Kunmunella
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''Kunmunella'' is an extinct raoellid
artiodactyl Artiodactyls are placental mammals belonging to the order Artiodactyla ( , ). Typically, they are ungulates which bear weight equally on two (an even number) of their five toes (the third and fourth, often in the form of a hoof). The other t ...
which inhabited what is now northern
India India, officially the Republic of India, is a country in South Asia. It is the List of countries and dependencies by area, seventh-largest country by area; the List of countries by population (United Nations), most populous country since ...
during the middle-upper
Eocene The Eocene ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...
(
Ypresian In the geologic timescale the Ypresian is the oldest age (geology), age or lowest stage (stratigraphy), stratigraphic stage of the Eocene. It spans the time between , is preceded by the Thanetian Age (part of the Paleocene) and is followed by th ...
). There are two species of ''Kunmunella'': ''K. kalakotensis'' and ''K. transversa''. The former species was named after the town of
Kalakote Kalakote is a town located in Rajouri district of Jammu region in the Indian union territory of Jammu and Kashmir. Kalakote is noted for its coal mines and has first Thermal Power Plant of North India. Additionally, the town serves as the head ...
within
Jammu and Kashmir Jammu and Kashmir may refer to: * Jammu and Kashmir (union territory), a region administered by India as a union territory since 2019 * Jammu and Kashmir (state), a region administered by India as a state from 1952 to 2019 * Jammu and Kashmir (prin ...
, the place from which it was found, while the latter species was named after the transverse nature of the upper teeth. ''K. kalakotensis'' is known from a
palate 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 sep ...
with its respective teeth, a left maxillary ramus, a left
mandible 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 ...
, two poorly preserved maxillas, and two isolated
molars The molars or molar teeth are large, flat tooth, teeth at the back of the mouth. They are more developed in mammal, mammals. They are used primarily to comminution, grind food during mastication, chewing. The name ''molar'' derives from Latin, '' ...
. ''K. transversa'' is known from only a right maxilla.


Discovery

The remains of ''Kunmunella'' were first found by Ranga Rao, 1971. He found only a left mandible, and assigned it to a new species of ''
Indohyus ''Indohyus'' (Meaning "India's pig" from the Greek words ''Indos'', "from India" and ''hûs'', "pig") is an extinct genus of artiodactyl known from Eocene fossils in Asia. This small chevrotain-like animal found in the Himalayas is among the clo ...
'', ''I. kalakotensis'', after the region in which he discovered the animal. In the following year, an isolated upper third molar would be found by Sahni and Khare, which would be assigned to the new
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 ...
''Kunmunella''. This new
taxon In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
would be named ''K. rajouriensis'', named after the
Rajouri district Rajouri is a district in the Jammu division of Indian union territory of Jammu and Kashmir (union territory), Jammu and Kashmir, in the larger Kashmir#Dispute, disputed Kashmir region. The application of the term "administered" to the various re ...
where the fossil was found. In 1985, a right maxilla with its respective teeth was found and assigned to the new species ''K. transversa''.


