Gorgas's Rice Rat
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''Oryzomys gorgasi'', also known as Gorgas's oryzomysMusser and Carleton, 2005, p. 1148 or Gorgas's rice rat, is a
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 ...
in the genus ''
Oryzomys ''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 (''O. palustris'') of the United States and ...
'' of 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, and has m ...
. First recorded in 1967, it is known from only a few localities, including a freshwater swamp in the lowlands of northwestern
Colombia Colombia (, ; ), officially the Republic of Colombia, is a country in South America with insular regions in North America—near Nicaragua's Caribbean coast—as well as in the Pacific Ocean. The Colombian mainland is bordered by the Car ...
and a mangrove islet in northwestern
Venezuela Venezuela (; ), officially the Bolivarian Republic of Venezuela ( es, link=no, República Bolivariana de Venezuela), is a country on the northern coast of South America, consisting of a continental landmass and many islands and islets in th ...
. It reportedly formerly occurred on the island of
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 ...
off northwestern Venezuela; this extinct population has been described as a separate species, ''Oryzomys curasoae'', but does not differ morphologically from mainland populations. ''Oryzomys gorgasi'' is a medium-sized, brownish species with large,
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 Semiaquatic animals include: * Verte ...
ally specialized feet. It differs from other ''Oryzomys'' species in several features of its skull. Its diet includes
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group ...
s,
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pairs ...
s, and plant material. The species is listed as "Endangered" by the
International Union for Conservation of Nature The International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of natu ...
as a result of destruction of its habitat and competition with the introduced
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 ...
(''Rattus rattus'').


Taxonomy

''Oryzomys gorgasi'' was first found in
Antioquia Department ) , anthem = Himno de Antioquia , image_map = Antioquia in Colombia (mainland).svg , map_alt = , map_caption = Antioquia shown in red , image_ma ...
of northwestern Colombia in 1967 during an expedition by the
U.S. Army Medical Department The Army Medical Department of the U.S. Army (AMEDD), formerly known as the Army Medical Service (AMS), encompasses the Army's six medical Special Branches (or "Corps"). It was established as the "Army Hospital" in July 1775 to coordinate the me ...
and the Gorgas Memorial Laboratory. In 1971,
Field Museum 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 ...
zoologist
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 ...
described a new species, ''Oryzomys gorgasi'', on the basis of the single known specimen, an old male. He named the animal after physician
William Crawford Gorgas William Crawford Gorgas KCMG (October 3, 1854 – July 3, 1920) was a United States Army physician and 22nd Surgeon General of the U.S. Army (1914–1918). He is best known for his work in Florida, Havana and at the Panama Canal in abating the ...
, the namesake of the Gorgas Memorial Laboratory.Hershkovitz, 1971, p. 700 Hershkovitz considered the new species most closely related to ''Oryzomys palustris'', which at the time included North and Central American populations now divided into several species, including 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'') and '' O. couesi''. The species was not recorded again until 2001, when Venezuelan zoologist J. Sánchez H. and coworkers reported on 11 specimens collected in coastal northwestern Venezuela in 1992, from the Colombian locality.Sánchez et al., 2001, p. 206 They confirmed that ''O. gorgasi'' is a distinct species related to the ''O. palustris'' group.Sánchez et al., 2001, p. 210 In 2001, Donald McFarlane and Adolphe Debrot described a new ''Oryzomys'' species from the
