Oryzomys couesi

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Oryzomys couesi
Temporal range: Late Pleistocene to Recent

Least Concern  (IUCN 3.1)[1]
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Rodentia
Family: Cricetidae
Subfamily: Sigmodontinae
Genus: Oryzomys
Species:
O. couesi
Binomial name
Oryzomys couesi
(Alston, 1877)[fn 1]
See text.
Distribution of Oryzomys couesi (in red) and other species of Oryzomys.
Synonyms
  • Hesperomys couesi Alston, 1877[6]
  • Oryzomys couesi: Thomas, 1893[7]
  • Oryzomys palustris couesi: Hall, 1960[8]

and see below.

Oryzomys couesi, also known as Coues's rice rat, is a

hantaviruses
.

The species was first described in 1877, the first of many related species from the region described until the 1910s. In 1918,

DNA sequence data, found evidence to recognize separate species from the Pacific and eastern sides of the distribution of O. couesi and two additional species from Panama and Costa Rica
. Generally, Oryzomys couesi is common and of no conservation concern, and it is even considered a plague species in places, but some populations are threatened.

Taxonomy

Oryzomys couesi and at least six more narrowly distributed species with peripheral distributions together form the O. couesi group within the genus

classified as a separate species, O. texensis).[10] 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 forty species from the genus.[11] All are placed in the tribe Oryzomyini ("rice rats"), a diverse assemblage of over a hundred species,[12] and on higher taxonomic levels in the subfamily Sigmodontinae of the family Cricetidae, along with hundreds of other species of mainly small rodents.[13]

History

Hesperomys, and noted similarities to the marsh rice rat (then called Hesperomys palustris) and two species now placed in Tylomys.[14] The specific name, couesi, honors American naturalist Elliott Coues, who had done much work on North American rodents.[6] In 1893, Oldfield Thomas wrote that the species, by then placed in the genus Oryzomys as Oryzomys couesi, had caused much confusion about its identity, because the three specimens (one from Cobán, Guatemala, and two from Mexico) used by Alston in fact belonged to two or three different species. He restricted the name couesi to the animal from Guatemala, and introduced the new name Oryzomys fulgens for one of the Mexican animals.[7] Several other related species were described from the early 1890s onwards[3] and in 1901 Clinton Hart Merriam united many of those into a palustris-mexicanus group of species, which also included the marsh rice rat.[15]

Edward Alphonso Goldman revised North American Oryzomys in 1918 and consolidated many forms into a single species Oryzomys couesi, with ten subspecies distributed from southern Texas and western Mexico south to Costa Rica. He placed it in an Oryzomys palustris group with the marsh rice rat and several species with more limited distributions, which he regarded as related to O. couesi but distinctive enough to be classified as separate species.[16] In the 1930s, a few more forms related to O. couesi were described.[3] As then recognized, the ranges of the marsh rice rat, a United States species, and Oryzomys couesi meet in southern Texas. In 1960, Raymond Hall reviewed specimens from this contact zone and found no grounds on which to separate the two species; thus, he reduced O. couesi to a subspecies of the marsh rice rat.[17] Other workers continued this lumping and by 1971 all other species Goldman had placed in the O. palustris group were classified under the marsh rice rat, together with Oryzomys azuerensis from Panama, described as a species in 1937.[3]

See text.
Distribution of Oryzomys couesi and related species according to Goldman (1918): 1, Oryzomys couesi couesi; 2, O. c. richmondi; 3, O. c. zygomaticus; 4, O. c. mexicanus; 5, O. c. aztecus; 6, O. c. crinitus; 7, O. c. regillus; 8, O. c. albiventer; 9, O. c. peragrus; 10, O. c. aquaticus; 11, O. gatunensis; 12, O. cozumelae; 13, O. antillarum; 14, O. peninsulae; 15, O. nelsoni.[18]

Additional studies of the palustris–couesi contact zone in Texas published in 1979, using more specimens and characters, indicated that the two species are in fact easily distinguishable there; therefore, O. couesi has since been regarded as a species distinct from the marsh rice rat.

