Casea Temporal range: Early Permian, | |
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C. broilii skeleton in the Field Museum of Natural History | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Synapsida |
Clade: | † Caseasauria |
Family: | † Caseidae |
Genus: | † Casea Williston, 1910 |
Type species | |
Casea broilii Williston, 1910 |
Casea is a genus of herbivorous caseid synapsids that lived during the late Lower Permian (Kungurian) in what is now Texas, United States. The genus is only represented by its type species, Casea broilii, named by Samuel Wendell Williston in 1910. [1] The species is represented by a skull associated with a skeleton (the holotype FMNH UC 656), a second skull (FMNH UC 698), a partial skull with a better preserved dentition than that of the preceding skulls (FMNH UC 1011), and several incomplete postcranial skeletons. [2] Three other Casea species were later erected, but these are considered today to be invalid or belonging to different genera. [3] [4] [5] Casea was a small animal with a length of about 1.20 m and a weight of around 20 kg. [6] [7]
The genus name and specific epithet honor paleontologists Ermine C. Case and Ferdinand Broili. [1]
The skull, relatively small compared to the size of the body, shows the typical morphology of the caseids with a snout tilted forward, a skull roof decorated with many small pits, and a large pineal opening. The external nares are smaller than those of more derived caseids. The orbits are very large and are directed outwards and slightly forward. In dorsal view, the end of the snout is wider and more rounded than that of the more derived caseidae. [8] [6] [9] The palate is broad and plate-like. A narrow interpterygoid vacuity divides the posterior portions of the palate at the midline. [3] The bones of the palate are almost completely covered with teeth, the largest being on the margins, and the smallest in the center of the palate. The upper jaws had two teeth on each premaxilla and 11 teeth on each maxilla. 12 teeth were present on each hemimandible, some being positioned on the coronoid eminence, a primitive character. The first six teeth of the upper and lower jaws are very strong, conical, almost circular at their base, but more rounded at the apex, and somewhat compressed medio-laterally. Those of the upper jaws are vertical, while the first six teeth of the mandible are directed outward and forward at an angle of forty degrees or more. Few details are visible on the teeth of the holotype FMNH UC 656 and paratype FMNH UC 698 due to very rough preparation of these specimens. However, more careful preparation of the maxillary teeth of the specimen FMNH UC 1011 revealed the presence of tricuspid crowns. [8] [6] [2]
Casea is a lightly-built caseid with rather short limbs compared to the length of the animal. The vertebral column has 24 or 25 presacral vertebrae while the sacrum consists of three vertebrae. The tail is not fully known. Three specimens have preserved an articulated caudal series including 18 to 22 vertebrae. On this basis, the total number of caudal vertebrae is estimated to be around fifty. [8] [6] The ribs form a barrel-shaped rib cage, typical of herbivorous caseids. The pelvis presents an ilium with a flared dorsal margin, in the shape of a very wide fan. Its medial surface is flat and smooth with minor streaks along the dorsal margin. The sacral ribs form a single, continuous contact with the ilium which is formed by the overlap of the sacral ribs one and two, as well as between ribs two and three. [8] [6] [7] The tibia is distinguished by its moderately enlarged proximal end, the latter being slightly wider than the distal end, as in Eocasea . In more derived caseids, the proximal end of the tibia is considerably enlarged. [7] An incomplete skeleton of Casea broilii (FMNH UR 2514), from the type locality and only described in 2014, shows an astragalus still articulated with the tibia. Thus, contrary to the descriptions by Romer and Price, and Olson, the astragalus of Casea broilii is an elongate element in which the articulation for the fibula is separated from the articulation with the tibia by a long neck. This difference in interpretation results from a misidentification of a partial and isolated foot (FMNH UC 657) attributed to Casea but which probably belongs to a different animal. [7]