Dutch Dutch commonly refers to: * Something of, from, or related to the Netherlands * Dutch people () * Dutch language () Dutch may also refer to: Places * Dutch, West Virginia, a community in the United States * Pennsylvania Dutch Country People E ...
island of
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 ...
off northwestern Venezuela. For their description, they used
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 ...
material from owl pellets, including two partial
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, the ...
s and several hemi
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. They referred the species to ''
Oecomys ''Oecomys'' is a genus of rodent within the tribe Oryzomyini of family Cricetidae. It contains about 17 species, which live in trees and are distributed across forested parts of South America, extending into Panama and Trinidad Trinidad i ...
'',McFarlane and Debrot, 2001, p. 184 a group of
arboreal Arboreal locomotion is the Animal locomotion, locomotion of animals in trees. In habitats in which trees are present, animals have evolved to move in them. Some animals may scale trees only occasionally, but others are exclusively arboreal. Th ...
(tree-living), mainly South American rodents related to ''Oryzomys''. ''O. curasoae'' has also been known as the "Curaçao Rice Rat"Lamoreux, 2008 and the "Curaçao Oryzomys". Marcelo Weksler and colleagues removed most of the species then placed in ''Oryzomys'' from the genus in 2006, retaining only the marsh rice rat and related species, including ''O. gorgasi''. They also kept ''O. curasoae'' in the genus and suggested that it may not be distinct from ''O. gorgasi''.Weksler et al., 2006, table 1, footnote e In a 2009 paper, R.S. Voss and Weksler examined the two and concluded that they represented the same species on the basis of direct comparisons and a
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
analysis.Voss and Weksler, 2009, p. 78 The resultant
tree In botany, a tree is a perennial plant with an elongated stem, or trunk, usually supporting branches and leaves. In some usages, the definition of a tree may be narrower, including only woody plants with secondary growth, plants that are ...
placed ''O. curasoae'' and ''O. gorgasi'' sister to each other and closer to ''O. couesi'' than to the marsh rice rat.Voss and Weksler, 2009, fig. 1 Accordingly, they placed ''O. curasoae'' as a
junior synonym The Botanical and Zoological Codes of nomenclature treat the concept of synonymy differently. * In botanical nomenclature, a synonym is a scientific name that applies to a taxon that (now) goes by a different scientific name. For example, Linna ...
of the earlier described ''O. gorgasi''.Voss and Weksler, 2009, p. 73 ''Oryzomys gorgasi'' is the southeasternmost representative of the genus ''Oryzomys'', which extends north into the eastern
United States The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territorie ...
(marsh rice rat, ''O. palustris'').Carleton and Arroyo-Cabrales, 2009, p. 106 ''O. gorgasi'' is further part of the ''O. couesi'' section, which is centered on the widespread Central American ''O. couesi'' and also includes six other species with more limited and peripheral distributions.Carleton and Arroyo-Cabrales, 2009, p. 117 Many aspects of the
systematics Biological systematics is the study of the diversification of living forms, both past and present, and the relationships among living things through time. Relationships are visualized as evolutionary trees (synonyms: cladograms, phylogenetic tre ...
of the ''O. couesi'' section remain unclear and it is likely that the current classification underestimates the true diversity of the group.Carleton and Arroyo-Cabrales, 2009, p. 107 ''Oryzomys'' is classified in the tribe
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 S ...
, a diverse assemblage of American rodents of over a hundred species,Weksler, 2006, p. 3 and on higher taxonomic levels in 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 specie ...
of 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, and has m ...
, along with hundreds of other species of mainly small rodents.Musser and Carleton, 2005


