UPDATED CHECKLIST OF ENDEMIC TERRESTRIAL MAMMALS OF THE WEST INDIES

Abstract

The list of terrestrial mammals of the West Indies in the orders Pilosa, Soricomorpha (Insectivora in part), Primates, Carnivora, Perissodactyla, and Rodentia is updated. Comments on taxonomy, systematics, new synonyms, distribution and conservation status are included. RESUMEN Se actualiza la lista de los mamíferos terrestres de las Antillas, exceptuando a los murciélagos. Se incluyen nuevas sinonimias. Se realizan comentarios sobre su taxonomía, sistemática, distribución, sinonimia y conservación de los órdenes Pilosa (Xenarthra), Soricomorpha (Insectivora), Primates, Perissodactyla y Rodentia, representados en las Antillas.

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CHAPTER 7. CHECKLIST
UPDATED CHECKLIST OF ENDEMIC TERRESTRIAL
MAMMALS OF THE WEST INDIES
Rafael Borroto-Páez, Carlos A. Mancina, Charles A. Woods and C. William Kilpatrick
ABSTRACT The list of terrestrial mammals of the West Indies in the orders Pilosa, Soricomorpha (Insectivora
in part), Primates, Carnivora, Perissodactyla, and Rodentia is updated. Comments on taxonomy, systematics,
new synonyms, distribution and conservation status are included.
KEY WORDS: terrestrial mammals, West Indies, synonymy, taxonomy
RESUMEN Se actualiza la lista de los mamíferos terrestres de las Antillas, exceptuando a los murciélagos. Se
incluyen nuevas sinonimias. Se realizan comentarios sobre su taxonomía, sistemática, distribución, sinonimia y
conservación de los órdenes Pilosa (Xenarthra), Soricomorpha (Insectivora), Primates, Perissodactyla y
Rodentia, representados en las Antillas.
PALABRA CLAVES: mamíferos terrestres, Antillas, sinonimias, taxonomía
INTRODUCTION
The islands of the West Indies are notorious for
having undergone proportionately more
mammalian extinctions in the late Quaternary than
any other region on Earth despite occupying a
small portion of the world's land surface. In the
past 500 years nearly 40% of the all species of
West Indian land mammals have become extinct
(Morgan and Woods 1986; MacPhee and Fleming
1999). These large-scale extinctions have occurred
in two waves: the first, concentrating around 5,000
years ago, led to the extinction of large mammals
(e.g. large ground sloths and giant rodents). The
second wave of extinctions coincided with the
arrival of Europeans in the New World and resulted
in the disappearance of most small mammals (e.g.
smaller rodents and insectivores). Both waves of
extinction have been associated with human
activity, either through direct predation or through
the introduction of exotic mammals and their
parasites and diseases (Morgan and Woods 1986;
MacPheeand Iturralde-Vinent 2000; Borroto-Páez
and Woods, this volume).
In the last 10 years, the estimated date for the
first invasions of the West Indies by mammals has
changed radically as a result of new paleontological
and molecular discoveries (MacPhee and Iturralde-
Vinent 1995b; MacPhee and Grimaldi 1996;
Domming et al. 1997; Roca et al. 2004). Most of
the known lineages of Antillean land-mammals
may have reached the islands comparatively early
in the Cenozoic, perhaps as early as the late
Palaeogene (Iturralde-Vinent and MacPhee 1999).
An unexpected feature of the Antillean record is
that, in several cases, the oldest fossils from the
islands are as old as, or even older than the oldest
mainland fossils attributed to the same major clades
(MacPhee 2005).
In the last 30 years, following the publication of
“Catálogo de los Mamíferos Vivientes y
Extinguidos de las Antillas” by Varona (1974b),
the taxonomic arrangement of West Indian
terrestrial mammals has changed considerably at all
levels of classification. Systematics (traditional
and molecular) and paleontological studies of the
area have led to the discovery of new taxa, and
redefined the taxonomic affinity between closely
related taxa.
In this checklist, we have relied on the
following sources: the two editions of
Biogeography of the West Indies (Woods 1989d;
Woods and Sergile 2001), Woods et al. (2001),
White and MacPhee (2001), Ottenwalder (2001),
Whidden and Asher (2001), the extensive
palaeontological explorations of the American
Museum of Natural History, the Florida Museum of
Natural History and the Museum of Natural History
in Havana (and their Contributions Series “Origin
of the Antillean Land Mammals Fauna” published
in various papers of MacPhee, Iturralde-Vinent,
Flemming and other researchers), the systematic
research of the Institute of Ecology and
Systematics in Havana (Borroto-Páez et al. 1992;
Borroto-Páez 2002; Borroto-Páez et al. 2005). We

TERRESTRIAL MAMMALS OF THE WEST INDIES
390
have also relied on the recent compilation of Cuban
terrestrial mammals (with extensive taxonomic
information about recent and extinct species) by
Silva-Taboada et al. (2007) of the Natural History
Museum of Havana.
‘West Indies’ is used in a biogeographical sense
following the classification of Hershkovitz (1958),
Morgan and Woods (1986), and Morgan (2001).
The West Indies includes the Greater Antilles
(Cuba, Jamaica, Hispaniola, and Puerto Rico), as
well as the Bahamas, Cayman Islands, Swan
Islands, Virgin Islands and the Lesser Antilles
south to St. Vincent and Barbados. Grenada and
the Grenadines, Trinidad, Tobago, Margarita,
Aruba, Bonaire and Curaçao are excluded from this
definition of the West Indies, as their biota is more
characteristic of continental South America. The
terms ‘Antillean’, ‘Caribbean’, and ‘West Indian’,
used in a faunal sense, are intended to be
synonyms, and thus encompass all the islands
defined above as part of the West Indies (Morgan
2001).
One of our goals in creating the checklist is to
simplify the taxonomic arrangement of West Indian
land mammals as much as possible. We focus on
described and undescribed taxa of the orders Pilosa,
Soricomorpha, Primates, Perissodactyla, and
Rodentia. In all cases, we have carefully analyzed
the validity of the diagnoses and evaluated the list
of characters used in the original species
descriptions and we have checked the validity of
the material used in the hypodigms. We have
selected as criterion to simplify and lump species in
justified synonymies. Our list excludes grouping
taxa at the subgeneric level because we are
concerned that there is too much subjectivity in this
category. We have decided to include information
on undescribed species and subspecies with the
intention of identifying gaps in our knowledge, and
to promote awareness, research and the
conservation of these potential taxa. We have
followed the format of Wilson and Reader (2005),
and included names of the reviewers for each group
(primarily from Woods and Kilpatrick 2005) and
taxonomic information above genera. We have
tried to include updated information at every
taxonomic level where possible. Introduced
mammals are excluded.
Table 1 summarizes the terrestrial mammal taxa
of the West Indies. The updated checklist that
follows recognizes 87 species (including 16
undescribed fossil and three probable recent extinct
species), belonging to six orders, nine families, and
33 genera described. We recognize 74 fossil
species (85%), and 13 recent species (15%).
Today, we count 77 extinct species (88.5%) and 10
living species (11.5%).
CHECKLIST
Magnorder Xenarthra Cope, 1889
Order Pilosa Flower, 1883
Suborder Phyllophaga Owen, 1842
Superfamily Megatherioidea Gray, 1821
Family Megalonychidae Gervais, 1855. Histoire
naturelle des mammifères. Vol. 2: 44.
Sloths were very diverse in the Pleistocene of the
Greater Antilles (Cuba, Hispaniola and Puerto
Rico). Fossil remains indicate that sloths colonized
the Antilles at least as early as the Oligocene
(MacPhee and Iturralde-Vinent 1994, 1995b;
MacPhee 2005), and most fossil forms that have
been found are of Quaternary age. This long
geological record has been used to explain the large
number of identified lineages (e.g. Arredondo and
Rivero 1997). All Antillean sloths are members of
the family Megalonychidae, but there is much
disagreement among taxonomists concerning genus
and species validity. According to White and
MacPhee (2001), a relatively large number of
species have been designated based on few and
isolated cranial and limb fragments and non-
comparable type specimens, leading to plethora of
taxa. We follow the systematic arrangement of
White and MacPhee (2001) who made the last
systematics revision of this group.
Acratocnus Anthony, 1916: 195. Ann. New. York
Acad. Sci., 27: 195.
TYPE SPECIES Acratocnus odontrigonus
Anthony, 1916
SYNONYMS Miocnus Matthew, 1931: 3;
Habanocnus Mayo, 1978: 688; Galerocnus
Arredondo and Rivero, 1997; Paramiocnus
Arredondo and Arredondo, 2000
COMMENT The presence of Acratocnus on Cuba,
Hispaniola, and Puerto Rico can only reasonably
be interpreted as having resulted from relatively
late Quaternary inter-island dispersals (Rega et al.
2002). Numerous materials assigned to
Acratocnus have been previously misidentified
and confused with other species (see White and
MacPhee 2001). Following these authors in
recognizing as few taxonomic categories as
possible and lumping taxa that have been
described based on single limb elements, we place
the genera Galerocnus and Paramiocnus
(respectively described from a single limb bone –
the distal half of left femur and left humerus) as
synonyms of Acratocnus. Both were recognized
similar to Miocnus and Habanocnus considered as
Acratocnus after White and MacPhee (2001).

UPDATED CHECKLIST – Borroto-Páez, Mancina, Woods and William Kilpatrick
391
Table 1. List of West Indies land mammal taxa
Order
Number
of species
perOrder
Family
Genera
Number of taxa
Species and subspecies
Pilosa
16
Megalonychidae
Acratocnus
4
Neocnus
4
Megalocnus
2
Parocnus
2
Imagocnus
1
Indeterminate
2 undescribed
Soricomorpha
11
Nesophontidae
Nesophontes
6 (2 undescribed)
Solenodontidae
Solenodon
4, 2 subspecies
Indeterminate (in
amber)
1 undescribed
Primates
6
Pitheciidae
Antillothrix
1
Paralouatta
Insulacebus
2
1
Xenothrix
1
Indeterminate
2 undescribed
Carnivora
1
Indeterminate (in
amber)
Indeterminate
1 undescribed
Perissodactyla
1
Rhinocerontidae
Hyrachyus
1
Rodentia
51
Muridae
Megalomys
5 (2 undescribed)
Oryzomys
2 (1 undescribed)
Oligoryzomys
1
Echimyidae
Boromys
2
Brotomys
2
Heteropsomys
2
Puertoricomys
1
Capromyidae
Capromys
5 (1 undescribed), 9
subspecies (4 undescribed)
Geocapromys
6 (2 undescribed), 3
subspecies
Mesocapromys
8
Mysateles
1, 3 subspecies
Hexolobodon
2 (1 undescribed)
Isolobodon
2
Zazamys
1
Plagiodontia
3, 2 subspecies
Rhizoplagiodontia
1
Heptaxodontidae
Clidomys
1
Amblyrhiza
1
Elasmodontomys
1
Tainotherium
1
Quemisia
1
Incertae sedis
Xaymaca
1
Caviida incertae
sedis
Indeterminate
1 undescribed

TERRESTRIAL MAMMALS OF THE WEST INDIES
392
Acratocnus odontrigonus Anthony, 1916. Ann.
New. York Acad. Sci., 27: 195.
COMMON NAME Puerto Rican sloth
TYPE LOCALITY Cueva de la Ceiba, near
Utuado, Puerto Rico
DISTRIBUTION Mainland of Puerto Rico
STATUS Extinct
SYNONYM Acratocnus major Anthony, 1918
(Paula Couto 1967; White and MacPhee 2001)
COMMENT See White and MacPhee (2001: 214).
Acratocnus antillensis (Matthew, 1931). Amer.
Mus. Novitates 511: 4.
COMMON NAME Cuban sloth
TYPE LOCALITY Casimba in Sierra de
Jatibonico, Cuba
DISTRIBUTION Cuba
STATUS Extinct
SYNONYMS Habanocnus hoffstetteri Mayo,
1978; Habanocnus paulacoutoi Mayo, 1978;
Galerocnus jaimezi Arredondo, 1997; Paramiocnus
riveroi Arredondo and Arredondo 2000
COMMENT See White and MacPhee (2001: 214)
and Silva-Taboada et al. (2007). The description of
G. jaimezi was based on insufficient material (a
damaged femur). In the case of P. riveroi, the
description material consisted in a single very
fragmented distal part of the left humerus. Both
came from the same locality (Cueva del Mono
Fosil, Sierra de Galeras, Pinar del Río). Arredondo
and Arredondo (2000) recognized riveroi
resembles antillensis and odontrigonus but with a
larger size, and they rejected the possibility of this
humerus belonging to jaimezi, although size
variation within population was not considered. In
their results, the humerus is larger than the femur
for antillensis, although the specimens analysed are
unclear. Silva-Taboada et al. (2007) support the
synonymy of jaimezi and riveroi.
Acratocnus ye MacPhee, White and Woods, 2000.
Amer. Mus. Novitates 3303: 11.
COMMON NAME Haitian Macaya sloth
TYPE LOCALITY Trouing Vapè Deron, Plain
Formon, Département du Sud, Haiti
DISTRIBUTION Hispaniola
STATUS Extinct
COMMENT see White and MacPhee (2001: 215).
Acratocnus simorhynchus Rega et al., 2002. Carib.
J. Sci. 38: 14.
COMMON NAME Dominican flat-nosed sloth
TYPE LOCALITY Cueva del Perezoso, Jaragua
National Park, Pedernales Province, Dominican
Republic
DISTRIBUTION Only known from the type
locality
STATUS Extinct
Neocnus Arredondo, 1961. Boletín Grupo
Exploraciones Científicas 1: 29.
TYPE SPECIES Neocnus gliriformis (Matthew,
1931)
SYNONYMS Microcnus (Paula Couto, 1967);
Cubanocnus Kretzoi, 1968
COMMENT Neocnus gliriformes was originally
described as Microcnus gliriformis, but this generic
was preoccupied, thus Kretzoi (1968) established
Cubanocnus. However, Neocnus had priority
(Varona 1976). Neocnus is a genus of small body
sizedsloths, see comments of White and MacPhee
(2001: 21).
Neocnus gliriformis (Matthew, 1931). Amer. Mus.
Novitates 511: 4.
COMMON NAME Cuban rodent-like sloth
TYPE LOCALITY Casimba in the Sierra de
Jatibonico, Cuba
DISTRIBUTION Cuba
STATUS Extinct
SYNONYM Cubanocnus gliriformis (of Varona,
1974b); Neocnus major Arredondo, 1961; Neocnus
minor Arredondo, 1961; Neocnus baireiensis
Mayo, 1980, Neocnus amplus Arredondo and
Arredondo 1999
COMMENT See comment for the genus as well as
White and MacPhee (2001). The synonymy of N.
amplus is based on the following reasons. The
material for the description of this taxon is a single
bone (a very fragmented distal part of the left
femur). The proximal part of the femur (normally
with the most diagnostic characters) is absent
(Arredondo and Arredondo 1999). The characters
in the diagnosis are weak and distinctiveness is
based on comparisons of femura from two species
(minor and baireiensis) that are considered
synonym of N. major (White and MacPhee 2001).
Therefore, the morphological variation in the femur
in Neocnus is significant. The type locality of
amplus is Cueva Paredones, in Caimito, Havana
province. It is a cave with an extensive history of
exploration dating from the 1950s. Silva-Taboada
et al. (2007) suggest that sexual dimorphism and
chronoclines are important factors in Cuban sloths,
and consider N. major to be N. gliriformis. White
and MacPhee (2001), however, consider that there
is no evidence (based on an analysis of a large
number of individuals) for sexual dimorphism in
living or fossil sloths.
Neocnus comes (Miller, 1929). Smithsonian Misc.
Coll., 81: 26.
COMMON NAME Lesser Haitian ground sloth
TYPE LOCALITY Cave near St. Michel de
l'Atalaye, Haiti
DISTRIBUTION Hispaniola
STATUS Extinct