Baja California Peninsula and O. albiventer from interior Mexico as species.[20] Still, O. couesi included 22 synonyms,[21] and Carleton and Arroyo-Cabrales wrote that further research on O. couesi and related species would certainly result in the recognition of additional species.[22]

A 2010 study by Delton Hanson and colleagues used DNA sequence data from the

basal to the other specimens, which fell into two large subclades—one containing animals from the Pacific seaboard from western Mexico to El Salvador and the other containing rats from the eastern seaboard from Texas to Nicaragua. The Panamanian and Costa Rican populations differed by 6.53% to 11.93% from the others and the western and eastern subclades differed by 4.41% on average.[24] Data from both of the slower-evolving nuclear markers Rbp3 and Adh1-I2 also placed examples of Oryzomys in two main clades, but did not recover the western and eastern groups of O. couesi as separate clades. In addition, Adh1-I2 placed the Costa Rican population within the marsh rice rat clade and placed some western O. couesi specimens closer to the marsh rice rat than to the O. couesi group.[26] The combined dataset supported the western and eastern clades within O. couesi and placed the Costa Rican population marginally closer to the marsh rice rat than to O. couesi.[27] Using the genetic species concept, the authors suggested that the four groups they found within O. couesi should be recognized as distinct species. If this suggestion is followed, the eastern subclade would retain the name Oryzomys couesi, the western group would be named Oryzomys mexicanus, and the appropriate names for the Panamanian and Costa Rican species remain unclear.[28]

Western Mexico to El Salvador

Taxonomic synonyms

Populations of Oryzomys couesi from

hantavirus, and according to Merriam (1901) have more robust skulls, with larger molars, stronger zygomatic arches (cheekbones), and better developed ridges along the margins of the interorbital region of the skull (between the eyes).[36] Within the "Oryzomys mexicanus" clade, Cytb sequence differences average 2.06% and western (Jalisco to Oaxaca) and eastern (Chiapas and El Salvador) groups form distinct subclades; Hanson and colleagues recognized these as different subspecies, mexicanus in the west and zygomaticus in the east.[37]

As defined by Carleton and Arroyo-Cabrales in 2009, the subspecies Oryzomys couesi mexicanus occurs along the Pacific coast from central Sonora to southeastern

Osgoodomys banderanus and has been recognized as a distinct biogeographic zone in some reviews.[39] O. c. mexicanus occurs close to three other Oryzomys species—O. albiventer, O. peninsulae, and O. nelsoni—which are larger and different in some proportions and details of coloration.[40]

nominate couesi, but usually with paler fur; the upperparts are more buffy than in couesi and the underparts are usually white, but may be buffy, the normal color in couesi.[43]

Oryzomys zygomaticus was first described by Merriam in 1901 as a separate species[33] similar to mexicanus, but with the zygomatic arches broadly spreading and curved downward.[44] Goldman, who reduced it to a subspecies of couesi, recorded it from southwestern Guatemala and nearby Chiapas and described it as slightly paler than O. c. couesi but darker than O. c. mexicanus.[45] Three specimens from central El Salvador have Cytb sequences similar to those of zygomaticus,[46] but in The Mammals of El Salvador (1961), Burt and Stirton recorded only the subspecies couesi from the country, while noting that specimens from some localities were slightly paler than others.[47]

Interior Mexico

Taxonomic synonyms

Goldman grouped four subspecies of couesi from the interior plateaus of central Mexico together—albiventer, crinitus, aztecus, and regillus.[51] Three of those (albiventer from Jalisco,[52] crinitus from the Distrito Federal,[49] and aztecus from Morelos)[48] were described by Merriam in 1901, and Goldman had himself described regillus from Michoacán in 1915.[50] According to Goldman, aztecus is pale and large-toothed,[53] crinitus is large, dark and large-toothed,[54] regillus is large and dark,[51] and albiventer is large and relatively pale.[55]

Skull, seen from above
Skull, seen from below
Skull of Oryzomys from Los Reyes, Michoacán (regillus Goldman, 1915)[56]

In their 2009 review of western Mexican Oryzomys, Carleton and Arroyo-Cabrales classified

Baja California Peninsula. They suggested that regillus and aztecus may represent no more than robust upland populations of mexicanus, but could not exclude the possibility that they represent a different species. That crinitus, which occurs at over 2,000 m (6,600 ft) altitude in the Valley of Mexico, was the same species as peninsulae from the lowlands of the Baja California Peninsula they could not accept and they recommended further research to determine the relationships of crinitus.[58] A specimen from inland Michoacán has Cytb data characteristic of mexicanus,[59] but Hanson and colleagues did not have data for other interior Mexican Oryzomys.[60]