Description

''Oryzomys gorgasi'' is a medium-sized
oryzomyine 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 ...
Hershkovitz, 1971, p. 701 with small ears and large feet,Hershkovitz, 1971, p. 702 and is similar to the marsh rice rat in general appearance. The long and coarse fur is brownish above and ochraceous below. At the base of the tail, the upper and lower sides differ in color and at the end is a short tuft of hairs. The scales on the tail are well-developed. As in other ''Oryzomys'', the hindfeet exhibit specializations for life in the water. The plantar (lower) surface of the
metatarsus The metatarsal bones, or metatarsus, are a group of five long bones in the foot, located between the tarsal bones of the hind- and mid-foot and the phalanges of the toes. Lacking individual names, the metatarsal bones are numbered from the med ...
is naked. Two of the
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 ...
are very small.
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, tufts of hair at the bases of the claws, are poorly developed.
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, ...
is present, but extends along less than half of the first
phalange The phalanges (singular: ''phalanx'' ) are digital bones in the hands and feet of most vertebrates. In primates, the thumbs and big toes have two phalanges while the other digits have three phalanges. The phalanges are classed as long bones. ...
s. In specimens from El Caimito, total length is , averaging (measured in 6 specimens); tail length is , averaging (measured in 8 specimens); hindfoot length is , averaging (measured in 10 specimens); ear length is , averaging (measured in 7 specimens); and condylo-incisive length (a measure of total skull size) is , averaging (measured in 5 specimens). In 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 several ...
from Colombia, an old male, total length is ; tail length is ; ear length is ; and condylo-incisive length is .Sánchez et al., 2001, table 1 The collector recorded the holotype's hindfoot as being long, but Sánchez and colleagues remeasured it as .Sánchez et al., 2001, p. 207 The rostrum (front part of the
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, the ...
) is short. The broad
zygomatic plate In rodent anatomy, the zygomatic plate is a bony plate derived from the flattened front part of the zygomatic arch (cheekbone). At the back, it connects to the front (maxillary) root of the zygomatic arch, and at the top it is connected to the ...
develops a prominent notch, but not a spine, on its front end, and its back margin is in front of the first molars. The
interorbital region The interorbital region of the skull is located between the eyes, anterior to the braincase. The form of the interorbital region may exhibit significant variation between taxonomic groups. In oryzomyine rodents, for example, the width, form, an ...
, located between the eyes, is narrowest towards the front and is flanked by beadings along its margins. The
interparietal bone An interparietal bone (os interparietale or Inca bone or ''os inca var.'') is a dermal bone situated between the parietal and supraoccipital. It is homologous to the postparietal bones of other animals. In humans, it corresponds to the upper p ...
is relatively long. The
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 palate between the incisors and the molars, are narrow and long and taper towards the end. The palate itself is also long, extending beyond the molars, and includes prominent
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 ...
near the third molars, which are excavated into deep fossae. The roof of the mesopterygoid fossa, the opening behind the palate, is not perforated by
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 ...
. ''O. gorgasi'' lacks an
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 ...
; in some other oryzomyines, this extension of the alisphenoid bone separates two openings in the skull, the masticatory–buccinator foramen and the foramen ovale accessorium. The squamosal bone lacks a suspensory process that contacts the tegmen tympani, the roof of the
tympanic cavity The tympanic cavity is a small cavity surrounding the bones of the middle ear. Within it sit the ossicles, three small bones that transmit vibrations used in the detection of sound. Structure On its lateral surface, it abuts the external auditory ...
, a defining character of oryzomyines. The
subsquamosal fenestra In some rodents, the subsquamosal fenestra is an opening between two parts of the squamosal bone, at the back of the skull. It can be seen in lateral view. Most Oryzomyini have the fenestra, but some species, including those in the genera ''Nectomys ...
, an opening at the back of the skull determined by the shape of the squamosal, is almost absent.Sánchez et al., 2001, p. 209 In the
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 ...
(lower jaw), the upper and lower
masseteric ridge Masseteric is an adjective meaning "of or pertaining to the Masseter muscle", such as: * Masseteric artery * Masseteric nerve The masseteric nerve is a nerve of the face. It is a branch of the mandibular nerve (V3). It crosses the mandibular n ...
s come close together below the first molars, but do not fuse. The back end of the lower incisor root is in a
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 behind the molars. The upper incisors have yellowish enamel and are
opisthodont In rodents, incisor procumbency refers to the orientation of the upper incisor, defined by the position of the cutting edge of the incisor relative to the vertical plane of the incisors. Proodont incisors have the cutting edge in front of the ver ...
, with the cutting edge inclined backwards. The molars are relatively small and are brachydont (low-crowned) and bunodont (with the cusps higher than the connecting crests). They are similar to those of the marsh rice rat in structural details. The upper and lower first molars have small accessory roots, as in many other oryzomyines, and the second and third lower molar each have two roots only. ''Oryzomys gorgasi'' is distinguished from other ''Oryzomys'' species by its short rostrum, the form of its incisive foramina, the absence of sphenopalatine vacuities, and the near absence of a subsquamosal fenestra. Within the species, the Colombian specimen differs from the Venezuelan animals in being larger in some measurements, but having smaller teeth, and in having oddly shaped wear facets of the incisors. The Colombian animal was probably kept in captivity for some time after it was caught, which would explain its large size and odd wear facets. There are no substantial differences between mainland ''O. gorgasi'' and material from Curaçao.Voss and Weksler, 2009, p. 77