UPDATED CHECKLIST – Borroto-Páez, Mancina, Woods and William Kilpatrick
393
SYNONYMS Acratocnus comes Miller, 1929;
Synocnus comes (Paula Couto, 1967)
COMMENT See White and MacPhee (2001: 218).
Neocnus dousman MacPhee, White and Woods,
2000. Amer. Mus. Novitates 3303: 13.
COMMON NAME Haitian pine forest sloth
TYPE LOCALITY Trouing de la Scierie, Morne
La Visite, Haiti
DISTRIBUTION Hispaniola
STATUS Extinct
COMMENT See White and MacPhee (2001: 219).
Neocnus toupiti MacPhee, White and Woods, 2000.
Amer. Mus. Novitates 3303: 15.
COMMON NAME Haitian rak bwa sloth
TYPE LOCALITY Trouing Jérémie #5, Plain
Formon, Département du Sud, Haiti
DISTRIBUTION Hispaniola
STATUS Extinct
COMMENT See White and MacPhee (2001: 220).
Megalocnus Leidy, 1868. Proc. Acad. Nat. Sci.
Philadelphia, 20: 180.
TYPE SPECIES Megalocnus rodens Leidy, 1868
SYNONYMS Myomorphus Pomel, 1868;
Megalocuus Miller, 1922; Oryctotherius Spencer,
1895; Megalonyx Allen, 1911
COMMENT See generic characters in White and
MacPhee (2001: 221).
Megalocnus rodens Leidy, 1868. Proc. Acad. Nat.
Sci. Philadelphia, 20: 180.
COMMON NAME Cuban giant sloth
TYPE LOCALITY Ciego Montero, near
Cienfuegos province, Cuba
DISTRIBUTION Cuba
STATUS Extinct
SYNONYMS Myomorphus cubensis Pomel, 1868;
Megalocnus rodens rodens Leidy, 1868;
Oryctotherius cubensis Spencer, 1895; Megalonyx
rodens Allen, 1911; Megalocnus rodens casimbae
Matthew, 1959; Megalocnus ursulus Matthew,
1959; Megalocnus junius Matthew, 1959;
Megalocnus intermedius Mayo, 1969
COMMENT THIS sloth is the largest known
Antillean megalonychid and was the first sloth to
be described in the West Indies. Its fossil remains
are very common throughout Cuba. This species
exhibits an extensive range of morphometric
variations (Arredondo 2000b). See comments in
White and MacPhee (2001: 221).
Megalocnus zile MacPhee, White and Woods,
2000, Amer. Mus. Novitates 3303: 7.
COMMON NAME Large Ile Tortue sloth
TYPE LOCALITY Trou Gallery, Île de la Tortue,
Département du Nord-Ouest, Haiti
DISTRIBUTION Hispaniola
STATUS Extinct
SYNONYM Megalocuus sp. [lapsus calami]
(Miller, 1922)
COMMENT See White and MacPhee (2001: 222).
Parocnus (Miller, 1929). Smithsonian Misc. Coll:
28.
TYPE SPECIES Parocnus serus Miller, 1929
SYNONYMS Barnumia Torre, 1911; “Megalocuus
sp.” [lapsus calami] (Miller, 1922); Mesocnus
Matthew, 1931; Neomesocnus Arredondo, 1961
COMMENT White and MacPhee (2001: 222)
consider Mesocnus a synonym of Parocnus. Both
species are so similar in all important diagnostic
features that a generic separation between them can
no longer be sustained. See the list of generic
characters in White and MacPhee (2001: 223).
Silva-Taboada et al. (2007) consider Neomesocnus
to be Parocnus.
Parocnus serus Miller, 1929, Smithsonian Misc.
Coll., 81: 29.
COMMON NAME Greater Haitian ground sloth
TYPE LOCALITY St. Michel de l'Atalaye, Haiti
DISTRIBUTION Hispaniola, including the islands
of La Tortue and La Gonâve
STATUS Extinct
SYNONYM “Megalocuus sp.” [lapsus calami]
(Miller, 1922)
COMMENT see White and MacPhee (2001: 223).
Parocnus browni (Matthew, 1931). Amer. Mus.
Novitates 511: 2.
COMMON NAME Matthew’s ground sloth
TYPE LOCALITY Ciego Montero, near
Cienfuegos Province, Cuba
DISTRIBUTION Cuba
STATUS Extinct
SYNONYMS Barnumia browni Torre, 1911;
Mesocnus browni Matthew, 1931; Mesocnus torrei
Matthew, 1931; Mesocnus herrerai Arredondo,
1977; Neomesocnus brevirostris Arredondo 1961.
COMMENT See White and MacPhee (2001: 224).
Neomesocnus brevirostris was considered a
synonym of Megalocnus rodens (Paula Couto
1967, Mayo 1969, Fischer 1971, White and
MacPhee 2001). Arredondo (2000b) re-evaluated
the type material (rediscovered recently) and
considered that enough diagnostic traits are present
to consider it as a valid taxon. However, Silva-
Taboada et al. (2007) argue for the synonymy of
this form with P. browni.
Imagocnus MaPhee and Iturralde-Vinent, 1994.
Amer. Mus. Novitates 3094:3.
TYPE SPECIES Imagocnus zazae MaPhee and
Iturralde-Vinent, 1994
SYNONYMS None

TERRESTRIAL MAMMALS OF THE WEST INDIES
394
COMMENT From early Miocene, this
megalonychid is the oldest known in Cuba. See
White and MacPhee (2001: 224).
Imagocnus zazae MacPhee and Iturralde-Vinent,
1994. Amer. Mus. Novitates 3094:3.
COMMON NAME Zaza sloth
TYPE LOCALITY Domo de Zaza, Sancti Spíritus
Province, Cuba
DISTRIBUTION Central Cuba, early Miocene
STATUS Extinct
COMMENT See White and MacPhee (2001: 224).
Megalonychidae, gen. et sp. indeter.
COMMENT Undescribed species from Puerto
Rico; MacPhee and Iturralde-Vinent (1995b)
reported a sloth from early Oligocene deposit near
Yauco in southwestern Puerto Rico (see comment
of White and MacPhee 2001: 225).
Megalonychidae, gen. et sp. indeter.
COMMENT Undescribed species from Finca
Horizonte, Corralillo, Villa Clara as part of the
Carlos de la Torre Collection now at the Institute of
Ecology and Systematics. The materials are left
ulna, fragmented left tibia and a diaphysis of a right
humerus belonging to an animal larger than both
M. Rodens and P. browni (Silva-Taboada et al.
2007).
Order Soricomorpha Gregory 1910
The relationship of West Indian soricomorphs with
other insectivore like mammals is a topic of much
discussion and analysis. Despite many similarities
between Solenodontidae and Nesophontidae, the
different patterns of their molar cusps place them in
distinct families. Solenodon has a zalambdodont
pattern of upper molars (like tenrecids and
chrysochlorids) and Nesophontes has a
dilambdodont pattern of upper molars (like soricids
and talpids) (Whidden and Asher 2001). Whidden
and Asher (2001: 248) considered four
biogeographical hypotheses for the origin of the
Greater Antillean “insectivorans”. The earliest
fossil of an “insectivore” in West Indies is the
partial axial skeleton embedded in a piece of late
Oligocene or early Miocene Dominican amber
(MacPhee and Grimaldi 1996). Molecular data
(Roca et al. 2004) indicates a Miocene origin for
the solenodons. However, there is no fossil record
as such for either solenodons or nesophontids prior
to the Quaternary (MacPhee 2005). The invasion
of “insectivora” into the West Indies must be from
a different source (North America) in comparison
with other groups of mammals with a South
American origin, and therefore at least two
dispersal patterns are necessary to explain the
origin of West Indian mammals.
Superfamily Soricoidea Fischer de Waldheim
1817
Family Nesophontidae Anthony, 1916. Bull.
Amer. Mus. Nat. Hist., 35: 725.
These small soricomorphs likely became extinct
after the arrival of the Spanish in the West Indies,
as their bones have been found together with those
of Rattus and of Mus in caves. Their extinctionis
attributed to the introduction of exotic species,
especially rats, cats and dogs (Morgan and Woods
1986, Morgan 1994b, Woods and Ottenwalder
1992, MacPhee et al. 1999b, Borroto-Páez and
Woods this volume). The Nesophontidae are
reviewed by McDowell (1958) and Whidden and
Asher (2001). See also Borroto-Páez (2011a) and
Fernández (2011).
Sexual dimorphism (males larger than females)
considered for some species (Antohny 1925,
Arredondo 1970a) is a potential variation that has
not been taken into consideration in the description
of most species. However, there is no sexual
dimorphism in other living soricomorphs.
Chronoclinal variation is present in Puerto Rico
(Choate and Birney 1968). Nesophontes on
Hispaniola and the Cayman Islands are
statigraphically superficial, while in Puerto Rico
they occur in deeper strata, suggesting an earlier
extinction there (McFarlane 1999).
Nesophontes Anthony, 1916. Bull. Amer. Mus.
Nat. Hist., 35: 725.
COMMON NAME West Indian island shrew,
West Indian nesophont, musarañas
TYPE SPECIES Nesophontes edithae Anthony,
1916
COMMENT A character of the genus is the
dilambdodont pattern in the upper molars.
Nesophontes edithae Anthony, 1916. Bull. Amer.
Mus. Nat. Hist., 35: 725.
COMMON NAME Puerto Rican nesophont,
Puerto Rican shrew
TYPE LOCALITY Cueva Catedral, near Morovis,
Puerto Rico
DISTRIBUTION Puerto Rico, Viequez, St. John,
and St. Thomas
STATUS Extinct
COMMENT The largest species in the genus.
Woods and Eisenberg (1989) attribute the large
size to the lack of competition with Solenodon,
absent in Puerto Rico. Estimated weight is 150 g.
The specimens from St. John and St. Thomas are
usually considered as Nesophontes sp.

