Marsh rice rat
Marsh rice rat Temporal range: Rancholabrean (300,000 years before present) – present
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Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Order: | Rodentia |
Family: | Cricetidae |
Subfamily: | Sigmodontinae |
Genus: | Oryzomys |
Species: | O. palustris
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Binomial name | |
Oryzomys palustris | |
Current (blue) and approximate former (light blue) distribution of the marsh rice rat in the eastern United States. A small part of the distribution of Oryzomys couesi is also shown (red). | |
Synonyms[13] | |
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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, from New Jersey and Kansas south to Florida and northeasternmost Tamaulipas, Mexico; its range previously extended further west and north, where it may have been a commensal in corn-cultivating communities. Weighing about 40 to 80 g (1.4 to 2.8 oz), the marsh rice rat is a medium-sized rodent that resembles the common black and brown rat. The upperparts are generally gray-brown, but are reddish in many Florida populations. The feet show several specializations for life in the water. The skull is large and flattened, and is short at the front.
John Bachman discovered the marsh rice rat in 1816, and it was formally described in 1837. Several subspecies have been described since the 1890s, mainly from Florida, but disagreement exists over their validity. The Florida Keys population is sometimes classified as a different species, the silver rice rat (Oryzomys argentatus). Data from the mitochondrial cytochrome b gene indicate a deep divergence between populations east of Mississippi and those further west, which suggests that the western populations may be recognized as a separate species, Oryzomys texensis. The species is part of the genus Oryzomys, which also includes several others occurring further south in Mexico, Central America, and northwestern South America, some of which have previously been regarded as subspecies of the marsh rice rat. One, Oryzomys couesi, occurs with the marsh rice rat in Tamaulipas and southern Texas.
The marsh rice rat is active during the night, makes nests of sedge and grass, and occasionally builds runways. Its diverse diet includes plants, fungi, and a variety of animals. Population densities are usually below 10 per ha (four per acre) and home ranges vary from 0.23 to 0.37 ha (0.57 to 0.91 acres), depending on sex and geography. Litters of generally three to five young are born after a pregnancy around 25 days, mainly during the summer. Newborns are helpless at birth, but are weaned after a few weeks. Several animals prey on the marsh rice rat, including the
Taxonomy
The marsh rice rat is classified as one of eight species in the genus Oryzomys, which is distributed from the eastern United States (marsh rice rat) into northwestern South America (O. gorgasi).[14] Oryzomys previously included many other species, which were reclassified in various studies culminating in contributions by Marcelo Weksler and coworkers in 2006 that removed more than 40 species from the genus.[15] All are placed in the tribe Oryzomyini ("rice rats"), a diverse assemblage of over 100 species,[16] and on higher taxonomic levels in the subfamily Sigmodontinae of the family Cricetidae, along with hundreds of other species of mainly small rodents, most of which occur in South and Central America. In the United States, the marsh rice rat is the only oryzomyine rodent except for Oryzomys couesi in a small area of southern Texas; the only other sigmodontines present are several species of cotton rats (Sigmodon) in the southern half of the country.[17]
Early history
The marsh rice rat was discovered in 1816 in
In 1854, in The quadrupeds of North America, Bachman redescribed it as Arvicola oryzivora, considering it more closely related to the
Species boundaries and subspecies
In the 1890s, several
Merriam and Goldman had recognized that a number of Central American species, including Oryzomys couesi and numerous forms with more limited distributions, are related to the marsh rice rat.[27] O. couesi ranges north to southernmost Texas, where its distribution meets that of the marsh rice rat. In 1960, Raymond Hall argued that specimens from the contact zone were intermediate between the local forms of O. couesi and the marsh rice rat, and accordingly included the former in the marsh rice rat.[28] While reporting on the ecology of Texan O. couesi in 1979, Benson and Gehlbach noted that populations of O. couesi and the marsh rice rat there were in fact distinct, with the latter being smaller and less brown and more gray in color; their karyotypes were also distinct.[29] Since then, the two have generally been retained as distinct species, as supported by further research; a 1994 study even found the two to occur at some of the same places (in sympatry) in southern Texas and nearby Tamaulipas, Mexico.[30]
In 1973, rice rats were discovered at
Among the described subspecies, a 1989
In 2010, Delton Hanson and colleagues published a study of the relationships among populations of Oryzomys on the basis of data from three genes—the
Common names
Many common names have been proposed for the marsh rice rat. Early describers used "rice meadow-mouse"[3] and "rice-field mouse"[43] and in the early 1900s, name such as "rice rat", "marsh mouse", and "swamp rice rat" came into use.[44] Some of the subspecies received their own common names, such as "Florida marsh mouse",[45] "swimming rice rat",[46] and "Central Florida rice rat" for O. p. natator;[47] "Bangs' marsh mouse",[45] "Cape Sable rice rat",[46] and "Everglades rice rat" for O. p. coloratus;[48] and "Texas rice rat" for O. p. texensis.[49] The species is now usually known as the "marsh rice rat",[50] although "marsh oryzomys" has also been in recent use.[51] The Florida Keys form (O. p. argentatus) is known as the "silver rice rat".[34]
Description
Population | n | Total length | Tail length | Hindfoot length |
---|---|---|---|---|
O. p. palustris (New Jersey)[52] | 4 | 242 (237–245) | 112 (109–116) | 31 (30–31.5) |
O. p. natator (Florida)[53] | 10 | 281.2 (246–318) | 140.6 (122–173) | 33.1 (28–37) |
O. p. coloratus (Florida)[53] | 11 | 283.0 (250–326) | 143.5 (123–171) | 33.4 (31–38) |
O. p. texensis (Texas)[49] | 8 | 242 (226–279) | 120 (108–133) | 29 (28.5–30.5) |
O. p. planirostris (Florida)[53] | 14 | 247.5 (226–266) | 129.6 (108–128) | 31 (29–33) |
O. p. sanibeli (Florida)[53] | 11 | 257.5 (233–274) | 123.6 (111–138) | 31.0 (29–33) |
O. argentatus (Florida Keys)[12] | 2 | 251, 259 | 121, 132 | 32, 32 |
Measurements are all in millimeters and are in the form "average (minimum–maximum)", except those of the Florida Keys population.
