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List of rodents of the Caribbean

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Desmarest's hutia (Capromys pilorides), a member of a rodent family known only from the Caribbean.
Desmarest's hutia (Capromys pilorides), a member of a rodent family known only from the Caribbean.

The Caribbean region is home to a diverse and largely endemic rodent fauna. This includes the endemic family Capromyidae (hutias), which are largely limited to the Greater Antilles, and two other groups of endemic hystricognaths, the heteropsomyines and giant hutias, including the extinct bear-sized[1] Amblyrhiza inundata. Lesser Antillean rodent faunas mostly consist of oryzomyines, members of a distantly related group of rodents, and include two of the largest known oryzomyines, Megalomys desmarestii and "Oryzomys hypenemus". Various other rodents are limited to land-bridge islands such as Trinidad, which were connected to the mainland during glacial-period lowered sea levels in the Pleistocene, or to smaller portions of the Caribbean archipelago. Much of the native rodent fauna of the Caribbean are extinct because of human influences, particularly following the introduction of invasive species such as Old World rats.

For the purposes of this article, the "Caribbean" includes all islands in the Caribbean Sea (except for small islets close to the mainland) and the Bahamas, Turks and Caicos Islands, and Barbados, which are not in the Caribbean Sea but biogeographically belong to the same Caribbean bioregion.

Overview

The Caribbean rodent fauna is dominated by oryzomyines (rice rats), members of the family Cricetidae, and by three groups of hystricognaths: hutias (family Capromyidae), giant hutias (taxonomy unclear), and heteropsomyines (subfamily Heteropsomyinae), which are part of the spiny rat family (Echimyidae). Many other rodent groups are known from the Caribbean, but these are either introduced or limited to islands outside the limits of the West Indian faunal region as defined by Koopman.[2]

So far, no rodents have been recorded from Navassa Island, the Turks and Caicos Islands, the British Virgin Islands, Saint Barthélemy, Redonda, Saba, the federal dependencies of Venezuela, or the Insular Region of Colombia.

Oryzomyines are a diverse group, consisting over a hundred species found from the eastern United States south to Tierra del Fuego, including also the Galápagos Islands. In the Caribbean, they are found on many islands close to the mainland, from the Florida Keys to Tobago, but also on Jamaica and throughout the Lesser Antilles north to Anguilla. Their invasion of the Lesser Antilles must have been relatively recent, as sigmodontine rodents only reached South America from North America about 5 million years ago.[3] In the Lesser Antilles, most species have not been named or described, and the true diversity of the group in that region remains unclear. All endemic Caribbean oryzomyines are now extinct, though several island populations of mainland species survive.[4]

Hutias are the most diverse endemic Caribbean hystricognath group and the only one that still survives, although many species are extinct. Most species are known from Cuba and Hispaniola, but hutias are also indigenous to Jamaica and some smaller islands and they have been introduced to several other parts of the Caribbean. The hutia family is divided into four subfamilies, of which one or two are now extinct.[5] They have been present in the Caribbean for a considerable span of time, as documented by the find of a hutia, Zazamys, in early Miocene sediments on Cuba.[6] This represents more than half the age (~32 million years) of the earliest known hystricognath rodent in South America (a continent hystricognaths colonized by rafting across the Atlantic from their ancestral home in Africa).[7]

Giant hutias are a heterogeneous group of hystricognaths that include not only truly giant forms like Amblyrhiza, but also the small Xaymaca. All are now extinct. Some have previously been included in a family Heptaxodontidae, but there is little evidence that the group is truly natural. Giant hutias are known from Jamaica (Clidomys, Xaymaca, and possibly another, very large form), Hispaniola (Quemisia), Puerto Rico (Elasmodontomys, Tainotherium), and Anguilla and Saint Martin in the northern Lesser Antilles (Amblyrhiza).[8]

Heteropsomyines form a distinct subfamily of the widespread Neotropic family of spiny rats, which is known only from Cuba, Hispaniola, and Puerto Rico. On each island, a distinct genus with two species has been found, but all are now extinct. A fourth genus, Puertoricomys, is known from an older, perhaps Pliocene, deposit on Puerto Rico.[9] These animals exhibited characters intermediate between hutias and spiny rats and have sometimes been treated as a subfamily of the Capromyidae.[10]

Among introduced rodents, the house mouse (Mus musculus), brown rat (Rattus norvegicus), and black rat (Rattus rattus) are most common; these three species, originally from the Old World, have been introduced to all parts of the world and occur throughout the Caribbean.[11] They are not mentioned specifically in the faunal lists in this article.