UPDATED CHECKLIST – Borroto-Páez, Mancina, Woods and William Kilpatrick
395
Nesophontes hypomicrus Miller, 1929. Smithsonian
Misc. Coll., 81(9): 4.
COMMON NAME Hispaniolan [Atalaye]
nesophont, Hispaniola island shrew
TYPE LOCALITY Cave four miles E of St. Michel
de l'Atalaye, Haiti
DISTRIBUTION Haiti including the island of La
Gonâve
STATUS Extinct
COMMENT Estimated size is 25-30 grams (Woods
and Ottenwalder 1992).
Nesophontes micrus Allen, 1917. Bull. Mus. Comp.
Zool., 61: 5.
COMMON NAME West Cuban nesophont, West
Cuban island shrew.
TYPE LOCALITY Sierra de Hato Nuevo,
Matanzas, Cuba
DISTRIBUTION Cuba including Isla de la
Juventud (former Isla de Pinos) and Cayo
Guillermo, Archipelago Camagüey, Hispaniola
STATUS Extinct
SYNONYMS superstes Fischer, 1977; submicrus
Arredondo, 1970; longirostris Anthony, 1919;
paramicrus Miller, 1929; major Arredondo, 1970
COMMENT We agree with Patterson(1962) that
N. paramicrus is not distinguishable from N.
micrus, and with Varona (1974b) that N.
paramicrusis a synonym of N. micrus. Hutterer
(1993) and MacPhee et al. (1999b) included
Hispaniola in the distribution of N. micrus but
continued to recognize N. paramicrus as a valid
Hispaniolan species. This could be true if N.
paramicrus was part of an early radiation of
Nesophontes on Hispaniola, or part of the “South
Island” Nesophontes radiation, and if N.micrus
dispersed to Hispaniola at a later time (perhaps as
part of a “North Island” radiation). It is an
intriguing idea. However, until careful
biogeographical and systematics analyses of the
distribution and relationship of N. micrus/N.
paramicrus sized Nesophontes on Hispaniola is
completed, we propose the synonymy of N.
paramicrus. We are, therefore, removing it from
the list of Hispaniolan taxa. Both N. superstes and
N. longirostris were described based on a single
specimen. Hutterer (1993) consider eight species
as valid. Several authors have proposed the
synonymy of N. longirostris with (as) N. micrus
(Morgan 1977; Condis-Fernández et al. 2005;
Silva-Taboada et al. 2007). Condis-Fernández et
al. (2005) proposed the synonymy of N. superstes
and N.submicrus as N. micrus, and considered N.
major as valid, and reported N. micrus in Cayo
Guillermo. Morgan (1977) considered N. major to
be N. micrus like Silva-Taboada et al. (2007).
Estimated weight is 40-50 grams. The true
significance in statigraphy, cronoclinal and sexual
dimorphism and variations must be reviewed for
specimens classified as N. micrus.
Nesophontes zamicrus Miller, 1929. Smithsonian
Misc. Coll., 81(9): 7.
COMMON NAME Haitian nesophont, Haitian
island shrew
TYPE LOCALITY Cave four miles E of St. Michel
de l'Atalaye, Haiti
DISTRIBUTION Hispaniola
STATUS Extinct
COMMENT The smallest species in the genus.
Estimated weight is 8-10 grams (Woods and
Ottenwalder 1992).
Nesophontes sp. 1
Remains of this genus were reported by Patton
(1966) on the Cayman Islands. This species, from
several caves from Cayman Brac, is in the process
of being described (Morgan 1994a; Morgan
personal communication).
Nesophontes sp. 2
This fossil species from Grand Cayman is also in
the process of being described (Morgan 1994a;
Morgan personal communication).
COMMENT These two Nesophontes forms are
very closely related species and very similar in
morphology but do not overlap in size. The fossils
from Cayman Islands appear to have been derived
from Cuban species, probably N. micrus, based on
several shared dental features (Morgan 1994a).
The Cayman Nesophontes are distinguished from
all other members of the genus by the total loss of
cingula on the upper molars and by the reduction of
the cingulids on the lower molars (Morgan 1994a).
Family Solenodontidae Dobson, 1882. A
monograph of Insectivora, syst. and anat. Part I: 82
The two species of solenodons (Solenodon
paradoxus and S. cubanus) comprise the only two
surviving species of West Indian “insectivores”.
The affinity of solenodons with other mammals has
been debated by several authors (e.g. Hedges et al.
1992). Some authors have suggested a close
relationship with soricids (McDowell 1958) or with
the tenrecids (Van Valen 1967). The divergence
from other placental mammals for Solenodon dates
to 76 million years ago, comparable to or older than
the estimated dates of divergence of some
interordinal splits in mammals and considerably
older than the basal divergence of most mammalian
orders (Roca et al. 2004). Based on molecular
analysis, solenodons may turn out to be the earliest
mammalian residents of the Greater Antilles (Roca
et al. 2004). At the present time, solenodons are
highly endangered by deforestation, increasing
human activities, and the negative impacts of
predation by invasive species. Dobson (1882: 82)

TERRESTRIAL MAMMALS OF THE WEST INDIES
396
was the first to raise the subfamily Solenodontinae
to the family level. The Antillean insectivores
were reviewed by McDowell (1958) and more
recently by Whidden and Asher (2001) and
Ottenwalder (2001).
Solenodon Brandt, 1833. Mem. Acad. Imp. Sci.,
St. Petersbourg, ser. 6, 2: 459.
TYPE SPECIES Solenodon paradoxus Brandt,
1833
SYNONYMS Antillogale Patterson, 1962,
Atopogale Cabrera, 1925
COMMENT Most authorities consider the large
insectivores from Cuba and Hispaniola to be part of
the same overall clade, and group them together in
the single genus Solenodon. However, Allen
(1910) felt that there were enough distinct
characters to justify separating them at the
subgeneric level. Cabrera went even further in
1925, splitting off the Cuban forms into the genus
Atopogale. While almost all authorities (except
Hall 1981) have continued to classify Hispaniolan
and Cuban solenodons as distinct species within the
genus Solenodon without subgeneric distinction,
molecular analyses suggest that both living species
could be two different genera (Roca et al. 2004).
Solenodon arredondoi Morgan and Ottenwalder,
1993. Ann. Carnegie Mus., 62: 154.
COMMON NAME Giant solenodon
TYPE LOCALITY Cueva Paredones, 3 km
southwest of Ceiba del Agua, Caimito, La Habana
province, Cuba
DISTRIBUTION Known from three localities in
western Cuba
STATUS Extinct
COMMENT The first account for the species was
published in Morgan et al. (1980) although
unidentified then and believed from the late
Pleistocene. Later, Morgan and Ottenwalder
(1993) working with additional materials
considered S. arredondoi to be more recent, and
attributedits extinction to habitat destruction by
humans and domestic dog predation.
Solenodon cubanus Peters, 1861. Monatsb. Akad.
Wiss. Berlin: 169.
COMMON NAME Cuban solenodon, almiquí
TYPE LOCALITY In Aguilera, on the slope of the
Sierra Maestra, between Cabo Maisí and Cabo
Cruz, eastern Cuba (data from collector and locality
are from the original label on the specimen).
DISTRIBUTION Eastern Cuba.
STATUS Endangered (IUCN 2004)
SYNONYMS ¨Solenodon paradojo¨ Poey, 1838;
S. paradoxus (of Poey 1851); Atopogale cubana
Cabrera, 1925; Solenodon poeyanus Barbour, 1944;
S. cubanus poeyanus Aguayo, 1950; Atopogale
cubanus Kratochvíl, 1976.
COMMENT This species is currently restricted to
a few localities in Alejandro de Humboldt National
Park and in the Sierra del Cristal. Fossil records
have been found throughout Cuba.
Solenodon marcanoi (Patterson, 1962). Breviora,
165: 2.
COMMON NAME Marcano’s solenodon
TYPE LOCALITY Nameless cave two km SE of
Rancho La Guardia, Hondo Valle, Elias Piña
Province, Dominican Republic
DISTRIBUTION Known from the massifs of La
Hotte and La Selle in Haiti, and from Sierra de
Neiba in the Dominican Republic
STATUS Extinct
SYNONYMS Antillogale marcanoi Patterson,
1962.
COMMENT This species apparently survived long
after European colonization. Woods and Eisenberg
(1989) postulate that it might still survive; Woods
and Ottenwalder (1992) believe it survived into the
1970’s.
Solenodon paradoxus Brandt, 1833. Mém. Acad.
Imp.Sci., St. Pétersbourg, ser. 6, Sci. Math. Phys.
Nat., 2: 459.
COMMON NAME Hispaniolan solenodon, nez-
long
TYPE LOCALITY Port-au-Prince, Haiti
DISTRIBUTION Hispaniola
STATUS Endangered (IUCN 2004)
Solenodon paradoxus paradoxus Brandt, 1833.
Mém. Acad. Imp.Sci., St. Pétersbourg, ser. 6,
Sci. Math. Phys. Nat., 2: 459.
COMMON NAME Hispaniolan solenodon,
nez-long, nen-long
TYPE LOCALITY Port-au-Prince, Haiti
DISTRIBUTION Dominican Republic, north
of the Neiba Valley. Apparently this subspecies
is a recent invader to the south island of
Hipaniola (see Ottenwalder 2001)
STATUS Endangered (IUCN 2004)
Solenodon paradoxus woodi Ottenwalder,
2001. Biogeography of the West Indies:
Patterns and Perspectives: 299.
COMMON NAME Hispaniolan solenodon,
nez-long, nen-long
TYPE LOCALITY Bucan de Tuí, Península
de Barahona, Pedernales Province, Dominican
Republic
DISTRIBUTION South Hispaniola; including
Peninsula de Barahona and Sierra de Baoruco
in Dominican Republic and Tiburon Peninsula,
Haiti
STATUS Endangered (IUCN 2004)

UPDATED CHECKLIST – Borroto-Páez, Mancina, Woods and William Kilpatrick
397
COMMENT This subspecies is distributed in
Haiti and southwestern Dominican Republic,
although it is a possible invader of the North
Island in the Dominican Republic
(Ottenwalder 2001).
Gen. and sp. indeterminate
COMMENT Indeterminate Solenodontid
(Nesophontes-like sized insectivore) from La Toca
mines, La Cumbre, northern Dominican Republic.
Thematerial represents the first mammal bones
preserved in Dominican amber, and consists of six
vertebrae and portions of several ribs in partial
articulation. Amber dated from this region
indicates that it cannot be older than late
Oligocene/early Miocene (MacPhee and Grimaldi
1996).
Order Primates Linnaeus, 1758
The first account of a “fossil” monkey in the West
Indies was reported in 1888, and described by
Ameghino (1910a) as Montaneia anthropomorpha.
Miller (1916b) considered itindistinguishable from
Ateles. Arredondo and Varona (1983) argued the
species validity. Radiocarbon dating of the canine
determined the holotype age to be approximately
70 ± 70 yrBP. In conclusion the teeth belonged to
a living species of Ateles (MacPhee and Rivero de
la Calle 1996). However, recent evidence has
demonstrated the presence of monkeys in the West
Indies in the early Miocene based on a specimen
from Cuba (MacPhee and Iturralde-Vinent 1995a,
1995b). During the late Quaternary, endemic
platyrrhines lived on three islands of the Greater
Antilles (Cuba, Hispaniola and Jamaica) (MacPhee
1996). They became extinct during the early
European colonization period (MacPhee and
Flemming 1999). Authorities are still working on
the phylogeny of Antillean monkeys as new
materials are being discovered (Rivero and
Arredondo 1991; Horovitz and MacPhee 1999;
MacPhee et al. 1995) for example, an Eocene fossil
petrosal from Seven Rivers, Jamaica is suspected to
belong to a primate (MacPhee et al. 1999a);
however, MacPhee (personal communication) no
longer considers the evidence sufficient to assign
the bone to a primate. For a review of Greater
Antillean monkeys see Horovitz and MacPhee (this
volume).
Infraorder Platirrhini Geoffroy Saint-Hilaire,
1812
Superfamily Ateloidae Rosenberger, Setoguchi
and Shigehara, 1990
Family Pitheciidae Mivart, 1865. Proc Zool Soc
Lond 1865: 547.
Subfamily Callicebinae Pocock, 1925. Proc.
Zool. Soc. Lond., 43.
Tribe Xenotrichini (Hershkovitz, 1970).
Folia Primat., 12: 3.
MacPhee and Horovitz (2004) proposed the
new tribe Xenotrichinito place the three Greater
Antillean taxa.
Antillothrix MacPhee, R. D. E., I. Horovitz, O.
Arredondo, and O. Jiménez Vázquez. 1995.
American Museum Novitates 3134: 3.
TYPE SPECIES Antillothrix bernensis (Rímoli,
1977)
COMMENT MacPhee and Woods (1982)
described additional materials and questioned the
affinity with Saimiri. MacPhee et al. (1995)
reviewed the status of “Saimiri” bernensis and
concluded that the specimen is distinctive enough
to be placed in the new genus Antillothrix. A
cladistics analysis based on craniodental characters
indicates an affinity with Paralouatta varonai.
Antillothrix bernensis (Rímoli, 1977). Cuad. del
CENDIA 242: 8
COMMON NAME Hispaniolan monkey
TYPE LOCALITY Cueva de Berna near Boca de
Yuma, 18°23’N and 68°35’W, La Altigracia,
Dominican Republic
DISTRIBUTION Known only from the type
locality
STATUS Extinct
COMMENT Described as Saimiri, Rosenberger
(1978) doubted the validity of the genus. MacPhee
et al. (1995) placed it in Antillothrix. See Horovitz
and MacPhee (this volume).
Insulacebus Cooke, S. B., A. L. Rosenberger and
S. Turvey. 2011. Proceedings National Academy
of Sciences 108: 2699-2704.
TYPE SPECIES Insulacebus toussaintiana (Cooke,
Rosenberger and Turvey, 2011)
COMMENT Cooke et al. (2011) discuss the
morphological features of this form as well as
Xenothrix and Antillothrix in the hope of resolving
their possible affinities. They note a possible close
affinity between Xenothrix and this newly
described form Insulacebus.
Insulacebus toussaintiana (Cooke, Rosenberger
and Turvey, 2011). Proc. Nat. Acad. Sci. 108:2699
COMMON NAME La Hotte Monkey
TYPE LOCALITY Trouing Jéremie number 5, a
sinkhole on the Plain of Formon, Department du
Sud, 17 km west of Camp Perrin in far
southwestern Haiti (18°20’N and 74°03’W).

