Reigomys
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''Reigomys primigenus'' is an
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
oryzomyine
rodent Rodents (from Latin , 'to gnaw') are mammals of the order Rodentia (), which are characterized by a single pair of continuously growing incisors in each of the upper and lower jaws. About 40% of all mammal species are rodents. They are na ...
known from
Pleistocene The Pleistocene ( , often referred to as the ''Ice age'') is the geological Epoch (geology), epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fina ...
deposits in
Tarija Department Tarija () is a department in Bolivia. It is located in south-eastern Bolivia bordering with Argentina to the south and Paraguay to the east. According to the 2012 census, it has a population of 482,196 inhabitants. It has an area of . The city ...
, southeastern
Bolivia , image_flag = Bandera de Bolivia (Estado).svg , flag_alt = Horizontal tricolor (red, yellow, and green from top to bottom) with the coat of arms of Bolivia in the center , flag_alt2 = 7 × 7 square p ...
. It is known from a number of isolated
jaw The jaw is any opposable articulated structure at the entrance of the mouth, typically used for grasping and manipulating food. The term ''jaws'' is also broadly applied to the whole of the structures constituting the vault of the mouth and serv ...
s and
molars The molars or molar teeth are large, flat teeth at the back of the mouth. They are more developed in mammals. They are used primarily to grind food during chewing. The name ''molar'' derives from Latin, ''molaris dens'', meaning "millstone to ...
which show that its molars were almost identical to those of the living ''
Lundomys ''Lundomys molitor'', also known as Lund's amphibious ratMusser and Carleton, 2005, p. 1124 or the greater marsh rat, is a semiaquatic rat species from southeastern South America. Its distribution is now restricted to Uruguay and nearby Ri ...
''. On the other hand, the animal possesses a number of
derived Derive may refer to: * Derive (computer algebra system), a commercial system made by Texas Instruments * ''Dérive'' (magazine), an Austrian science magazine on urbanism *Dérive, a psychogeographical concept See also * *Derivation (disambiguatio ...
traits of the
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 sepa ...
which document a closer relationship to living ''
Holochilus ''Holochilus'' is a genus of semiaquatic rodents in the tribe Oryzomyini of family Cricetidae,* sometimes called marsh rats. It contains five living species, '' H. brasiliensis'', '' H. chacarius'', '' H. nanus'', '' H. oxe'', and '' H. sciureus' ...
'', the genus of South American marsh rats, and for this reason it was placed in the genus ''Holochilus'' when it was first described in 1996. The subsequent discoveries of ''
Noronhomys ''Noronhomys vespuccii'', also known as Vespucci's rodent, is an extinct rat species from the islands of Fernando de Noronha off northeastern Brazil. Italian explorer Amerigo Vespucci may have seen it on a visit to Fernando de Noronha in 1503, ...
'' and ''
Carletonomys ''Carletonomys cailoi'' is an extinction, extinct rodent from the Pleistocene (Ensenadan) of Buenos Aires Province, Argentina. Although known only from a single maxilla (upper jaw) with the first molar (tooth), molar, its features are so distinct ...
'', which may be more closely related to extant ''Holochilus'' than ''H. primigenus'' is, have cast its placement in ''Holochilus'' into doubt, and it was ultimately made the type species of a separate genus, ''Reigomys''.