Agoutis, members of a mainland American hystricognath genus that is also indigenous to some Caribbean islands, have also been widely introduced, resulting, among others, in the presence of the red-rumped agouti (Dasyprocta leporina) on most of the Lesser Antilles. Several other hystricognaths have also been introduced in parts of the Caribbean.[5]

The genera of Caribbean rodents are classified as follows:[12]

Greater Antilles

The four islands of the Greater Antilles, Cuba, Hispaniola, Jamaica, and Puerto Rico, are home to a diverse indigenous hystricognath fauna. Jamaica also harbors an indigenous oryzomyine and various introduced rodents now occur on all islands.

Cuba

Desmarest's hutia is the most common species of hutia on Cuba.

Cuba, the largest of the Antilles, and its surrounding islands, of which the Isla de la Juventud is the most significant, harbor a diverse hutia fauna, including many species with very limited distributions. Various other hutias and an endemic genus of heteropsomyine became extinct during the Quaternary and the oldest known hutia, Zazamys, has been found in Miocene Cuban deposits.

Jamaica

The rodent fauna of Jamaica is relatively poor at the species level, including only five indigenous species, but diverse at higher taxonomic levels, including the only oryzomyine of the Greater Antilles and several distantly related hystricognaths.

  • Clidomys osborni, an extinct giant hutia from Jamaica. Numerous morphological variants have been described under separate names, but these are now all recognized as synonyms.[25]
  • Geocapromys brownii, a hutia known only from the island.[22]
  • A femur of a large rodent, somewhat similar to that of Amblyrhiza, provides evidence for the possible occurrence of a large hystricognath, perhaps a heptaxodontine, on Jamaica.[26]
  • Oryzomys antillarum, a medium-sized oryzomyine closely related to extant Central American Oryzomys which probably became extinct around 1880. Its taxonomic status as either a subspecies of the mainland species Oryzomys couesi or Oryzomys palustris or as a separate species has been disputed, but it is now recognized as a separate species.[27] It is also found in late Pleistocene deposits.[28]
  • Xaymaca fulvopulvis, a small hystricognath that may be most closely related to the giant hutias of Hispaniola, Puerto Rico, and the Leeward Islands.[26]

Hispaniola

Hispaniola, the second largest of the Antilles, is politically divided into Haiti and the Dominican Republic. Various hutias and other hystricognaths are known from both the main island and several surrounding islands, including Gonâve Island.

Puerto Rico

Several hystricognaths are known from Puerto Rico, the easternmost of the Greater Antilles, which is under United States sovereignty. Hutias are notably absent from the indigenous fauna, but giant hutias and heteropsomyines are present.

Leeward Islands

The Leeward Islands, the northern segment of the Lesser Antilles, have various indigenous oryzomyines and a single indigenous hystricognath in addition to a series of introduced rodents.

U.S. Virgin Islands

The red-rumped agouti has been introduced to the U.S. Virgin Islands as well as many other islands in the Lesser Antilles

The United States Virgin Islands are a group of islands east of Puerto Rico, centered around the three main islands of Saint Thomas, Saint John and Saint Croix. No native rodents are known, but several species have been introduced.

Anguilla

Anguilla is a British island with some native rodents, including the giant Amblyrhiza.

