Casea Rutena
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''Euromycter'' is an
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of caseid synapsids that lived in what is now southern
France France (), officially the French Republic ( ), is a country primarily located in Western Europe. It also comprises of Overseas France, overseas regions and territories in the Americas and the Atlantic Ocean, Atlantic, Pacific Ocean, Pac ...
during the Early Permian (late Artinskian) about 285 million years ago. The
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...
and only known specimen of ''Euromycter'' ( MNHN.F.MCL-2) includes the complete
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 ...
with lower jaws and hyoid apparatus, six
cervical vertebrae In tetrapods, cervical vertebrae (singular: vertebra) are the vertebrae of the neck, immediately below the skull. Truncal vertebrae (divided into thoracic and lumbar vertebrae in mammals) lie caudal (toward the tail) of cervical vertebrae. In ...
with
proatlas In anatomy, the atlas (C1) is the most superior (first) cervical vertebra of the spine (anatomy), spine and is located in the neck. It is named for Atlas (mythology), Atlas of Greek mythology because, just as Atlas supported the globe, it suppo ...
, anterior part of interclavicle, partial right
clavicle The clavicle, or collarbone, is a slender, S-shaped long bone approximately 6 inches (15 cm) long that serves as a strut between the shoulder blade and the sternum (breastbone). There are two clavicles, one on the left and one on the rig ...
, right posterior coracoid, distal head of right
humerus The humerus (; ) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius and ulna, and consists of three sections. The humeral upper extremity consists of a roun ...
, left and right
radius In classical geometry, a radius ( : radii) of a circle or sphere is any of the line segments from its center to its perimeter, and in more modern usage, it is also their length. The name comes from the latin ''radius'', meaning ray but also the ...
, left and right
ulna The ulna (''pl''. ulnae or ulnas) is a long bone found in the forearm that stretches from the elbow to the smallest finger, and when in anatomical position, is found on the medial side of the forearm. That is, the ulna is on the same side of t ...
, 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 Grès was a French haute couture fashion house founded by Madame Grès in 1942. Parfums Grès is the associated perfume house, which still exists, and is now based in Switzerland. History Germaine Émilie Krebs (1903–1993), known as Alix B ...
, near the village of Valady (département of
Aveyron Aveyron (; oc, Avairon; ) is a department in the region of Occitania, Southern France. It was named after the river Aveyron. Its inhabitants are known as ''Aveyronnais'' (masculine) or ''Aveyronnaises'' (feminine) in French. The inhabitants o ...
),
Rodez Rodez ( or ; oc, Rodés, ) is a small city and commune in the South of France, about 150 km northeast of Toulouse. It is the prefecture of the department of Aveyron, region of Occitania (formerly Midi-Pyrénées). Rodez is the seat of the ...
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 Robert Rafael Reisz is a Canadian paleontologist and specialist in the study of early amniote and tetrapod evolution. Research career Reisz received his B.Sc. (1969), M.Sc. (1971) and Ph.D. (1975) from McGill University as Robert L. Carroll ...
, Hillary C. Maddin, Jörg Fröbisch and Jocelyn Falconnet in
2011 File:2011 Events Collage.png, From top left, clockwise: a protester partaking in Occupy Wall Street heralds the beginning of the Occupy movement; protests against Libyan dictator Muammar Gaddafi, who was killed that October; a young man celebrate ...
. 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.Spindler F., Falconnet J. and Fröbisch J. (2016). ''Callibrachion'' and ''Datheosaurus'', two historical and previously mistaken basal caseasaurian synapsids from Europe. ''Acta Palaeontologica Polonica'' 61. .


Etymology

The generic name refers to the location of the taxon in
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
, and “mycter” = nose, refers to the enlarged external naris that characterizes the caseids. The species epithet refers to
ruteni The Ruteni were a Gallic tribe dwelling in the southern part of the Massif Central, around present-day Rodez, during the Iron Age and the Roman period. Name They are mentioned as ''Rutenos'' by Caesar (mid-1st c. BC), ''R̔outēnoì'' (Ῥου ...
(Les Rutènes in French) the
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
name of the Gallic tribe that lived in the Rodez area.


Description

The skull is well-preserved but has suffered of a slight flattening as a result of a
tectonic Tectonics (; ) are the processes that control the structure and properties of the Earth's crust and its evolution through time. These include the processes of mountain building, the growth and behavior of the strong, old cores of continents k ...
thrust exerted toward the right side and the front. As in other caseid, the skull is small compared to the skeleton (here mainly the forelimbs), and shows very large external nares, a short facial region, and a dorsal surface of the skull dotted of numerous small depressions. In addition, they are many palatal teeth, and the teeth of the upper jaws are numerous (four premaxillary and eleven maxillary teeth) and spatulated with crowns having 5 to 8 cusps. The front teeth, fairly long and slightly recurved, were probably suited to aid in gathering vegetation into the mouth, whereas it is presumed that palatal teeth had to worked in conjunction with a tough and massive muscular
tongue The tongue is a muscular organ (anatomy), organ in the mouth of a typical tetrapod. It manipulates food for mastication and swallowing as part of the digestive system, digestive process, and is the primary organ of taste. The tongue's upper surfa ...
as indicated by the presence of a very well-developed hyoid apparatus. Characteristically, ''Euromycter'' shows an unusually broad skull, large
temporal fenestra An infratemporal fenestra, also called the lateral temporal fenestra or simply temporal fenestra, is an opening in the skull behind the orbit in some animals. It is ventrally bordered by a zygomatic arch. An opening in front of the eye sockets, ...
, and lack of expansion of the axial neural spine. It can be distinguished from other caseids by the presence of a supernumerary blade-like intranarial bone located posteromedially to the septomaxilla, proportional differences in forelimb and manus, presence of an accessory proximal articulation between metacarpals 3 and 4, medial recurvature of metacarpal, and its manual phalangeal formula of 2-3-4-4-3.