n=Number of specimens measured. |
The marsh rice rat is a medium-sized rodent that looks much like the common
The fore feet have four and the hind feet five digits.[59] On the fore feet, the ungual tufts (tufts of hair on the digits) are absent.[60] The hind feet are broad and have a short fifth digit. Many of the pads are reduced, as are the ungual tufts, but small interdigital webs are present.[61] The Florida Keys form, P. o. argentatus, has even more reduced ungual tufts.[62] Many of these traits are common adaptations to life in the water in oryzomyines.[63]
Some geographic variation in fur color occurs; western populations (P. o. texensis) are lighter than those from the east (
Total length is 226 to 305 mm (8.9 to 12.0 in), tail length 108 to 156 mm (4.3 to 6.1 in), hind foot length 28 to 37 mm (1.1 to 1.5 in),
The
Male reproductive anatomy
The glans penis is long and robust,[80] averaging 7.3 mm (0.29 in) long and 4.6 mm (0.18 in) broad, and the baculum (penis bone) is 6.6 mm (0.26 in) long.[81] As is characteristic of the Sigmodontinae, the marsh rice rat has a complex penis, with the distal (far) end of the baculum ending in three digits.[82] The central digit is notably larger than those at the sides.[80] The outer surface of the penis is mostly covered by small spines, but a broad band of nonspinous tissue is seen.[83] The papilla (nipple-like projection) on the dorsal (upper) side of the penis is covered with small spines, a character the marsh rice rat shares only with Oligoryzomys and Oryzomys couesi among oryzomyines examined.[84] On the urethral process, located in the crater at the end of the penis,[85] a fleshy process (the subapical lobule) is present; it is absent in all other oryzomyines with studied penes except O. couesi and Holochilus brasiliensis.[86] The baculum is deeper than it is wide.[80]
Some features of the accessory glands in the male genital region vary among oryzomyines. In the marsh rice rat, a single pair of
Skull
The marsh rice rat has a large, flattened skull
The
In the mandible, the mental foramen, an opening just before the first molar, opens sidewards, not upwards as in a few other oryzomyines.[101] The upper and lower masseteric ridges, which anchor some of the chewing muscles, join at a point below the first molar and do not extend forward beyond that point.[102] The capsular process, a raising of the bone of the back of the mandible that houses the back end of the incisor, is present, but not as large as in O. couesi.[103]
Teeth
The
The upper molars have two longitudinal rows of cusps, not three as in the black and brown rats.
