''Carletonomys cailoi'' is an
extinct 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 lower jaws. About 40% of all mammal species are roden ...
from the
Pleistocene
The Pleistocene ( , often referred to as the '' Ice age'') is the geological 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 finally confirmed ...
(
Ensenadan
The Ensenadan age is a period of geologic time (1.2–0.8 Ma) within the Early Pleistocene epoch of the Quaternary used more specifically with South American Land Mammal Ages. It follows the Uquian and precedes the Lujanian
The Lujanian age is a ...
) of
Buenos Aires Province,
Argentina
Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the second-largest country in South America after Brazil, t ...
. Although known only from a single
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 ...
(upper jaw) with the first
molar, its features are so distinctive that it is placed in its own
genus
Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial n ...
, ''Carletonomys''. Discovered in 1998 and formally described in 2008, it is part of a well-defined group of
oryzomyine
Oryzomyini is a 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 United States to the southernmost parts of S ...
rodents that also includes ''
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 ...
'', ''
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, ...
'', ''
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 ...
'', and ''
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 ...
''. This group is characterized by progressive
semiaquatic
In biology, semiaquatic can refer to various types of animals that spend part of their time in water, or plants that naturally grow partially submerged in water. Examples are given below.
Semiaquatic animals
Semi aquatic animals include:
* Ve ...
specializations and a reduction in the complexity of molar
morphology
Morphology, from the Greek and meaning "study of shape", may refer to:
Disciplines
*Morphology (archaeology), study of the shapes or forms of artifacts
*Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies, ...
.
The single known molar is high-crowned (
hypsodont
Hypsodont is a pattern of dentition with high-crowned teeth and enamel extending past the gum line, providing extra material for wear and tear. Some examples of animals with hypsodont dentition are cows and horses; all animals that feed on grit ...
) and flat-crowned (planar) and is distinctive in lacking the ridge that connects the front to the middle part of the molar, the
anterior mure
Many different terms have been proposed for features of the tooth crown in mammals.
The structures within the molars receive different names according to their position and morphology. This nomenclature was developed by Henry Fairfield Osborn ...
, and in the configuration of another ridge, the
mesoloph
Many different terms have been proposed for features of the tooth crown in mammals.
The structures within the molars receive different names according to their position and morphology. This nomenclature was developed by Henry Fairfield Osborn ...
. ''Carletonomys'' was probably
herbivorous
A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...
and lived in a wet habitat.
Taxonomy
''Carletonomys cailoi'' was discovered in 1998 in a
silt
Silt is granular material of a size between sand and clay and composed mostly of broken grains of quartz. Silt may occur as a soil (often mixed with sand or clay) or as sediment mixed in suspension with water. Silt usually has a floury feel wh ...
deposit in
San Cayetano Partido, southeastern
Buenos Aires Province. The
stratigraphic
Stratigraphy is a branch of geology concerned with the study of rock layers ( strata) and layering (stratification). It is primarily used in the study of sedimentary and layered volcanic rocks.
Stratigraphy has three related subfields: lithostr ...
context suggests this locality is slightly over 1 million years old (
Ensenadan
The Ensenadan age is a period of geologic time (1.2–0.8 Ma) within the Early Pleistocene epoch of the Quaternary used more specifically with South American Land Mammal Ages. It follows the Uquian and precedes the Lujanian
The Lujanian age is a ...
South American Land Mammal Age), making ''Carletonomys'' the oldest known oryzomyine.
[Pardiñas, 2008, p. 1276] The single known specimen is now in the collections of the
Museo de La Plata
The La Plata Museum ( es, Museo de la Plata) is a natural history museum in La Plata, Argentina. It is part of the (Natural Sciences School) of the UNLP (National University of La Plata).
The building, long, today houses 3 million fossils and ...
. It was initially referred to the genus ''
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, ...
'', which is currently known only from the 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 i ...
off northeastern
Brazil
Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...
, but in 2008 Argentinean mammalogist Ulyses Pardiñas established it as the
holotype
A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of seve ...
of a new genus and species of rodent in a publication in the ''
Journal of Mammalogy
The ''Journal of Mammalogy'' is a bimonthly peer-reviewed scientific journal published by Oxford University Press on behalf of the American Society of Mammalogists. Both the society and the journal were established in 1919. The journal covers rese ...