Taxonomy

Material of ''Reigomys primigenus'' was collected in 1924 and 1927 by Elmer Riggs of the
Field Museum of Natural History The Field Museum of Natural History (FMNH), also known as The Field Museum, is a natural history museum in Chicago, Illinois, and is one of the largest such museums in the world. The museum is popular for the size and quality of its educational ...
, but the animal was not described until 1996, when Scott Steppan formally named it as a new species of the genus ''Holochilus'' and diagnosed it, contrasting it to related species.Steppan, 1996, p. 523 The material Riggs collected includes nine
mandible In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
s, three
maxilla The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The t ...
e, and five isolated molars. The specific name Steppan gave to the animal, ''primigenus'', means "primitive" in Latin and refers to the primitive features of the animal when compared to its relatives ''Holochilus'' and ''Lundomys''.Steppan, 1996, p. 524 In order to determine the relationships of his new species, Steppan carried out a
cladistic Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups (" clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived char ...
analysis, in which he also included the oryzomyines ''
Holochilus ''Holochilus'' is a genus of semiaquatic rodents in the tribe Oryzomyini of family Cricetidae,* sometimes called marsh rats. It contains five living species, '' H. brasiliensis'', '' H. chacarius'', '' H. nanus'', '' H. oxe'', and '' H. sciureus' ...
'', ''
Lundomys ''Lundomys molitor'', also known as Lund's amphibious ratMusser and Carleton, 2005, p. 1124 or the greater marsh rat, is a semiaquatic rat species from southeastern South America. Its distribution is now restricted to Uruguay and nearby Ri ...
'', ''
Pseudoryzomys ''Pseudoryzomys simplex'', also known as the Brazilian false rice rat or false oryzomys, is a species of rodent in the family Cricetidae from south-central South America. It is found in lowland palm savanna and thorn scrub habitats. It is a me ...
'', and '' Cerradomys'', as well as the non-oryzomyine '' Sigmodon''. His results supported a close relation between ''R. primigenus'' and extant ''Holochilus'', with ''Lundomys'' and ''Pseudoryzomys'' more distantly related. In 1999, another rodent from the same group was described: ''
Noronhomys ''Noronhomys vespuccii'', also known as Vespucci's rodent, is an extinct rat species from the islands of Fernando de Noronha off northeastern Brazil. Italian explorer Amerigo Vespucci may have seen it on a visit to Fernando de Noronha in 1503, ...
'', a recently extinct species from the Brazilian island of
Fernando de Noronha Fernando de Noronha () is an archipelago in the Atlantic Ocean, part of the State of Pernambuco, Brazil, and located off the Brazilian coast. It consists of 21 islands and islets, extending over an area of . Only the eponymous main island is inha ...
. Michael Carleton and
Storrs Olson Storrs Lovejoy Olson (April 3, 1944 – January 20, 2021) was an American biologist and ornithologist who spent his career at the Smithsonian Institution, retiring in 2008. One of the world's foremost avian paleontologists, he was best known ...
, who described the animal, argued that ''Holochilus primigenus'' was probably outside 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, ...
of ''Noronhomys'' and extant ''Holochilus'' and that ''H. primigenus'' should probably be excluded from the genus. When Ulyses Pardiñas described another extinct rodent from this group, ''
Carletonomys ''Carletonomys cailoi'' is an extinction, extinct rodent from the Pleistocene (Ensenadan) of Buenos Aires Province, Argentina. Although known only from a single maxilla (upper jaw) with the first molar (tooth), molar, its features are so distinct ...
'' from the Pleistocene of Argentina, he suggested that ''H. primigenus'' should be placed in its own genus because of its mosaic of ''Holochilus''- and ''Lundomys''-like features. The phylogenetic analysis conducted by Machado ''et al.'' (2014) confirmed that living members of the genus ''Holochilus'' are more closely related to ''Noronhomys'' and ''Carletonomys'' than to ''H. primigenus''. The authors moved ''H. primigenus'' to its own genus, which they named ''Reigomys''.


Description

''Reigomys primigenus'' was a large rat, though smaller than both ''Lundomys'' and living ''Holochilus'', characterized by an S-shaped ("sigmodont") crown on the third lower molar. Features of the molars are nearly indistinguishable from those of ''Lundomys'' and include cusps located opposite each other, enamel valleys that barely reach the midline of the molars, and short mesolophs and mesolophids (accessory crests). Other notable features include a robust mandible (lower jaw) with a steeply rising coronoid process, short
incisive foramina In the human mouth, the incisive foramen (also known as: "''anterior palatine foramen''", or "''nasopalatine foramen''") is the opening of the incisive canals on the hard palate immediately behind the incisor teeth. It gives passage to blood vesse ...
(perforations of the front part of the
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 sepa ...
) that barely extend between the first molars, and a short bony palate that hardly extends beyonds the third molars, all of which are shared with extant species of ''
Holochilus ''Holochilus'' is a genus of semiaquatic rodents in the tribe Oryzomyini of family Cricetidae,* sometimes called marsh rats. It contains five living species, '' H. brasiliensis'', '' H. chacarius'', '' H. nanus'', '' H. oxe'', and '' H. sciureus' ...
'' to the exclusion of ''Lundomys''. Also unlike in ''Lundomys'', there is an additional small root present at the labial (outer) side of the first upper molar. In eight specimens that could be measured, the length of the lower toothrow is 6.79 to 7.58 mm, averaging 7.28 mm; the first lower molar is 2.62 to 3.08 mm long, averaging 2.89 mm, and 1.75 to 1.93 mm broad, averaging 1.85 mm in seven measured teeth; the only preserved complete upper toothrow is 6.64 mm long; and the first upper molar is 2.63 to 2.70 mm long and 2.03 mm broad in two specimens.