TERRESTRIAL MAMMALS OF THE WEST INDIES
398
DISTRIBUTION Known only from the type
locality
STATUS Extinct
COMMENT Collected by Charles Woods and
Daniel Cordier in a sinkhole along with the very
locally distributed rodent Rhizoplagiodontia lemkei
and sloth Neocnus toupiti. These forms have not
turned up in other parts of Hispaniola, and may
have been true endemics to the southern peninsula
of Haiti, and particularly the upland forests of the
Massif la Hotte. Neocnus toupiti is the smallest
known West Indian sloth and may have been quite
squirrel-like (White, this volume). McAfee (2011)
concluded that this for was a small arboreal
folivore. Rhizoplagiodontia may have been quite
arboreal as well.
Paralouatta Rivero de la Calle and Arredondo,
1991. Jour. Hum. Evol. 21:1.
TYPE SPECIES Paralouatta varonai Rivero de la
Calle and Arredondo, 1991
COMMENT Important characters are the relative
large size of the orbits and the position of foramen
magnum.
Paralouatta varonai, Rivero de la Calle and
Arredondo, 1991. Jour. Hum. Evol. 21:1.
COMMON NAME Galeran monkey, Cuban
monkey
TYPE LOCALITY Cueva del Mono Fósil, located
at the slope South of Sierra de Galeras, Cordillera
de Guaniguanico, Municipality of Viñales in Pinar
del Río, Cuba
DISTRIBUTION Known only from the type
locality and Cueva Alta
STATUS Extinct
COMMENT A late Quaternary monkey judging by
the associated fauna and considered semi terrestrial
(MacPhee and Meldrum 2006). See Horovitz and
MacPhee (this volume).
Paralouatta marianae MacPhee, Iturralde-Vinent
and Gaffney, 2003. Amer. Mus. Novitates 3394: 15
COMMON NAME Zaza monkey.
TYPE LOCALITY Domo de Zaza, in deposit near
south end of section, south-central Cuba. Sancti
Spíritus
DISTRIBUTION Known only from the type
locality
STATUS Extinct
COMMENT Early Miocene Monkey (MacPhee et
al. 2003) described from a large astragalus. It is the
indeterminate genus and species A of McPhee and
Iturralde-Vinent (1995b).
Xenothrix Williams and Koopman, 1952. Am.
Mus. Novitates 1546: 12.
TYPE SPECIES Xenothrix mcgregori Williams
and Koopman, 1952.
COMMENT Important characters of the skull are
the substantial width of the nasal fossa and
maxillary sinus, which extend into the roots of the
zygomatic arch.
Xenothrix mcgregori Williams and Koopman,
1952. Am. Mus. Novitates 1546: 12, 1952
COMMON NAME Jamaican monkey
TYPE LOCALITY Long Mile Cave, Trelawney
Parish, Jamaica
DISTRIBUTION Long Mile cave, Lloyd’s Cave.
Skeleton Cave, Somerville Cave, Jamaica
STATUS Extinct
COMMENT See Horovitz and MacPhee (this
volume). Considered a large and slow-moving
arboreal monkey. Species remains are associated
with Rattus suggesting that this monkey survived
into European times (MacPhee and Flemming
1997).
Platyrrhine gen. et sp. Indeter. 1.
COMMENT Undescribed species from Coco Ree
in St. Catherine, Jamaica.
Platyrrhine gen. et sp. Indeter. 2.
COMMENT Undescribed species from Sheep Pen
in Trelawny, Jamaica.
COMMENT Femora of both of the undescribed
species listed above differ from the primate femur
assigned to Xenothrix (Ford and Morgan 1986;
Ford 1990; MacPhee and Fleagle 1991). Until
platyrrhine dentition materialis recovered to
unambiguously differenciate from Xenothrix, these
intriguing specimens are left as indeterminate (see
Horovitz and MacPhee, this volume).
Order Carnivora Bowdich, 1821
Familia Canidae Gray, 1821
The existence of endemic carnivores in the West
Indies has been debated for many years. Two
species have been described in Cuba: Cubacyon
transversidens Arredondo and Varona, 1974, from
Cueva del Túnel, La Salud, Habana (the single type
material has been lost from the collection) and
Indocyon caribensis (Arredondo 1981), from Cueva
de Bélica, Güirabo, Holguín originally described as
Paracyon caribensis (Arredondo 1981a), generic
preoccupied and later corrected (Arredondo 1981b).
The latter taxon is frequently found in burial remains
throughout Cuba. Dog remains in the West Indies
are known from burial sites of the Dominican
Republic, Cuba, Puerto Rico, Jamaica, Barbados and
the Bahamas (Lawrence 1977; Wing 1991; Wing,
this volume) and are sporadically found in Caicos,
the Lesser Antilles (Martinique, Anguilla, Antigua)
and the Virgin Islands (see Wing, this volume). The
first European chroniclers documented the presence
of mute dogs in the life of Antillean aborigines.

UPDATED CHECKLIST – Borroto-Páez, Mancina, Woods and William Kilpatrick
399
However, the validity of these two species has been
questioned and frequently they are synonymized as
Canis lupus familiaris (Morgan and Woods 1986).
Domestic dogs are known to be highly variable in
size and morphology. However, more recently a
fossil hair preserved in amber from the Dominican
Republic has been attributed to a carnivore (Poinar
and Poinar 1999), indicating that the possible
presence of West Indian endemic carnivores must be
reconsidered.
Order Perissodactyla, Owen, 1848.
Family Rhinocerontidae Gray, 1821. London
Med. Repos., 15: 306.
Hyrachyus Leidy, 1871. Proc. Acad. Nat. Sci. Phil.
23: 229.
TYPE LOCALITY Near Seven Rivers, Parish of
St. James
DISTRIBUTION Early or middle Eocene of
Jamaica
STATUS Extinct
COMMENT The specimen is a right dentary,
considered to be at least 12 million years older than
the next oldest Antillean mammal. The presence of
Hyrachyus in Jamaica late early to early middle
Eocene is the earliest evidence of colonization of
the Greater Antilles region by land mammals
(Domning et al. 1997; see also Portell et al. 2001).
However, there is evidence that the entire island of
Jamaica was inundated from middle Eocene to the
lateMiocene (Robinson 1994). The genus
Hyrachyus is well known from the Eocene of
Eurasia and North America (McKenna and Bell
1997). Order Rodentia Bowdich 1821
Rodentia is the most diverse order of terrestrial
mammals in the West Indies.
Suborder Myomorpha Brandt, 1855
Family Muridae Illiger, 1815. Abhandl. K. Akad.
Wiss. , Berlin for 1804-11, p. 46, 129.
Subfamily Sigmodontinae Wagner, 1843. In
Schreber, Dien Säugethiere, Suppl., 3: 398.
COMMENT The two genera Megalomys and
Oryzomys represent two distinct dispersals into the
West Indies. According to Musser and Carleton
(2005) Megalomys has a suite of characters that
associate the taxon with the Sigmodontinae sensu
stricto, and with the Oryzomyini in particular.
Fossil information of this radiation in the Lesser
Antilles are scattered [see G. M. Allen (1942) for
documentation of the historical records].
Megalomys Trouessart, 1881. Le Naturaliste 1: 357
TYPE SPECIES Mus desmarestii Fischer, 1829
SYNONYMS Moschomys (of Trouessart 1903),
Moschophoromys (of Elliot 1904)
COMMENT Originally named as a subgenus, and
associated with Oryzomys by Major (1901). It is
typically recognized as an oryzomyine genus
endemic to the Lesser Antilles (Ellerman 1941; Tate
1932; Hall 1981). Both species are extinct but
persisted perhaps until the late 1800s (Ray 1962;
Woods 1989a).
Megalomys audreyae Hopwoods, 1926. Ann Mag.
Nat. Hist. ser. 9, 17: 329
COMMON NAME Barbuda giant rice rat, Barbuda
musk-rat
TYPE LOCALITY Cave in the Island of Barbuda,
West Indies
DISTRIBUTION Known only from the type
locality.
STATUS Extinct
SYNONYMS M. majori (of Trouessart, 1904)
COMMENT Known only from a mandibular
fragment and an upper incisor. The extinction has
been attributed to European colonization on the
island. The species is considered extinct but is not
listed by Musser and Carleton (2005).
Megalomys desmarestii (Fischer, 1829). Synopsis
Mamm., 316
COMMON NAME Martinique giant rice rat,
Martinique muskrat, pilorie
TYPE LOCALITY Martinique, Lesser Antilles
DISTRIBUTION Known only from the type
locality
STATUS Extinct
SYNONYMS pilori (of Rochefort 1658, Geoffroy
and Cuvier 1830), piloris (of Zimmermann 1777,
Forsyth Major 1901), and pilorides (of Desmarest
1826, Trouessart 1881) (see Forsyth Major 1901;
Friant 1941; Allen 1942)
COMMENT Described as Mus desmarestii. Its
extinction has been attributed to the 1902 volcanic
eruption of Mont Pelé (Nowak 1999).
Megalomys luciae Forsyth Major, 1901. Ann. Mag.
Nat. Hist. ser. 7, 7: 206.
COMMON NAME St. Lucia giant rice rat, St.
Lucia muskrat
TYPE LOCALITY St. Lucia, Lesser Antilles
DISTRIBUTION Known only from the type
locality
STATUS Extinct
COMMENT Extinction occurred in the 19th
Century. One captive animal may have lived from
1849 to 1852 at the London Zoo (Nowak 1999).
Megalomys sp.1
Small extinct oryzomyine rodent from Montserrat,
Anguilla, St. Eustatius and St. Kitts (Steadman et al.

TERRESTRIAL MAMMALS OF THE WEST INDIES
400
1984; Morgan and Woods 1986; Woods 1989a,
1990).
Megalomys sp. 2
Large extinct oryzomyine rodent from Montserrat,
Anguilla, Barbuda, Guadeloupe, and Marie Galante
(Steadman et al. 1984; Morgan and Woods 1986;
Woods 1989a, 1990).
Oryzomys Baird, 1858. Mammalia in Repts. U. S.
Expl. Surv., 8, 1: 458.
TYPE SPECIES Mus palustris Harlan, 1837.
Oryzomys couesi (Alston,1877). Proc. Zool. Soc.
Lond., 1876: 756 [1877].
COMMON NAME Coues’ oryzomys.
TYPE LOCALITY Guatemala, Alta Verapaz
Dept., Cobán.
Oryzomys couesi antillarum (Thomas, 1898).
Ann. Mag. Nat. Hist., ser 7, 1: 177.
COMMON NAME Jamaica rice rat
TYPE LOCALITY Jamaica
DISTRIBUTION Known only from the type
locality
STATUS Extinct
SYNONYMS Oryzomys antillarum Thomas,
1898
COMMENT Considered Oryzomys antillarum by
Allen (1942), Varona (1974b), Woods (1989a),
Woods et al. (2001). The extinction is estimated
to the 1880s, five years after the introduction of
the mongoose (Herpestes auropunctatus) (Allen
1911, 1942).
Oryzomys sp. 1
COMMENT Undescribed extinct species from
Barbados (Marsh 1985)
Oligoryzomys Bangs, 1900. New England Zool.
Club., 194.
TYPE SPECIES Oryzomys navus Bangs 1900
(= Hesperomys fulvescens Sausurre, 1860)
COMMENT Described as a subgenus of Oryzomys
but the diagnosis was amended at the generic level
by Carleton and Musser (1989).
Oligoryzomys victus (Thomas, 1898). Ann, Mag.
Nat. Hist., ser 7, 1: 178.
COMMON NAME Pygmy rice rat
TYPE LOCALITY Lesser Antilles, Saint Vincent
DISTRIBUTION Known only from the type
locality
STATUS Extinct
COMMENT Considered Oryzomys victusby
Ellerman (1941), Goldman (1918), Woods (1989a)
and Woods et al. (2001). Thomas (1898) and Ray
(1962) emphasized its relationship with
Oligoryzomys, and Musser and Carleton (1993)
made this new combination.
Suborder Hystricomorpha Brandt, 1855
Infraorder Hystricognathi Tullberg, 1899
The Infraorder Hystrcognathi is the most diverse
group of terrestrial mammals in the West Indies.
Family Echimyidae Gray, 1825. Ann. Philos., n.s.,
10:341.
This family in addition to being the most diverse is
also the most ancientwithin the hystricognath,
dating from the early Oligocene of Patagonia. The
endemic West Indiansubfamily Heteropsomyinae
derived from this massive South American group,
and the Capromyidae may also originate from
echimyids. Time and location of their radiation
and relationships between the two groups is a
matter of interest and future resolution.
Subfamily Heteropsomyinae Anthony, 1917.
Bull. Am. Mus. Nat. Hist., 37(4):18.
Named by Anthony to reflect perceived
relationship between Heteropsomys and
Dasyprocta, it was later expanded by Kraglievich
(1965) and Patterson and Pascual (1968b) to
include Proechimys and associated genera.
Modified by Woods (1982:386) wherein West
Indian Spiny rats were classified with the
Capromyidae. McKenna and Bell (1997) included
the genera of the Eumysopinae within the
Heteropsomyinae. West Indian spiny rats are
transitional in characters between Echimyidae and
Capromyidae, and here are placed in their own
subfamily within the Echimyidae until futher
revisions. Molecular data of Leite and Patton
(2002) suggest that the sister taxa of both the
myocastorids and capromyids is a group of the
Eumysopinae that includes the genera Clyomys and
Euryzygomatomys, but do not allow an evaluation
of the distinctiveness of a West Indian clade of
Spiny rats (Heteropsominae) suggested by Woods
(1982). See Kilpatrick et al. (this volume) for
another molecular approach. Varona (1974b)
placed all West Indian spiny rats in the genus
Heteropsomys. However, there are clear
differences between the forms on each island, so
the original generic names are maintained, except
for Homopsomys which is combined with
Heteropsomys (Woods 1989a, b). All species in
the West Indies became extinct after the
introduction of rats, dogs and cats by Europeans.
Boromys Miller, 1916. Smithson. Misc. Coll.,
66(12):7.
TYPE SPECIES Boromys offella Miller, 1916.
SYNONYMS Heteropsomys (of Varona, 1974b),
Geoboromys Arredondo, 1958

