Oryzomys Nitida
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''Oryzomys'' is a genus of semiaquatic rodents in the tribe Oryzomyini living in southern North America and far northern South America. It includes eight species, two of which—the
marsh rice rat The marsh rice rat (''Oryzomys palustris'') is a semiaquatic North American rodent in the family Cricetidae. It usually occurs in wetland habitats, such as swamps and salt marshes. It is found mostly in the eastern and southern United States, fr ...
(''O. palustris'') of the United States and '' O. couesi'' of Mexico and Central America—are widespread; the six others have more restricted distributions. The species have had eventful
taxonomic Taxonomy is the practice and science of categorization or classification. A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...
histories, and most species were at one time included in the marsh rice rat; additional species may be recognized in the future. The name ''Oryzomys'' was established in 1857 by
Spencer Fullerton Baird Spencer Fullerton Baird (; February 3, 1823 – August 19, 1887) was an American naturalist, ornithologist, ichthyologist, Herpetology, herpetologist, and museum curator. Baird was the first curator to be named at the Smithsonian Institution. He ...
for the marsh rice rat and was soon applied to over a hundred species of American rodents. Subsequently, the genus gradually became more narrowly defined until its current contents were established in 2006, when ten new genera were established for species previously placed in ''Oryzomys''. Species of ''Oryzomys'' are medium-sized rats with long, coarse fur. The upperparts are gray to reddish and the underparts white to buff. The animals have broad feet with reduced or absent
ungual tufts In mammals, ungual tufts are tufts of hairs at the base of claws of the forefeet and hindfeet. Their presence has been used as a character in cladistic studies of the Cricetidae, a large family of rodents.Weksler, 2006, p. 19 Rice rats Membe ...
of hair around the claws and, in at least some species, with webbing between the toes. The rostrum (front part of the skull) is broad and the
braincase In human anatomy, the neurocranium, also known as the braincase, brainpan, or brain-pan is the upper and back part of the skull, which forms a protective case around the brain. In the human skull, the neurocranium includes the calvaria or skul ...
is high. Both the marsh rice rat and ''O. couesi'' have 56  chromosomes, lack a gall bladder, and have a complex penis (as is characteristic of the Sigmodontinae) with some traits that are rare among oryzomyines; these characteristics are unknown in the other species of this genus. The habitat includes various kinds of wetlands, such as lakes, marshes, and rivers. ''Oryzomys'' species swim well, are active during the night, and eat both plant and animal food. They build woven nests of vegetation. After a
gestation period In mammals, pregnancy is the period of reproduction during which a female carries one or more live offspring from implantation in the uterus through gestation. It begins when a fertilized zygote implants in the female's uterus, and ends once it ...
of 21 to 28 days, about four young are born. Species of ''Oryzomys'' are infected by numerous parasites and carry at least three hantaviruses, one of which (
Bayou virus ''Bayou orthohantavirus'' (BAYV) is a species of ''Orthohantavirus'' first identified in 1993 in Louisiana. and later confirmed by other investigators. In 1996, the marsh rice rat was identified as the natural reservoir of the virus, indicating t ...
) also infects humans. Two, maybe three, species have gone extinct over the last two centuries and at least one other is endangered, but the widespread marsh rice rat and ''O. couesi'' are not threatened.