The first lower molar is rounded at the front end and the labial and lingual conules of the
Postcranial skeleton
As usual in oryzomyines, 12 ribs are present. The first rib
Physiology
In poor conditions, the weight of the adrenal gland may increase up to 200%,[123] and rice rats are unable to conserve water well when dehydrated,[76] and in water contaminated with oil, they swim less and their mortality increases.[124] The median amount of radiation needed to kill a marsh rice rat is 5.25 Gy and the lethal dose of potassium cyanide is 7.20 mg/kg; both values are relatively low for cricetid rodents.[125] In one study, wild rice rats in radioactively contaminated areas did not show signs of disease.[126] Exposure to more daylight and higher food availability cause increased development of the gonads in both adult and juvenile rice rats.[127] When the pineal gland is removed or melatonin is administered in male rice rats, the testes are reduced and tend to regress into the body.[128]
Distribution and habitat
The marsh rice rat currently occurs in much of the eastern and southern United States, northeast to southern New Jersey, and south to southeastern Texas and far northeastern
Cave and archeological remains indicate that the range of the marsh rice rat has extended substantially further north and west earlier in the
In Tamaulipas and southern Texas, the ranges of the marsh rice rat and the related Oryzomys couesi meet;
The marsh rice rat occurs in several habitats, ranging from coastal salt marshes to mountain streams and clearings. It is semiaquatic, spending much time in the water, and usually occurs in wetland habitats. It prefers areas where the ground is covered with grasses and sedges, which protect it from predators.[142] In southern Illinois, marsh rice rats are more likely to occur in wetlands with more herbaceous cover, visual obstruction, and nearby grasslands.[143] The species also occurs in drier uplands, which serve as sinks for young, dispersing animals and as refuges during high tide.[144] Rice rats are adept overwater dispersers; studies on islands off Virginia's Delmarva Peninsula show that they readily cross 300-m (1000 ft) channels between islands.[145]
Behavior and ecology
Marsh rice rats are active during the night, so are rarely seen, although they may be among the most common small mammals in part of their range. They build nests of sedge and grass, about 13 cm (5 in) large, which are placed under debris, near shrubs, in short burrows, or high in aquatic vegetation. They may also use old nests of
Many animals prey on marsh rice rats. The
Population dynamics
The population density of the marsh rice rat usually does not reach 10 per ha (4 per acre).[158] The weather may influence population dynamics;[123] in the Everglades, densities may exceed 200 per ha (80 per acre) when flooding concentrates populations on small islands,[159] In the Florida Keys, population density is less than 1 per ha (0.4 per acre).[160] On Breton Island, Louisiana, perhaps an atypical habitat, home ranges in males average about 0.37 hectares (0.91 acres) and in females about 0.23 hectares (0.57 acres). A study in Florida found male home ranges to average 0.25 hectares (0.62 acres) and female 0.33 hectares (0.82 acres).[161]
Population size is usually largest during the summer and declines during winter,[159] although populations in Texas and Louisiana may be more seasonally stable.[162] Animals also often lose weight during winter.[163] Population size varies dramatically from year to year in southern Texas.[92] In coastal Mississippi, storms probably do not cause the population to decline substantially, and in Texas, inundation of its habitat did not significantly influence population density.[164] However, in Mississippi, flooding did cause a marked decline in rice rat abundance.[165]
In the northern part of its range, the species often occurs with the
Diet
The marsh rice rat takes both vegetable and animal food, and is more carnivorous than most small rodents are;
Animals that are important to the marsh rice rat's diet include insects,
Laboratory studies have found that rice rats assimilate 88% to 95% of the energy in their food. They lose weight when fed on Spartina, fiddler crabs, or sunflower seeds alone, but a diet consisting of several of those items or of mealworms is adequate to maintain weight.[59] In an experiment, marsh rice rats did not show hoarding behavior, but wild rice rats have been observed carrying food to a nest.[74] Even when they live in uplands, they mostly eat water plants and animals, although they consume some upland plants.[144]
Reproduction and lifecycle
Breeding occurs mostly during the summer. Some studies report that breeding ceases entirely during the winter, but winter breeding occurs as far north as Virginia, primarily because photoperiod influences their circadian rhythm which determines breeding. In both Texas and Virginia, variation in reproductive activity in females is less than in males. In the south of its range, animals may breed less when the summer is at its warmest.[175] The duration of the estrous cycle ranges from 6 to 9 days, with an average of 7.72 days. Estrus occurs again after a litter is born.[74] Copulatory behavior in the marsh rice rat is similar to that in laboratory brown rats. Before mating starts, "the male pursues the running female from behind."[176] The male then repeatedly mounts and dismounts the female; not all mounts result in an ejaculation. Penetrations only last for about 250 ms, but during mating, the penetrations[176] and the intervals between them become longer.[177] Even when a male is satiated after mating, it is able to copulate again when a new female is introduced (the Coolidge effect).[178] Partly because of resistance by the female, the frequency of ejaculation during mating is rather low in marsh rice rats as compared to laboratory rats, hamsters, and deermice.[179]
Age (days) | Body mass (g) | Body mass (oz) |
---|---|---|
10 | 8–17 | 0.3–0.6 |
20 | 18–27 | 0.6–1.0 |
40 | 27–40 | 1.0–1.4 |
60 | 40–60 | 1.4–2.1 |
120 | 50–80 | 1.8–2.8 |
After a gestation of about 25 days, three to five young are usually born, although litter sizes vary from one to seven. Females may have up to six litters a year. Newborns weigh 3 to 4 g (about 0.10 to 0.15 oz) and are blind and almost naked. About as many males as females are born. The external ears (pinnae) soon unfold and on the first day, claws are visible and the young emit high-pitched squeaks. On the second day, they are able to crawl, and during the third to fifth days, the whiskers and eyelids develop. On the two subsequent days, the mammae and incisors become visible and the animals become more active. Between the eighth and 11th days, the eyes open, the fur develops, and the young begin to take solid food. Weaning occurs on the 11th to 20th day, according to different studies. Considerable variation is reported in body masses at different ages, perhaps because of geographic variation. Sexual activity commences when the animals are about 50 to 60 days old.[180] In the wild, rice rats usually live for less than a year;[158] one study suggested that the average lifespan is only seven months.[181]
Human interactions
The marsh rice rat is generally of little importance to humans, which is perhaps why it is not as well studied as some other North American rodents.
The marsh rice rat is the primary host of the
The 2016
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Literature cited
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