Distribution and ecology

As far as known, ''Oryzomys gorgasi'' has a disjunct distribution in northwestern South America, including Colombia, Venezuela, and Curaçao. In a 2009 paper, Carleton and Arroyo-Cabrales speculated that its distribution may extend into Central America. The Colombian population is known from the holotype only, caught at Loma Teguerre (7°54'N, 77°W) in Antioquia Department, northwestern Colombia, near the
Río Atrato The Atrato River () is a river of northwestern Colombia. It rises in the slopes of the Western Cordillera and flows almost due north to the Gulf of Urabá (or Gulf of Darién), where it forms a large, swampy delta. Its course crosses the C ...
, at about 1 m above sea level. The location is apparently a freshwater swamp, and Hershkovitz suggested that ''O. gorgasi'' probably occurred throughout the swamp forests in the Río Atrato basin. On Curaçao, it is known from cave faunas at Tafelberg Santa Barbara, Noordkant, Ser'i Kura, and Hermanus. At Tafelberg Santa Barbara, it was found in association with introduced
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 ...
s (''Rattus rattus''), indicating that the population persisted at least until the first European contact in 1499.McFarlane and Debrot, 2001, p. 182 In Venezuela, it was found on
El Caimito EL, El or el may refer to: Religion * El (deity), a Semitic word for "God" People * EL (rapper) (born 1983), stage name of Elorm Adablah, a Ghanaian rapper and sound engineer * El DeBarge, music artist * El Franco Lee (1949–2016), American p ...
, a small () islet just east of the outlet of
Lake Maracaibo Lake Maracaibo (Spanish: Lago de Maracaibo; Anu: Coquivacoa) is a lagoon in northwestern Venezuela, the largest lake in South America and one of the oldest on Earth, formed 36 million years ago in the Andes Mountains. The fault in the northern se ...
in the state of Zulia, where the only other native non-flying mammal is the
opossum Opossums () are members of the marsupial order Didelphimorphia () endemic to the Americas. The largest order of marsupials in the Western Hemisphere, it comprises 93 species in 18 genera. Opossums originated in South America and entered North ...
'' Marmosa robinsoni''.Sánchez et al., 2001, p. 211 El Caimito is separated from the mainland by a narrow, brackish channel and contains sand banks with xerophytic vegetation surrounded by marshy lagoons with '' Rhizospora mangle'' mangroves. ''Oryzomys gorgasi'' was caught in all habitats on the islet, but has not been found in other similar sites in northwestern Venezuela, where the introduced black rat is the only rodent collected. Analysis of stomach contents of El Caimito specimens indicates that the species is an
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 ...
, with a diet including
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group ...
s,
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pairs ...
s, plant seeds, and other plant material. The crustaceans may include fiddler crabs (''Uca'') and a mangrove tree crab of the genus '' Aratus''; the insects include flies (Diptera); and the plants include grass seeds. 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, ''
Litomosoides sigmodontis ''Litomosoides'' is a genus of nematodes belonging to the family Onchocercidae. The species of this genus are found in Europe and America. Species Species: *''Litomosoides anguyai'' *''Litomosoides bonaerensis'' *''Litomosoides brasilien ...
'' (family Onchocercidae) and an undetermined species of ''
Pterygodermatites ''Pterygodermatites'' is a genus of parasitic nematodes in the family Rictulariidae. Their life-cycle is complex. Species include: *'' Pterygodermatites baiomydis'' Lynggaard, García-Prieto, Guzmán-Cornejo & Osorio-Sarabia, 2014 *'' Pterygo ...
'' (family
Rictulariidae Rictulariidae is a family of nematodes belonging to the order Rhabditida. Genera: * '' Pseudorictularia'' Dollfus & Desportes, 1945 * ''Pterygodermatites ''Pterygodermatites'' is a genus of parasitic nematodes in the family Rictulariidae. ...
), are known to infect ''O. gorgasi''. The 2009 IUCN Red List tersely indicates that the species has been found in a second Venezuelan locality.


Conservation status

On the 2017 IUCN Red List, ''O. gorgasi'' is listed as "endangered" and ''O. curasoae'' as "data deficient". The species may be threatened by competition with introduced black ratsSánchez et al., 2001, p. 211; Weksler and Timm, 2018 and destruction of its habitat,Sánchez et al., 2001, p. 205 but does occur in at least one
protected area Protected areas or conservation areas are locations which receive protection because of their recognized natural, ecological or cultural values. There are several kinds of protected areas, which vary by level of protection depending on the ena ...
. Displacement by the black rat has caused the species to become locally extinct in parts of its Venezuelan range. Suitable habitats for ''O. gorgasi'' exist in inland Venezuela, and further study is needed to determine whether it is present there. The extinction of the Curaçao population may also have been caused by competition with the black rat, which has been found together with ''Oryzomys'' in subfossil deposits.


References


Literature cited

* * * * * * * * * {{Featured article Mammals of Colombia Mammals of Venezuela Oryzomys Taxa named by Philip Hershkovitz Taxonomy articles created by Polbot Mammals described in 1971