''. The
generic name, ''Carletonomys'', combines the name of American mammalogist Michael Carleton with the
Ancient Greek
Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Dark Ages (), the Archaic p ...
μυς ''mys'' "mouse"
[Pardiñas, 2008, p. 1271] and the
specific name, ''cailoi'', honors Argentinean biologist Carlos "Cailo" Galliari.
[Pardiñas, 2008, p. 1272]
The fossil has a number of features that suggest a relation to a group of
oryzomyine
Oryzomyini is a 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 United States to the southernmost parts of S ...
rodents that includes the South American marsh rat ''
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 ...
'', its living relatives ''
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 ...
'' and ''
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 ...
'', and the extinct ''
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 ''
Holochilus primigenus
''Reigomys primigenus'' is an extinct oryzomyine rodent known from Pleistocene deposits in Tarija Department, southeastern Bolivia. It is known from a number of isolated jaws and molars which show that its molars were almost identical to those ...
''.
[Pardiñas, 2008, p. 1275] They share high-crowned (
hypsodont
Hypsodont is a pattern of dentition with high-crowned teeth and enamel extending past the gum line, providing extra material for wear and tear. Some examples of animals with hypsodont dentition are cows and horses; all animals that feed on grit ...
) molars and several simplifications of molar morphology, as well as other features that cannot be assessed in ''Carletonomys'', which indicate specializations towards a
semiaquatic
In biology, semiaquatic can refer to various types of animals that spend part of their time in water, or plants that naturally grow partially submerged in water. Examples are given below.
Semiaquatic animals
Semi aquatic animals include:
* Ve ...
lifestyle. It shows the most similarity to ''Noronhomys'' and ''Holochilus'', so much so that Pardiñas considered placing it in either of these two genera, but its distinctive morphological features justify placement in a separate genus.
This group of genera encompasses only a small part of the diversity of the
tribe
The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide usage of the term in English is in the discipline of anthropology. This definition is contested, in part due to confl ...
Oryzomyini
Oryzomyini is a 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 United States to the southernmost parts of ...
, a group of over a hundred species distributed mainly in South America, including nearby islands such as the
Galápagos Islands
The Galápagos Islands ( es, Islas Galápagos) are an archipelago of volcanic islands in the Eastern Pacific, located around the Equator west of the mainland of South America. They form the Galápagos Province of the Republic of Ecuador, with ...
and some of the
Antilles
The Antilles (; gcf, label=Antillean Creole, Antiy; es, Antillas; french: Antilles; nl, Antillen; ht, Antiy; pap, Antias; Jamaican Patois: ''Antiliiz'') is an archipelago bordered by the Caribbean Sea to the south and west, the Gulf of Mex ...
. Oryzomyini is one of several tribes recognized within the
subfamily Sigmodontinae
The rodent subfamily Sigmodontinae includes New World rats and mice, with at least 376 species. Many authorities include the Neotominae and Tylomyinae as part of a larger definition of Sigmodontinae. When those genera are included, the speci ...
, which encompasses hundreds of species found across South America and into southern North America. Sigmodontinae itself is the largest subfamily of the family
Cricetidae
The Cricetidae are a family of rodents in the large and complex superfamily Muroidea. It includes true hamsters, voles, lemmings, muskrats, and New World rats and mice. At almost 608 species, it is the second-largest family of mammals, a ...
, other members of which include
vole
Voles are small rodents that are relatives of lemmings and hamsters, but with a stouter body; a longer, hairy tail; a slightly rounder head; smaller eyes and ears; and differently formed molars (high-crowned with angular cusps instead of lo ...
s,
lemming
A lemming is a small rodent, usually found in or near the Arctic in tundra biomes. Lemmings form the subfamily Arvicolinae (also known as Microtinae) together with voles and muskrats, which form part of the superfamily Muroidea, which also incl ...
s,
hamster
Hamsters are rodents (order Rodentia) belonging to the subfamily Cricetinae, which contains 19 species classified in seven genera.Fox, Sue. 2006. ''Hamsters''. T.F.H. Publications Inc. They have become established as popular small pets. The ...
s, and
deermice
''Peromyscus'' is a genus of rodents. They are commonly referred to as deer mice or deermice, not to be confused with the chevrotain or "mouse deer". They are New World mice only distantly related to the common house and laboratory mouse, ''Mu ...
