varanopids after all, with the characters supporting a more derived position for caseasaurs being the result of convergent evolution between caseids and more derived synapsids. [4] [7] The diadectomorphs, conventionally regarded as anamniote tetrapods, may prove to be synapsids even more basal than Caseasauria. [8] [9]

Eocasea martini

Martensius bromackerensis

Casea broili

Oromycter dolesorum

Trichasaurus texensis

Casea nicholsi

Euromycter rutenus

Ennatosaurus tecton

Angelosaurus romeri

Alierasaurus ronchii

Cotylorhynchus romeri

Cotylorhynchus bransoni

Cotylorhynchus hancocki

Phylogeny of Caseidae, after Berman et al. 2020 [10]

Most caseasaurs are divided into two families, Eothyrididae and Caseidae. The affinities of the earliest definitive caseasaur, Eocasea, are uncertain, with some phylogenetic analyses finding it to be a caseid and others finding it to be a basal caseasaur belonging to neither family. [11] [12]

Three genera are typically regarded as belonging to the family Eothyrididae: Eothyris, Oedaleops, and Vaughnictis. However, some phylogenetic analyses have failed to resolve the eothyridids as a clade, instead finding them to be paraphyletic with respect to Caseidae. [13] [11] Asaphestera has been provisionally regarded as an eothyridid as well, without being included in a phylogenetic analysis. [14]

The remaining caseasaurs belong to the family Caseidae.

List of species

Species of Caseasauria
GenusSpeciesYear NamedFamilyAgeLocationNotes
Eocasea E. martini2014incertae sedis Gzhelian May be a caseid
Asaphestera A. platyris1934 Eothyrididae Bashkirian Synapsida incertae sedis; may be an eothyridid
Eothyris E. parkeyi1937
Oedaleops O. campi1965
Vaughnictis V. smithae1965 AsselianSakmarian Originally assigned to the genus Mycterosaurus
Callibrachion C. gaudryi1893 Caseidae
Datheosaurus D. macrourus1904
Oromycter O. dolesorum2005
Phreatophasma P. aenigmatum1954 Roadian
Martensius M. bromackerensis2020
Ruthenosaurus R. russellorum2011
Casea C. broilii1910
C. halselli1954
C. nicholsi1954
Euromycter E. rutenus1974Originally described as Casea rutena
Ennatosaurus E. tecton1956The geologically youngest known caseasaur
Caseopsis C. agilis1962
Caseoides C. sanangelensis1953
Arisierpeton A. simplex2019
Alierasaurus A. ronchii2014
Cotylorhynchus C. romeri1937
C. hancocki1953
C. bransoni1962
Angelosaurus A. dolani1953
A. greeni1962
A. romeri1962
Trichasaurus T. texensis1910

Paleoecology

The paleoecology of caseids is debated. They are typically interpreted as terrestrial animals of dry, upland habitats. However, Caseids exhibit a similar bone microstructure to cetaceans and pinnipeds, which has led to the hypothesis that they led an aquatic lifestyle. [15] [16] This hypothesis has been challenged on the grounds that their bone microstructure specifically resembles fully pelagic animals, and is unlike the bone microstructure of semiaquatic animals, but that the body plan of caseids is inconsistent with a pelagic lifestyle. [16] Moreover, caseid fossils are predominantly associated with arid upland deposits.

See also

Related Research Articles

Synapsid Clade of tetrapods

Synapsids are one of the two major groups of animals that evolved from basal amniotes, the other being the sauropsids, the group that includes reptiles and birds. The group includes mammals and every animal more closely related to mammals than to sauropsids. Unlike other amniotes, synapsids have a temporal fenestra, an opening low in the skull roof behind each eye orbit, leaving a bony arch beneath each; this accounts for their name. The distinctive temporal fenestra developed about 318 million years ago during the Late Carboniferous period, when synapsids and sauropsids diverged, but was subsequently merged with the orbit in early mammals.

Diadectomorpha Extinct clade of tetrapods

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 positioned close to, but outside of the clade Amniota, though some recent research has recovered them as the sister group to the traditional Synapsida within Amniota, based on inner ear anatomy. They include both large carnivorous and even larger herbivorous forms, some semi-aquatic and others fully terrestrial. The diadectomorphs seem to have originated during late Mississippian times, although they only became common after the Carboniferous rainforest collapse and flourished during the Late Pennsylvanian and Early Permian periods.