All Casea broilii specimens come from a single fossiliferous pocket known as Cacops bone bed, located in Baylor County, Texas. [10] [8] This locality, discovered and excavated by Paul Miller in 1909 and 1910, is no longer accessible today because it was submerged in the 1920s after the construction of the Lake Kemp dam. [11] The fossil pocket was about 3 m long, 1.8 m wide, and 60 cm thick. [10] [8] It was part of a red clay level interspersed between strata of the upper Arroyo Formation and the coarser sediments of the lower Vale Formation (two formations of the Clear Fork Group). [10] [8] [7] Ammonoids faunas found in marine strata present at the base and top of the Clear Fork Group indicate that the three formations (Arroyo, Vale, and Choza) that compose it represent a relatively short geological duration corresponding to part of the Kungurian. [12] [13] The location and faunal composition of the Cacops bone bed indicate the existence of a distinct fauna that lived in a geographical area far from the floodplains and deltas of coastal regions, where lived the well known fauna from the more classical Lower Permian localities of North America. [14] The Casea specimens were in association with very numerous specimens of the armoured and entirely terrestrial amphibian Cacops (more than 50 individuals are listed including ten skulls, hence the name of the bone bed), a dozen skeletons of the Varanopidae Varanops , and fragments of Seymouria and Captorhinus . Many of the bones were covered with a very thin layer of cemented clay; others were more or less cemented together in nodular masses. Most of the skeletons were lying on their belly, but some were found on their backs. On most of the skeletons the limbs were articulated. In others, the phalanges of the feet were more or less dispersed, and the tail or the entire limbs were disarticulated. [10] [8] It appears from the position of the skeletons and the conditions of deposition that the bodies underwent very little disturbance after the death of the animals. For Williston, these animals would have died in a pool of stagnant and perennial water. The corpses piled on top of each other in successive layers would correspond to an accumulation spread over several generations. [10] However, no taphonomic study of the site has been published, and given the inaccessibility of the locality today it is difficult to say more.
Currently, the genus Casea contains only the species Casea broilii. In the past, three other species were assigned to the genus, but these represent today separate genera and/or are considered invalid. In 1954, Everett C. Olson reported two new species found in the Clear Fork Group in Texas, Casea nicholsi and C. halselli. [15] In 1974, Denise Sigoneau-Russell and Donald E. Russell established the species Casea rutena for a specimen from southern France. [16] These three species are known from the following material :
In 2008, the first phylogenetic analysis of Caseidae revealed for the first time the paraphyly of the genus Casea, the French species Casea rutena representing a distinct and more derived genus, still not named in this study. [3] Three years later, Casea rutena was removed from the genus Casea and placed in a new genus, Euromycter , with the new combination Euromycter rutenus. [4] In 2015, another study published by Romano and Nicosia again resolved the genus Casea as paraphyletic. Casea nicholsi is identified as a taxon more closely related to the genera Caseoides and Caseopsis than to Casea broilii. Thus, C. nicholsi certainly belongs to a different genus, which is however not sufficiently well known to receive a name. Romano and Nicosia also consider the fragmentary species Casea halselli as a nomen dubium , although important differences with the type species, in the shape and robustness of the femur and tibia, suggest that they belong to a genus other than Casea. [5] According to Werneburg and colleagues, C. halselli is a problematic taxon of uncertain, possibly sphenacomorph affinity. [17]
In the first phylogenetic analysis of Caseidae published in 2008, Casea broilii occupies a basal position within the caseidae, but is however more derived than Oromycter . [3]
Below the first cladogram of Caseidae published by Maddin et al. in 2008. [3]
Caseasauria |
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A phylogenetic analysis made by Benson shows a similar position for Casea broilii. This analysis also confirms the paraphyly of the genus Casea. [18]
Below the phylogenetic analysis of Caseasauria published by Benson in 2012. [18]
Caseasauria |
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A study published in 2015 by Romano & Nicosia, and including almost all the Caseidae (with the exception of Alierasaurus ronchii from Sardinia, considered too fragmentary), shows a similar position for Casea broilii. C. nicholsi is recovered as a more derived taxon, closer to the genera Caseoides and Caseopsis than to Casea broilii. [5]
Below the most pasimonious phylogenetic analysis published by Romano & Nicosia in 2015. [5]
Caseasauria |
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In the phylogenetic analysis published in 2022 by Werneburg and colleagues, Casea broilii is positioned between Oromycter and “Casea” nicholsi. The latter occupies a more basal position than in the cladogram of Romano and Nicosia, being recovered as a more basal taxon than the genus Euromycter. [17]
Below is the cladogram published by Werneburg and colleagues in 2022. [17]