Taxonomy

''Oryzomys'' is one of about thirty genera within the tribe Oryzomyini, a diverse group of well over a hundred species, many of which were formerly also included in ''Oryzomys''. Oryzomyini is one of several tribes within the subfamily Sigmodontinae of the family Cricetidae, which includes hundreds of other species of mainly small rodents, distributed mainly in the Americas and Eurasia. Within Oryzomyini, a 2006 phylogenetic analysis by Marcelo Weksler which used both morphological and
DNA sequence DNA sequencing is the process of determining the nucleic acid sequence – the order of nucleotides in DNA. It includes any method or technology that is used to determine the order of the four bases: adenine, guanine, cytosine, and thymine. Th ...
data found some evidence that ''Oryzomys'' is most closely related to a group including '' Holochilus'', ''
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 ...
'', 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 ...
''. Although analyses based on morphological and combined data supported this relationship, sequences of the ''Rbp3'' gene alone instead placed ''Oryzomys'' among a group that included '' Nectomys'', ''
Sigmodontomys ''Sigmodontomys'' is a genus of rodent in the tribe Oryzomyini of family Cricetidae. It is related to ''Nectomys'' and ''Melanomys'' and used to be included in ''Nectomys''. It includes two species, ''Sigmodontomys alfari'' and the much rarer ''S ...
'', and a few other genera. In all analyses, ''Oryzomys'' appeared within clade D of Oryzomyini. The relationship between ''Oryzomys'' and the ''Holochilus'' group was supported by five
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 have ...
(shared derived characters)—absence or reduction of both the hypothenar and interdigital
pads Pads (also called leg guards) are a type of protective equipment used in a number of sports and serve to protect the legs from the impact of a hard ball, puck, or other object of play travelling at high speed which could otherwise cause injuries t ...
; reduction of
ungual tuft In mammals, ungual tufts are tufts of hairs at the base of claws of the forefeet and hindfeet. Their presence has been used as a character in cladistic studies of the Cricetidae, a large family of rodents.Weksler, 2006, p. 19 Rice rats Membe ...
s of hairs surrounding the claws; having the back margin of the zygomatic plate of the skull at the same level as the front of the first upper molar; and the
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 ...
(front cusp) of the first upper molar divided by an
anteromedian fossette 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 ...
. The first three are adaptations to the semiaquatic lifestyle that ''Oryzomys'' and the members of the ''Holochilus'' group share, and may thus be examples of convergent evolution.