Eupelycosauria Clade of synapsids

Eupelycosauria is a large clade of animals characterized by the unique shape of their skull, encompassing all mammals and their closest extinct relatives. They first appeared 308 million years ago during the Early Pennsylvanian epoch, with the fossils Archaeothyris and perhaps an even earlier genus, Protoclepsydrops, representing just one of the many stages in the evolution of mammals, in contrast to their earlier amniote ancestors.

Varanopidae Extinct family of tetrapods

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 Carboniferous. However, some recent studies have recovered them being taxonomically closer to diapsid reptiles. A varanopid from the latest Middle Permian Pristerognathus Assemblage Zone is the youngest known varanopid and the last member of the "pelycosaur" group of synapsids.

Eothyrididae Extinct family of synapsids

Eothyrididae is an extinct family of very primitive, insectivorous synapsids. Only three genera are known, Eothyris, Vaughnictis and Oedaleops, all from the early Permian of North America. Their main distinguishing feature is the large caniniform tooth in front of the maxilla.

Caseidae Extinct family of synapsids

Caseidae is an extinct family of synapsids ranging from the Late Carboniferous to Middle Permian. While the most primitive known caseid, Eocasea, was a small carnivore, the rest of the group were medium-large sized herbivores with barrel shaped bodies and proportionally small heads. They were one of the first groups of tetrapods to evolve herbivory. Some members of the group reached lengths exceeding four metres and weights of several hundred kilograms, making them amongst the largest terrestrial animals of the Permian period.

Casea is an extinct genus of medium to large-bodied, herbivorous, pelycosaur synapsids from the late Carboniferous until the middle Permian. The name Casea references its appearance from the Caseasauria which developed new morphology of their external naris and snout. Casea were known to be about 1.2 meters long. It weighed between 150 kg to 200 kg. It was slightly smaller than the otherwise very similar Caseoides. Casea was one of the first amniote herbivores, sharing its world with animals such as Dimetrodon and Eryops. It was possibly also aquatic.

<i>Eothyris</i> Extinct genus of synapsids

Eothyris is a genus of extinct synapsid in the family Eothyrididae from the early Permian. It was a carnivorous insectivorous animal, closely related to Oedaleops. Only the skull of Eothyris, first described in 1937, is known. It had a 6-centimetre-long (2.4-inch) skull, and its total estimated length is 30 centimetres. Eothyris is one of the most primitive synapsids known and is probably very similar to the common ancestor of all synapsids in many respects. The only known specimen of Eothyris was collected from the Artinskian-lower.

<i>Oedaleops</i> Extinct genus of synapsids

Oedaleops is an extinct genus of caseasaur synapsids from the Early Permian of the southwestern United States. Fossils have been found in the Cutler Formation in New Mexico, which dates back to the Wolfcampian stage of the Early Permian. All remains belong to the single known species Oedaleops campi. Oedaleops was closely related to Eothyris, and both are part of the family Eothyrididae. Like Eothyris, it was probably an insectivore.

<i>Cotylorhynchus</i> Extinct genus of synapsids

Cotylorhynchus is an extinct genus of very large herbivorous synapsids that lived in the southern part of what is now North America during the Early Permian period. It is the best known member of the synapsid clade Caseidae, usually considered the largest terrestrial vertebrates of the Early Permian.

<i>Phreatophasma</i> Extinct genus of synapsids

Phreatophasma is an extinct genus of synapsids from the Middle Permian of European Russia. It includes only one species, Phreatophasma aenigmatum, which is itself known from a single femur found in a mine near the town of Belebei in Bashkortostan. Phreatophasma comes from a fossil assemblage that is latest Ufimian to earliest Kazanian in age under the Russian stratigraphic scheme, correlating with the Roadian Age under the international stratigraphic timescale. Because the species is based on a single specimen with few diagnostic anatomical features, uncertainty remains as to where it belongs in tetrapod phylogeny; originally interpreted in 1954 as an enigmatic "theromorph" synapsid by Soviet paleontologist Ivan Yefremov, Phreatophasma was later described as a therapsid incertae sedis by American paleontologist Alfred Romer in 1956 and then as a member of a basal synapsid family called Caseidae starting with Everett C. Olson in 1962. Olson's classification was later supported by Canadian paleontologist Robert Reisz in 1986 and American paleontologist Robert L. Carroll in 1988. Ivakhneneko et al. (1997) and Maddin et al. (2008) both considered Phreatophasma an indeterminate synapsid.