The holotype of the species Oryzomys fulgens, which Thomas had described in 1893, has no more precise locality than "Mexico", but the Valley of Mexico has been suggested as its origin.[61] It is a large, coarse-furred, bright reddish,[7] long-tailed species with a broad skull with widely spreading zygomatic arches.[62] Goldman wrote that it was similar to crinitus, but more intensely colored, and differed in the form of the interorbital region; he retained it as a separate species pending further investigations.[63] Carleton and Arroyo-Cabrales noted that archival research may yet uncover the precise origin of O. fulgens, which could establish it as an older name for one of the other central Mexican Oryzomys.[64]

Texas to Nicaragua

Taxonomic synonyms

Oryzomys populations from Texas to Nicaragua form a single Cytb clade, within which the average sequence divergence is 1.28%,[76] and Hanson and colleagues proposed that the name Oryzomys couesi be restricted to this clade.[28] These populations correspond to two subspecies recognized by Goldman (O. c. aquaticus and O. c. couesi) and an island form he retained as a species (O. cozumelae). Two other subspecies Goldman recognized, O. c. richmondi and O. c. peragrus, and a third, O. c. pinicola, that was described after Goldman's paper occur in the same region, but have not been studied genetically.[77]

Skull, seen from above
Skull, seen from below
Skull of Oryzomys from Brownsville, Texas (aquaticus Allen, 1891)[56]

The northernmost populations of Oryzomys couesi, those in southernmost Texas and nearby

sympatric (occur in the same places). In the contact zone, couesi occurs further inland, while the marsh rice rat lives along the coast.[80] In experimental conditions, the two fail to interbreed[81] and genetic analysis yields no evidence of gene flow or hybridization in the wild.[82] Compared to populations further to the south, aquaticus is larger and paler and has a more robust skull.[83] Specimens from Tamaulipas are slightly darker than those from Texas.[84] The Cytb sequences of specimens of aquaticus form a separate group, but cluster among specimens of O. c. couesi from further south.[85]

The form peragrus is known from further south in Mexico, in the

San Luis Potosi, the state of Hidalgo, and far northern Veracruz.[86] Late Pleistocene fossils of this form have been found in Cueva de Abre, Tamaulipas.[87] According to Goldman, it is intermediate in color between O. c. aquaticus and O. c. couesi, but has a skull similar to that of aquaticus.[88]

Goldman united populations ranging from northern Veracruz through eastern Mexico, Guatemala, Honduras, and Nicaragua south to far northwestern Costa Rica in the

type locality in Guatemala—Oryzomys jalapae Allen and Chapman, 1897, from Veracruz; Oryzomys jalapae rufinus Merriam, 1901, from Veracruz; Oryzomys teapensis Merriam, 1901, from Tabasco; Oryzomys goldmani Merriam, 1901, from Veracruz; Oryzomys jalapae apatelius Eliot, 1904, from Veracruz; and Oryzomys richardsoni Allen, 1910, from Nicaragua.[89] According to Goldman, individual variation within the subspecies is large, which has led to the large number of published synonyms, but populations from all parts of its range are essentially similar.[90]

Skull, seen from above
Skull, seen from below
Skull of Oryzomys from Cozumel, Mexico (cozumelae Merriam, 1901)[56]

The subspecies Oryzomys couesi pinicola was described in 1932 from a pine ridge in western British Honduras (now Belize); it is smaller and darker than nominate couesi, which also occurs in Belize, and has a more delicate skull.[91] In 1901, Merriam described the Oryzomys of the island of Cozumel as a separate species, Oryzomys cozumelae, and Goldman kept it as such because of its large size, dark fur, and long tail.[92] In 1965, however, Knox Jones and Timothy Lawlor judged the differences between cozumelae and mainland couesi trivial and found that cozumelae was inside the range of variation of mainland Oryzomys populations; accordingly, they demoted the island form to a subspecies.[93] Mark Engstrom and colleagues, writing in 1989, reaffirmed this conclusion.[94] For an island form, this population is highly genetically variable.[95] In its Cytb sequence data, it falls among populations of nominate couesi.[60] Oryzomys couesi is also found on Turneffe Atoll off the coast of Belize[96] and Roatán off Honduras.[97]