  • Amblyrhiza inundata, an extremely large giant hutia also known from nearby Saint Martin. It is known from Quaternary fossils but may not have survived until its first contact with humans.[37]
  • Both Mus musculus and Rattus have been recorded from archeological sites.[38]
  • An oryzomyine, similar in size to "Undescribed species A" from Montserrat, has been recorded from several Amerindian archeological sites on Anguilla.[39]

Saint Martin

The island of Saint Martin is divided into a French and a Dutch part. Its rodent fauna is similar to that of nearby Anguilla.

  • Amblyrhiza inundata, a giant hutia also known from Anguilla (see there).[37]
  • An oryzomyine has been recorded at an Amerindian site at Hope Estate on the French side of the island which dates from about 500 BCE.[40]

Sint Eustatius

Sint Eustatius is a small island near Saint Kitts that is part of the Netherlands.

Saint Kitts

Saint Kitts forms the nation of Saint Kitts and Nevis together with nearby Nevis.

  • Dasyprocta leporina, an introduced northern South American agouti. The subspecies on Saint Kitts is D. l. noblei.[43] This agouti has also been round in an archeological site.[44]
  • Pennatomys nivalis, an extinct oryzomyine, is known from archeological sites.[42]

Nevis

Nevis is the second main island of Saint Kitts and Nevis. Its rodents are known from archeological finds only.

  • Dasyprocta leporina, a northern South American agouti. It was recorded on the basis of remains from several archeological sites ranging in age from 100 BCE to 1200 CE.[45]
  • Pennatomys nivalis, an extinct oryzomyine, is known from archeological sites.[42] Ages for these sites range from 790 BCE to 1200 CE.[45]

Antigua

Antigua, which together with Barbuda forms Antigua and Barbuda, was home to a giant oryzomyine which has been described from a site named Mill Reef under the invalid name "Oryzomys hypenemus".[46] Subsequently, rice rats were also reported from two other Antigua sites, Burma Quarry and Indian Creek; although not explicitly equated with "O. hypenemus", the oryzomyines from the two new sites and Mill Reef were reported under the same species.[47] A more detailed study on the remains from Burma Quarry, an Amerindian site which is about 4500 to 2500 years old, reported the presence of a large rice rat, known as "Undescribed species B", which also occurred on nearby Barbuda, Montserrat, Guadeloupe, and Marie Galante. The femora from this site fell into three different size classes, however, suggesting the presence of more than one species of oryzomyine.[48] At another archeological site, Indian Creek, dated at 1 to 600 CE, a large oryzomyine was found. Its weight was estimated at 408 g on the basis of an average femur depth of 4.8 mm.[49] An unidentified Rattus, the agouti Dasyprocta leporina, and the guinea pig (Cavia porcellus), all of which were introduced to the island by humans, have also been found in archeological sites.[50]

Barbuda

Barbuda is the other main island of Antigua and Barbuda. In addition to "Oryzomys hypenemus", another, smaller oryzomyine has also been recorded, Megalomys audreyae, though on the basis of very limited material.[51] It has been suggested that this species is from Barbados instead.[52] A later study reported "Undescribed species B", which was apparently widespread in the region, from archeological sites on Barbuda (see under Antigua).[53]

Montserrat

Montserrat is a small British island which has hosted several rodents.

  • Dasyprocta leporina, an introduced northern South American agouti. The subspecies on Montserrat is D. l. noblei.[43] It is also known from archeological deposits.[54]
  • The Trant's and Radio Antilles Amerindian sites on Montserrat yielded possibly four different oryzomyines, each represented by limited and fragmentary material. A relatively small form, designated "Undescribed species A", is similar in size to oryzomyines from Saint Kitts and Anguilla, and a larger form is apparently identical to "Undescribed species B", described from Antigua and other islands. Two other possible species, one larger than the second and the other smaller than the first species, are documented by even more fragmentary material. Some of these may have been introduced by the Amerindians.[55]

Guadeloupe

The double island of Guadeloupe, which consists of two parts separated only by a narrow channel, is the core of the French overseas department of Guadeloupe. Several rodents are known.