, all mainly from Eurasia and North America.
Description

The holotype is a right
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 ...
(upper jaw) with the upper first
molar (M1) in it. It is broken off behind the M1, but much of the front part is preserved, including the
zygomatic plate, the flattened front portion of the
zygomatic arch
In anatomy, the zygomatic arch, or cheek bone, is a part of the skull formed by the zygomatic process of the temporal bone (a bone extending forward from the side of the skull, over the opening of the ear) and the temporal process of the zygom ...
(cheekbone). The M1 is moderately worn, indicating that it is from an adult individual.
[ With an M1 length of 3.59 mm and width of 2.53 mm, ''C. cailoi'' was one of the largest oryzomyines known, rivaled only by '']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 ...
'' and the extinct Antillean ''Megalomys
''Megalomys'' is a genus of rodent in the family Cricetidae, part of the tribe Oryzomyini. The genus contains five large rodents from various Caribbean islands, of which two are known to have survived into modern times, but all of which are now e ...
'' and "''Ekbletomys
"''Ekbletomys hypenemus''" is an extinct oryzomyine rodent from the islands of Antigua and Barbuda, Lesser Antilles. It was described as the only species of the subgenus "''Ekbletomys''" of genus ''Oryzomys'' in a 1962 Ph.D. thesis, but that na ...
''". The height of the M1 is 1.37 mm and it has four roots, including a large one in front, another large one on the inner (lingual) side, and two smaller ones on the outer (labial) side. The presence of a second labial root is a variable character among oryzomyines, occurring among others in ''Holochilus'' and ''Pseudoryzomys'' but not in ''Lundomys''. The maxilla itself shows few significant characters. The back margin of the incisive foramen
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 ves ...
, which perforates 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 s ...
between the upper incisor
Incisors (from Latin ''incidere'', "to cut") are the front teeth present in most mammals. They are located in the premaxilla above and on the mandible below. Humans have a total of eight (two on each side, top and bottom). Opossums have 18, w ...
s and the molars, is not visible, suggesting that the foramen was short, as in ''Holochilus''. The configuration of the zygomatic plate shows features that distinguish ''C. cailoi'' from some of its relatives.[Pardiñas, 2008, p. 1273]
The molar is plane and hypsodont: the crowns are relatively high and the main cusps are about as high as the other parts of the crown,[ as they are in ''Holochilus''. Most other oryzomyines have bunodont and brachydont molars, in which the crowns are lower and the cusps are higher than the rest of the crown. As in closely related species, the front part of the molar is relatively simple, lacking an ]anteroloph
Many different terms have been proposed for features of the tooth crown in mammals.
The structures within the molars receive different names according to their position and morphology. This nomenclature was developed by Henry Fairfield Osborn i ...
, an additional ridge that is well-developed in most oryzomyines. A shallow anteromedian flexus
Many different terms have been proposed for features of the tooth crown in mammals.
The structures within the molars receive different names according to their position and morphology. This nomenclature was developed by Henry Fairfield Osborn i ...
is present, superficially dividing the front cusp (anterocone
Many different terms have been proposed for features of the tooth crown in mammals.
The structures within the molars receive different names according to their position and morphology. This nomenclature was developed by Henry Fairfield Osborn i ...
).[ Uniquely, the ]anterior mure
Many different terms have been proposed for features of the tooth crown in mammals.
The structures within the molars receive different names according to their position and morphology. This nomenclature was developed by Henry Fairfield Osborn ...
, which connects the anterocone to the rest of the crown, is absent; although this structure is sometimes missing in young individuals of other oryzomyines, it usually develops as a result of wear in adults. The two cusps on the middle part of the molar, the paracone
A paracone is a 1960s atmospheric reentry or spaceflight mission abort concept using an inflatable ballistic cone.[protocone
A cusp is a pointed, projecting, or elevated feature. In animals, it is usually used to refer to raised points on the crowns of teeth.
The concept is also used with regard to the leaflets of the four heart valves. The mitral valve, which has two ...]
, are broadly connected. The median mure
Many different terms have been proposed for features of the tooth crown in mammals.
The structures within the molars receive different names according to their position and morphology. This nomenclature was developed by Henry Fairfield Osborn ...
, which connects the middle to the back pair of cusps, is attached to the back of the paracone. A complete mesoloph
Many different terms have been proposed for features of the tooth crown in mammals.