The Oryzomys of the eastern lowlands of Nicaragua was described as a separate species, Oryzomys richmondi, by Merriam in 1901, and Goldman retained it as a subspecies of O. couesi on the basis of its distinctly dark fur.[98] In reviewing Nicaraguan Oryzomys in 1986, Jones and Engstrom did not keep richmondi as separate, because they thought the difference in color too small for the recognition of subspecies.[99] Oryzomys dimidiatus, a small, dark Oryzomys with gray underparts, occurs with O. couesi in southeastern Nicaragua.[100] According to Jones and Engstrom, rice rats from the island of Ometepe in Lake Nicaragua are distinctive in their large skull and small external measurements, with an especially short tail, soft fur that is orange-brown above and buffish below, and lack of sphenopalatine vacuities (openings in the roof of the mesopterygoid fossa, the gap behind the end of the bony palate). They considered that this population probably represented a separate subspecies, but declined to propose a new name because they had only one adult specimen.[101] In Nicaragua, O. couesi occurs up to an altitude of 1,250 m (4,100 ft).[102]

Costa Rica, Panama, and Colombia

Taxonomic synonyms

Oryzomys from Costa Rica have historically been referred to O. c. couesi,[105] but Hanson and colleagues found that two specimens from Refugio Nacional de Vida Silvestre Mixto Maquenque, northeastern Costa Rica, differed as much from other O. couesi (11.93% Cytb sequence divergence) as O. couesi differed from the marsh rice rat (11.30%). They suggested that these animals represented a species distinct from O. couesi, but were unable to resolve the correct name for the species because they could not examine samples of dimidiatus or richmondi.[106]

Skull, seen from above
Skull, seen from below
Skull of Oryzomys from Gatún, Panama (gatunensis Goldman, 1912)[56]

Oryzomys is rare in Panama.

braincase), and the long nasal bones.[108] In 1937, Bole described another species of Panamanian Oryzomys, Oryzomys azuerensis from Paracoté, Veraguas Province.[103] It is a brown form, lacking the reddish tones of nearby populations, and has a broad skull with a short rostrum (front part) and ridges on the interorbital region like those of gatunensis.[109] Although Goldman recommended to him that gatunensis and azuerensis both be treated as subspecies of couesi, Bole described azuerensis as a species because it did not seem intermediate between the geographically closest forms, gatunensis and couesi, and was separated by a large gap from the nearest known populations of O. couesi in northwestern Costa Rica and southeastern Nicaragua.[110] In a 1966 review of Panamanian mammals, Charles Handley reduced both gatunensis and azuerensis to subspecies of the marsh rice rat (in which O. couesi was included at the time),[111] and when O. couesi was reinstated as a separate species these forms went with it.[3] Specimens from near the type locality of azuerensis differ by about 7% in their Cytb sequences from other O. couesi, which suggests that they may represent a separate species. However, Hanson and colleagues did not reinstate azuerensis as a species, because they could not examine samples of gatunensis.[36]

Oryzomys couesi was first reported from Colombia in 1987, when

Chocó in western Colombia.[113]

Common names

Several common names have been proposed for Oryzomys couesi and the synonyms currently associated with it. Eliot in 1905[114] and Goldman in 1918 gave separate common names for each of the species and subspecies they recognized.[115] Many authors have used "Coues' Rice Rat" or some variation thereof for O. couesi,[116] but "Coues' Oryzomys" has also been used.[3]

Description

Measurements of different populations of Oryzomys couesi
Population n[fn 3] Total length Tail Hindfoot
Western Mexico to El Salvador
San José de Guaymas, Sonora (lambi)[30][fn 4] 4 227 113 29
Escuinapa, Sinaloa (mexicanus)[43] 10 251.4 (239–273) 137.4 (127–165) 28.9 (27–35)
Jalisco (mexicanus)[117] 58 245.7 (195–301) 132.3 (102–160) 30.2 (26–34)
Nayarit (mexicanus)[117] 62 244.8 (210–288) 125.1 (105–150) 30.5 (27–33)
Nenton, Guatemala (zygomaticus)[45] 1 290 152 33
El Salvador[118] 87 190–304 109–194 25–33
Interior Mexico
Morelos (aztecus)[53] 3 307 (297–318) 161 (154–170) 34 (33–35)
Mexico City (crinitus)[54] 2 307, 280 161, 148 37, 35
Michoacán (regillus)[51] 3 308 (285–320) 168 (155–180) 35 (34–36)
Unknown (fulgens)[119] 1 311 151 37.5[fn 5]
Texas to Nicaragua
Brownsville, Texas (aquaticus)[83] 5 297 (283–310) 161 (138–180) 34.5 (32–38)
Rio Verde, San Luis Potosí (peragrus)[88] 3 281 (265–294) 157 (143–167) 34 (33–35)
Orizaba, Veracruz (couesi)[120] 7 263 (248–294) 148 (139–174) 33.1 (32–34.5)
Tumbala, Chiapas (couesi)[121] 4 252 (242–265) 130 (127–135) 30.7 (30–31)
El Cayo, Belize (pinicola)[122] 18 108 (96–128)[fn 6] 122 (107–146) 27 (24.6–29)
Cozumel (cozumelae)[92] 6 306 (285–327) 172 (163–177) 34.3 (33–35.5)
Yaruca, Honduras (couesi)[121] 10 267.5 (255–280) 138 (130–145) 29.1 (28–32)
San Antonio, Nicaragua (couesi)[123] 25 264.9 (242–292) 135.6 (127–150) 28.8 (27–31)
Southeastern Nicaragua (richmondi)[98] 10 275.8 (255–295) 137 (124–151) 30.9 (29–33.5)
Isla de Omotepe, Nicaragua[123] 1 260 121 30
Costa Rica, Panama, and Colombia
Azuero, Panama (azuerensis)[109] 1 203 107.5 30
Gatún, Panama (gatunensis)[124] 1 224 115 31.5
Measurements are in millimeters and are in the form "average (minimum–maximum)", except where there are only one or two measured specimens.