Stratigraphic range

The holotype of ''Euromycter'' comes from the top of the red pelitic beds of the M1 Member, in the basal part of the Grès Rouge (“Red Sandstone”) Group, a sedimentary sequence subdivided into five hectometric members (M1 to M5) localized in the western
Rodez Rodez ( or ; oc, Rodés, ) is a small city and commune in the South of France, about 150 km northeast of Toulouse. It is the prefecture of the department of Aveyron, region of Occitania (formerly Midi-Pyrénées). Rodez is the seat of the ...
basin. The deposits are interpreted as a playa-lake environment (or Sabkha) under a semi-arid, hot climate. The age of the Grès Rouge Group is uncertain but it is regarded as contemporaneous to the Saxonian Group of the neighbouring
Lodève Lodève (; oc, Lodeva ) is a commune in the département of Hérault, in the Occitanie region in southern France. It is a sub-prefecture of the department. The derivation of the city's name is from Gaulish ''Luteva'', composed of lut-, swamp, ...
basin, where
radiometric Radiometry is a set of techniques for measuring electromagnetic radiation, including visible light. Radiometric techniques in optics characterize the distribution of the radiation's power in space, as opposed to photometric techniques, which cha ...
and
magnetostratigraphic Magnetostratigraphy is a geophysical correlation technique used to date sedimentary and volcanic sequences. The method works by collecting oriented samples at measured intervals throughout the section. The samples are analyzed to determine their ' ...
data suggested previously an age between the late Sakmarian (middle of the Early Permian) and the early
Lopingian The Lopingian is the uppermost series/last epoch of the Permian. It is the last epoch of the Paleozoic. The Lopingian was preceded by the Guadalupian and followed by the Early Triassic. The Lopingian is often synonymous with the informal terms l ...
(early Late Permian). However, new
chronostratigraphic Chronostratigraphy is the branch of stratigraphy that studies the ages of rock strata in relation to time. The ultimate aim of chronostratigraphy is to arrange the sequence of deposition and the time of deposition of all rocks within a geological ...
and magnetostratigraphic data for the Saxonian Group indicate an age between the Artinskian (for the Rabejac Formation and the Octon Member of the Salagou Formation) and the late Roadian- Wordian and possibly early Capitanian (for La Lieude Formation). A more precise stratigraphic correlation of the Permian Rodez basin with that of Lodève was proposed in 2022 by Werneburg and colleagues. In the Rodez basin, the FII Formation of the Salabru Group, underlying the Grès Rouge Group, yielded palynomorphs,
conchostracans Clam shrimp are a group of bivalved branchiopod crustaceans that resemble the unrelated Bivalvia, bivalved molluscs. They are extant and also known from the fossil record, from at least the Devonian period and perhaps before. They were originally ...
, and a
tetrapod 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 theraps ...
footprints assemblage equivalent to that of the Viala Formation of the Lodève basin. The lower part of the Viala Formation yielded a radiometric age of 290.96 ± 0.19 Ma corresponding to the late Sakmarian. It has also been determined that the M1 and M2 megasequences of the Rodez basin are equivalent to the Rabéjac Formation of the Lodève basin (and also of the Combret Member of the Saint-Pierre Formation of the Saint-Affrique basin in southern
Aveyron Aveyron (; oc, Avairon; ) is a department in the region of Occitania, Southern France. It was named after the river Aveyron. Its inhabitants are known as ''Aveyronnais'' (masculine) or ''Aveyronnaises'' (feminine) in French. The inhabitants o ...
). Above the Rabéjac Formation, the lower two-thirds of the Octon Member of the Salagou Formation yielded four
tuff Tuff is a type of rock made of volcanic ash ejected from a vent during a volcanic eruption. Following ejection and deposition, the ash is lithified into a solid rock. Rock that contains greater than 75% ash is considered tuff, while rock cont ...
horizons radiometrically dated. The oldest tuff horizon provided an age of 284.40 ± 0.07 Ma corresponding to the latest Artinskian. Based on this dating, the Rabéjac Formation and the correlative M1 and M2 megasequences of the Grès Rouge Group can be dated to the late Artinskian. A conclusion consistent with the magnetostratigraphy which suggests that the entire Grès Rouge Group (members M1 to M5) would have an age between the late Artinskian and the early Wordian.