The structures within the molars receive different names according to their position and morphology. This nomenclature was developed by Henry Fairfield Osborn ...
is present, descending from the median mure slightly behind the paracone.[ The configuration of the paracone–median mure–mesoloph complex is unique to ''Carletonomys''.][ The two posterior cusps, the ]hypocone
A cusp is a pointed, projecting, or elevated feature. In animals, it is usually used to refer to raised points on the crowns of teeth.
The concept is also used with regard to the leaflets of the four heart valves. The mitral valve, which has two ...
and the metacone, are connected at the back margin of the molar. Unlike in most oryzomyines, no posteroflexus
Many different terms have been proposed for features of the tooth crown in mammals.
The structures within the molars receive different names according to their position and morphology. This nomenclature was developed by Henry Fairfield Osborn ...
is present, so that the metacone is situated directly at the back margin.[Pardiñas, 2008, fig. 1; p. 1273]
Ecology
''Carletonomys'' was found in association with remains of several other animals, including fishes, chelid
Chelidae is one of three living families of the turtle suborder Pleurodira, and are commonly called Austro-South American side-neck turtles. The family is distributed in Australia, New Guinea, parts of Indonesia, and throughout most of South Amer ...
turtles, frogs, birds, armadillos, and several rodents, including ''Reithrodon auritus
The bunny rat, or hairy-soled conyrat (''Reithrodon auritus'') is a species of rodent in the family Cricetidae, native to southern South America.
Description
The bunny rat is a heavily built rat-like rodent, with a total length of , including the ...
'', the coypu
The nutria (''Myocastor coypus''), also known as the coypu, is a large, herbivorous, semiaquatic rodent.
Classified for a long time as the only member of the family Myocastoridae, ''Myocastor'' is now included within Echimyidae, the family of ...
(''Myocastor''), both of which still live in the area, the extinct echimyid
Echimyidae is the family of neotropical spiny rats and their fossil relatives. This is the most species-rich family of hystricognath rodents. It is probably also the most ecologically diverse, with members ranging from fully arboreal to terres ...
'' Dicolpomys'', and unidentified caviid
Caviidae, the cavy family, is composed of rodents native to South America and includes the domestic guinea pig, wild cavies, and the largest living rodent, the capybara. They are found across South America in open areas from moist savanna t ...
s and octodontid
Octodontidae is a family of rodents, restricted to southwestern South America. Fourteen species of octodontid are recognised, arranged in seven genera. The best known species is the common degu, ''Octodon degus''.
Octodontids are medium-sized ...
s. ''C. cailoi'' probably lived in a wetland
A wetland is a distinct ecosystem that is flooded or saturated by water, either permanently (for years or decades) or seasonally (for weeks or months). Flooding results in oxygen-free (anoxic) processes prevailing, especially in the soils. The ...
habitat under relatively warm and moist climatic conditions. Although the limited material known permits few inferences as to the animal's natural history, it likely fed on hard plant material, as do related, morphologically similar extant species.
References
Bibliography
* Musser, G.G. and Carleton, M.D. 2005. Superfamily Muroidea. Pp. 894–1531 in Wilson, D.E. and Reeder, D.M. (eds.)
Mammal Species of the World: a taxonomic and geographic reference. 3rd ed
Baltimore: The Johns Hopkins University Press,
* 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.
* Ray, C.E. 1962. The Oryzomyine Rodents of the Antillean Subregion. Doctor of Philosophy thesis, Harvard University, 211 pp.
* Voss, R.S. and Carleton, M.D. 1993
A new genus for ''Hesperomys molitor'' Winge and ''Holochilus magnus'' Hershkovitz (Mammalia, Muridae) with an analysis of its phylogenetic relationships
''American Museum Novitates'' 3085:1–39.
* Weksler, M. 2006
Phylogenetic relationships of oryzomyine rodents (Muroidea: Sigmodontinae): separate and combined analyses of morphological and molecular data
''Bulletin of the American Museum of Natural History'' 296:1–149.
{{Good article
Oryzomyini
Prehistoric rodent genera
Pleistocene rodents
Pleistocene genus extinctions
Pleistocene mammals of South America
Ensenadan
Pleistocene Argentina
Fossils of Argentina
Fossil taxa described in 2008