Circumscription

The name ''Oryzomys'' was introduced in 1857 by
Spencer Fullerton Baird Spencer Fullerton Baird (; February 3, 1823 – August 19, 1887) was an American naturalist, ornithologist, ichthyologist, Herpetology, herpetologist, and museum curator. Baird was the first curator to be named at the Smithsonian Institution. He ...
for the
marsh rice rat The marsh rice rat (''Oryzomys palustris'') is a semiaquatic North American rodent in the family Cricetidae. It usually occurs in wetland habitats, such as swamps and salt marshes. It is found mostly in the eastern and southern United States, fr ...
(now ''Oryzomys palustris'') of the eastern United States,Baird, 1857, p. 482 which had been first described twenty years earlier by Richard Harlan.Musser and Carleton, 2005, p. 1144 The name combines the Greek ''oryza'' "rice" and ''mys'' "mouse" and refers to the feeding habits of the marsh rice rat.Schwartz and Schwartz, 2001, p. 192 Baird placed ''Oryzomys'' as a
subgenus In biology, a subgenus (plural: subgenera) is a taxonomic rank directly below genus. In the International Code of Zoological Nomenclature, a subgeneric name can be used independently or included in a species name, in parentheses, placed between t ...
of the now-defunct genus ''
Hesperomys Vesper mice are rodents belonging to a genus ''Calomys''. They are widely distributed in South America. Some species are notable as the vectors of Argentinian hemorrhagic fever and Bolivian hemorrhagic fever. The genus was originally named ''H ...
'' and included only the marsh rice rat in it, a classification which was followed by Elliott Coues in 1874 and 1877. In 1890, ''Oryzomys'' was raised to generic rank, and in subsequent years numerous additional species were ascribed to it, many of which were soon moved to separate genera. In the 1898 ''Catalogus Mammalium'', Édouard Louis Trouessart listed 67 species of ''Oryzomys'', including some that are now placed in ''
Calomys Vesper mice are rodents belonging to a genus ''Calomys''. They are widely distributed in South America. Some species are notable as the vectors of Argentinian hemorrhagic fever and Bolivian hemorrhagic fever. The genus was originally named ''He ...
'', '' Necromys'', ''
Thomasomys ''Thomasomys'' is a genus of rodent in the family Cricetidae, named after British zoologist Oldfield Thomas. Nuclear DNA sequence analysis has indicated that it is a sister taxon to ''Rhagomys''. It contains the following species: * Anderson ...
'', and other genera unrelated to ''Oryzomys''. Some of the new genera proposed were soon subsumed in ''Oryzomys'' again, and in ''The Families and Genera of Living Rodents'' (1941), John Ellerman listed '' Microryzomys'', ''
Oligoryzomys ''Oligoryzomys'' is a genus of rodents in the tribe Oryzomyini of family Cricetidae. Many species are known as pygmy rice rats or colilargos.Musser and Carleton, 2005 The genus is found from Mexico to Tierra del Fuego and includes approximately 1 ...
'', ''
Melanomys ''Melanomys'' is a genus of rodent in the tribe Oryzomyini of family Cricetidae, which is distributed in northern South America and adjacent Central America. It contains three species, two of which—'' Melanomys robustulus'' and '' Melanomys ...
'', ''
Nesoryzomys ''Nesoryzomys'' is a genus of rodent in the tribe Oryzomyini of family Cricetidae, endemic to the Galápagos Islands. Five species have been described, with two of them considered extinct Extinction is the termination of a kind of organism ...
'', and '' Oecomys'' as
synonyms A synonym is a word, morpheme, or phrase that means exactly or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are all ...
of ''Oryzomys''Ellerman, 1941, p. 340 and included about 127 species in it. In 1948,
Philip Hershkovitz Philip Hershkovitz (12 October 1909 – 15 February 1997) was an American mammalogy, mammalogist. Born in Pittsburgh, he attended the Universities of Pittsburgh and Michigan and lived in South America collecting mammals. In 1947, he was appointed ...
suggested that other oryzomyines like ''Nectomys'' and '' Megalomys'' could as well be included in ''Oryzomys'', and
Clayton Ray Clayton may refer to: People * Clayton (name) * Clayton baronets * The Clayton Brothers, Jeff and John, jazz musicians *Clayton Brothers, Rob and Christian, painter artists * Justice Clayton (disambiguation), the judges Clayton Places Canada * C ...