Classification


External Classification

The first remains of ''Kunmunella'' discovered were first assigned to ''Indohyu''s. When ''Kunmunella'' was first named, it was assigned to
Helohyidae Helohyidae were a group of artiodactyl mammals. They were most prominent in the mid-to-upper Eocene (~50 to 39 million years ago). Description Helohyidae share vague similarities to present-day pigs, though were slimmer in build. They possessed ...
, which was at that time within
Dichobunidae Dichobunidae is an extinct family of basal artiodactyl mammals from the early Eocene to late Oligocene of North America, Europe, and Asia. The Dichobunidae include some of the earliest known artiodactyls, such as ''Diacodexis''. Description T ...
, due to the morphology of its molar, which was at the time the only remains classified under the genus ''Kunmunella''. In 1977, ''Kunmunella'' was moved from Dichobunidae to Helohyidae. In the same year, Helohyidae was moved from Dichobunoidea to Anthracotherioidea (now
Ancodonta Ancodonta is an infraorder of artiodactyl ungulates including modern hippopotamus and all mammals closer to hippos than to cetaceans (whales). Ancodonts first appeared in the Middle Eocene, with some of the earliest representatives found in fossi ...
) and by extension ''Kunmunella'' was moved to Anthracotherioidea. In 1981, ''Kunmunella'', along with ''Raoella'' (now ''Indohyus'') and ''
Khirtharia ''Khirtharia'' is an extinct genus of raoellid artiodactyl that inhabited what is now northern India and Pakistan during the middle-upper Eocene (early Lutetian, 48 to 45 million years ago). There are three species of ''Khirtharia'': ''K. dayi,' ...
'', were found to all share various
synapomorphies In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to ...
and were grouped together in the new family Raoellidae. However, Thewissen et al. (1987) determined that ''Kunmunella'', along with ''Raoella'', were synonymous with ''Indohyus''. This change was reverted by Thewissen, Williams & Hussain (2001), who found ''Kunmunella'' to be a valid genus. In 2011, Orliac and Ducrocq through a
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 ...
determined ''Kunmunella'' to be the most basal raoellid. In 2020, a study found that ''Kunmunella'' was a junior synonym of ''Indohyus'', although this result was not widely reconginzed.After the discovery of ''
Rajouria ''Rajouria'', named after the Rajouri district in the Indian province of Jammu and Kashmir, is an extinct raoellid artiodactyl from the middle-upper Eocene (early Lutetian). There is only a single species of ''Rajouria'', ''Rajouria gunnelli'', w ...
'' in 2021, Rana et al., through a phylogenetic analysis, determined that the newly found ''Rajouria'' was the most basal member of Raoellidae, with ''Kunmunella'' being the second most basal.


Internal Classification

The first species of ''Kunmunella'' to be named was ''K. rajouriensis'', although the first remains of what is now known to be Kunmunella were found earlier and assigned to ''I. kalakotensis''. In 1985, Kumar and Sahni found that ''I. kalakotensis'' was a synonym of ''K. rajouriensis'' and that ''Kunmunella'' was a distinct species. Since older names have precedent in
taxonomy image:Hierarchical clustering diagram.png, 280px, Generalized scheme of taxonomy Taxonomy is a practice and science concerned with classification or categorization. Typically, there are two parts to it: the development of an underlying scheme o ...
, they changed the name of ''K. rajouriensis'' to ''K. kalakotensis.'' The same study found that there was another species of ''Kunmunela'', known only from a right maxilla: ''K. transversa''. Rana et al. (2021) reaffirmed that the two species of ''Kunmunella'' were most closely related to each other than to any other animal, and therefore formed a valid genus.


Description

''Kunmunella'' has more transverse upper
molars The molars or molar teeth are large, flat tooth, teeth at the back of the mouth. They are more developed in mammal, mammals. They are used primarily to comminution, grind food during mastication, chewing. The name ''molar'' derives from Latin, '' ...
, a labially (towards the lips) shifted hypercone (a large
cusp A cusp is the most pointed end of a curve. It often refers to cusp (anatomy), a pointed structure on a tooth. Cusp or CUSP may also refer to: Mathematics * Cusp (singularity), a singular point of a curve * Cusp catastrophe, a branch of bifu ...
in the molars), a triangular third molar, and more prominent lingual (tongue side) enamel ridges. The third molar has a large conical paracone (a type of cusp) connected to a smaller metacone by an enamel ridge. The protocone is large and in the forward part of the molars while the hypocone, connected to the metacone by a ridge, is small. Labial cingulum (enamel ridges) are larger than posterior cingulum. Upper molars have bunolophodont cusps, the paracone is the largest cusp, and the mesocone is absent, while the
premolar The premolars, also called premolar Tooth (human), teeth, or bicuspids, are transitional teeth located between the Canine tooth, canine and Molar (tooth), molar teeth. In humans, there are two premolars per dental terminology#Quadrant, quadrant in ...
s are non-molarized.