Distribution and ecology

Remains of ''Reigomys primigenus'' come from several localities in river sediments of the
Tarija Formation Tarija or San Bernardo de la Frontera de Tarixa is a city in southern Bolivia. Founded in 1574, Tarija is the largest city and capital and municipality within the Tarija Department, with an airport (Capitán Oriel Lea Plaza Airport, (TJA)) offe ...
in
Tarija Department Tarija () is a department in Bolivia. It is located in south-eastern Bolivia bordering with Argentina to the south and Paraguay to the east. According to the 2012 census, it has a population of 482,196 inhabitants. It has an area of . The city ...
, which have been paleomagnetically dated to about 0.7 to 1.0 million years ago ( chrons Clr.ln to early Cln,
Ensenadan The Ensenadan age is a period of geologic time (1.2–0.8 Ma) within the Early Pleistocene epoch of the Quaternary The Quaternary ( ) is the current and most recent of the three periods of the Cenozoic Era in the geologic time scale of the ...
South American land mammal age The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Mya (unit), Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referre ...
). Other sigmodontine rodents found there include '' Andinomys'', '' Calomys'', ''
Kunsia The woolly giant rat (''Kunsia tomentosus'') is a species of large burrowing rodent native to South America. No subspecies are currently recognised. It is the only member of the genus ''Kunsia''. Description The woolly giant rat is an exceptio ...
'', ''
Nectomys ''Nectomys'' is a genus of rodent in the tribe Oryzomyini of family Cricetidae. Musser and Carleton, 2005. It is closely related to ''Amphinectomys'' and was formerly considered congeneric with ''Sigmodontomys''. It consists of five species, which ...
'', ''
Oxymycterus ''Oxymycterus'' is a genus of rat-like rodents commonly known as hocicudos. They are endemic to South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively sma ...
'', '' Phyllotis'', and another akodontine, probably ''
Akodon ''Akodon'' is a genus consisting of South American grass mice. They mostly occur south of the Amazon Basin and along the Andes north to Venezuela, but are absent from much of the basin itself, the far south of the continent, and the lowlands west ...
'', ''
Necromys ''Necromys'' is a genus of South American sigmodontine rodents allied to ''Akodon''. This genus has also been known as ''Cabreramys'' or more recently ''Bolomys'', and the northern grass mouse (''N. urichi'') has recently been transferred from ' ...
'', or a related genus. The deposits were deposited by a river and the paleoenvironment was probably a floodplain or channel. ''Reigomys primigenus'' is not known from any other localities and is considered to be extinct.Steppan, 1996, p. 527


References


Bibliography

* Carleton, M.D. and Olson, S.L. 1999
Amerigo Vespucci and the rat of Fernando de Noronha: a new genus and species of Rodentia (Muridae, Sigmodontinae) from a volcanic island off Brazil's continental shelf
''American Museum Novitates'' 3256:1–59. * Pardiñas, U.F.J. 2008
A new genus of oryzomyine rodent (Cricetidae: Sigmodontinae) from the Pleistocene of Argentina
(subscription required). ''Journal of Mammalogy'' 89(5):1270–1278. * Steppan, S.J. 1996
A new species of ''Holochilus'' (Rodentia: Sigmodontinae) from the Middle Pleistocene of Bolivia and its phylogenetic significance
(subscription required). ''Journal of Vertebrate Paleontology'' 16(3):522–530. * {{Good article Oryzomyini Prehistoric rodent genera Pleistocene rodents Pleistocene mammals of South America Ensenadan Pleistocene Bolivia Fossils of Bolivia Fossil taxa described in 1996