Holochilus Primigenus
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''Reigomys primigenus'' is an
extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
oryzomyine
rodent Rodents (from Latin , 'to gnaw') are mammals of the Order (biology), order Rodentia ( ), which are characterized by a single pair of continuously growing incisors in each of the upper and Mandible, lower jaws. About 40% of all mammal specie ...
known from
Pleistocene The Pleistocene ( ; referred to colloquially as the ''ice age, Ice Age'') is the geological epoch (geology), epoch that lasted from to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fin ...
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 2024 census, it has a population of 534,348 inhabitants. It has an area of . The city ...
, southeastern
Bolivia Bolivia, officially the Plurinational State of Bolivia, is a landlocked country located in central South America. The country features diverse geography, including vast Amazonian plains, tropical lowlands, mountains, the Gran Chaco Province, w ...
. It is known from a number of isolated
jaw The jaws are a pair of opposable articulated structures 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 ...
s and
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, '' ...
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 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 sep ...
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'' and '' Carletonomys'', 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 educationa ...
, 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 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 ...
s, three
maxilla In vertebrates, the maxilla (: maxillae ) 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 two maxil ...
e, and five isolated molars. The
specific name Specific name may refer to: * in Database management systems, a system-assigned name that is unique within a particular database In taxonomy, either of these two meanings, each with its own set of rules: * Specific name (botany), the two-part (bino ...
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 ( ; from Ancient Greek 'branch') 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 ...
analysis, in which he also included the
oryzomyines Oryzomyini is a tribe (taxonomy), tribe of rodents in the subfamily Sigmodontinae of the family Cricetidae. It includes about 120 species in about thirty genera,Weksler et al., 2006, table 1 distributed from the eastern List of mammals of the Uni ...
''
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 List of mammals of South America, South America. It is found in lowland Tropical and sub ...
'', 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'', a recently extinct species from the Brazilian island of
Fernando de Noronha Fernando de Noronha (), officially the State District of Fernando de Noronha () and formerly known as the Federal Territory of Fernando de Noronha () until 1988, is an archipelago in the Atlantic Ocean, part of the state of Pernambuco, Brazil, and ...
. 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 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 ...
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'' 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 (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 sep ...
) 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 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 2024 census, it has a population of 534,348 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
South American land mammal age The South American land mammal ages (SALMA) establish a geologic timescale for prehistoric South American fauna beginning 64.5 Ma during the Paleocene and continuing through to the Late Pleistocene (0.011 Ma). These periods are referred to as age ...
). Other sigmodontine rodents found there include '' Andinomys'', ''
Calomys Vesper mice are rodents belonging to the genus ''Calomys''. They are widely distributed in South America. Some species are notable as the Vector (epidemiology), vectors of Argentinian hemorrhagic fever and Bolivian hemorrhagic fever. The genus w ...
'', '' Kunsia'', '' Nectomys'', '' Oxymycterus'', ''
Phyllotis ''Phyllotis'' is a genus of rodent in the family Cricetidae. These mice are commonly confused with ''Auliscomys, Graomys'' and ''Loxodontomys.'' ''.'' In order to tell these genera apart, one must look at the tail. Species in the genus ''Phyllo ...
'', 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'', 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 Monotypic prehistoric rodent genera Pleistocene rodents Pleistocene mammals of South America Ensenadan Pleistocene Bolivia Fossils of Bolivia Fossil taxa described in 1996