  • Dasyprocta leporina, an introduced northern South American agouti. The subspecies on Guadeloupe is D. l. noblei.[43] It has also been recorded from archeological deposits.[56]
  • A large oryzomyine, designated as "Undescribed species B" (see also Antigua) is known from several archeological sites on Guadeloupe.[57]
  • An unidentified Rattus has been recorded from an archeological deposit on Guadeloupe.[56]

Marie Galante

Marie Galante is a smaller island that politically belongs to nearby Guadeloupe.

  • Dasyprocta leporina is known from an archeological deposit.[44]
  • A large oryzomyine, later designated as "Undescribed species B" (see under Antigua) has been described from an archeological site.[58]

Dominica

Dominica, the southernmost of the Leeward Islands, is an independent state. Unlike in most nearby islands, no oryzomyines have been recorded, but several introduced rodents are known.

Windward Islands

The rodent fauna of the Windward Islands is more diverse than that of the Leeward Islands, reflecting the islands' location closer to the South American mainland.

Martinique

Martinique is a French overseas department. In addition to a very large oryzomyine, some introduced hystricognaths are known.

  • Dasyprocta leporina, an introduced northern South American agouti. The subspecies on Martinique is D. l. fulvus.[43]
  • Makalata didelphoides, a northern South American spiny rat also known from Trinidad and Tobago, has been recorded from Martinique, but the record is probably erroneous.[59] In 1878, as museum curator, True [60] received an animal captured on the island, but believed it to have been a single individual transported on a sailing vessel and not part of a local breeding population. This individual is represented by a single skin (NMNH 13039), but no cranial material.[61]
  • Megalomys desmarestii, a very large oryzomyine that became extinct by about 1900. It was closely related to Megalomys luciae, which occurred further south on Saint Lucia.[62] It was as large as a cat.[63]

Saint Lucia

The island of Saint Lucia is an independent state. Its rodent fauna is similar to that of its northern neighbor, Martinique.

Saint Vincent and Grenadines

Saint Vincent and the Grenadines is an independent state, composed of the main island of Saint Vincent and the northern portion of the Grenadines. A few rodents are known.

  • Dasyprocta leporina, an introduced northern South American agouti on Saint Vincent. The subspecies on Saint Vincent is D. l. albida.[43]
  • Oligoryzomys victus, a small oryzomyine related to South American Oligoryzomys. Known only from a single specimen captured on Saint Vincent around 1892, it is now presumed extinct.[65]

Grenada

The island of Grenada, the southernmost of the main island chain of the Lesser Antilles, is part of an independent state that also comprises the southern Grenadines, including Carriacou. In addition to some Recent rodents, a small Pliocene fauna is known.

  • An unidentified hutia (reported simply as "Capromyidae") is known from archeological deposits on Carriacou.[66]
  • Dasyprocta leporina, an introduced northern South American agouti. The subspecies on Grenada is D. l. albida.[43] An unidentified agouti has been recorded from archeological deposits on Carriacou.[66]
  • Hydrochoerus gaylordi, an endemic capybara known from a few teeth of late Pliocene age. The same fauna also contained some sloth remains.[67]
  • Oryzomyines have been found in archeological deposits on Carriacou (reported as Oryzomys sp.)[66] and Grenada itself, where two forms occur, which correspond in size to "Undescribed species A" and "B" recorded from Montserrat.[68]

Barbados

Barbados lies east of the main island chain of the Lesser Antilles. There are some anecdotal records of small indigenous mouse-like rodents, probably oryzomyines, from historic times and an oryzomyine ("Oryzomys new sp.") has been recorded in archeological deposits.[69] It has been suggested that Megalomys audreyae, described from similarly named Barbuda, actually came from Barbados.[70] The Barbados rice rat was described as Megalomys georginae in 2012.[71]

Trinidad and Tobago

The Brazilian porcupine is one of many species widespread on mainland South America that is also found on Trinidad and Tobago

Trinidad and Tobago, two islands off northeastern Venezuela, form an independent country. Trinidad has a more diverse fauna than smaller and more remote Tobago, but as Pleistocene land-bridge islands, both have faunas that are more diverse than those of most Caribbean islands. On the other hand, endemism is much more limited than on other islands, with only one definite endemic species known.