''K. kalakotensis''


Upper dentition

''Kunmunella kalakotensis'' has canines separated from
premolar The premolars, also called premolar Tooth (human), teeth, or bicuspids, are transitional teeth located between the Canine tooth, canine and Molar (tooth), molar teeth. In humans, there are two premolars per dental terminology#Quadrant, quadrant in ...
s by a small space. The first premolar is very similar to the canines; it is single cusped (only has one, large bump) and is sharp. The second premolar is larger than the first and elongated but lingo-lingually compressed. There are two large ridges which descend from the highpoint of the tooth. The third molar is the largest. It has a posterior ridge which splits into two near the top of the tooth. It has transverse upper molars with conical cusps and large lophs. Cingula are prominent and present throughout the entirety of the molar. ''K. kalakotensis'' also has an anterior ridge which is undivided. The lingual enamel ridges are larger than the labial ridges. The fourth premolar is similar to the previous teeth. The paracone is large and nearly twice the height of the protocone. The first molar is rectangular and wider than it is long. The paracone is the largest and tallest cusp. There are grooves of varying deepness between the four large cusps. The occlusal surface of the enamel is smooth, although the sides of the tooth are rugose. The cingulum are prominent throughout the entire tooth. The second molar is larger with more prominent cusps than the first molar. There is no metaconule on the second molar. The third molar is triangular. The paracone is high and conical, while the hypocone is reduced. Similar to the second molar, the cingulum are prominent.


Lower dentition

''Kunmunella kalakotensis'' has lower dentition known only by the last two molars. The third molar is labio-lingualy compressed, similar to the second premolar of the upper dentition. The metaconid is the largest cusp, while the entoconid is the smallest and the paraconid is absent. There are ridges connecting the cusps. Anterior and lingual cingula are well developed.


''K. transversa''

The fourth premolar is high, transverse, and almost the same size as the first molar. As opposed to ''K. kalakotensis'' which has elongated paracone, ''K. transversa'' has a conical paracone. The first molar is rectangular and wider than it is long. It has a larger hypocone than the first molar in ''K. kalakotensis''. There is no metacone and cingula are prominent. The second molar is transverse. The protocone is displaced towards the lingual side of the tooth. The third molar is rectangular (unlike the triangular molar in ''K. kalakotensis'') and has a paracone displaced lingually. The posterior cingululm of the third molar forms a shelf-like structure.


Paleobiology

Being a raoellid, ''Kunmunella'' likely had a similar lifestyle to ''Indohyus''. ''Kunmunella'' was likely an
omnivore An omnivore () is an animal that regularly consumes significant quantities of both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize t ...
because related genus ''Indohyus'' had teeth that allowed it to grasp and hold
prey Predation is a biological interaction in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not ki ...
, while also having molars adapted to processing plant matter. ''Kunmunella'' and other raoellids were almost certainly
semiaquatic In biology, being semi-aquatic refers to various macroorganisms that live regularly in both aquatic and terrestrial environments. When referring to animals, the term describes those that actively spend part of their daily time in water (in ...
. This is due to higher
bone density Bone density, or bone mineral density, is the amount of bone mineral in bone tissue. The concept is of mass of mineral per volume of bone (relating to density in the physics sense), although medicine#Clinical practice, clinically it is measured by ...
than fully terrestrial
mammal A mammal () is a vertebrate animal of the Class (biology), 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 ...
s and
auditory bulla The tympanic part of the temporal bone is a curved plate of bone lying below the squamous part of the temporal bone, in front of the mastoid process, and surrounding the external part of the ear canal. It originates as a separate bone (tympanic b ...
, both of which were found in Indohyus.


References

{{Taxonbar, from=Q134350811 Whippomorphs Prehistoric Artiodactyla genera Fossil taxa described in 1972