Oryzomys couesi is a medium-sized to large rat

mammae.[131] Head and body length is 98 to 142 mm (3.9 to 5.6 in), tail length is 107 to 152 mm (4.2 to 6.0 in), hindfoot length is 27 to 33 mm (1.1 to 1.3 in), ear length is 13 to 18 mm (0.51 to 0.71 in), and body mass is 43 to 82 g (1.5 to 2.9 oz).[127] Studies in Texas and El Salvador found that males are slightly larger than females.[132]

The

acrocentric, with a long and a short arm, or subtelocentric, with a long and a vestigial arm.[25] The form of the sex chromosomes has been used to distinguish the marsh rice rat from Oryzomys couesi, but there are no consistent differences between the two.[136]

As is characteristic of Sigmodontinae, Oryzomys couesi has a complex penis, with the baculum (penis bone) ending in three cartilaginous digits at its tip.[137] The outer surface of the penis is mostly covered by small spines, but there is a broad band of nonspinous tissue.[138] The papilla (nipple-like projection) on the dorsal (upper) side of the penis is covered with small spines, a character Oryzomys couesi shares only with Oligoryzomys and the marsh rice rat among oryzomyines examined.[139] On the urethral process, located in the crater at the end of the penis,[140] a fleshy process (the subapical lobule) is present; it is absent in all other oryzomyines with studied penes except the marsh rice rat and Holochilus brasiliensis.[141]

Skull

Skull, seen from above
Skull, seen from below
Skull of Oryzomys from Yaruca, Honduras (couesi Alston, 1877)[56]

The nasal and

maxillary bones meet.[142] The zygomatic plate, the flattened front part of the zygomatic arch, is broad and develops a notch at its front end.[143] The plate's back margin is located before the first upper molar.[144] The jugal bone, part of the zygomatic arch, is reduced, as usual in oryzomyines.[145] The sphenopalatine foramen, a foramen (opening) at the side of the skull above the molars, is small; it is much larger in the marsh rice rat.[146] The narrowest part of the interorbital region is towards the front and the edges are lined by prominent shelves.[147] The parietal bones extend to the sides of the braincase.[148] The interparietal bone is narrow and wedge-shaped, so that the parietal and squamosal bones meet extensively.[149]

The

mastoid bone.[157]

In the mandible (lower jaw), the mental foramen, an opening just before the first molar, opens sidewards, not upwards as in a few other oryzomyines.[158] 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.[159] The capsular process, a raising of the bone of the back of the mandible that houses the back end of the incisor, is large.[160]

Teeth

The

mesolophid on the lower molars, are present, another trait O. couesi shares with most but not all other oryzomyines.[164] The flexi and flexids (valleys between the cusps and crests) at the labial (outer) side of the molars are closed by cingula (ridges).[165]

On the first and second upper molars, the flexi do not extend to the midline of the molars.

anteroloph, is present behind the labial cuspule.[166] As in most oryzomyines, the upper molars all have one root on the inner (lingual) side and two on the outer (labial) side; in addition, the first upper molar usually has another small labial root.[162]