  • Coendou prehensilis, a porcupine widespread on mainland South America that also occurs on Trinidad.[72]
  • Cuniculus paca, a large hystricognath widespread in Central and South America that also occurs on Trinidad.[73]
  • Dasyprocta leporina, an agouti from northern South America that occurs on Trinidad, as well as on Tobago; it has also been introduced to much of the Lesser Antilles.[74]
  • Heteromys anomalus, a heteromyid that occurs on both Trinidad and Tobago and also in northern South America and on Isla Margarita.[75]
  • Hylaeamys megacephalus, an oryzomyine that reaches the northern limit of its distribution in Trinidad and northern Venezuela and that occurs south to Paraguay.[76]
  • Makalata didelphoides, a spiny rat from northern South America that is found on both main islands.[59] See also Martinique.
  • Megalomys, or an animal similar to it, was described from Tobago in two 17th-century accounts, but no evidence to confirm its occurrence there has ever been found and the records may have referred to agoutis instead or have been borrowed from accounts of Martinique Megalomys.[77]
  • Necromys urichi, an akodontine from both Trinidad and Tobago that also occurs in the highlands of Venezuela and adjacent countries. It was formerly placed in Akodon.[78]
  • Nectomys palmipes, an oryzomyine from Trinidad and the nearby mainland.[79]
  • Oecomys speciosus, an oryzomyine from Trinidad and nearby Colombia and Venezuela.[80]
  • Oecomys trinitatis, an oryzomyine from both islands with a large distribution in northern South America and into Central America. The taxonomic status of many populations is in doubt, however.[81]
  • Oligoryzomys fulvescens, an oryzomyine widespread in Central and northern South America and also reported from Trinidad.[82]
  • Proechimys trinitatus, a spiny rat known only from Trinidad that is part of a group containing several northern South American Proechimys.[83]
  • Rhipidomys couesi, a thomasomyine from lowland forests from Colombia east to Trinidad.[84]
  • Rhipidomys venezuelae, a thomasomyine that is found on Trinidad and Tobago and in the mountains of northern Venezuela and nearby Colombia.[85]
  • Sciurus granatensis, a squirrel from both main islands, also known from northern South America and southern Central America, as well as from Isla Margarita.[86]
  • Zygodontomys brevicauda, an oryzomyine with a distribution that spans northern South America and southern Central America, and also Trinidad and Tobago.[87] Pleistocene fossils are known from Aruba.
  • Zygodontomys sp., an extinct and undescribed oryzomyine from Trinidad. It was mentioned in a 1962 paper, but has received no attention since and no detailed information has been published.[88]

ABC islands

The ABC islands are three islands off northwestern Venezuela that are part of the Kingdom of the Netherlands. They have a diverse rodent fauna, including many oryzomyines and several others, notably including many from Pleistocene faunas.

Bonaire

Bonaire is the easternmost of the ABC islands. Two species of oryzomyines in a single genus, restricted to Bonaire, are known from Pleistocene fossil deposits, together with an indeterminate third species:

  • Agathaeromys donovani is known from four sites that are probably 900 000 to 540 000 years old.[89]
  • Agathaeromys praeuniversitatis is known from a single site that is probably 540 000 to 230 000 years old.[90]
  • An indeterminate sigmodontine is known from a single edentulous mandible from one of the sites that also yielded A. donovani.[91]

Curaçao

Capybara remains have been found at a fossil site on Curaçao. The species is not currently present on the island.

Curaçao is the largest of the ABC islands and it has received the most mammalogical attention. Its known rodent fauna includes several species, some with restricted distributions.