Caseidae |
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Casea represents one of the first large and highly successful herbivores among terrestrial synapsids. [19] Among vertebrates this feeding strategy can be subdivided into many categories, including folivory, frugivory, and granivory but among early terrestrial vertebrates, it is feeding on leaves, stems, roots and rhizomes. Herbivores use massive crushing dentition on the palate and mandibles. [3] Caseids belong to the most basal clade of synapside, the Caseasauria, which also includes the small carnivorous eothyrdids. [3] In the case of Caseids, herbivory is indicated by the presence of a massive rib cage in the thoracic and dorsal regions, and the expanded trunk extends posteriorly to the pelvic girdle, with large ribs fused to the lumbar vertebrae. This suggests that this feeding strategy originated sometime between the late Pennsylvanian and the Early Permian. [3] Some Caseids show dental specializations, with leaf-like large serrations being present in the marginal dentition. [20]
The locomotion of Casea involves a three-vertebra sacrum in early synapsids and no apparent link to body size. LeBlanc and Reisz argue that this sacral anatomy was related to more efficient terrestrial locomotion than to increased weight bearing. [7] Selective pressures for weight-bearing or more efficient locomotory styles and increasingly terrestrial lifestyles may have promoted the repeated acquisition of three sacral vertebrae in Synapsida. [7] The development of the third sacral rib attachment to the pelvis in Synapsids may support this hypothesis. [7]
Seymouria is an extinct genus of seymouriamorph from the Early Permian of North America and Europe. Although they were amphibians, Seymouria were well-adapted to life on land, with many reptilian features—so many, in fact, that Seymouria was first thought to be a primitive reptile. It is primarily known from two species, Seymouria baylorensis and Seymouria sanjuanensis. The type species, S. baylorensis, is more robust and specialized, though its fossils have only been found in Texas. On the other hand, Seymouria sanjuanensis is more abundant and widespread. This smaller species is known from multiple well-preserved fossils, including a block of six skeletons found in the Cutler Formation of New Mexico, and a pair of fully grown skeletons from the Tambach Formation of Germany, which were fossilized lying next to each other.
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.
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 are an extinct family of basal synapsids that lived from the Late Carboniferous to Middle Permian between about 300 and 265 million years ago. Fossils of these animals come from the south-central part of the United States, from various parts of Europe, and possibly from South Africa if the genus Eunotosaurus is indeed a caseid as some authors proposed in 2021. Caseids show great taxonomic and morphological diversity. The most basal taxa were small insectivorous and omnivorous forms that lived mainly in the Upper Carboniferous and Lower Permian, such as Eocasea, Callibrachion, and Martensius. This type of caseid persists until the middle Permian with Phreatophasma and may be Eunotosaurus. During the early Permian, the clade is mainly represented by many species that adopted a herbivorous diet. Some have evolved into gigantic forms that can reach 6–7 metres (20–23 ft) in length, such as Cotylorhynchus hancocki and Alierasaurus ronchii, making them the largest Permian synapsids. Caseids are considered important components of early terrestrial ecosystems in vertebrate history because the numerous herbivorous species in this family are among the first terrestrial tetrapods to occupy the role of primary consumer. The caseids experienced a significant evolutionary radiation at the end of the early Permian, becoming, with the captorhinid eureptiles, the dominant herbivores of terrestrial ecosystems in place of the edaphosaurids and diadectids.
Secodontosaurus is an extinct genus of "pelycosaur" synapsids that lived from between about 285 to 272 million years ago during the Early Permian. Like the well known Dimetrodon, Secodontosaurus is a carnivorous member of the Eupelycosauria family Sphenacodontidae and has a similar tall dorsal sail. However, its skull is long, low, and narrow, with slender jaws that have teeth that are very similar in size and shape—unlike the shorter, deep skull of Dimetrodon, which has large, prominent canine-like teeth in front and smaller slicing teeth further back in its jaws. Its unusual long, narrow jaws suggest that Secodontosaurus may have been specialized for catching fish or for hunting prey that lived or hid in burrows or crevices. Although no complete skeletons are currently known, Secodontosaurus likely ranged from about 2 to 2.7 metres (7–9 ft) in length, weighing up to 110 kilograms (250 lb).
