Raranimus Dashankouensis (2)
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''Raranimus'' is an extinct
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 therapsids of the
Middle Permian The Guadalupian is the second and middle series/epoch of the Permian. The Guadalupian was preceded by the Cisuralian and followed by the Lopingian. It is named after the Guadalupe Mountains of New Mexico and Texas, and dates between 272.95 ± 0. ...
. It was described in 2009 from a partial skull found in 1998 from the Dashankou locality of the Qingtoushan Formation, outcropping in the
Qilian Mountains The Qilian Mountains (), together with the Altyn-Tagh sometimes known as the Nan Shan, as it is to the south of the Hexi Corridor, is a northern outlier of the Kunlun Mountains, forming the border between Qinghai and the Gansu provinces of n ...
of
Gansu Gansu is a provinces of China, province in Northwestern China. Its capital and largest city is Lanzhou, in the southeastern part of the province. The seventh-largest administrative district by area at , Gansu lies between the Tibetan Plateau, Ti ...
,
China China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...
. The genus is the most basal known member of the
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 ...
Therapsida, to which the later
Mammalia A mammal () is a vertebrate animal of the class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three middle ear bon ...
belong.


Description

''Raranimus'' shares a number of features with later therapsids and ancestral
Sphenacodontia Sphenacodontia is a stem-based taxon, stem-based clade of derived synapsids. It was defined by Amson and Laurin (2011) as "the largest clade that includes ''Haptodus baylei'', ''Haptodus garnettensis'' and ''Sphenacodon ferox'', but not ''Edaphos ...
. The skull consists of a well preserved
rostrum Rostrum may refer to: * Any kind of a platform for a speaker: **dais **pulpit ** podium * Rostrum (anatomy), a beak, or anatomical structure resembling a beak, as in the mouthparts of many sucking insects * Rostrum (ship), a form of bow on naval ...
. The teeth suggest a carnivorous lifestyle for ''Raranimus'', as the incisors are recurved and the second canines are serrated on their posterior edges. The incisors are morphologically similar to those seen in more derived
theriodont The theriodonts (clade Theriodontia) are a major group of therapsids which appeared during the Middle Permian and which includes the gorgonopsians and the eutheriodonts, itself including the therocephalians and the cynodonts. Naming In 1876, ...
s. The presence of two linguo−labially compressed canines is a diagnostic feature of ''Raranimus''. The presence of two functional canines is characteristic of sphenacodontids, and this condition is seen in no other therapsid other than ''Rananimus''. However, the slender, compressed shape of these canines is a derived characteristic of therapsids, with the canines of similarly sized sphenacodontids being more massively built. The precanines are small and anteriorly serrated, similar to what is seen in the synapsids ''
Dimetrodon ''Dimetrodon'' ( or ; ) is an extinct genus of sphenacodontid synapsid that lived during the Cisuralian (Early Permian) Epoch (geology), epoch of the Permian period, around 295–272 million years ago. With most species measuring long and ...
'' and '' Tetraceratops''. In the palate region of the skull, the anterior process of the
vomer The vomer (; ) is one of the unpaired facial bones of the skull. It is located in the midsagittal line, and articulates with the sphenoid, the ethmoid, the left and right palatine bones, and the left and right maxillary bones. The vomer forms ...
ventrally overlies the
premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammals h ...
at the anterior margin of the
choana The choanae (: choana), posterior nasal apertures or internal nostrils are two openings found at the back of the nasal passage between the nasal cavity and the pharynx, in humans and other mammals (as well as crocodilians and most skinks). They ...
. This overlap is also seen in
dinocephalia Dinocephalians (terrible heads) are a clade of large-bodied early therapsids that flourished in the Early and Middle Permian between 279.5 and 260 million years ago (Ma), but became extinct during the Capitanian mass extinction event. ...
ns. However, unlike any other therapsid, the choanae are short and extend only from the level of the fourth incisor back to the first canine.


Phylogenetics

According to a phylogenetic analysis conducted along with its initial description, ''Raranimus'' is considered to be the basalmost therapsid. There has been some controversy as to whether ''Tetraceratops'' is a therapsid or a more basal
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 ...
. If ''Tetraceratops'' is a therapsid, as has recently been proposed, it would be the oldest and most basal one known, surpassing ''Raranimus'' in age by several million years. However, later studies have questioned the placement of ''Tetraceratops'' within Therapsida, and the 2009 phylogenetic analysis using ''Raranimus'' places the genus outside of the clade. ''Raranimus'' occurs in strata that were deposited during the early Roadian stage of the
Middle Permian The Guadalupian is the second and middle series/epoch of the Permian. The Guadalupian was preceded by the Cisuralian and followed by the Lopingian. It is named after the Guadalupe Mountains of New Mexico and Texas, and dates between 272.95 ± 0. ...
.The hypothesized age for this locality is supported by the presence of the dissorophoid
temnospondyl Temnospondyli (from Greek language, Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') or temnospondyls is a diverse ancient order (biology), order of small to giant tetrapods—often considered Labyrinth ...
'' Anakamacops'', the bolosaurid '' Belebey'', and the basal therapsids '' Biseridens'', ''Sinophoneus'', and ''Stenocybus''.
The sphenacodontids were most diverse before the Roadian in 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 ...
, yet therapsids did not appear as a diverse group until near the Roadian-Wordian boundary. This has left a morphological and temporal gap in the fossil record during which the origin of therapsids must have occurred called Olson’s Extinction. With the general absence of therapsid remains from Olson's Extinction, different hypotheses have developed in order to explain the group's origins and initial diversification. One theory suggests that therapsids diversified quickly through rapid
apomorphy In phylogenetics, an apomorphy (or derived trait) is a novel Phenotypic trait, character or character state that has evolution, evolved from its ancestral form (or Plesiomorphy and symplesiomorphy, plesiomorphy). A synapomorphy is an apomorphy sh ...
accumulation sometime during the gap, while the other proposes that therapsids evolved gradually over the course of up to 35 Ma. Only recently have remains of basal therapsids such as ''Raranimus'' been found from China that occur during Olson's Extinction. Other therapsids that are known to have existed during the gap include '' Sinophoneus'' and '' Stenocybus''. Below is a
cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...
modified from Liu ''et al.'', 2009 depicting the phylogenetic relationships of ''Raranimus'' with therapsids that occur in Olson's Gap highlighted:


References

{{Taxonbar, from=Q2044958 Permian synapsids of Asia Guadalupian synapsids Prehistoric therapsid genera Fossil taxa described in 2009 Guadalupian genus first appearances Guadalupian genus extinctions