followed this suggestion in 1962. Hershkovitz and Ray's classification was never widely followed, and from 1976 on authors started to reinstate some of the other groups lumped in ''Oryzomys'' as separate genera. The genus was reduced to 43 species (out of 110 in Oryzomyini) in the third edition (2005) of ''Mammal Species of the World'',Musser and Carleton, 2005, p. 900 but it was still not a natural,
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
group; rather, it mostly united those oryzomyines that lacked the conspicuous specializations of other genera.Weksler, 2006, p. 82 In 2006, Marcelo Weksler's comprehensive phylogenetic analysis produced further evidence that the genus was polyphyletic, as species of ''Oryzomys'' were dispersed all over the oryzomyine tree. He proposed that eleven new genera should be created to accommodate those species that were not closely related to the type species of ''Oryzomys'', the marsh rice rat;Weksler, 2006, p. 75 he considered other options that would require fewer new genera, but argued that that would result in less meaningful genus-level groups in Oryzomyini. Later in the same year, Weksler, Percequillo, and Voss created ten new genera—''
Aegialomys ''Aegialomys'' is a genus of oryzomyini, oryzomyine rodents from the lowlands and mountains of western Peru and Ecuador, including the Galápagos Islands. The species in this genus have historically been placed in ''Oryzomys'', but according to c ...
'', ''
Cerradomys ''Cerradomys'' is a genus of oryzomyine rodents from eastern Bolivia, Paraguay, and central Brazil found in cerrado, Caatinga and Gran Chaco habitats. The species in this genus have historically been placed in ''Oryzomys'', but according to cl ...
'', ''
Eremoryzomys ''Eremoryzomys polius'', also known as the gray rice rat or the Marañon oryzomys,Musser and Carleton, 2005, p. 1153 is a rodent species in the tribe Oryzomyini of the family Cricetidae. Discovered in 1912 and first described in 1913 by Wilfr ...
'', ''
Euryoryzomys ''Euryoryzomys'' is a genus of rodents in the tribe Oryzomyini of family Cricetidae. It includes six species, which are distributed in South America. Until 2006, its members were included in the genus ''Oryzomys'', but they are not closely related ...
'', ''
Hylaeamys ''Hylaeamys'' is a genus of South American oryzomyine rodents found principally in humid forested areas east of the Andes. The species in this genus have historically been placed in ''Oryzomys''. They are most closely related to ''Euryoryzomys'', ...
'', ''
Mindomys ''Mindomys'' is a genus of sigmodontine rodents in the family Cricetidae. It includes two species known only from Ecuador, Hammond's rice rat ''Mindomys hammondi'', also known as Hammond's rice rat or Hammond's oryzomys,Musser and Carleton, ...
'', ''
Nephelomys ''Nephelomys'' is a genus of South American oryzomyine rodents found in the Andes from Bolivia to Venezuela, with a westward extension into the mountains of Costa Rica. Its generic name is derived from the Ancient Greek word ''nephelê'' "mist", ...
'', '' Oreoryzomys'', '' Sooretamys'', and ''
Transandinomys ''Transandinomys'' is a genus of rodents in the tribe Oryzomyini of family Cricetidae. It includes two species—'' T. bolivaris'' and '' T. talamancae''—found in forests from Honduras in Central America south and east to southwester ...
''—for species formerly placed in ''Oryzomys'' and placed six more species related to '' "Oryzomys" alfaroi'' in ''
Handleyomys ''Handleyomys'' is a genus of Central and South American rodents in the tribe Oryzomyini of family Cricetidae. It was first described in 2002 to include two species from the Colombian Andes which were previously included in distinct and unrelat ...
'' pending the description of more new genera for them. They left only five species in ''Oryzomys'', which was now finally a natural, monophyletic group. Because of subsequent taxonomic work, the number of species has since increased to at least eight. Some problems remain: ?''Oryzomys pliocaenicus'', a Miocene fossil from Kansas, is of uncertain identity but may belong in '' Bensonomys'',Weksler, 2006, p. 87 and fossils from the Miocene of Oregon and Pliocene of New Mexico have also been ascribed to ''Oryzomys'', but probably incorrectly. A possible ''Oryzomys'' has been recorded from the
Irvingtonian The Irvingtonian North American Land Mammal Age on the geologic timescale is the North American faunal stage according to the North American Land Mammal Ages chronology (NALMA), spanning from 1.9 million – 250,000 years BP. ...
(Pleistocene) of Saskatchewan.