Mycterosaurus is an extinct genus of synapsids belonging to the family Varanopidae. It is classified in the varanopid subfamily Mycterosaurinae. Mycterosaurus is the most primitive member of its family, existing from 290.1 to 272.5 MYA, known to Texas and Oklahoma. It lacks some features that its advanced relatives have.
Varanops is an extinct genus of Early Permian varanopid known from Texas and Oklahoma of the United States. It was first named by Samuel Wendell Williston in 1911 as a second species of Varanosaurus, Varanosaurus brevirostris. In 1914, Samuel W. Williston reassigned it to its own genus and the type species is Varanops brevirostris.
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.
Cotylorhynchus is an extinct genus of herbivorous caseid synapsids that lived during the late Lower Permian (Kungurian) and possibly the early Middle Permian (Roadian) in what is now Texas and Oklahoma in the United States. The large number of specimens found make it the best-known caseid. Like all large herbivorous caseids, Cotylorhynchus had a short snout sloping forward and very large external nares. The head was very small compared to the size of the body. The latter was massive, barrel-shaped, and ended with a long tail. The limbs were short and robust. The hands and feet had short, broad fingers with powerful claws. The barrel-shaped body must have housed large intestines, suggesting that the animal had to feed on a large quantity of plants of low nutritional value. Caseids are generally considered to be terrestrial, though a semi-aquatic lifestyle has been proposed by some authors. The genus Cotylorhynchus is represented by three species, the largest of which could reach more than 6 m in length. However, a study published in 2022 suggests that the genus may be paraphyletic, with two of the three species possibly belonging to separate genera.
Angelosaurus is an extinct genus of herbivorous caseid synapsids that lived during the late Lower Permian (Kungurian) and early Middle Permian (Roadian) in what is now Texas and Oklahoma in the United States. Like other herbivorous caseids, it had a small head, large barrel-shaped body, long tail, and massive limbs. Angelosaurus differs from other caseids by the extreme massiveness of its bones, particularly those of the limbs, which show a strong development of ridges, processes, and rugosities for the attachment of muscles and tendons. Relative to its body size, the limbs of Angelosaurus were shorter and wider than those of other caseids. The ungual phalanges looked more like hooves than claws. The few known cranial elements show that the skull was short and more robust than that of the other representatives of the group. Angelosaurus is also distinguished by its bulbous teeth with shorter and wider crowns than those of other caseids. Their morphology and the high rate of wear they exhibit suggests a diet quite different from that of other large herbivorous caseids, and must have been based on particularly tough plants. A study published in 2022 suggests that the genus may be paraphyletic, with Angelosaurus possibly only represented by its type species A. dolani.
Ennatosaurus is an extinct genus of caseid synapsid that lived during the Middle Permian in northern European Russia. The genus is only represented by its type species, Ennatosaurus tecton, which was named in 1956 by Ivan Antonovich Efremov. The species is known from at least six skulls associated with their lower jaws, as well as from the postcranial bones of several juvenile individuals. Ennatosaurus has the typical caseid skull with a short snout tilted forward and very large external nares. However, it differs from other derived caseids by its postcranial skeleton with smaller proportions compared to the size of the skull. As with other advanced caseids, the teeth of Ennatosaurus were well suited for slicing and cutting vegetation. The presence of a highly developed hyoid apparatus indicates the presence of a massive and mobile tongue, which had to work in collaboration with the palatal teeth during swallowing. With a late Roadian - early Wordian age, Ennatosaurus is one of the last known caseids.
Echinerpeton is an extinct genus of synapsid, including the single species Echinerpeton intermedium from the Late Carboniferous of Nova Scotia, Canada. The name means 'spiny lizard' (Greek). Along with its contemporary Archaeothyris, Echinerpeton is the oldest known synapsid, having lived around 308 million years ago. It is known from six small, fragmentary fossils, which were found in an outcrop of the Morien Group near the town of Florence. The most complete specimen preserves articulated vertebrae with high neural spines, indicating that Echinerpeton was a sail-backed synapsid like the better known Dimetrodon, Sphenacodon, and Edaphosaurus. However, the relationship of Echinerpeton to these other forms is unclear, and its phylogenetic placement among basal synapsids remains uncertain.