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401
COMMENT Characteristics of the genus are:
mesopterygoid fossa (palatine notch) moderate and
extending to the level of the second upper molar,
incisive foramen moderate in size and without a
slight lip, postorbital process lacking.
Boromys offella Miller, 1916. Smithson Misc.
Coll., 66(12):8.
COMMON NAME Oriente cave rat, larger Cuban
spiny rat
TYPE LOCALITY Cuba, Oriente Province,
Baracoa, Maisí
DISTRIBUTION Cuba and Isla de la Juventud
(Isle of Pines)
STATUS Extinct
COMMENT Large Boromys.
Boromys torrei Allen, 1917. Bull. Mus. Comp.
Zool., 61:6.
COMMON NAME Torre’s cave rat, small Cuban
spiny rat
TYPE LOCALITY Cuba, Matanzas Province,
Cave in Sierra de Hato-Nuevo
DISTRIBUTION Cuba and Isla Juventud (Isle of
Pines)
STATUS Extinct
COMMENT Small Boromys.
Brotomys Miller, 1916. Smithson. Misc. Coll.,
66(12):6.
TYPE SPECIES Brotomys voratus Miller, 1916
SYNONYMS Heteropsomys (of Varona, 1974b)
COMMENT Mesopterygoid fossa shallow and
extending to the level of the middle of the third
upper molar tooth. The incisive foramen is very
narrow and without a lip. The postorbital process
is slight.
Brotomys contractus Miller, 1929. Smithson Misc.
Coll., 81(9):13.
COMMON NAME Haitian edible rat, Hispaniola
spiny rat, mohuy
TYPE LOCALITY Haiti, Département de
l’Artibonite, St. Michel de l’Atalye (small cave)
(19o22’N and 72o20’W)
DISTRIBUTION Hispaniola
STATUS Extinct
COMMENT Frontopalatal depth less than 88% of
interorbital breadth.
Brotomys voratus Miller, 1916. Smithson Misc.
Coll., 66(12):7. = Miller 1916a
COMMON NAME Hispaniolan edible rat,
Hispaniola spiny rat, mohuy
TYPE LOCALITY Dominican Republic, San
Pedro de Macoris, kitchen midden
DISTRIBUTION Hispaniola (Haiti and Dominican
Republic), and La Gonave Island
STATUS Extinct (within the last 60 years)
COMMENT Frontopalatal depth more than 88% of
interorbital breadth.
Heteropsomys Anthony, 1916. Ann. New York
Acad. Sci., 27:203.
TYPE SPECIES Heteropsomys insulans Anthony,
1916.
SYNONYM Homopsomys Anthony, 1917.
COMMENT Includes Homopsomys; see comments
under the subfamily. The two species from Puerto
Rico are much larger in body size than
heterosomyines from Hispaniola and Cuba (size
approaching Plagiodontia aedium), and the
probable explanation is that capromyid rodents did
not naturally occur in Puerto Rico and the species
evolved without competition and fill hutia like
niches (Woods 1996). A character of the genus is
the upper cheek teeth have a single conspicuous
internal fold with three separate, transverse,
enamel-surrounded lakes.
Heteropsomys antillensis (Anthony, 1917). Bull.
Am. Mus. Nat. Hist., 37:187.
COMMON NAME Antillean cave rat, large Puerto
Rican spiny rat
TYPE LOCALITY Puerto Rico, Cave at Utuado
DISTRIBUTION Puerto Rico
STATUS Extinct
COMMENT Described as Homopsomys.
Heteropsomys insulans Anthony, 1916. Ann New
York Acad. Sci., 27:202.
COMMON NAME Puerto Rican cave rat, hutia
like Puerto Rican spiny rat
TYPE LOCALITY Puerto Rico, Cueva de la
Ceiba, Hacienda Jobo (near Utuado)
DISTRIBUTION Puerto Rico and Vieques Island
in cave deposits
STATUS Extinct
COMMENT Nearly as large in body size as the
Hispaniolan hutia Plagiodontia aedium.
Puertoricomys Woods, 1989, Biogeography of the
West Indies, p. 748.
TYPE SPECIES Proechimys corozalus Williams
and Koopman, 1951.
COMMENT Characters of the genus include: teeth
with one long fold passing entirely across occlusal
surface; lower incisor very deep with width less
than 50% of depth.
Puertoricomys corozalus (Williams and Koopman,
1951). Am. Mus. Novit., 1515: 4.
COMMON NAME Old Puerto Rican spiny rat
TYPE LOCALITY Puerto Rico, Corozal
Limestone Cave
DISTRIBUTION Known only from the type
locality
STATUS Extinct

TERRESTRIAL MAMMALS OF THE WEST INDIES
402
COMMENT Known only from a fragmentary
mandible. Specimen is mineralized and associated
with old looking material, and may be older than
Pleistocene. Described as Proechimys corozalus
but placed in its own monotypic genus
Puertoricomys by Woods (1989a).
Family Capromyidae Smith, 1842. The
Naturalist’s Library (London), 15:308.
Excludes Myocastor, and apparently represents an
entirely intra-Caribbean radiation (Woods and
Howland 1979; Woods 1989a, b). Woods
(1982:386-387) included West Indian
Heteropsomyinae in this family, but here this group
is assigned to the Echimyidae (see comments under
Heteropsomyinae). Varona (1974b), and Corbet
and Hill (1991:203) classified most of the
following species in either of two genera
(Capromys or Plagiodontia), whereas Woods
(1989a, b) separated them into several distinct
subfamilies and genera. Fossil records of the
¨Capromyidae¨ occurring during middle Miocene
in Quebrada Honda, Bolivia, South America
(MacFadden and Wolf 1981; McKenna and Bell
1997) is not well supported.
Subfamily Capromyinae Smith, 1842. The
Naturalist’s Library (London), 15:30.
Reviewed by Kratochvíl et al. (1978), Varona and
Arredondo (1979), Borroto-Páez (2002) and
Borroto-Páez et al. 2005). Borroto-Páez and
Borroto-Páez et al. provided synonymies and group
all the subgenera of Capromys, Geocapromys,
Mesocapromys, Mysateles, which are accepted as
valid by the most recent revisions (Kratochvíl et al.
1978; Woods and Kilpatrick 2005). On the basis of
molecular analyses Woods et al. (2001), Borroto-
Páez et al. (2005) and Kilpatrick et al. (this
volume) obtained clades that are in concordance
with the four proposed genera and demonstrated
that Mysateles is paraphyletic, and placed
melanurus within the genus Mesocapromys. There
are seven forms to be described, three species from
Cayman Islands and four subspecies from Cuba.
Silva-Taboada et al. (2007) estimate several species
are synonyms.
Capromys Desmarest, 1822. Bull. Sci. Soc. Philom.
Paris, p. 185.
TYPE SPECIES Capromys fourniere Desmarest,
1822 (= Isodon pilorides Say, 1822)
SYNONYMS Procapromys Chapman, 1901,
Macrocapromys Arredondo, 1958, Paleocapromys
Varona and Arredondo, 1979
COMMENT “Capromys” geayi (Pousarges, 1899)
was described from Venezuela, where it was
collected in the mountains between La Guayra and
Caracas. It was placed in its own genus
(Procapromys) by Chapman (1901:322). The type
specimen is Mus. Nat. Hist. Nat. (Paris) No. 1898-
1785 (1834A). No further specimens had been
collected, and it is possible that the collection
locality was incorrectly recorded. This specimen is
likely a juvenile Capromys from Cuba erroneously
reported from Venezuela. Characters of the genus
include: the ratio tail/head and body is 0.50, tail is
not prehensile, liver multilobulated with reticulate
appearance, blackish skin in palm and genitals,
small corpora lutea (mean of 1.6 mm), ratio vertical
diameters preorbital/orbital larger than one,
preorbital arc is wide, larger body size. Mass is
3,783g, but old adults can weigh up 6 kg (Borroto-
Páez 2002; Borroto-Páez et al. 2005).
Silva-Taboada et al. (2007) maintain the genus
Macrocapromys (of Arredondo, 1958) with two
species: M. acevedo (including C. antiquus as a
synonym) and M. latus (including C. robustus). We
believe that characters used are not sufficiently
distinct to separate these species from Capromys at
the generic level. Although, Silva-Taboada et al.
(op. cit.) studied more material assigned to
Macrocapromys, it is still fragmentary (two
fragmented crania and the hemi-mandibules). All
measurements of the material assigned to
Macrocapormys are within the range of Capromys.
Robust skeletal elements and large mass are
common among fossil specimens of Capromys
pilorides (see data in Silva-Taboada et al. op cit.).
Capromys acevedo Arredondo 1958. El Cartero
Cubano 17 (12):10.
COMMON NAME Acevedo’s hutia
TYPE LOCALITY Cueva Lamas, about 2 km SW
of Playa Santa Fé, La Habana
DISTRIBUTION Cuba
STATUS Extinct
SYNONYMS acevedoi Varona 1974; antiguus
Varona and Arredondo 1979; artiguus Woods 1989
STATUS Extinct
COMMENT See comment in Capromys. Described
as Macrocapromys acevedo (Arredondo 1958).
Silva-Taboada et al (2007) support the validly of
Macrocapromys but the change could be
premature. The species could be a synonymous
with pilorides. The additional material assigned to
acevedo in Silva-Taboada et al. (2007) are in the
range of C. pilorides. Arredondo's spelling was
acevedo, but this was noted as [sic] by Varona
(1974b) who changed the spelling to acevedoi.
This spelling was followed by Hall (1981) but it
should be noted that acevado is the correct usage
according to Brodkorb (1961).
Capromys garridoi Varona, 1970. Poeyana, ser. A,
74:2.
COMMON NAME Garrido’s hutia

UPDATED CHECKLIST – Borroto-Páez, Mancina, Woods and William Kilpatrick
403
TYPE LOCALITY Cayo Majá, northwestern
extremity of Cayo Largo, Banco de los Jardines y
Jardinillos, Los Canarreos Archipelago, Cuba. See
comments.
DISTRIBUTION Known only from a single
specimen from a small unnamed islet northwest of
Cayo Largo, Cuba
STATUS Critically endangered (IUCN 2008)
COMMENT Placed in the genus Mysateles,
subgenus Leptocapromys by Kratochvíl et al.
(1978:15), retained as Capromys by Varona
(1974b) and Woods et al. (2001) and considered
Mysateles by Borroto-Páez (2002) and Borroto-
Páez et al. (2005). Silva-Taboada et al (2007)
consider it is a non-adult Capromys pilorides and
species inquerenda. We believe that the specimen
is an adult since the M3 is completely erupted but
nearly unworn. Borroto-Páez et al. (2005) by
multivarite analyses of principal components of 33
cranial characters (without mandible) found more
similarities with Mysateles and the validity of the
species. The analyses included subadult and young
adult specimens of Mysateles prehensilis (5) and
Capromys pilorides (4) and results demonstrated
that garridoi was distinct. The blackish skin in
palm and genitals pointed out in the description is
considered here to maintain in Capromys. The
descriptions of excrement pellets assigned to
garridoi are peculiar and differ from pellets
produced by Capromys pilorides and Mysateles
prehensilis (Garrido 1971). Similar excrement
pellets were found on Cayo Largo del Sur during
the 1990 expedition. Only one specimen (skull
without mandible) is in the collection.
The exact location of the type locality is
unclear. The small key without a name where the
animal was found is not part of Cayos Majáes, but
rather faces a small pier in the northwestern part of
Cayo Largo in the Canarreo Archipelago. Varona
(1970a) incorrectly applied the name Cayo Majá to
this islet. It is known that this hutia does not occur
on the adjacent Cayo Majá (three cayos to the
northwest of Cayo Largo) based on a 1990
expedition to this region by R. Borroto-Páez. The
single known specimen may not be from the area,
but was probably collected by fishermen and left on
site. Additional research would be required to find
more specimens, determine the distribution and
status of this species and expand the analysis.
Capromys latus Varona and Arredondo, 1979.
Poeyana 195:5.
COMMON NAME Robust hutia
TYPE LOCALITY Cueva Lamas, about 2 km SW
of Playa Santa Fé, Havana City
DISTRIBUTION Type locality; Cueva de Pio
Domingo, Sumidero, Pinar del Río; Cueva del
Tunel and Cueva Insunsa, La Salud, Habana;
Cueva de Paredones, San Antonio de los Baños,
Habana
STATUS Extinct
SYNONYMS robustus Varona and Arredondo,
1979
COMMENT The synonymy of robustus is based
on the following reasons: description is based on
very fragmentary material, insufficient
comparisons with other species, distinctiveness of
many of the characters are not convincing,
measurements of some characters are in the range
of variation of C. pilorides. Silva-Taboada et al
(2007) proposed a new combination with
Macrocapromys latus (including C. robustus) and
arredondoi as synonym of C. pilorides. Varona
and Arredondo (1979) and Varona (1980a, 1984)
consider robust molariform teeth to be
characteristic of the majority of fossil species.
Fossil specimens of Capromys from Cayman
Islands also have robust characteristics of the skull.
Capromys pilorides (Say, 1822). Jour. Acad. Nat.
Sci. Philadelphia, 2:333.
COMMON NAME Desmarest’s hutia, conga hutia
TYPE LOCALITY South America or one of the
West Indian islands
DISTRIBUTION Island of Cuba, Isla de la
Juventud, many islands and keys of the Doce
Leguas, Sabana and Cuban archipelagos
STATUS Common (extremely abundant in some
areas such as keys, coastal and mangrove zones.
SYNONYMS arredondoi Varona, 1984; fourniere
Desmarest, 1822; geayi Pousarges, 1899;
intermedius (Arredondo, 1958), megas (Varona y
Arredondo 1979), pappus Varona, 1984 (see
comment in C. p. relictus). The subspecies C. p.
relictus Allen, 1911; C. p. doceleguas Varona,
1980; C. p. gundlachianis Varona, 1983; C. p.
ciprianoi Borroto-Páez, Camacho and Ramos,
1992.
COMMENT This species was first mentioned as
Mus pilorides Pallas, 1778; Tate (1935) noted that
it is not associated with this genus. Sometimes
placed in the subgenus Capromys (see Hall
1981:863). This species is very variable in cranial
and mandibular characteristics, size, coloration, and
habits. There are five named subspecies: ciprianoi,
doceleguas, gundlachianus, pilorides, and relictus
(see Allen 1911; Varona 1980b, 1983a; Borroto-
Páez et al. 1992). No genetic differentiation of
cytochrome b was reported for the two subspecies
ciprianoi and relictus on the Isla de la Juventud
(Woods et al. 2001). See Díaz-Franco (2001) for
synonymy of megas and Silva–Taboada (2007) for
synonymy of arredondoi. See synonymy of
pappus in C. p. relictus. Karyptype has 2n=40;
FN=64 (Hsu and Benirschke 1971; Milišnikov et
al. 1990).