Species

The current concept of ''Oryzomys'' derives from the ''palustris-mexicanus'' group recognized within a much larger genus ''Oryzomys'' by Merriam (1901) and the ''palustris'' group proposed by Goldman (1918).Carleton and Arroyo-Cabrales, 2009, p. 116 Merriam recognized 21 species within his group, but Goldman consolidated them into eight—the marsh rice rat in the United States, '' O. couesi'' in far southern Texas, Mexico, and Central America, and six others with small distributions. In 1960,
Raymond Hall Raymond is a male given name. It was borrowed into English from French (older French spellings were Reimund and Raimund, whereas the modern English and French spellings are identical). It originated as the Germanic ᚱᚨᚷᛁᚾᛗᚢᚾᛞ ( ...
united ''O. couesi'' and the marsh rice rat into a single species, ''Oryzomys palustris'', and thereafter, other localized forms were also included in ''O. palustris''.Musser and Carleton, 2005, p. 1147 Hershkovitz described another species in the group, '' O. gorgasi'' from Colombia, in 1970 and the next year he noted that '' O. dimidiatus'', previously classified as a ''Nectomys'', was similar to ''O. palustris''. After 1979, the marsh rice rat and ''O. couesi'' were again regarded as separate as a result of further work in Texas, where their ranges meet. While reviewing ''O. gorgasi'' in 2001, J. Sánchez H. and colleagues redefined and characterized the ''O. palustris'' group and listed ''O. couesi'', ''O. dimidiatus'', ''O. gorgasi'', and the marsh rice rat as its members; Guy Musser and Michael Carleton in the 2005 third edition of ''Mammal Species of the World'' additionally listed '' O. nelsoni'' from María Madre Island in western Mexico. In 2006, Weksler and colleagues followed the 2001 definition by Sánchez and others for the restricted genus ''Oryzomys'', but added '' O. antillarum'' from Jamaica as a species. Carleton and Joaquin Arroyo-Cabrales reviewed ''Oryzomys'' from western Mexico in 2009 and in this context provided an extended diagnosis of ''Oryzomys''. They recognized eight species: the six previously mentioned plus '' O. albiventer'' and '' O. peninsulae''. Also in 2009, Robert Voss and Weksler identified the
subfossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved in ...
''Oryzomys curasoae'' from
Curaçao Curaçao ( ; ; pap, Kòrsou, ), officially the Country of Curaçao ( nl, Land Curaçao; pap, Pais Kòrsou), is a Lesser Antilles island country in the southern Caribbean Sea and the Dutch Caribbean region, about north of the Venezuela coast ...
as an island population of ''O. gorgasi''.Voss and Weksler, 2009, p. 73 The next year, Delton Hanson and colleagues published a study using DNA sequence data from the cytochrome ''b'',
interphotoreceptor retinoid-binding protein Retinol-binding protein 3, interstitial (RBP3), also known as IRBP is a protein that in humans is encoded by the ''RBP3'' gene. RBP3 orthologs have been identified in most eutherians except tenrecs and armadillos. Function The inter- photore ...
, and alcohol dehydrogenase 1 genes to assess relationships within ''Oryzomys''. They recommended that the marsh rice rat be split into two species and that ''O. couesi'' be split into four species on the basis of the observed sequence divergence and other data. Merriam divided his ''palustris-mexicanus'' group in two "series" according to the color of the underparts (white or fulvous). Goldman divided his ''palustris'' group in two "sections"—a ''couesi'' section with ''O. couesi'' and six related species, and a ''palustris'' section with ''O. palustris'' only. He noted that the latter differed from the former in the generally darker, more brownish, longer fur, and larger
sphenopalatine vacuities In rodents, sphenopalatine vacuities are perforations of the roof of the mesopterygoid fossa, the open space behind the palate, in between the parapterygoid fossae. They may perforate the presphenoid or basisphenoid bone. Their development and ...
(openings in the mesopterygoid fossa, the gap behind the end of the palate).Goldman, 1918, p. 20 As Weksler's 2006 analysis included only ''O. couesi'' and the marsh rice rat among species of ''Oryzomys'' in the strict sense, he could not test those groups. Carleton and Arroyo-Cabrales concurred with Goldman's division, listing additional characters, and noted that the ''palustris'' group may be more semiaquatically adapted than the members of the ''couesi'' group are. In the latter, the fur is usually reddish-brown, as opposed to grayish-brown in the ''palustris'' group. Members of the ''couesi'' group have smaller sphenopalatine vacuities and a smaller sphenopalatine foramen, a foramen (opening) in the side of the skull above the molars, and a more highly developed
anterolabial cingulum 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 ...
on the third lower molar (a crest at the front of the tooth). The hypothenar pad of the hindfoot, located on the sole far from the fingers, is present in the ''couesi'' group, but absent in the ''palustris'' group.
Interdigital webbing Interdigital webbing is the presence of membranes of skin between the digits. Normally in mammals, webbing is present in the embryo but resorbed later in development, but in various mammal species it occasionally persists in adulthood. In humans, ...
may be more highly developed in the ''palustris'' group.Carleton and Arroyo-Cabrales, 2009 Using morphological data, Voss and Weksler found a closer relationship between ''O. couesi'' and ''O. gorgasi'' to the exclusion of ''O. palustris'', but with low confidence. The DNA sequence data of Hanson and colleagues supported a deep separation between the ''palustris'' and ''couesi'' groups, but a Costa Rican sample (assigned to ''O. couesi'') was about as distant from the two groups as they were from each other. The genus currently includes the following species:


Description

''Oryzomys'' contains medium-sized, semiaquatically specialized oryzomyine rodents. They have long, coarse fur that is grayish to reddish on the upperparts and white to buff on the underparts. The marsh rice rat superficially resembles the introduced species
black rat The black rat (''Rattus rattus''), also known as the roof rat, ship rat, or house rat, is a common long-tailed rodent of the stereotypical rat genus ''Rattus'', in the subfamily Murinae. It likely originated in the Indian subcontinent, but is n ...
and brown rat, but has larger differences in color between the upper- and underparts. The vibrissae (whiskers) are short and the ears are small and well-haired. The tail is usually as long as or longer than the head and body and is sparsely haired, but the hairs on the lower side are longer than those above. Females have eight mammae, as in most oryzomyines. The hindfeet are broad and have the first and fifth digits notably shorter than the middle three. The upper surface is hairy, but the underside is naked and covered with small irregularities (
squamae In some rodents, squamae are small tubercles resembling scales on the sole of the hindfeet. Among oryzomyine rodents, their development is variable; most have well-developed squamae, but in others they are indistinct or entirely absent. '' Delomys ...
). The pads are generally poorly developed, as are the ungual tufts. Interdigital webbing may be present, but its development is variable within the genus. The
karyotype A karyotype is the general appearance of the complete set of metaphase chromosomes in the cells of a species or in an individual organism, mainly including their sizes, numbers, and shapes. Karyotyping is the process by which a karyotype is disce ...
has been recorded in various populations of the marsh rice rat and ''O. couesi'' and is apparently stable within the genus at 56  chromosomes, with the fundamental number of chromosomal arms ranging from 56 to 60 (2n = 56, FN = 56–60). In both species, the stomach has the characteristic pattern of sigmodontines (
unilocular A locule (plural locules) or loculus (plural loculi) (meaning "little place" in Latin) is a small cavity or compartment within an organ or part of an organism (animal, plant, or fungus). In angiosperms (flowering plants), the term ''locule'' usu ...
-hemiglandular): it is not split in two chambers by an
incisura angularis The angular incisure (or angular notch) is a small notch on the stomach. It is located on the lesser curvature of the stomach near the pyloric The pylorus ( or ), or pyloric part, connects the stomach to the duodenum. The pylorus is considered a ...
and the front part ( antrum) is covered by a glandular epithelium. Furthermore, the gall bladder is absent, a synapomorphy of Oryzomyini. ''Oryzomys'' species have a large skull with a short rostrum and high
braincase In human anatomy, the neurocranium, also known as the braincase, brainpan, or brain-pan is the upper and back part of the skull, which forms a protective case around the brain. In the human skull, the neurocranium includes the calvaria or skul ...
. The interorbital region, located between the eyes, is narrowest to the front and is flanked by well-developed beads at its margins. The zygomatic plate is broad and has a well-developed zygomatic notch at its front. The zygomatic arch is robust and contains a small but distinct jugal bone. The interparietal bone, part of the roof of the braincase, is narrow and short;Goldman, 1918, p. 19; Carleton and Arroyo-Cabrales, 2009, p. 116 its narrowness is a synapomorphy for ''O. couesi'' plus the marsh rice rat according to Weksler's analysis.Weksler, 2006, p. 131 The incisive foramina are long, with their back margin at the front of the first molars or further back. The palate is also long, extending beyond the back margin of the maxillary bone, and is perforated near the third molars by well-developed
posterolateral palatal pits In anatomy, posterolateral palatal pits are gaps at the sides of the back of the bony palate, near the last molars.Weksler, 2006, p. 34 Posterolateral palatal pits are present, in various degrees of development, in several members of the roden ...