Discovery

During a prospecting survey carried out in the Permian red sandstones outcropping in
badlands Badlands are a type of dry terrain where softer sedimentary rocks and clay-rich soils have been extensively eroded."Badlands" in ''Chambers's Encyclopædia''. London: George Newnes, 1961, Vol. 2, p. 47. They are characterized by steep slopes, m ...
on the western flank of the Cayla Hill ( commune of Valady, northwest of
Rodez Rodez ( or ; oc, Rodés, ) is a small city and commune in the South of France, about 150 km northeast of Toulouse. It is the prefecture of the department of Aveyron, region of Occitania (formerly Midi-Pyrénées). Rodez is the seat of the ...
), the paleontologists Denise Sigogneau-Russell and Donald Eugene Russell discovered in the summer of 1970 the skeletons of two herbivorous reptiles. An eroded vertebra picked up on the western slope of the hill led the scientists to explore the surrounding canyons where they discovered a large articulated skeleton still in place in the sediments but damaged by erosion. The skull, neck, most of the limbs, and the tail had been washed away and destroyed. On the southeastern flank of the same hill, the same team discovered a bone fragment from a different animal. The systematic exploration of surrounding slopes permits the discovery of bones still in place in the rock belonging to a smaller animal than the first. The whole back of the skeleton had already destroyed by erosion. The right forearm was exposed at the surface of the rock, then was found the skull, several articulated cervical vertebrae, the left forearm articulated with the complete left manus, a piece of the right humerus and parts of the right shoulder. Both specimens were identified as caseids pelycosaurs. The smaller specimen found at about two kilometers horizontally from the first skeleton, but 120 meters lower stratigraphically, was described in 1974 and assigned to a new species of the genus '' Casea'', ''Casea rutena''. This animal was celebrated as the first caseid found in
Western Europe Western Europe is the western region of Europe. The region's countries and territories vary depending on context. The concept of "the West" appeared in Europe in juxtaposition to "the East" and originally applied to the ancient Mediterranean ...
, making of this species a geographical link with other specimens of the family that were previously known only in the south central
United States The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territorie ...
(
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 ...
) and in the northern
European Russia European Russia (russian: Европейская Россия, russian: европейская часть России, label=none) is the western and most populated part of Russia. It is geographically situated in Europe, as opposed to the cou ...
. In 2008, a
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
analysis of Caseidae demonstrates the
paraphyly In taxonomy, a group is paraphyletic if it consists of the group's last common ancestor and most of its descendants, excluding a few monophyletic subgroups. The group is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
of the genus ''Casea'', the French species representing a distinct unnamed genus. Three years later, the species ''Casea rutena'' was removed of the genus ''Casea'' and assigned to a new genus, ''Euromycter'', in the new combination ''Euromycter rutenus''. In the same article the authors described the larger and stratigraphycally younger skeleton from the Cayla Hill, and assigned it to a new genus named ''
Ruthenosaurus ''Ruthenosaurus'' is an extinct genus of caseid synapsids that lived in what is now southern France during the Early Permian (late Artinskian) about 285 million years ago. It is known from the holotype MNHN.F.MCL-1 an articulated partial pos ...
''.


Classification

In the first phylogenetic analysis of the caseids published in 2008, ''Euromycter'', then still designated as ''“Casea” rutena'', was recovered as the
sister taxon In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and t ...
to a derive
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
containing ''
Ennatosaurus tecton ''Ennatosaurus'' is an extinct genus of caseid synapsid that lived during the Middle Permian (late Roadian - early Wordian) in northern European Russia. The genus is only represented by its type species, ''Ennatosaurus tecton'', which was named ...
'', ''
Cotylorhynchus romeri ''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 ...
'' and ''
Angelosaurus dolani ''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 casei ...
''. Below the first phylogenetic analysis of Caseidae published by Maddin et al. in 2008. A phylogenetic analysis made by Benson shows a similar position for ''Euromycter'' (again regarded as “''Casea''” ''rutena''). Below the phylogenetic analysis of Caseasauria published by Benson in 2012. In 2015, Romano & Nicosia found a similar position for ''Euromycter'' in their most
parsimonious Occam's razor, Ockham's razor, or Ocham's razor ( la, novacula Occami), also known as the principle of parsimony or the law of parsimony ( la, lex parsimoniae), is the problem-solving principle that "entities should not be multiplied beyond neces ...
analysis including nearly all caseids (to the exclusion of the very fragmentary '' Alierasaurus ronchi'' from
Sardinia Sardinia ( ; it, Sardegna, label=Italian, Corsican and Tabarchino ; sc, Sardigna , sdc, Sardhigna; french: Sardaigne; sdn, Saldigna; ca, Sardenya, label=Algherese and Catalan) is the second-largest island in the Mediterranean Sea, after ...
). Below the most parsimonious phylogenetic analysis published by Romano & Nicosia in 2015.


References

{{Taxonbar, from=Q5411994 Caseasaurs Prehistoric synapsid genera Fossil taxa described in 2011 Fossils of France