Raranimus dashankouensis
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''Raranimus'' is an extinct genus of therapsids of the Middle Permian. It was described in 2009 from a partial skull found in 1998 from the Dashankou locality of the
Xidagou Formation The Qingtoushan Formation is a Middle Permian-age geologic formation in the Qilian Mountains of Gansu, China. It is known for its diverse tetrapod fauna known as the Dashankou fauna, which likely dates to the Roadian, and includes some of the ol ...
, outcropping in the Qilian Mountains of
Gansu Gansu (, ; alternately romanized as Kansu) is a province in Northwest China. Its capital and largest city is Lanzhou, in the southeast part of the province. The seventh-largest administrative district by area at , Gansu lies between the Tibet ...
,
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...
. The genus is the most
basal Basal or basilar is a term meaning ''base'', ''bottom'', or ''minimum''. Science * Basal (anatomy), an anatomical term of location for features associated with the base of an organism or structure * Basal (medicine), a minimal level that is nec ...
known member of the
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, ...
Therapsida, to which the later Mammalia belong.


Description

''Raranimus'' shares a number of features with later therapsids and ancestral Sphenacodontia. The skull consists of a well preserved rostrum. 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 theriodonts. 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 ,) meaning "two measures of teeth,” is an extinct genus of non-mammalian synapsid that lived during the Cisuralian (Early Permian), around 295–272 million years ago (Mya). It is a member of the family Sphenacodontid ...
'' and '' Tetraceratops''. In the palate region of the skull, the anterior process of the
vomer The vomer (; lat, vomer, lit=ploughshare) 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 maxill ...
ventrally overlies the premaxilla at the anterior margin of the choana. This overlap is also seen in dinocephalians. 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. 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 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') is a diverse order (biology), order of small to giant tetrapods—often considered Labyrinthodontia, primitive amphi ...
''
Anakamacops ''Anakamacops'' (meaning "similar to ''Kamacops''" in Greek) is a genus of dissorophid temnospondyl from the early Middle Permian of China. It is known from the right side of a snout that was described in 1999 from the Dashankou locality of the ...
'', the Bolosauridae, bolosaurid ''Belebey (genus), Belebey'', and the basal therapsids ''Biseridens'', ''Sinophoneus'', and ''Stenocybus''.
The sphenacodontids were most diverse before the Roadian in the Early Permian, 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 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 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