. There is no
alisphenoid strut In some rodents, the alisphenoid strut is an extension of the alisphenoid bone that separates two foramina in the skull, the masticatory–buccinator foramen and the foramen ovale accessorium. The presence or absence of this strut is variable be ...
, an extension of the
alisphenoid bone The greater wing of the sphenoid bone, or alisphenoid, is a bony process of the sphenoid bone; there is one on each side, extending from the side of the body of the sphenoid and curving upward, laterally, and backward. Structure The greater wi ...
that in some other oryzomyines separates two foramina in the skull. The auditory bullae are large. The condition of the arteries in the head is highly
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 ...
. In the mandible (lower jaw), the
coronoid process The Coronoid process (from Greek , "like a crown") can refer to: * The coronoid process of the mandible, part of the ramus mandibulae of the mandible * The coronoid process of the ulna The coronoid process of the ulna is a triangular process proj ...
, a process at the back, is well developedGoldman, 1918, p. 19 and the
capsular process In rodents, the capsular process or projection is a bony capsule that contains the root of the lower incisor. It is visible on the labial (outer) side of the mandible In anatomy, the mandible, lower jaw or jawbone is the largest, strongest ...
, a raising of the mandibular bone housing the root of the lower incisor, is conspicuous. As usual in oryzomyines, the molars are pentalophodont (have 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 i ...
s and
mesolophid 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 ...
s, accessory crests, well developed) and bunodont, with the cusps higher than the connecting crests. The cusps on the upper molars are arranged in two longitudinal series, not three as in the black and brown rats. The front cusps of the first upper and lower molar (anterocone and
anteroconid 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 ...
, respectively) are broad and not divided completely by an
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 ...
or flexid. Behind the anterocone, the
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 ...
(a smaller crest) is complete and separated from the anterocone. On both the second and third lower molars, the
anterolophid 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 ...
(a crest on the inner front corner) is present, a putative synapomorphy of the genus. The first molars have additional small roots in addition to the main ones, so that the upper first molar has four and the lower has three or four roots. As is characteristic of Sigmodontinae, the marsh rice rat and ''O. couesi'' have a complex penis, with the
baculum The baculum (also penis bone, penile bone, or ''os penis'', ''os genitale'' or ''os priapi'') is a bone found in the penis of many placental mammals. It is absent from the human penis, but present in the penises of some primates, such as the ...
(penis bone) displaying large protuberances at the sides. The outer surface of the penis is mostly covered by small spines, but there is a broad band of nonspinous tissue. The papilla (nipple-like projection) on the dorsal (upper) side of the penis is covered with small spines, a character these two species share only with ''Oligoryzomys'' among oryzomyines examined. On the urethral process, located in the crater at the end of the penis, a fleshy process (the
subapical lobule A subapical consonant is a consonant made by contact with the underside of the tip of the tongue. The only common subapical articulations are in the postalveolar Postalveolar or post-alveolar consonants are consonants articulated with the tongue ...
) is present; it is absent in all other oryzomyines with studied penes except '' Holochilus brasiliensis''. Both traits are recovered as synapomorphies of ''O. couesi'' plus the marsh rice rat in Weksler's analysis.