<i>Ianthodon</i> Extinct genus of synapsids

Ianthodon is an extinct genus of basal haptodontiform synapsids from the Late Carboniferous about 304 million years ago. The taxon was discovered and named by Kissel & Reisz in 2004. The only species in the taxon, Ianthodon schultzei, was found by separating it from a block that also contained the remains of Petrolacosaurus and Haptodus. The evolutionary significance of the taxon wasn't realized until a publication in 2015. The fossil of this organism was discovered in Garnett, Kansas.

<i>Datheosaurus</i> Extinct genus of synapsids

Datheosaurus is an extinct genus of caseasaur. It was at least 1.5 metres (5 ft) in length. It lived during the Latest Carboniferous to Early Permian in Poland.

<i>Callibrachion</i> Extinct genus of synapsids

Callibrachion is an extinct genus of caseasaur. It was at least 1.5 metres (5 ft) in length. It lived during the Early Permian.

Olson's Extinction was a mass extinction that occurred 273 million years ago in the late Cisuralian or early Guadalupian of the Permian period and which predated the Permian–Triassic extinction event. It is named after Everett C. Olson. There was a sudden change between the early Permian and middle/late Permian faunas. Some authors also place a hiatus in the continental fossil record around that time, but others disagree. Since then this event has been realized across many groups, including plants, marine invertebrates, and tetrapods.

<i>Euromycter</i> Extinct genus of synapsids

Euromycter is an extinct genus of medium-sized caseid synapsids from Early (Artinskian-Kungurian) or Middle (Roadian-Wordian) Permian deposits of Southern France. The holotype and only known specimen of Euromycter (MNHN.F.MCL-2) includes the complete skull with lower jaws and hyoid apparatus, six cervical vertebrae with proatlas, anterior part of interclavicle, partial right clavicle, right posterior coracoid, distal head of right humerus, left and right radius, left and right ulna, and complete left manus. It was collected by D. Sigogneau-Russell and D. Russell in 1970 at the top of the M1 Member, Grès Rouge Group, near the village of Valady, Rodez Basin. It was first assigned to the species "Casea" rutena by Sigogneau-Russell and Russell in 1974. More recently, it was reassigned to its own genus, Euromycter, by Robert R. Reisz, Hillary C. Maddin, Jörg Fröbisch and Jocelyn Falconnet in 2011. The preserved part of the skeleton suggests a size between 1,70 m (5,5 ft) and 1,80 m (5,9 ft) in length for this individual.

Tambacarnifex is an extinct genus of varanodontine synapsids known from the Early Permian Tambach Formation of Free State of Thuringia, central Germany. It was first named by David S. Berman, Amy C. Henrici, Stuart S. Sumida, Thomas Martens and Valerie Pelletier in 2013 and the type species is Tambacarnifex unguifalcatus.

Dendromaia is an extinct genus of varanopid from the Carboniferous of Nova Scotia. It contains a single species, Dendromaia unamakiensis. Dendromaia is the oldest known varanopid, likely the oldest known synapsid, and the only member of the family Varanopidae to be discovered in Nova Scotia. Known from a large partial skeleton preserved with its tail wrapped around a much smaller partial skeleton, Dendromaia may also represent the oldest known occurrence of parental care in the fossil record. While the larger skeleton possessed certain mycterosaurine-like features, the smaller skeleton resembled basal varanopids such as Archaeovenator and Pyozia, creating uncertainty over whether characteristics at the base of Varanopidae have legitimate phylogenetic significance or instead reflect the immaturity of basal varanopid specimens.

Hillary Catherine Maddin is a Canadian paleontologist and developmental biologist known for her work on development in extinct and extant amphibians. She is currently an associate professor in the Department of Earth Sciences at Carleton University.

References

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