Euromycter
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''Euromycter'' is an
extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
caseid 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 (Texas, Oklaho ...
synapsid Synapsida is a diverse group of tetrapod vertebrates that includes all mammals and their extinct relatives. It is one of the two major clades of the group Amniota, the other being the more diverse group Sauropsida (which includes all extant rept ...
s that lived in what is now
Southern France Southern France, also known as the south of France or colloquially in French as , is a geographical area consisting of the regions of France that border the Atlantic Ocean south of the Marais Poitevin,Louis Papy, ''Le midi atlantique'', Atlas e ...
during the
Early Permian 01 or 01 may refer to: * The year 2001, or any year ending with 01 * The month of January * 1 (number) Music * '01 (Richard Müller album), ''01'' (Richard Müller album), 2001 * 01 (Urban Zakapa album), ''01'' (Urban Zakapa album), 2011 * ''01011 ...
(late
Artinskian In the geologic timescale, the Artinskian is an age (geology), age or stage (stratigraphy), stage of the Permian. It is a subdivision of the Cisuralian Epoch (geology), Epoch or series (stratigraphy), Series. The Artinskian likely lasted between ...
) about 285 million years ago. The
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
and only known specimen of ''Euromycter'' ( MNHN.F.MCL-2) includes the complete
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 ...
with
lower jaw In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone i ...
s and
hyoid apparatus The hyoid apparatus is the collective term used in veterinary anatomy for the bones which suspend the tongue and larynx. It consists of pairs of stylohyoid, thyrohyoid, epihyoid and ceratohyoid bones, and a single basihyoid bone. The hyoid appar ...
, six
cervical vertebrae In tetrapods, cervical vertebrae (: 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 saurop ...
with
proatlas The proatlas is a paired bone in the skeleton of many vertebrates that occurs between the skull and the first cervical vertebra. It ossifies endochondrally. A number of different interpretations have been made of the proatlas. The most common i ...
, anterior part of
interclavicle An interclavicle is a bone which, in most tetrapods, is located between the clavicles. Therian mammals ( marsupials and placentals) are the only tetrapods which never have an interclavicle, although some members of other groups also lack one. In ...
, partial right
clavicle The clavicle, collarbone, or keybone is a slender, S-shaped long bone approximately long that serves as a strut between the scapula, shoulder blade and the sternum (breastbone). There are two clavicles, one on each side of the body. The clavic ...
, right posterior
coracoid A coracoid is a paired bone which is part of the shoulder assembly in all vertebrates except therian mammals (marsupials and placentals). In therian mammals (including humans), a coracoid process is present as part of the scapula, but this is n ...
, distal head of right
humerus The humerus (; : humeri) 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 (bone), radius and ulna, and consists of three sections. The humeral upper extrem ...
, left and right
radius In classical geometry, a radius (: radii or radiuses) of a circle or sphere is any of the line segments from its Centre (geometry), center to its perimeter, and in more modern usage, it is also their length. The radius of a regular polygon is th ...
, left and right
ulna The ulna or ulnar bone (: ulnae or ulnas) is a long bone in the forearm stretching from the elbow to the wrist. It is on the same side of the forearm as the little finger, running parallel to the Radius (bone), radius, the forearm's other long ...
, 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 (département of
Aveyron Aveyron (; ) is a Departments of France, department in the Regions of France, region of Occitania (administrative region), Occitania, Southern France. It was named after the river Aveyron (river), Aveyron. Its inhabitants are known as ''Aveyro ...
),
Rodez Rodez (, , ; , ) 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 communau ...
Basin. It was first assigned to the species ''"
Casea ''Casea'' is a genus of Herbivore, herbivorous Caseidae, 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 Sam ...
" 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 year marked the start of a Arab Spring, series of protests and revolutions throughout the Arab world advocating for democracy, reform, and economic recovery, later leading to the depositions of world leaders in Tunisia, Egypt, and Yemen ...
. 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 continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...
, 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'' (Ῥ ...
(Les Rutènes in French) the
Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
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 ( via Latin ) are the processes that result in the structure and properties of the Earth's crust and its evolution through time. The field of ''planetary tectonics'' extends the concept to other planets and moons. These processes ...
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 The palate () is the roof of the mouth in humans and other mammals. It separates the oral cavity from the nasal cavity. A similar structure is found in crocodilians, but in most other tetrapods, the oral and nasal cavities are not truly sepa ...
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 work in conjunction with a tough and massive muscular
tongue The tongue is a Muscle, muscular organ (anatomy), organ in the mouth of a typical tetrapod. It manipulates food for chewing and swallowing as part of the digestive system, digestive process, and is the primary organ of taste. The tongue's upper s ...
as indicated by the presence of a very well-developed
hyoid apparatus The hyoid apparatus is the collective term used in veterinary anatomy for the bones which suspend the tongue and larynx. It consists of pairs of stylohyoid, thyrohyoid, epihyoid and ceratohyoid bones, and a single basihyoid bone. The hyoid appar ...
. Characteristically, ''Euromycter'' shows an unusually broad skull, large
temporal fenestra Temporal fenestrae are openings in the temporal region of the skull of some amniotes, behind the orbit (eye socket). These openings have historically been used to track the evolution and affinities of reptiles. Temporal fenestrae are commonly (al ...
, and lack of expansion of the axial
neural spine Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spinal ...
. It can be distinguished from other caseids by the presence of a
supernumerary Supernumerary means "exceeding the usual number". Supernumerary may also refer to: * Supernumerary actor, a performer in a film, television show, or stage production who has no role or purpose other than to appear in the background, more common ...
blade-like intranarial bone located posteromedially to the septomaxilla, proportional differences in forelimb and manus, presence of an accessory proximal articulation between