Distribution, ecology, and behavior

The range of ''Oryzomys'' extends from New Jersey in the eastern United States through Mexico and Central America south to northwestern Colombia and east to northwestern Venezuela and Curaçao. Species of ''Oryzomys'' usually live in wet habitats such as marshes, streams, and mangroves, but both the marsh rice rat and ''O. couesi'' are also occasionally encountered in drier habitats. They occur or occurred on many continental-shelf islands and one oceanic island, Jamaica; their adeptness at colonizing islands may be caused by their close association with water and frequent occurrence in coastal wetlands. The oldest fossils date to the Rancholabrean of the United States, about 300,000 years ago; although there have been some earlier North American records, those are not in fact referable to ''Oryzomys'' or even Oryzomyini. Oryzomyines likely evolved in South America east of the Andes; the presence of ''Oryzomys'' in Central America and other trans-Andean regions is thought to be the result of one of several independent invasions of this region by oryzomyines.Weksler, 2006, p. 88 Alternatively, ''Oryzomys'' may have evolved from the Pliocene North American '' Jacobsomys''. ''O. antillarum'' may have reached Jamaica during the last glacial period while sea levels were low. Behavior is known mainly from the marsh rice rat and ''O. couesi'', with some scattered data from the other species. ''Oryzomys'' are semiaquatic, spending much time in the water, and otherwise mainly live on the ground;Reid, 2009, p. 205 both the marsh rice rat and ''O. couesi'' are known to be excellent swimmers and will flee into the water when disturbed. Both are also active during the night and build nests of interwoven vegetation, which may be suspended above the water. Breeding may occur throughout the year in both species, but is known to be seasonally variable in the marsh rice rat. In both, gestation takes about 21 to 28 days and litter size is usually one to seven, averaging three to five. Young marsh rice rats and ''O. couesi'' become reproductively active when about 50 days old. The marsh rice rat, ''O. couesi'', and ''O. gorgasi'' are known to be
omnivore An omnivore () is an animal that has the ability to eat and survive on both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize the nutr ...
s, eating both plant and animal material. They eat both seeds and green plant parts and consume a variety of animals, including insects, crustaceans, and many others. The
barn owl The barn owl (''Tyto alba'') is the most widely distributed species of owl in the world and one of the most widespread of all species of birds, being found almost everywhere except for the polar and desert regions, Asia north of the Himalaya ...
(''Tyto alba'') is a major predator on the marsh rice rat and remains of ''O. antillarum'', ''O. couesi'', and ''O. gorgasi'' have been found in
owl pellet A pellet, in ornithology, is the mass of undigested parts of a bird's food that some bird species occasionally regurgitate. The contents of a bird's pellet depend on its diet, but can include the exoskeletons of insects, indigestible plant ma ...
deposits. Several other animals are known to prey on ''Oryzomys''. A variety of parasites are known from ''O. couesi'' and the marsh rice rat and two parasitic
nematode The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant-Parasitism, parasitic nematodes also known as eelworms. They are a diverse animal phylum inhab ...
s have been found in ''O. gorgasi''.


Human interactions

Two species of ''Oryzomys'', ''O. antillarum'' and ''O. nelsoni'', have gone extinct since the 19th century, and a third, ''O. peninsulae'', is unlikely to be still extant. Their extinction may have been caused by habitat destruction and by introduced species such as the
small Asian mongoose Small Asian mongoose is a common name applied to two mammals which were formerly considered to be a single species: *Javan mongoose *Small Indian mongoose The small Indian mongoose (''Urva auropunctata'') is a mongoose species native to Iraq and ...
and the brown and black rat. These same causes may threaten ''O. gorgasi'', which the IUCN Red List assesses as " Endangered". ''O. albiventer'' has been affected by human alteration of its habitat, but likely still survives.Carleton and Arroyo-Cabrales, 2009, p. 115 In contrast, the widespread species, the marsh rice rat and ''O. couesi'', are common and of no conservation concern—indeed, both have been considered a pest—but some populations are threatened. Like these two species, ''O. dimidiatus'' is assessed as " Least Concern" by the Red List. The marsh rice rat is the natural reservoir of the
Bayou virus ''Bayou orthohantavirus'' (BAYV) is a species of ''Orthohantavirus'' first identified in 1993 in Louisiana. and later confirmed by other investigators. In 1996, the marsh rice rat was identified as the natural reservoir of the virus, indicating t ...
, the second most common cause of hantavirus pulmonary syndrome in the United States. Two other hantaviruses,
Catacamas virus Catacamas virus is a single-stranded, enveloped novel RNA virus in the genus ''Orthohantavirus'' of the order ''Bunyavirales'' isolated in ''Oryzomys couesi'' near the town of Catacamas in eastern Honduras. It is a member virus of ''Bayou orthoh ...
and
Playa de Oro virus Playa de Oro virus (OROV) is a probable species of orthohantavirus found in the rodents ''Oryzomys couesi'' and ''Sigmodon mascotensis'' in the Mexican state of Colima. The former is thought to be the main host (biology), host. The DNA sequence, ...
, occur in ''O. couesi'' in Honduras and western Mexico, respectively, but are not known to infect humans.Milazzo et al., 2006; Chu et al., 2008


Notes


References


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