  • Calomys hummelincki, a small phyllotine also known from nearby Aruba and the adjacent mainland. It has been suggested to have been introduced to the area, but this is no longer accepted.[92]
  • Dushimys larsi, an oryzomyine found at a single Middle Pleistocene site near Duivelsklip.[93]
  • Hydrochoerus hydrochaeris, the capybara, which is widespread in South America. It is known from fragmentary remains from Tafelberg Santa Barbara on southeastern Curaçao, a site that may be of considerable antiquity.[94]
  • Megalomys curazensis, a large oryzomyine from several Pleistocene sites that may also have occurred on Aruba.[94]
  • Oryzomys gorgasi, a rare oryzomyine also known from mainland Colombia and Venezuela that formerly occurred on Curaçao, but that is now extinct there. It was described as a separate species, Oryzomys curasoae, but it cannot be distinguished from mainland examples of O. gorgasi.[95]
  • Oryzomys sp. Several different oryzomyines from Curaçao have been recorded under this name from locations including Tafelberg Santa Barbara, Bullenbaai, and Noordkant.[96] Some may be referable to O. curasoae (currently Oryzomys gorgasi).[97]

Aruba

Aruba is the smallest of the ABC islands and the closest to the mainland. Several rodents are known, none of which are definitely endemic. Most are from Pleistocene deposits, but Calomys hummelincki is extant.

  • Calomys hummelincki, a phyllotine also known from nearby Curaçao (see there).[92]
  • Megalomys curazensis. A few molars from a cave at Seroe Canashito form the only record of this species from Aruba; it has been suggested that they were transported there by raptors.[96]
  • Oligoryzomys sp. An unnamed species of this genus occurs in the Pleistocene of Aruba.[98]
  • Oryzomys sp. An extinct oryzomyine from Seroe Canashito has been recorded under this name.[96]
  • Sigmodon hirsutus, a widespread Central American and northern South American species known from Isla and Seroe Canashito.[99]
  • Zygodontomys brevicauda, another widespread Central American and northern South American species, but known only from Seroe Canashito.[96]

Miscellaneous

Florida Keys

The Florida Keys are a group of islands near the Florida mainland. The rodent fauna includes two species which both also occur on the adjacent mainland.

  • Cricetomys gambianus, an introduced nesomyid from Africa. It has established a breeding population on Grassy Key.[100]
  • Neotoma floridana, a widespread woodrat from the eastern United States. The Florida Keys population has been recognized as a separate subspecies, N. f. smalli.[101]
  • Oryzomys palustris, another species that is widespread in the eastern U.S. Different portions of the Florida Keys population have been recognized as the subspecies O. p. natator and as a separate species, O. argentatus; the latter is not currently accepted.[102]

Bahamas

The Bahamas are a large archipelago north of Cuba and east of Florida. The only indigenous rodent is a hutia.

Cayman Islands

The Cayman Islands are a group of British islands west of Jamaica. Their native rodent fauna consisted of several species of hutia, which are now extinct.

Cozumel

Cozumel is a large island near the mainland of the Mexican state of Quintana Roo. The rodent fauna includes several species, all with close relations to forms from the adjacent mainland. The pocket gopher Orthogeomys hispidus has also been recorded, but most likely in error.[107]

Honduran islands

Sigmodon hispidus or another cotton rat close to it is among the rodents recorded from the Bay Islands of Honduras.

Several Caribbean islands are under the sovereignty of Honduras. These islands harbor two endemic rodents, one of which is now extinct. Several other species also occur on the Central American mainland.

Nueva Esparta

Nueva Esparta is a Venezuelan state that comprises Isla Margarita and some nearby islands. Like Trinidad and Tobago, Isla Margarita is a land-bridge island with a relatively diverse rodent fauna.

