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''Ruthenosaurus'' 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
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 Synapsids + (, 'arch') > () "having a fused arch"; synonymous with ''theropsids'' (Greek, "beast-face") are one of the two major groups of animals that evolved from basal amniotes, the other being the sauropsids, the group that includes reptil ...
s 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 regions and territories in the Americas and the Atlantic, Pacific and Indian Oceans. Its metropolitan area ...
during the Early Permian (late
Artinskian In the geologic timescale, the Artinskian is an age or stage of the Permian. It is a subdivision of the Cisuralian Epoch or Series. The Artinskian likely lasted between and million years ago (Ma) according to the most recent revision of th ...
) about 285 million years ago. It is known from 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 sever ...
MNHN.F.MCL-1 an articulated partial
postcrania Postcrania (postcranium, adjective: postcranial) in zoology and vertebrate paleontology is all or part of the skeleton apart from the skull. Frequently, fossil remains, e.g. of dinosaurs or other extinct tetrapods, consist of partial or isolated sk ...
l skeleton. It was collected by D. Sigogneau-Russell and D. Russell in 1970 in the upper part of the M2 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 ...
, in the Rodez Basin, 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 ...
), in
Occitanie Occitanie may refer to: *Occitania, a region in southern France called ''Occitanie'' in French *Occitania (administrative region) Occitania ( ; french: Occitanie ; oc, Occitània ; ca, Occitània ) is the southernmost administrative region of ...
Region In geography, regions, otherwise referred to as zones, lands or territories, are areas that are broadly divided by physical characteristics ( physical geography), human impact characteristics ( human geography), and the interaction of humanity an ...
. It was first named by Robert R. Reisz, Hillary C. Maddin, Jörg Fröbisch and Jocelyn Falconnet in 2011, and 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 ...
is ''Ruthenosaurus russellorum''.


Etymology

Ruthenosaurus is named after a Gallic regional tribe, Ruthenie (in
Medieval Latin Medieval Latin was the form of Literary Latin used in Roman Catholic Western Europe during the Middle Ages. In this region it served as the primary written language, though local languages were also written to varying degrees. Latin functione ...
), that have also given ''Ruthénois'', the name of the inhabitants of the town of Rodez; and the
Ancient Greek Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Dark Ages (), the Archaic p ...
''sauros'', "lizard". The type species is named in honour of Drs Denise Sigogneau-Russell and Donald E. Russell, the original collectors of the holotype.


Discovery

The holotype of ''Ruthenosaurus'' was discovered in the summer 1970 by the paleontologists Denise Sigogneau-Russell and Donald Eugene Russell, during a prospecting survey carried out in 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 near the
commune A commune is an alternative term for an intentional community. Commune or comună or comune or other derivations may also refer to: Administrative-territorial entities * Commune (administrative division), a municipality or township ** Communes of ...
of Valady, northwest of Rodez. 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 were missing, probably destroyed by erosion. The known material includes ribs and
vertebra 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 characteristi ...
e of 18 presacrals, three sacrals, and 12 anterior caudals; incomplete
scapulocoracoid The scapulocoracoid is the unit of the pectoral girdle that contains the coracoid and scapula. The coracoid itself is a beak-shaped bone that is commonly found in most vertebrates with a few exceptions. The scapula is commonly known as the ''shoulde ...
s and interclavicule; right humerus badly crushed and damaged; left humerus in two pieces, shaft damaged, but proximal and distal heads well preserved; complete left
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 nearly complete
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 ...
; complete right
femur The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates wit ...
, complete right
tibia The tibia (; ), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outside of the tibia); it connects ...
, and proximal portion of right
fibula The fibula or calf bone is a leg bone on the lateral side of the tibia, to which it is connected above and below. It is the smaller of the two bones and, in proportion to its length, the most slender of all the long bones. Its upper extremity i ...
; and the complete right pelvis overlain by vertebral column. On the southeastern flank of the same hill, but in older strata, the same team discovered the anterior part of a skeleton (including the skull) belonging to a smaller animal. This specimen was first assigned to a new species of the genus ''
Casea ''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 Willisto ...
'', ''Casea rutena''. But it is now regarded as a distinct genus named ''
Euromycter ''Euromycter'' 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. The holotype and only known specimen of ''Euromycter'' ( MNHN.F.MCL-2) incl ...
'', with the new combination ''Euromycter rutenus''. The larger skeleton, found stratigraphycally 120 meters above the ''Euromycter'' level, remained largely unprepared until 2003. The few overlapping elements with ''Euromycter'' suggested that it belongs to a different taxon named ''Ruthenosaurus russellorum'' in 2011.