TERRESTRIAL MAMMALS OF THE WEST INDIES
404
Capromys pilorides ciprianoi Borroto-Páez,
Camacho and Ramos, 1992. Misc. zool.
hungarica 7: 98.
COMMON NAME Cipriano's hutia, Conga
hutia from the south of Isla de la Juventud
TYPE LOCALITY Punta del Este, South of
Isla de la Juventud, Municipio Especial Isla de
la Juventud
DISTRIBUTION South of Isla de la Juventud.
The Lanier Swamp is the northern limit.
STATUS Common, but not evaluated by
IUCN.
COMMENT Woods et al. (2001) consider as C.
p. relictus based on limited molecular
divergence and Woods and Kilpatrick (2005)
consider it synonymous with C. pilorides.
However, Borroto-Páez et al. (2005) justified
maintaining it as a distinct subspecies based on
morphological differences. The population has
decreased in the last few years because of
hunting.
Capromys pilorides relictus Allen, 1911. Bull.
Mus. Comp. Zool. 54:207.
COMMON NAME Conga hutia from the north
of Isla de la Juventud
TYPE LOCALITY Casas Montain, Nueva
Gerona, Ile of Pines (now Isla de la Juventud),
Cuba
DISTRIBUTION North of Isla de la Juventud,
the northern border of the Lanier Swamp is the
southern limit.
STATUS Not assessed by IUCN. According to
Borroto-Páez and Ramos (this volume) it is
critically endangered, because of reduced
distribution and limited or small population
size.
SYNONYM pappus Varona, 1984.
COMMENT Initially all specimens from Isla
de la Juventud were believed to be C. p.
relictus. In the description of C. p. ciprianoi
Borroto-Páez et al., 1992, northern and southern
forms are distinct. Silva-Taboada et al. (2007)
propose pappus as a synonym. We agree for
the following reasons: the type is a mandible of
a young adult; in the hypodigm only a
fragmentary mandible, and few molars and
incisors are included (no part of skull is
present). The characters considered in the
diagnosis are not convincing; some comparable
measurement values of the mandible are in the
range of variation C. pilorides (sensu lato) and
C. p. relictus (Kratochvíl et al. 1978; Borroto-
Páez 2002; Borroto-Páez et al. 2005), and
known only from the type locality.
Capromys pilorides doceleguas Varona, 1980.
Mem. Soc. Cien. Nat. La Salle, 40 (114):142.
(Varona 1980d)
COMMON NAME Conga hutia from Doce
Leguas
TYPE LOCALITY Miraflores, Cayo Anclitas,
Doce Leguas Archipelago, Camagüey, Cuba
DISTRIBUTION Doce Leguas Archipelago,
Jardines de la Reina, Camagüey, Cuba, the
following keys: Cabeza del Este, Piedra
Grande, Boca Chica, Camposanto, Anclitas,
Caballones, and Cayo Balandra of the adjacent
Ana María Micro-Archipelago
STATUS Common, but not evaluated by IUCN.
SYNONYMS None
COMMENT The mangrove islets known as
Cayos Salinas, Ana María Archipelago, near
Pasa Seca channel are included. There, C.
pilorides is present and sympatric with
Mesocapromys angelcabrerai, in part of its
known distribution.
Capromys pilorides gundlachianus Varona,
1983. Carib. J. Sci. 19 (3-4):77.
COMMON NAME Conga hutia from Sabana-
Camaguey Archipelago, Mandinga hutia
TYPE LOCALITY In the mangrove of the
narrow channel, Bahia de Santa Clara, west of
Cayo Bahía de Cadiz, 2 km south of the mouth
of Canal Blanca (main channel), Sabana
Archipelago, Cuba
DISTRIBUTION Cayo Bahía de Cádiz region,
in the Sabana Archipelago, keys located NE of
Cardenas, as well as Cayo Fragoso, Cayo Santa
María, Cayo Guillermo, Cayo Patabán, Cayo
Jutía, and Cayo Punta Arena
STATUS Common, but not evaluated by
IUCN. Extirpated on Cayo Las Brujas.
SYNONYM sabana Kratochvíl et al. 1978
COMMENT Erroneously elevated to species
status based on sequence divergence of a
specimen from Cayo Ballenato del Medio
(Woods and Kilpatrick 2005), however, this
locality is outside the published range of
gundlachianus. On Cayo Fragoso, north of
Caibarién, occurs sympatricly with
Mesocapromys auritus. See Capromys pilorides
ssp. 2.
Capromys pilorides pilorides (Say, 1822) Jour.
Acad. Nat. Sci. Philadelphia, 2:333
COMMON NAME Desmarest’s hutia, Conga
hutia
TYPE LOCALITY South America or one of
the West Indian islands
DISTRIBUTION Only on Cuban mainland
(sensu strito)
STATUS Common.
COMMENT See comment in C. pilorides.

UPDATED CHECKLIST – Borroto-Páez, Mancina, Woods and William Kilpatrick
405
Capromys pilorides ssp. 1
Undescribed extant subspecies from Cayo
Campo, Los Canarreos Archipelago, south of
Cuba. The population is considered
autochthonous to forms on other keys where
C. pilorides have been introduced from south of
Isla de la Juventud and Zapata Swamp. This
subspecies is relatively smaller than the other
Demarest's hutias. It should be studied in detail
to be defined. Borroto-Páez and Ramos, (this
volume) propose critically endangered for
conservation status because of the limited
distribution and vulnerability to competition
with introduced monkeys as well as
overhunting.
Capromys pilorides ssp. 2
Undescribed extant subspecies from Cayo
Ballenato del Medio, north of Camagüey, Cuba.
Considered as C. p. gundlachianus in Woods et
al. (2001) and as Capromys gundlachianus in
Woods and Kilpatrick (2005), although outside
of the distribution of C. p. gundlachianus. The
specimen is morphologically identical to C.
pilorides but different in the sequence
divergence in cytochrome b and is considered a
cryptic species (Borroto-Páez et al. 2005). An
authoutonous population of hutias has always
lived at this locality, but on several occasions in
recent times, individuals from Cayo Sabinal
have been introduced onto Cayo Ballenato del
Medio. Therefore the origin of the specimen
from which sequence data is available (Borroto-
Páez et al. 2005; Kilpatrick et al. this volume)
cannot be presently traced. Analyses of
additional samples from Cayo Sabinal and from
Cayo Ballenato del Medio are needed.
However, Kilpatrick et al. (this volume) found
that the sequence from the specimen from Cayo
Ballenato del Medio formed a clade with a
sequence from an animal reported as C.
p.pilorides from a Zoo in the USA (locality of
origin in Cuba unknown.)
Capromys pilorides ssp. 3
Undescribed small extant subspecies from Cayo
Macío and Cayo Diego Perez, south of Zapata
Swamp (Borroto-Páez et al. 2007)
Capromys pilorides ssp. 4
Undescribed large extant subspecies from Cayo
Macío and Cayo Diego Perez, south of Zapata
Swamp (Borroto-Páez et al. 2007)
Capromys sp.1.
Undescribed extinct species from Grand Cayman,
Cayman Brac and Little Cayman, Cayman Islands
(Morgan 1994a). Scudder and Quitmyer (1998)
believe that capromyids from Cayman Brac could
belong to two species of different genera.
Geocapromys Chapman, 1901. Bull. Am. Mus.
Nat. Hist., 14:314.
TYPE SPECIES Capromys (Geocapromys)
brownii Fischer, 1830
SYNONYM Synodontomys
COMMENT Original use of the name was as a
subgenus of Capromys. Included in Capromys by
Mohr (1939:75), Varona (1974b), Hall (1981:865),
and Corbet and Hill (1991:203). Considered a
distinct genus by Miller (1929a), Woods and
Howland (1979:112), and Morgan (1985:30). Two
species are known only from Holocene fossils from
Cuba. Geocapromys is the most widespread genus
in the Capromyidae. Morgan (1985, 1989a) placed
the recent and extinct population of G. ingrahami,
and the extinct species from Cuba and Cayman
Islands in the ingrahami-group, whereas he placed
G. brownii and G. thoracatus in the brownii-group.
This taxonomic arrangement is based on cranial
characters, but all extinct populations from the
Bahamas could represent a closely related species
complex of two or more species. Characters of the
genus include: ratio tail/body size is 0.18 (Borroto-
Páez et al. 2005), short tail, origin of the root of the
upper incisor capsule high on maxilla above the
premolar, broad vertically or posteriorly oriented
superior zygomatic root of maxilla, anteriorly
convergent upper toothrow, 30% inclination of
occlusal surface on all cheek teeth, and presence of
a third anterolingual re-entrant (anteroflexid) fold
on the premolar (Morgan 1989b). The skin of the
palms and genitals is unpigmented, and the liver is
multilobulated and has the usual solid structure.
Geocapromys brownii (Fischer, 1830). Synopis.
Mamm., Addenda, p. 389 (=589).
COMMON NAME Jamaican hutia, Indian coney,
Browne’s hutia
TYPE LOCALITY “Jamaica”
DISTRIBUTION Jamaica
STATUS Vulnerable (IUCN 1996, Turvey and
Dávalos 2008)
SYNONYM Capromys brachyurus (Hill, 1851)
COMMENT Sometimes includes G. thoracatus
from Little Swan Island (see this account).
Reviewed by Anderson et al. (1983). Karyotype
has 2n=88; FN=136 (George and Weir 1972). It
has been found in archaological sites in Holguín,
Cuba (Silva-Taboada et al. 2007).
Geocapromys columbianus (Chapman, 1892).Bull.
Amer. Mus. Nat. Hist. 4: 314.
COMMON NAME Cuban coney, Columbian
hutia, Cuban short-tailed hutia
TYPE LOCALITY Coastal cave in Trinidad,
Sancti Spíritus, Cuba
DISTRIBUTION Cuba, Isla de la Juventud and
some island offshore

TERRESTRIAL MAMMALS OF THE WEST INDIES
406
STATUS Extinct
SYNONYM cubanus Allen, 1917 and
pleistocenicus Arredondo 1958
COMMENT Described as Capromys columbianus.
Allen (1918) recognized that the description
materials used for cubanus belonged to a young
animal and placed it in Geocapromys. Allen
(1917b) established the genus Synodontomys
(= Geocapromys) with this type specimen. Some
remains are post-Columbian. Arredondo (1958)
described Geocapromys columbianus
pleistocenicus without a type locality. In a footnote,
Arredondo (1970b) elevated it to the species level
and provided the type locality (Cueva Lamas, Santa
Fé, La Habana). It is the smallest Geocapromys
species, about the size of a rat. The species was
widely distributed and is abundant in fossil remains
in which some are post-Columbian. See Silva-
Taboada et al (2007) propose the synonymy with
columbianus.
Geocapromys ingrahami (J. A. Allen, 1891). Bull
Am. Mus. Nat. Hist., 3:329.
COMMON NAME Bahamian Hutia
TYPE LOCALITY East Plana Cay, Bahamas
DISTRIBUTION Known from the type locality
and introduced populations on Little Wax Cay
(1973) and Warderick Wells Cay (1981). For
information on introduction and ecology, see
Jordan (this volume). Fossil subspecies are found
in Great Abaco and Crooked Island.
STATUS Vulnerable (IUCN 2008). Two
subspecies are extinct.
SYNONYMS G. i. abacornis Lawrence, 1934 and
G. i. irrectus Lawrence, 1934
COMMENT Described as Capromys ingrahami.
Geocapromys ingrahami abaconis Lawrence,
1934. Occ. Pap. Boston Soc. Nat. Hist. 8:191.
COMMON NAME Abaco Island hutia
TYPE LOCALITY Imperial Lighthouse Caves,
Hole in the Wall, Great Abaco Island, Bahamas
DISTRIBUTION Great Abaco Island, Bahamas
STATUS Extinct.
Geocapromy ingrahami ingrahami (Allen,
1891). Bull Am. Mus. Nat. Hist., 3:329.
COMMON NAME Bahamian hutia.
TYPE LOCALITY East Plana Cay, Bahamas
DISTRIBUTION Known from the type locality
and introduced populations on Little Wax Cay
(1973) and Warderick Wells Cay (1981). For
reintroductions see Jordan (this volume).
STATUS Vulnerable (IUCN 2008).
Geocapromys ingrahami irrectus Lawrence,
1934. Occ. Pap. Boston Soc. Nat. Hist. 8:190.
COMMON NAME Crooked Island hutia,
Crooked Island coney.
TYPE LOCALITY Burial cave #1, Gordon Hill
caves, Crooked Island, Bahamas
DISTRIBUTION Type locality, Eleuthera
Island and Long Island
STATUS Extinct.
Geocapromys thoracatus (True, 1888). Proc. U.S.
Natl. Mus., 11:469.
COMMON NAME Swan Island hutia.
TYPE LOCALITY “Little Swan Island, one of the
two small islands lying at the entrance of the Gulf
of Honduras”.
DISTRIBUTION Little Swan Island.
STATUS Extinct (IUCN 2008); became extinct in
the 1950s (Clough 1976), probably as a result of cat
introduction on the island.
COMMENT Some times included as a subspecies
of G. brownii; see Mohr (1939:77), Hall
(1981:866), and Varona (1974b). Reviewed by
Morgan (1989b) although confirmed to be a
distinct species (Morgan 1985).
Geocapromys sp. 1
Undescribed large extinct species from Grand
Cayman, Cayman Islands (Morgan 1994a).
Geocapromys sp. 2
Undescribed small extinct species from Cayman
Brac, Cayman Islands (Morgan 1994a).
Mesocapromys Varona, 1970. Poeyana, 73:8.
TYPE SPECIES Capromys (Mesocapromys)
auritus Varona, 1970
SYNONYMS Paracapromys Kratochvíl,
Rodríguez, and Baruš, 1978; Pygmaeocapromys
Varona, 1979; Stenocapromys Varona and
Arredondo, 1979
COMMENT Originally a subgenus, which was
elevated to a genus by Kratochvíl et al. (1978:15).
Mesocapromys, unlike Mysateles, constructs large
nests made of sticks. The genus consists of four
extant and seven extinct species. Most
Mesocapromys species are very similar to M.
nanus. Morphological and geographic variations
have not been adequately studied because
specimens in collections are limited and
fragmented. All type specimens are fragmented
mandibles, some known only from their type
locality. The majority of species is from western
Cuba, and has small geographical ranges. The
diagnostic characters of some species are weak.
The small differences are most likely due to
geographic variations. We synonymize some
species and recommend that others need additional
systematic analysis.
Primary characters of the genus are: small body
size, weighing less than 800 g (except melanurus,
which averages 1,231g), ratio tail/body size ranges
between 0.70 to 0.80, tail is semi-prehensile, skin