metacarpals In human anatomy, the metacarpal bones or metacarpus, also known as the "palm bones", are the appendicular skeleton, appendicular bones that form the intermediate part of the hand between the phalanges (fingers) and the carpal bones (wrist, wris ...
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 (, , ; , ) 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 communau ...
basin. The deposits are interpreted as a playa-lake environment (or
Sabkha A sabkha () is a predominately coastal, supratidal mudflat or sandflat in which evaporite-saline minerals accumulate as the result of a semiarid to arid climate. Sabkhas are gradational between land and intertidal zone within restricted coast ...
) 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 (; , ; ) is a Communes of France, commune in the Departments of France, department of Hérault, in the Regions of France, region of Occitania (administrative region), Occitania, southern France. It is a Subprefectures in France, subprefe ...
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 ch ...
and magnetostratigraphic data suggested previously an age between the late
Sakmarian In the geologic timescale, the Sakmarian is an age or stage of the Permian period. It is a subdivision of the Cisuralian Epoch or Series. The Sakmarian lasted between 293.52 and million years ago (Ma). It was preceded by the Asselian and fo ...
(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 te ...
(early Late Permian). However, new chronostratigraphic and magnetostratigraphic data for the Saxonian Group indicate an age between the
Artinskian In the geologic timescale, the Artinskian is an age (geology), age or stage (stratigraphy), stage of the Permian. It is a subdivision of the Cisuralian Epoch (geology), Epoch or series (stratigraphy), Series. The Artinskian likely lasted between ...
(for the Rabejac Formation and the Octon Member of the Salagou Formation) and the late
Roadian In the geologic timescale, the Roadian is an age or stage of the Permian. It is the earliest or lower of three subdivisions of the Guadalupian Epoch or Series. The Roadian lasted between and million years ago (Ma). It was preceded by the Kun ...
-
Wordian In the geologic timescale, the Wordian is an age or stage of the Permian. It is the middle of three subdivisions of the Guadalupian Epoch or Series. The Wordian lasted between and million years ago (Ma). It was preceded by the Roadian and fo ...
and possibly early
Capitanian In the geologic timescale, the Capitanian is an age or stage of the Permian. It is also the uppermost or latest of three subdivisions of the Guadalupian Epoch or Series. The Capitanian lasted between and million years ago. It was preceded by t ...
(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
palynomorph Palynology is the study of microorganisms and microscopic fragments of mega-organisms that are composed of acid-resistant organic material and occur in sediments, sedimentary rocks, and even some metasedimentary rocks. Palynomorphs are the mic ...
s, conchostracans, and a
tetrapod A tetrapod (; from Ancient Greek :wiktionary:τετρα-#Ancient Greek, τετρα- ''(tetra-)'' 'four' and :wiktionary:πούς#Ancient Greek, πούς ''(poús)'' 'foot') is any four-Limb (anatomy), limbed vertebrate animal of the clade Tetr ...
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 Saint-Affrique (; Languedocien: ''Sant Africa'') is a commune in the Aveyron department in Southern France. History Saint-Affrique grew in the 6th century around the tomb of St. Africain, bishop of Comminges. In the 12th century a fortre ...
basin in southern
Aveyron Aveyron (; ) is a Departments of France, department in the Regions of France, region of Occitania (administrative region), Occitania, Southern France. It was named after the river Aveyron (river), Aveyron. Its inhabitants are known as ''Aveyro ...
). 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 co ...
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, ...
on the western flank of the Cayla Hill ( commune of Valady, northwest of
Rodez Rodez (, , ; , ) 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 communau ...
), 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 permitted 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 Caseidae are an Extinction, extinct Family (biology), family of Basal (phylogenetics), 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 so ...
pelycosaur Pelycosaur ( ) is an older term for basal or primitive Late Paleozoic synapsids, excluding the therapsids and their descendants. Previously, the term mammal-like reptile was used, and Pelycosauria was considered an order, but this is now thoug ...
s. 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'' is a genus of Herbivore, herbivorous Caseidae, 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 Sam ...
'', ''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 extent varies depending on context. The concept of "the West" appeared in Europe in juxtaposition to "the East" and originally applied to the Western half of 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 (USA), also known as the United States (U.S.) or America, is a country primarily located in North America. It is a federal republic of 50 U.S. state, states and a federal capital district, Washington, D.C. The 48 ...
(
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 ...
) and in the northern
European Russia European Russia is the western and most populated part of the Russia, Russian Federation. It is geographically situated in Europe, as opposed to the country's sparsely populated and vastly larger eastern part, Siberia, which is situated in Asia ...
. In 2008, a
phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...
analysis of Caseidae demonstrates the
paraphyly Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping 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''.


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 ...
to a derive
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
containing '' Ennatosaurus tecton'', '' Cotylorhynchus romeri'' and '' Angelosaurus dolani''. Below the first
phylogenetic analysis In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical data ...
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 In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical data ...
of Caseasauria published by Benson in 2012. In 2015, Romano & Nicosia found a similar position for ''Euromycter'' in their most
parsimonious In philosophy, Occam's razor (also spelled Ockham's razor or Ocham's razor; ) is the problem-solving principle that recommends searching for explanations constructed with the smallest possible set of elements. It is also known as the principle o ...
analysis including nearly all caseids (to the exclusion of the very fragmentary '' Alierasaurus ronchi'' from
Sardinia Sardinia ( ; ; ) is the Mediterranean islands#By area, second-largest island in the Mediterranean Sea, after Sicily, and one of the Regions of Italy, twenty regions of Italy. It is located west of the Italian Peninsula, north of Tunisia an ...
). Below the most parsimonious phylogenetic analysis published by Romano & Nicosia in 2015.


References

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