Description

''Ruthenosaurus'' is diagnosed by several
autapomorphies In phylogenetics, an autapomorphy is a distinctive feature, known as a derived trait, that is unique to a given taxon. That is, it is found only in one taxon, but not found in any others or outgroup taxa, not even those most closely related to t ...
including dorsal vertebrae with anteriorly tilting neural spines and a diamond-shaped outline in transverse section; a first
sacral Sacral may refer to: *Sacred Sacred describes something that is dedicated or set apart for the service or worship of a deity; is considered worthy of spiritual respect or devotion; or inspires awe or reverence among believers. The property ...
rib with robust distal head, twice that of the second sacral rib; and a short
iliac blade The crest of the ilium (or iliac crest) is the superior border of the wing of ilium and the superiolateral margin of the greater pelvis. Structure The iliac crest stretches posteriorly from the anterior superior iliac spine (ASIS) to the posteri ...
with prominent posterior
process A process is a series or set of activities that interact to produce a result; it may occur once-only or be recurrent or periodic. Things called a process include: Business and management *Business process, activities that produce a specific se ...
. It can be distinguished from ''Euromycter'', from older deposits of the same locality, by the shape of the distal part of the humerus, including an ectepicondylar notch rather than a fully enclosed
foramen In anatomy and osteology, a foramen (;Entry "foramen"
in
, the specific shape of the ulna, and the overall robustness of the specimen. The lack of fusion of the
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 ...
with their respective vertebral
centra Centra is a convenience shop chain that operates throughout Ireland. The chain operates as a symbol group owned by Musgrave Group, the food wholesaler, meaning the stores are all owned by individual franchisees. The chain has three different ...
and incomplete ossification of the ends of the limb elements, including the absence of an ossified
olecranon The olecranon (, ), is a large, thick, curved bony eminence of the ulna, a long bone in the forearm that projects behind the elbow. It forms the most pointed portion of the elbow and is opposite to the cubital fossa or elbow pit. The olecranon ...
on the ulna, show clearly that this specimen represents a juvenile individual. However, it is distinctly larger than the fully mature specimen of ''Euromycter'', suggesting a very large size for adult ''Ruthenosaurus''.


Stratigraphic range

The holotype of ''Ruthenosaurus'' was found in the uppert part of the red pelitic beds of the M2 Member, which belongs to the Grès Rouge (“Red Sandstone”) Group, a sedimentary sequence subdivided into five hectometric members (M1 to M5) localized in the western Rodez basin. Like the M1 Member that has yielded ''Euromycter'', the deposits of the M2 Member are interpreted as a playa-lake environment (or
Sabkha A sabkha ( ar, سبخة) is a coastal, supratidal mudflat or sandflat in which evaporite-saline minerals accumulate as the result of semiarid to arid climate. Sabkhas are gradational between land and intertidal zone within restricted coastal p ...
) 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 neighbourhing
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 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. 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 followed by t ...
(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 geologic ...
and magnetostratigraphic data for the Saxonian Group indicate an age between the
Artinskian In the geologic timescale, the Artinskian is an age or stage of the Permian. It is a subdivision of the Cisuralian Epoch or Series. The Artinskian likely lasted between and million years ago (Ma) according to the most recent revision of th ...
(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 ...
-
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 f ...
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 ...
(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 dust" (from grc-gre, παλύνω, palynō, "strew, sprinkle" and ''-logy'') or of "particles that are strewn". A classic palynologist analyses particulate samples collected from the air, from water, or from deposit ...
s, conchostracans, 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 t ...
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 (; 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 ...
). 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 ...
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.


Classification

The first phylogenetic analysis of the caseids containing ''Ruthenosaurus'' was made by Romano & Nicosia in 2015. 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 ...
), ''Ruthenosaurus'' is found very close to '' Cotylorhynchus'' and most specifically to the giant '' Cotylorhynchus hancocki''. However, Romano & Nicosia have concluded that given the large body size of the two taxa and considering the absence of cranial material for ''Ruthenosaurus'', it is possible that their close similarity is size related. Below the most parsimonious phylogenetic analysis published by Romano & Nicosia in 2015. A phylogenetic analysis published in 2022 by Werneburg and colleagues suggests a possible close relationship between ''Ruthenosaurus'' and '' Caseopsis''. ''Ruthenosaurus'' is also recovered as the sister taxon to a clade containing the genera ''Alierasaurus'', ''Cotylorhynchus'' and ''
Lalieudorhynchus ''Lalieudorhynchus'' is an extinct genus of caseid synapsids that lived during the Guadalupian (= Middle Permian) in what is now the south of France. The genus is only known by its type species, ''Lalieudorhynchus gandi'', which was named in 2022 ...
''. Below is the cladogram published by Werneburg and colleagues in 2022.


References

{{Taxonbar, from=Q7383306 Caseasaurs Prehistoric synapsid genera Fossils of France Fossil taxa described in 2011 Permian synapsids of Europe Taxa named by Robert R. Reisz