See also

References

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  2. ^ Koopman, 1959
  3. ^ Marshall, L. G.; Butler, R. F.; Drake, R. E.; Curtis, G. H.; Tedford, R. H. (1979-04-20). "Calibration of the Great American Interchange". Science. 204 (4390). AAAS: 272–279. doi:10.1126/science.204.4390.272. PMID 17800342. Retrieved 2008-04-15.
  4. ^ Musser and Carleton, 2005; Pregill et al., 1994
  5. ^ a b Woods and Kilpatrick, 2005
  6. ^ a b MacPhee et al., 2003; Woods et al., 2001, p. 337
  7. ^ Flynn et al., 2007
  8. ^ Woods and Kilpatrick, 2005; MacPhee and Flemming, 2003; Turvey et al., 2009
  9. ^ Woods and Kilpatrick, 2005; MacPhee et al., 2003, p. 18
  10. ^ Woods and Kilpatrick, 2005 p. 1591
  11. ^ Musser and Carleton, 2005
  12. ^ Woods and Kilpatrick, 2005; Musser and Carleton, 2005; Thorington and Hoffmann, 2005; Patton, 2005
  13. ^ Turvey et al., 2009, pp. 588–589
  14. ^ a b c d e f Woods and Kilpatrick, 2005, p. 1592
  15. ^ a b c d e f g h i j k l m n o Woods et al., 2001, p. 336
  16. ^ a b Woods and Kilpatrick, 2005, p. 1594
  17. ^ Díaz-Franco, 2001
  18. ^ Woods and Kilpatrick, 2005, p. 1559
  19. ^ a b c d Woods and Kilpatrick, 2005, p. 1558
  20. ^ Díaz-Franco and Jímenez Vázquez, 2008
  21. ^ Account in IUCN, 2009 (accessed September 13, 2009); cf. Woods and Kilpatrick, 2005, p. 1595
  22. ^ a b c d Woods and Kilpatrick, 2005, p. 1595
  23. ^ a b c Woods and Kilpatrick, 2005, p. 1596
  24. ^ a b c Woods and Kilpatrick, 2005, p. 1597
  25. ^ a b Woods and Kilpatrick, 2005, p. 1599
  26. ^ a b MacPhee and Flemming, 2003
  27. ^ Weksler et al., 2006, table 1; Musser and Carleton, 2005, p. 1147; Ray, 1962, pp. 27–47
  28. ^ McFarlane et al., 2002
  29. ^ Díaz-Franco and Jímenez-Vázquez, 2008, p. 44
  30. ^ Woods and Kilpatrick, 2005, pp. 1597–1598
  31. ^ a b c d e Woods and Kilpatrick, 2005, p. 1598
  32. ^ Woods and Kilpatrick, 2005, pp. 1598–1599
  33. ^ a b Woods and Kilpatrick, 2005, p. 1600
  34. ^ MacPhee et al., 2003, p. 18; Williams and Koopman, 1951
  35. ^ Turvey et al., 2009
  36. ^ Account in IUCN, 2009 (accessed September 13, 2009); Woods and Kilpatrick, 2005, p. 1557, mention the "Virgin Islands", but do not specify whether the U.K. or U.S. Virgins are meant
  37. ^ a b Woods and Kilpatrick, 2005, p. 1600; Biknevicus et al., 1993
  38. ^ Pregill et al., 1994, p. 10
  39. ^ Crock, 2000, p. 221; Steadman et al., 1984b, p. 23; Pregill et al., 1994, p. 10
  40. ^ Wing, 1995; Pregill et al., 1994, p. 14
  41. ^ Pregill et al., 1994, p. 15
  42. ^ a b c Turvey et al., 2010, p. 761
  43. ^ a b c d e f g h Woods and Kilpatrick, 2005, p. 1557
  44. ^ a b Wing, 1973
  45. ^ a b Kozuch and Wing, 2006
  46. ^ Ray, 1962, p. 109
  47. ^ Wing et al., 1968, p. 128; Steadman et al., 1984a
  48. ^ Pregill et al., 1988, p. 22
  49. ^ Rouse and Morse, 1999
  50. ^ Wing et al., 1968, pp. 128–129; Rouse and Morse, 1999; Woods and Kilpatrick, 2005, p. 1557 for nomenclature