UPDATED CHECKLIST – Borroto-Páez, Mancina, Woods and William Kilpatrick
407
on palm and genitals is unpigmented, liver is
multilobulated and has the usual solid structure,
large corpora lutea (mean 3.0 mm), crista
supraorbital present, fossa orbital bigger than fossa
preorbital, bullae relatively large.
Mesocapromys angelcabrerai (Varona, 1979).
Poeyana, 194:6.
COMMON NAME Conguino hutia, Cabrera’s
hutia
TYPE LOCALITY Cuba, Ciego de Ávila
Province, Cayos de Ana Maria (21o30’N, 78o40’W)
DISTRIBUTION Cayos de Ana Maria, Cuba
STATUS Critically endangered (IUCN 2008)
COMMENT Placed in the subgenus
Pygmaeocapromys by Varona (1979) and reviewed
by Camacho et al. (1994). Sexually dimorphic
unlike other capromyids. There are unconfirmed
reports of the presence of M. angelcabrerai on the
coast near Júcaro. This species and M. nanus are
the two smallest hutias in Cuba.
Mesocapromys auritus (Varona, 1970). Poeyana,
ser. A, 73:1.
COMMON NAME Large-eared hutia, eared hutia.
TYPE LOCALITY Cuba, Las Villas Province,
Sabana Archipelago, Cayo Fragoso (22o41’N,
79o27’W).
DISTRIBUTION Type locality and introduced on
Cayo Pasaje, Cayo La Sagra, and Cayo Pajonal,
close to Cayo Fragoso (Manójina et al. 1994).
STATUS Critically endangered (IUCN 2008)
COMMENT This form is known only from the
mangroves along channel edges passing across
Cayo Fragoso. M. auritus and M. angelcabrerai
construct similar nests. Karyotype is 2n=36
(Hernández and Sánchez 1987).
Mesocapromys gracilis (Varona and Arredondo,
1979). Poeyana 195: 17.
COMMON NAME Gracile small hutia
TYPE LOCALITY Cueva del Túnel, La Salud, La
Habana
DISTRIBUTION Type locality; Cueva de
Perfecto, Sumidero, Pinar del Río; Cueva de
Paredones, Habana
STATUS Extinct
COMMENT Described as Capromys
(Stenocapromys) gracilis. The elements for the
paratypes are missing in the collection. We
provisionally retain M. gracilis as a recognized
taxon taking into account its western distribution
and reported morphological distinctness.
Systematics revision is needed. Silva-Taboada et
al. (2007) consider this species as inquirenda.
Mesocapromys kraglievichi (Varona and
Arredondo, 1979). Poeyana 195: 19.
COMMON NAME Small hutia from Paredones
TYPE LOCALITY Cueva de Paredones, Ceiba del
Agua, La Habana
DISTRIBUTION Type locality; Cueva Lamas,
Santa Fé, Ciudad de la Habana
STATUS Extinct
SYNONYMS barbouri Varona and Arredondo,
1979
COMMENT Described as Capromys
(Mesocapromys) kraglievich. Silva-Taboada et al.
(2007) maintain this species and propose the
synonymy of barbouri. Pieces for the paratypes
are missing in the collection. All materials in the
description are fragmented mandibles. The species
must be reviewed.
Mesocapromys melanurus (Poey, 1865). In Peters,
1865, Monatsb. K. Preuss. Akad. Wiss. Berlin,
p. 384.
COMMON NAME Andaraz hutia, black-tailed
hutia, mono hutia, bushy-tailed hutia
TYPE LOCALITY Cuba, Manzanillo, Granma
Province (Oriente region)
DISTRIBUTION Eastern provinces of Cuba
STATUS Vulnerable (IUCN 2008)
SYNONYMS arboricolus Kratochvíl, Rodriguez,
and Barus, 1978; rufescens Mohr, 1839.
COMMENT The author and date of publication for
this species are usually given as Peters, 1864, but
Varona (1974b), established the correct author and
date as Poey in Peters, 1864 [1865]. Includes
arboricolus of Kratochvíl et al. (1978:48), who
placed it in the genus Mysateles, subgenus
Leptocapromys. However, this specimen is
considered to be a young female M. melanurus by
Varona (1986:7). We have examined this
specimen, and concur with Varona. Woods et al.
(2001) (based on analysis of cytochrome b
sequence data) concluded that Mysateles is
paraphyletic and that the affinities of melanurus are
with taxa of the genus Mesocapromys rather than
with Mysateles p. prehensilis or M. p. gundlachi.
Borroto-Páez (2002) and Borroto-Páez et al. (2005)
proposed a new combination under Mesocapromys.
Silva-Taboada et al. (2007) considered this change
to be premature. A new molecular analysis shows
more resolution in the relationship of melanururs
with angelcabrerai (Kilpatrick et al. this volume).
We agree that further analyses with additional
samples are needed. Reviewed by Borroto-Páez
and Camacho (this volume). Locally abundant in
Guisa, Granma Province, but uncommon in other
regions of the eastern provinces (Borroto-Páez et
al. 2001, Borroto-Páez and Camacho, this volume).
Mesocapromys minimus (Varona and Arredondo,
1979). Poeyana 195:1.
COMMON NAME Mayarí small hutia
TYPE LOCALITY Arroyo del Palo, Taino midden
at Mayarí, Holguín, Cuba

TERRESTRIAL MAMMALS OF THE WEST INDIES
408
DISTRIBUTION Type locality; Cueva de
Seboruco, Mayarí, Holguín, Cuba
STATUS Extinct
COMMENT Initially described as Capromys
(Pygmaeocapromys) minimus. The elements for
the paratypes are missing in the collection. We
provisionally maintain the species taking into
account its eastern distribution (and the
morphological variation that it represents) but
should be reviewed. Fossil materials for
Mesocapromys species are less abundant in the
eastern part of Cuba. Silva-Taboada et al. (2007)
consider this species as inquirenda.
Mesocapromys nanus (G. M. Allen, 1917). Proc.
New England Zool. Club, 6:54.
COMMON NAME Dwarf hutia
TYPE LOCALITY Cuba, Matanzas Province,
Sierra de Hato Nuevo
DISTRIBUTION Cienaga (swamp) de Zapata
(Matanzas Province, Cuba)
STATUS Critically endangered (IUCN 2008).
Some mammalogists consider this species extinct.
Collected last October 26, 1951 by J. E. Guilday in
central part of Zapata Swamp, Matanzas, and the
specimen is deposited at the Carnegie Museum of
Natural History, Pittsburgh; Catalogue number
57500. Last evidence dates March 23, 1978, when
a nest was found, an animal was detected and fecal
pellets were collected (Garrido 1980).
SYNONYMS beatrizae Varona and Arredondo,
1979; delicatus Varona and Arredondo, 1979;
silvai Varona and Arredondo, 1979.
COMMENT Reviewed by Allen (1918) and
Varona (1974a), then placed in the genus
Mesocapromys, subgenus Paracapromys by
Kratochvíl et al. (1978) and Rodriguez et al.
(1979), then in the subgenus Pygmaeocapromys by
Varona (1979). Retained in Capromys subgenus
Mysateles by Hall (1981) because of its long tail
and small body size, although Varona (1979) stated
that in spite of these similarities, there are
important cranial differences between nanus and
other Mysateles. Originally based on fossil
materials, live specimens were subsequently found
in Zapata Swamp. Silva-Taboada et al. (2007)
propose the synonymy of beatrizae. The
description of beatrizae (Varona and Arredondo
1979) is based on large amount of materials,
although all paratype materials are currently
missing from the collection. In the type specimen
illustration, M2 is erroneously present.
The type specimen of delicatus is a very
fragmented mandible without the coronoid,
condylar and angular processes and known only
from the type locality. Additional factors
contributing to placing delicatus as a synonym
include the nature of the other specimen in the
hypodigm (another incomplete mandible with
incisor and molars) and inconsistency with
characters used in the description.
The type specimen of silvai is a very
fragmented anterior part of a mandible with incisor
but without molars or coronoid, condilar and
angular processes. The other eight known
specimens (lost from the collection) are fragmented
mandibles without molars or processes, one
fragmented mandible with M1 only, and a very
fragmented skull of an immature animal.
Moreover, silvai is known only from the type
locality.
Mesocapromys sanfelipensis (Varona and Garrido,
1970). Poeyana, ser. A, 75:3.
COMMON NAME San Felipe hutia, little Earth
hutia
TYPE LOCALITY Cuba, Pinar del RíoProvince,
Los Canarreos Archipelago, Cayo Juan Garcia
(21o59’N, 83o31’W)
DISTRIBUTION Known only from four
specimens collected at the type locality
STATUS Critically endangered. Fire destroyed
much of its habitat on Cayo Juan García (Frias et
al. 1988). It might be extinct now.
COMMENT Placed in subgenus Mesocapromys by
Varona (1974b) and in genus Mesocapromys,
subgenus Paracapromys by Kratochvíl et al.
(1978:15). All known specimens were captured on
perennial glasswort (Batis maritima or yerba de
vidrio) rather than mangroves. The type locality
and other adjacent keys were recently assessed, but
there was no evidence of its presence probably
because of the precarious conditions of the habitat.
Mysateles Lesson, 1842. Nouv. Tabl. Règne
Animal Mammifères, p. 124.
TYPE SPECIES Capromys prehensilis Poeppig,
1824
SYNONYMS Leptocapromys Kratochvíl et al.
1978; Brachycapromys Varona and Arredondo,
1979
COMMENT Lesson (1842) classified Capromys
prehensilis as Mysateles poeppingi; see Varona
(1979). Kratochvíl et al. (1978) treated Mysateles
as a genus in their new classification. Mysateles
was included in Capromys by Woods (1989a), Hall
(1981), and Corbet and Hill (1991). However,
biochemical and morphological evidence supports
Mysateles as a valid genus (Camacho et al. 1995;
Borroto-Páez 2002). Woods et al. (2001) found
that Mysateles was paraphyletic in relation to
Mesocapromys angelcabrerai but was distinct from
Capromys. Characters of the genus include:
medium body size (mean body weight is 1,730 g),
ratio tail/body size is 0.78, prehensile tail,
unpigmented skin of the palm and genital area,

UPDATED CHECKLIST – Borroto-Páez, Mancina, Woods and William Kilpatrick
409
multilobulated liver showing the usual solid
structure, large corpora lutea (mean 4.6 mm), the
vertical diameter of the orbital and preorbital fossas
is approximately the same, the crista supraorbital is
absent.
Mysateles prehensilis (Poeppig, 1824). Jour. Acad.
Nat. Sci. Philadelphia, 4:11.
COMMON NAME Prehensile-tailed hutia, carabalí
hutia, mona hutia
TYPE LOCALITY Cuba, wooded south coast
DISTRIBUTION Cuba, mainly west of Camagüey
Province. The distribution of this species in the
eastern provinces (old Oriente Province) is unclear.
STATUS Near threatened (IUCN 2008)
SYNONYMS pallidus (Poey, 1865); poeppingi
Lesson, 1842; poeyi (Guerin, 1834), jaumei Varona
and Arredondo, 1979, gundlachi (Varona, 1986).
The subspecies M. p. gundlachi (Chapman, 1901),
and M. p. meridionalis (Varona, 1986) proposed by
Silva-Taboada et al. (2007)
COMMENT Placed in the subgenus Myseteles by
Mohr (1939) and Varona (1974b), and in the genus
and subgenus Mysateles by Kratochvíl et al.
(1978). Mysateles gundlachi was described on the
basis of the baculum, which is distinctly broad-
based unlike that of M. meridionalis (Varona
1986). Based on levels of sequence divergence,
Woods et al. (2001), Borroto-Páez (2002) and
Borroto-Páez et al. (2005) consider gundlachi as an
insular subspecies of M. prehensilis, which we
follow here. Karyotype is 2n=34; FN=54-56
(Milišnokov et al. 1990).
M. jaumei is a synonym because the single
known specimen consists in a very fragmented
rostral portion, illustrated in Varona and Arredondo
(1979) with the figure legend erroneously
identifying the species as brevirostris, and the
diagnosis characters are weak. Although it appears
to be an immature individual similar to C.
pilorides, the molariforms are well worn and there
is an erupted M3 indicating that the specimen was
adult. We agree with Silva-Taboada et al. (2007)
that the rostral variation could be teratologic.
These authors are proposing Mysateles prehensilis
meridionalis and consider jaumei as species
inquirenda.
Mysateles prehensilis gundlachi (Chapman,
1901). Bull. Amer. Mus. Nat. Hist. 14:317.
COMMON NAME Prehensile-tailed hutia from
north of Isla de la Juventud, carabalí hutia from
north of Isla de la Juventud, Chapman's
prehensile-tailed hutia, Gundlach's hutia
TYPE LOCALITY Nueva Gerona, Isla de la
Juventud, Cuba
DISTRIBUTION North of Isla de la Juventud,
with the Lanier Swamp as its southern limit
STATUS Vulnerable (see Borroto-Páez and
Ramos García, this volume.)
SYNONYM Capromys gundlachi (of Varona,
1986)
COMMENT Elevated to a full species based on
the morphology of the baculum by Varona
(1986), however, the molecular divergence is
insufficient to consider a valid species (Woods et
al. 2001; Borroto-Páez 2002; Borroto-Páez et al.
2005). This subspecies is reduced in distribution
to just a few localities with low population.
Mysateles prehensilis meridionalis (Varona,
1986).Poeyana, 315:4.
COMMON NAME Prehensile-tailed hutia from
south of Isla de la Juventud, carabalí hutia from
south of Isla de la Juventud, southern hutia,
meridionalis hutia
TYPE LOCALITY Cuba, north of Caleta
Cocodrilos (Jacksonville), SW Isla de la Juventud
(former Isle of Pines)
DISTRIBUTION Restricted to lowland forests
SW of the central savanna on Isla de la Juventud
STATUS Critically endangered (Borroto-Páez
and Ramos 2003)
COMMENT Silva-Taboada et al. (2007) believe
that this form has more affinity with M. p.
gundlachi than with M. p. prehensilis. Borroto-
Páez (2002) and Borroto-Páez et al. (2005)
demonstrated a close affinity with M. p.
prehensilis based on an analysis of 41 skull
characters, while an analysis of external
characters indicates an affinity with M. p.
gundlachi. We consider the relationship of body
size compared with tail length to be a singular
important character in capromyids at both the
species and generic levels. Silva-Taboada et al.
(op. cit.) consider this ratio to be more important
at the subspecific level. The ratio of the body to
tail is 62 percent in meridionalis, 73 percent in
gundlachi, and 79 percent in prehensilis (see
comments under prehensilis).
M. prehensilis prehensilis (Poeppig, 1824). See
account of M. prehensilis.
Subfamily Hexolobodontinae Woods, 1989. Los
Angeles Co. Mus. Nat. Hist., Sci. Ser., 33:76.
Hexolobodon was originally considered to be allied
with the genus Capromys and other members of the
subfamily Capromyinae because the pattern of the
reentrant folds of the molariform teeth is so similar.
However, the cranial features of this group are
distinct and more primitive than capromyines, thus
Woods (1989c) created this subfamily to recognize
their distinct heritage.