On the first lower molar, the labial and lingual conules of the

anterolabial cingulum) at the outer front (anterolabial) edge of the molar, before the protoconid.[167] The third lower molar also bears an anterolophid and an anterolabial cingulum.[169] The first lower molar has large roots at the front and back of the tooth and two smaller ones in between, at the labial and lingual side. The second and third lowers molars have two large roots, one at the front and one at the back.[162]

Postcranial skeleton

As usual in oryzomyines, there are twelve ribs. The first rib

hemal arches (small bones) are present with a spinous back border.[172] The entepicondylar foramen is absent, as in all members of the Sigmodontinae; if present, as in some other rodents, this foramen perforates the distal (far) end of the humerus (upper arm bone).[173]

Ecology and behavior

The distribution of Oryzomys couesi extends from southern Texas and central Sonora, but not the central plateau of Mexico, through Central America south and east to northwestern Colombia;[3] see under "Taxonomy" for details. The species has also been found in late Pleistocene cave deposits in Mexico and Honduras.[174] It is common in watery habitats, such as marshes and small streams, but also occurs in forests and shrublands with sufficient cover.[175] In addition, it is found in sugarcane and rice fields.[107] In Texas, it occurs in marsh vegetation along resacas (oxbow lakes)[176] and in Veracruz, it has even been found on the dry coastal plain among shrubs.[177] It occurs from 2,300 m (7,500 ft) altitude down to sea level.[125] On Cozumel, the proportion of juveniles and females is higher near roads that function as habitat edges.[178] Cozumel rice rats rarely cross roads, which may isolate subpopulations on the island.[179]

Oryzomys couesi lives on the ground and is

active during the night.[177] Oryzomys couesi builds globular nests of woven vegetation suspended among reeds, about 1 m (3.3 ft) above the water or the ground;[183] in Texas, larger individuals make larger nests.[184] It does not usually make its own runways in vegetation, but may use those of other rodents, such as cotton rats.[185]

Population densities range from 5 to 30 per ha (2 to 12 per acre).[1] On Cozumel, density is around 14.5 to 16.5 per ha (5.9 to 6.7 per acre), but shows large seasonal variation.[186] In western Mexico, one study found densities of 3 per ha (1.2 per acre) in cloud forest and 1 per ha (0.4 per acre) in a disturbed area.[187] In 24 hours, male Texas O. couesi move up to 153 m (502 ft) and females up to 126 m (413 ft).[188] The diet includes both plant material, including seeds and green parts, and animals, including small fish, crustaceans, snails, insects like ants and beetles, and other invertebrates.[189] It probably breeds around the year and after a pregnancy of 21 to 28 days,[125] the female produces litters of two to seven young, with an average of 3.8, according to Reid's Mammals of Central America & Southeast Mexico.[107] In 28 pregnant females from Nicaragua, litter size varied from one to eight, averaging 4.4.[123] The young become reproductively active when seven weeks old and the life cycle is short.[125]

The

Bayou virus infecting the marsh rice rat, a common cause of hantavirus infections in the United States. No hantavirus infections in humans have been linked to O. couesi hantaviruses, however.[196] Chiapas O. couesi easily survive experimental infection with several arboviruses, including the Venezuelan equine encephalitis virus, suggesting that the species may serve as a reservoir for that virus.[197]

Conservation status

The

Least Concern", because it is a widely distributed, common species with broad habitat tolerance that occurs in many protected areas. Habitat destruction, such as drainage of wetlands, may threaten some populations.[1] In many areas, it is so common that it is considered a plague species.[125] Populations even persist in the Valley of Mexico, as evidenced by a photograph published in 2006.[198] However, the species is listed as threatened in Texas, where its distribution is very limited, because of habitat loss.[199] In 1979, Benson and Gehlbach estimated the size of the Texas population to be about 15,000.[200] A 2001 study predicted that climate change would drive the Texas population to extinction, because no suitable habitats would continue to exist.[201] The Cozumel population has declined substantially since the mid-1980s, perhaps due to habitat disturbance and predation by introduced species.[186]

Footnotes

  1. Proceedings of the Zoological Society of London, but the parts of this journal were not always published in the volume year.[4] A 2005 review confirms that p. 756 of the 1876 volume was published in 1877.[5]
  2. ^ Merriam described Oryzomys cozumelae as new in both of his 1901 papers,[68] but his Proceedings of the Biological Society of Washington article was published earlier (July 19) than the one in the Proceedings of the Washington Academy of Sciences (July 26); thus, the former is the original publication of this name.[69]
  3. ^ Number of specimens measured.
  4. ^ Only averages available.
  5. ^ With claws.
  6. ^ Head and body length instead of total length.