  51. ^ Ray, 1962
  52. ^ Ray, 1962, pp. 90–92
  53. ^ Watters et al., 1984, p. 405
  54. ^ Steadman et al., 1984b, p. 24
  55. ^ Steadman et al., 1984b, pp. 23–24
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  67. ^ MacPhee et al., 2000; Haynes, 2009, table 9.2 for spelling of generic name as Hydrochoerus, not Hydrochaeris as in the original description
  68. ^ Pregill et al., 1994, p. 31
  69. ^ Ray, 1962, pp. 90–91; Pregill et al., 1994, p. 30
  70. ^ Ray, 1962, pp. 90–91
  71. ^ Turvey et al., 2012
  72. ^ Woods and Kilpatrick, 2005, p. 1547
  73. ^ Account in IUCN, 2009; also AMNH specimens Archived 2009-09-07 at the Wayback Machine 4764 and 6041 (both accessed September 13, 2009)
  74. ^ Account in IUCN, 2009; also AMNH specimens Archived 2009-09-07 at the Wayback Machine 4761, 4762, 4763, 4796, 6038, 6039, 6040, 31226, and 31227 (both accessed September 13, 2009)
  75. ^ a b Patton, 2005, p. 850
  76. ^ Musser and Carleton, 2005, p. 1151
  77. ^ Ray, 1962, pp. 64–65
  78. ^ Musser and Carleton, 2005, p. 1131
  79. ^ Musser and Carleton, 2005, p. 1132
  80. ^ Musser and Carleton, 2005, p. 1138
  81. ^ Musser and Carleton, 2005, p. 1139
  82. ^ Allen and Chapman, 1897, p. 17; Musser and Carleton, 2005, p. 1141 for nomenclature
  83. ^ Woods and Kilpatrick, 2005, p. 1588
  84. ^ Musser and Carleton, 2005, p. 1168
  85. ^ Musser and Carleton, 2005, p. 1171
  86. ^ a b Thorington and Hoffmann, 2005, p. 761
  87. ^ Musser and Carleton, 2005, p. 1186
  88. ^ Ray, 1962, table 1
  89. ^ Zijlstra et al., 2010, p. 863
  90. ^ Zijlstra et al., 2010, p. 866
  91. ^ Zijlstra et al., 2010, p. 867
  92. ^ a b Musser and Carleton, 2005, p. 1107
  93. ^ Zijlstra, 2012
  94. ^ a b Hooijer, 1959; De Buisonjé, 1974
  95. ^ Voss and Weksler, 2009
  96. ^ a b c d De Buisonjé, 1974
  97. ^ McFarlane and Debrot, 2001
  98. ^ Zijlstra, 2012, p. 62
  99. ^ De Buisonjé, 1974; Musser and Carleton, 2005, p. 1175 for referral to S. hirsutus instead of S. hispidus
  100. ^ Perry et al., 2006
  101. ^ Musser and Carleton, 2005, p. 1055
  102. ^ Musser and Carleton, 2005, p. 1152-1153
  103. ^ Woods, 1989, pp. 12–13
  104. ^ Morgan, 1994b, pp. 489–491
  105. ^ Morgan, 1994a
  106. ^ Morgan, 1994b, pp. 491–492
  107. ^ Jones and Lawlor, 1965, p. 418
  108. ^ a b c Jones and Lawlor, 1965, p. 416
  109. ^ Jones and Lawlor, 1965, p. 413; Musser and Carleton, 2005, p. 1147; Engstrom et al., 1989, p. 414
  110. ^ Engstrom et al., 1989, p. 414
  111. ^ Jones and Lawlor, 1965, pp. 413–415; Musser and Carleton, 2005, p. 1084; Engstrom et al., 1989, p. 414
  112. ^ Morgan, 1989
  113. ^ Koopman, 1959, p. 237
  114. ^ Koopman, 1959, p. 237, 238
  115. ^ Musser and Carleton, 2005, p. 1175
  116. ^ Wagenaar Hummelinck, 1940, p. 69
  117. ^ Woods and Kilpatrick, 2005, p. 1578
  118. ^ Emmons, 2005, pp. 282–283

Literature cited

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