Oromycter is an extinct genus of caseid synapsids from the Early Permian of Oklahoma. The sole and type species, Oromycter dolesorum, was named in 2005 by Robert R. Reisz.
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.
Callibrachion is an extinct genus of caseid synapsids that lived in east-central France during the Lower Permian (Asselian). The holotype and only known specimen (MNHN.F.AUT490) is represented by an almost complete postcranial skeleton associated with skull fragments discovered at the end of the 19th century in the Permian Autun basin in Saône-et-Loire department, in the Bourgogne-Franche-Comté region. It belongs to an immature individual measuring less than 1.50 m in length. Callibrachion was long considered a junior synonym of the genus Haptodus and classified among the sphenacodontid pelycosaurs. In 2015, a new study found that Callibrachion was a different animal from Haptodus and that it was a caseasaur rather than a sphenacodontid. This was confirmed in 2016 by a cladistic analysis which recovered Callibrachion as a basal caseid. Callibrachion's sharp teeth and unenlarged ribcage indicate that this animal was likely faunivorous.
Ruthenosaurus is an extinct genus of caseid synapsids that lived in what is now southern France during the Early Permian about 285 million years ago. It is known from the holotype MNHN.F.MCL-1 an articulated partial postcranial skeleton. It was collected by D. Sigogneau-Russell and D. Russell in 1970 in the upper part of the M2 Member, Grès Rouge Group, in the Rodez Basin, near the village of Valady, in Occitanie Region. It was first named by Robert R. Reisz, Hillary C. Maddin, Jörg Fröbisch and Jocelyn Falconnet in 2011, and the type species is Ruthenosaurus russellorum.
Euromycter is an extinct genus of caseid synapsids that lived in what is now southern France during the Early Permian about 285 million years ago. 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.
Alierasaurus is an extinct genus of caseid synapsid that lived during the early Middle Permian (Roadian) in what is now Sardinia. It is represented by a single species, the type species Alierasaurus ronchii. Known from a very large partial skeleton found within the Cala del Vino Formation, Alierasaurus is one of the largest known caseids. It closely resembles Cotylorhynchus, another giant caseid from the San Angelo Formation in Texas. The dimensions of the preserved foot elements and caudal vertebrae suggest an estimated total length of about 6 or 7 m for Alierasaurus. In fact, the only anatomical features that differ between Alierasaurus and Cotylorhynchus are found in the bones of the feet; Alierasaurus has a longer and thinner fourth metatarsal and it has ungual bones at the tips of the toes that are pointed and claw-like rather than flattened as in other caseids. Alierasaurus and Cotylorhynchus both have very wide, barrel-shaped rib cages indicating that they were herbivores that fed primarily on high-fiber plant material.
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.
Lalieudorhynchus is an extinct genus of caseid synapsids that lived during the Guadalupian in what is now the south of France. The genus is only known by its type species, Lalieudorhynchus gandi, which was named in 2022 by Ralf Werneburg, Frederik Spindler, Jocelyn Falconnet, Jean-Sébastien Steyer, Monique Vianey-Liaud, and Joerg W. Schneider. Lalieudorhynchus is represented by a partial postcranial skeleton discovered in the Lodève basin in the central part of the Hérault department in the Occitanie region. It belongs to an individual measuring approximately 3.75 m (12.3 ft) in length. The degree of ossification of its bones, however, indicates that it was a late juvenile or still growing young adult. Based on the internal structure of its bones, the describing authors interpreted Lalieudorhynchus as a semiaquatic animal that may have had a lifestyle similar to that of hippopotamus, spending part of its time in water but returning to land for food, though the idea that caseids were semi-aquatic has been previously contested by other authors. It is geologically one of the youngest known representatives of the caseids. The phylogenetic analysis proposed by Werneburg and colleagues identified Lalieudorhynchus as a derived caseid closely related to the North American species "Cotylorhynchus" hancocki.