References

  1. ^ a b c Linzey et al., 2016
  2. ^ Hanson et al., 2010, p. 336
  3. ^ a b c d e f g h Musser and Carleton, 2005, p. 1147
  4. ^ Dickinson, 2005, p. 427
  5. ^ Dickinson, 2005, table 1
  6. ^ a b c d Alston, 1877, p. 756
  7. ^ a b c d Thomas, 1893, p. 403
  8. ^ Hall, 1960, p. 173
  9. ^ a b Carleton and Arroyo-Cabrales, 2009, p. 117
  10. ^ Hanson et al., 2010, p. 342
  11. ^ Weksler et al., 2006, table 1
  12. ^ Weksler, 2006, p. 3
  13. ^ Musser and Carleton, 2005
  14. ^ Alston, 1877, pp. 756–757; Musser and Carleton, 2005, p. 1189, for placement in Tylomys
  15. ^ Merriam, 1901b, p. 275
  16. ^ Goldman, 1918, pp. 16, 28–29
  17. ^ Hall, 1960, pp. 172–173
  18. ^ Goldman, 1918, fig. 3
  19. ^ Musser and Carleton, 2005, p. 1152; Weksler et al., 2006, table 1, footnote e
  20. ^ Carleton and Arroyo-Cabrales, 2009, p. 94
  21. ^ Carleton and Arroyo-Cabrales, 2009, p. 116
  22. ^ Carleton and Arroyo-Cabrales, 2009, p. 107
  23. ^ Hanson et al., 2010, p. 337
  24. ^ a b Hanson et al., 2010, figs. 1–2, table 1
  25. ^ a b Hanson et al., 2010, p. 341
  26. ^ Hanson et al., 2010, figs. 1, 3–4
  27. ^ Hanson et al., 2010, fig. 5
  28. ^ a b Hanson et al., 2010, pp. 342–343
  29. ^ Allen, 1897, p. 53
  30. ^ a b c Burt, 1934, p. 107
  31. ^ Allen, 1897, p. 52
  32. ^ Merriam, 1901b, p. 287
  33. ^ a b c Merriam, 1901b, p. 285
  34. ^ Hanson et al., 2010, fig. 2, p. 343, appendix I
  35. ^ Hanson et al., 2010, table 1
  36. ^ a b Hanson et al., 2010, p. 343
  37. ^ Hanson et al., 2010, fig. 2, table 1, appendix I
  38. ^ Carleton and Arroyo-Cabrales, 2009, p. 119
  39. ^ a b Carleton and Arroyo-Cabrales, 2009, p. 120
  40. ^ Carleton and Arroyo-Cabrales, 2009, pp. 118, 121–122
  41. ^ Carleton and Arroyo-Cabrales, 2009, pp. 118–120; Goldman, 1918, p. 34
  42. ^ Burt, 1934, p. 108
  43. ^ a b Goldman, 1918, p. 34
  44. ^ a b Merriam, 1901b, p. 286
  45. ^ a b Goldman, 1918, p. 33
  46. ^ Hanson et al., 2010, figs. 1–2, appendix I
  47. ^ Burt and Stirton, 1961, p. 61
  48. ^ a b Merriam, 1901b, p. 282
  49. ^ a b Merriam, 1901b, p. 281
  50. ^ a b Goldman, 1915, p. 129
  51. ^ a b c Goldman, 1918, p. 37
  52. ^ Merriam, 1901b, p. 279
  53. ^ a b Goldman, 1918, p. 35
  54. ^ a b Goldman, 1918, p. 36
  55. ^ Goldman, 1918, p. 38
  56. ^ a b c d e Goldman, 1918, plate I
  57. ^ Carleton and Arroyo-Cabrales, 2009, p. 118
  58. ^ Carleton and Arroyo-Cabrales, 2009, p. 113
  59. ^ Hanson et al., 2010, fig. 1, appendix I
  60. ^ a b Hanson et al., 2010, fig. 1
  61. ^ Carleton and Arroyo-Cabrales, 2009, pp. 113–114
  62. ^ Thomas, 1893, p. 404
  63. ^ Goldman, 1918, p. 282
  64. ^ Carleton and Arroyo-Cabrales, 2009, p. 114
  65. ^ Eliot, 1904, p. 266
  66. ^ Allen, 1891, p. 289
  67. ^ Merriam, 1901a, p. 103
  68. ^ Merriam, 1901a, p. 103; Merriam, 1901b, p. 280
  69. ^ Poole and Schantz, 1942, p. 306
  70. ^ Merriam, 1901b, p. 288
  71. ^ Allen and Chapman, 1897, p. 206
  72. ^ Merriam, 1901b, p. 283
  73. ^ Murie, 1932, p. 1
  74. ^ Allen, 1910, p. 99
  75. ^ Merriam, 1901b, p. 284
  76. ^ Hanson et al., 2010, fig. 1, table 1
  77. ^ Goldman, 1918, fig. 3; Murie, 1932, p. 1; Handon et al., 2010, fig. 1
  78. ^ Goldman, 1918, pp. 39–40; Hanson et al., 2010, appendix A
  79. ^ Schmidt and Engstrom, 1994, p. 914
  80. ^ Schmidt and Engstrom, 1994, pp. 916–917
  81. ^ Schmidt and Engstrom, 1994, pp. 915–916
  82. ^ Schmidt and Engstrom, 1994, p. 920
  83. ^ a b Goldman, 1918, p. 40
  84. ^ Baker, 1951, p. 215
  85. ^ Hanson et al., 2010, fig. 2
  86. ^ Goldman, 1918, p. 39; Jones et al., 1983, p. 379; Hall and Dalquest, 1963, p. 289
  87. ^ Dalquest and Roth, 1970, pp. 220, 226
  88. ^ a b Goldman, 1918, p. 39
  89. ^ Goldman, 1918, pp. 29–30
  90. ^ Goldman, 1918, p. 31
  91. ^ Murie, 1932, pp. 1–2; Murie, 1935, p. 26
  92. ^ a b Goldman, 1918, p. 43
  93. ^ Jones and Lawlor, 1965, p. 413
  94. ^ Schmidt and Engstrom, 1989, p. 414
  95. ^ Vega et al., 2004, p. 210
  96. ^ Platt et al., 2000, p. 167
  97. ^ Koopman, 1959, p. 237
  98. ^ a b Goldman, 1918, p. 32
  99. ^ Jones and Engstrom, 1986, p. 10
  100. ^ Genoways and Jones, 1971, p. 834
  101. ^ Jones and Engstrom, 1986, pp. 10, 12
  102. ^ Jones and Engstrom, 1986, p. 7
  103. ^ a b Bole, 1937, p. 165
  104. ^ a b Goldman, 1912, p. 7
  105. ^ Goldman, 1918, p. 31; Harris, 1943, p. 12
  106. ^ Hanson et al., 2010, p. 343, appendix A
  107. ^ a b c d e Reid, 2009, p. 207
  108. ^ Goldman, 1918, pp. 42–43
  109. ^ a b Bole, 1937, p. 166
  110. ^ Bole, 1937, p. 167
  111. ^ Handley, 1966, p. 781
  112. ^ Hershkovitz, 1987, p. 152
  113. ^ Hershkovitz, 1987, p. 154
  114. ^ Eliot, 1905, pp. 182–186
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  116. ^ Eliot, 1905, p. 186; Goldman, 1918, p. 29; Linzey et al., 2008; Duff and Lawson, 2004, p. 54
  117. ^ a b Carleton and Arroyo-Cabrales, 2009, table 2
  118. ^ Burt and Stirton, 1961, p. 60
  119. ^ Goldman, 1918, p. 41
  120. ^ Goldman, 1918, pp. 30–31
  121. ^ a b Goldman, 1918, p. 30
  122. ^ Murie, 1932, table I
  123. ^ a b c Jones and Engstrom, 1986, p. 12
  124. ^ Goldman, 1918, p. 42
  125. ^ a b c d e f Medellín and Medellín, 2006, p. 710
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  127. ^ a b Reid, 2009, p. 206
  128. ^ Weksler, 2006, pp. 19, 23, table 5
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  130. ^ Weksler, 2006, pp. 79, 81
  131. ^ Weksler, 2006, p. 17, table 5
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  133. ^ Weksler, 2006, p. 59
  134. ^ Weksler, 2006, pp. 58–59
  135. ^ Haiduk et al., 1979, p. 612
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  139. ^ Hooper and Musser, 1964, p. 13; Weksler, 2006, p. 57
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  141. ^ Weksler, 2006, p. 57
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  169. ^ Weksler, 2006, p. 53
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  201. ^ Cameron and Scheel, 2001, p. 676

Literature cited

External links

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