TERRESTRIAL MAMMALS OF THE WEST INDIES
410
Hexolobodon Miller, 1929. Smithson. Misc. Coll.,
81:19.
TYPE SPECIES Hexolobodon phenax Miller, 1929
COMMENT Characters of the genus include: the
molariform teeth are high crowned and open rooted
in young individuals, but the roots close in mid-age
so that molars wear to become short and
prominently rooted. The upper cheekteeth have
two lingual reentrant folds (hypoflexus and
posterolingual flexus) that match the pattern of two
labial reentrant folds (paraflexus and mesoflexus)
giving the occlusal surface a distinctive six-lobed
appearance (hence the name Hexolobodon). Older
individuals may lose this pattern because of
toothwear.
Hexolobodon phenax Miller, 1929. Smithson.
Misc. Coll., 81:19.
COMMON NAME Imposter hutia, even–toothed
hutia
TYPE LOCALITY Haiti, Departement de
l’Artibonite, Small cave northeast of St. Michel de
l’Atalye
DISTRIBUTION Hispaniola and La Gonave Island
STATUS Extinct
SYNONYM poolei Rímoli, 1976 [1977]
COMMENT Includes H. poolei (Rímoli, 1976:21),
which is known only by the type specimen.
Hexolobodon sp.
COMMON NAME La Hotte even-toothed hutia
COMMENT Undescribed extinct species from
Trou Wòch Sa Wo, near Camp Perrin,
southwestern Haiti (Woods 1989a, Woods and
Ottenwalder 1992).
Subfamily Isolobodontinae Woods, 1989. Los
Angeles Co. Mus. Nat. Hist., Sci. Ser., 33:76.
Recognized as a distinct subfamily because of the
unique nature of the bands of cement surrounding
the molariform teeth. All isolobodontines have
extensive cement within the reentrant folds. Thick
cement also surrounds the cheekteeth in distinct
rings to produce clear growth bands. In addition
the enamel structure of the cheekteeth is thin and
soft.
Isolobodon Allen, 1916. Ann. New York Acad.
Sci., 27:19.
TYPE SPECIES Isolobodon portoricensis J. A.
Allen, 1916
SYNONYMS Aphaetreus Miller, 1922; Ithydontia
Miller, 1922
COMMENT Characters of the genus include:
labial and lingual reentrant folds of the upper
molariform teeth do not overlap in the central
occlusal area, and have a tendency to join each
other where they meet to form lamellar plates.
Isolobodon montanus (Miller, 1922). Smithson.
Misc. Coll., 74:3.
COMMON NAME Montane hutia
TYPE LOCALITY Haiti, Département. de
l’Artibonite, Cave northeast of St. Michel de
l’Atalaye
DISTRIBUTION Hispaniola and La Gonave Island
STATUS Extinct
COMMENT Originally described as a separate
genus from I. portoricensis because the enamel
folds of the cheekteeth connect to form separate
laminar plates, but an analysis of a large series of
each group indicates that this character is clinal,
and the two forms are very closely related. This
species is larger in mass and is more robust than I.
portoricensis, and the molariform teeth are much
broader.
Isolobodon portoricensis Allen, 1916. Ann. N.Y.
Acad. Sci., 27:19.
COMMON NAME Puerto Rican hutia, Indian
hutia
TYPE LOCALITY Puerto Rico, Jobo Dist., near
Utuado
DISTRIBUTION Hispaniola (Haiti and Dominican
Republic) and offshore islands. It is the only
known hutia on La Tortue Island, and was probably
introduced on Puerto Rico, Vieques Island, St.
Thomas, St. Croix, St John, and Mona Island.
STATUS Extinct (IUCN 2008), but possibly
surviving on La Tortue Island off the north coast of
Haiti (Woods et al. 1985) where there are fresh
looking remains and many stories of hutias having
been observed.
SYNONYM levir (Miller, 1922)
COMMENT Although the type locality is in Puerto
Rico, were Amerindians may have introduced the
species, the original natural range may have been
Hispaniola. Reported as extinct by Hall (1981:868),
but this species survived in Hispaniola and Puerto
Rico until the last few decades, and may still
survive in certain remote areas (Woods et al. 1985),
especially on the island of La Tortue, off the
northwestern coast of Haiti. Its remains are
abundant throughout Hispaniola, and are frequently
associated with kitchen midden sites.
Zazamys MacPhee and Iturralde-Vinent, 1995.
Amer. Mus. Novitates 3141:1-31.
TYPE SPECIES Zazamys veronicae MacPhee and
Iturralde-Vinent, 1995
COMMENT Characters of the genus include:
lower molariform teeth are distinct from
Isolobodon, i.e. the reentrant folds meet in the
center of the occlusal surface rather than being
lingually displaced, and the ridged bands of cement
on the outer surface of the molariform teeth are
barely visible rather than very distinctly banded.

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411
The presence of possible early Miocene
islobodontine rodent on Cuba suggests that
isolobodontine radiation was widespread in the
Greater Antilles, and not restricted to Hispaniola as
was previously thought.
Zazamys veronicae MacPhee and Iturralde-Vinent,
1995. Amer. Mus. Novitates 3141:1-31.
COMMON NAME Zaza hutia
TYPE LOCALITY Domo de Zaza; approximatly
21°45’N, 79°30’W, 18 km SE of Sancti Spíritus
city, Sancti Spíritus.
DISTRIBUTION Known only from the type
locality
STATUS Extinct
COMMENT Early Miocene species. The earliest
rodent in Cuba and the oldest recognized
capromyid (MacPhee et al. 2003).
Subfamily Plagiodontinae Ellerman, 1940.
The families and genera of living rodents, 1:25.
This subfamily includes at least six known species
from two different genera that range in mass from
500 g to an estimated 5 kg. The occlusal surfaces
of left and right toothrows tend to be on the same
horizontal plane and the reentrant folds are often
oblique and overlap significantly. Unlike
isolobodontines, there is no cement on the lateral
surfaces of the molariform teeth, which tend to
have strong enamel and robust structure. The range
in the degree of hypsodonty (high-crowned) and
hypselodonty (open rooted) is substantial (from
low-crowned with well-formed roots to high
crowned and open rooted throughout life). So far,
all recent and fossil records of plagiodontines are
restricted to the island of Hispaniola.
Plagiodontia Cuvier, 1836. Ann. Sci. Nat. Zool.
(Paris), ser 2, 6:347.
TYPE SPECIES Plagiodontia aedium F. Cuvier,
1836
SYNONYM Hyperplagiodontia Rímoli, 1977.
COMMENT Reviewed by Mohr (1939), Johnson
(1948), Anderson (1965), and Woods (1989a, b).
Characters of the genus include: labial and lingual
reentrant folds of upper molariform teeth overlap
significantly, reentrant folds dramatically oblique
in orientation, metaflexus present as major
posterolateral reentrant fold, enamal parts of
cheekteeth robust with no cement sourrounding the
molariform teeth. The deciduous molar is replaced
in older individuals.
Plagiodontia aedium Cuvier, 1836. Ann. Sci. Nat.
Zool. (Paris), ser 2, 6:347.
COMMON NAME Hispaniolan hutia, rat cayes,
zagouti
TYPE LOCALITY Saint-Domingue (probably
Haiti)
DISTRIBUTION Hispaniola, La Gonave Island
(not recorded from La Tortue Island)
STATUS Endangered (IUCN 2008)
SYNONYMS spelaeum Miller, 1929; Plagiodontia
aedium hylaeum (Miller, 1927)
COMMENT Includes P. a. hylaeum as a separate
subspecies (Anderson 1965). The liver is
multilobuled and is reticulated as C. pilorides but
the reticules are smaller. The synonymy of
spelaeum was based on the presence of only
fragmented material from a single location in
northern Haiti (St. Michel de l’Atalaye). It is likely
a young individual of Plagiodontia aedium, which
is abundant in the St. Michel region. It is the
smallest Plagiodontia.
Plagiodontia aedium aedium Cuvier, 1836
COMMON NAME Haitian hutia
TYPE LOCALITY Saint-Domingue (probably
Haiti)
DISTRIBUTION SW Haiti
STATUS Vulnerable (as Plagiodontia aedium).
SYNONYM spelaeum Miller, 1929
COMMENT This classic species of Hispaniolan
hutia was thought to be extinct in Haiti until it
was rediscovered in the mountains above
Miragoane. It is secretive and frequently
associated with karst outcrops where it lives in
deep crevices. There are viable populations
surviving in the Massif de la Selle, the Massif de
la Hotte and the surrounding foothills (such as
near Camp Perrin) and in the mountains above
Miragoane.
Plagiodontia aedium hylaeum (Miller, 1927)
COMMON NAME Dominican hutia
TYPE LOCALITY Guarabo, 10 miles east of
Jovero, Samana Province, Dominican Republic
DISTRIBUTION NE Dominican Republic
STATUS IUCN – Vulnerable (as Plagiodontia
aedium).
SYNONYM None
COMMENT Described as P. hylaeum. Proposed
as a subspecies by Anderson (1965).
Plagiodontia araeum Ray, 1964. Breviora, Mus.
Comp. Zool., 203:2.
COMMON NAME Narrow-toothed hutia
TYPE LOCALITY Dominican Republic, San
Rafael Province, unnamed cave 2 km SE Ranchola
Guardia (18o43’N, 71o39’W)
DISTRIBUTION Southern Hispaniola
STATUS Extinct
SYNONYM Hyperplagiodontia stenocoronalis
(Rímoli, 1976 [1977])
COMMENT Type description based on only left
upper cheek tooth. Subsequent fossil material
collected in Haiti and deposited at the Florida
Museum of Natural History includes complete

TERRESTRIAL MAMMALS OF THE WEST INDIES
412
cranial and dentary material, confirming the
validity of this taxon. Hyperplagiodontia
stenocoronalis of Rimoli (1976) is not distinct from
P. araeum. This species is a very large wide-
toothed hutia.
Plagiodontia ipnaeum Johnson, 1948. Proc. Biol.
Soc. Wash., 61:72.
COMMON NAME Samana hutia
TYPE LOCALITY Dominican Republic, Samana
Province, Anadel (19o12’N, 69o19’W)
DISTRIBUTION NE of Dominican Republic,
Hispaniola
STATUS Probably extinct (Woods et al., 1985)
SYNONYMS caletensis Rímoli, 1976 [1977];
velozi Rímoli, 1976 [1977]
COMMENT A large version of P. aedium.
Rhizoplagiodontia Woods, 1989. Los Angeles Co.
Mus. Nat. Hist., Sci. Ser., 33:62.
TYPE SPECIES Rhizoplagiodontia lemkei Woods,
1989
COMMENT This taxon differs from all other
capromyids in having the upper premolar with
three labial reentrant folds. It is most similar to
Plagiodontia. Differing from Plagiodontia in
having less oblique reentrant folds in maxillary
cheekteeth and with a slight covering of cement on
lateral surfaces of molars. Molariform teeth are
rooted from mid-life on.
Rhizoplagiodontia lemkei Woods, 1989. Los
Angeles Co. Mus. Nat. Hist., Sci. Ser., 33:62.
COMMON NAME Lemke’s hutia, La Hotte hutia
TYPE LOCALITY Haiti, Département du Sud, 17
km W Camp Perrin (18o20’N, 74o03’W)
DISTRIBUTION Endemic to SW Haiti in the
Massif de la Hotte
STATUS Extinct
COMMENT Remains of Rhizoplagiodontia lemkei
are abundant in deep ravines and high plateaus of
the Massif de la Hotte between Jérémie and Camp
Perrin, southwestern Haiti. This area was a dense
broadleaf forest with abundant rainfall. Despite
extensive collections done in other regions of Haiti
and the Dominican Republic, this taxon has been
found only in the southern peninsula of Haiti, west
of Miragoane. Carbon14 analyses from the type
locality indicate ages between 3,715 and 10,320
yBP for the layers where these remains were
recovered. None of the remains looked “recent”,
but there were numerous intact cranial and
mandibular remains. This is also the type locality
of the La Hotte monkey Insulacebus toussaintiana
(Cooke et al. 2011) and the sloth Neocnus toupiti,
suggesting the early presence of a rich endemic
assemblage of mammals in the forests of the La
Hotte Mountains of SW Haiti.
Family Heptaxodontidae Anthony, 1917. Bull.
Am. Mus. Nat. Hist., 37(4):183.
SYNONYMS Amblyrhizinae Schaub, 1951;
Elasmodontomyinae Anthony, 1917.
COMMENT Known only from sub-recent fossils
from the Greater Antilles and northern Lesser
Antilles (Woods 1989a). There are two named
extinct forms (Tetrastylomys and Pentastylomys)
from the Miocene or Pliocene of Argentina
(Kraglievich, 1926). Both genera are considered to
be heptaxodontids by McKenna and Bell (1997).
However, Landry (1957) believed that the laminar
dental pattern in these forms could be derived from
dinomyids, and that they were not heptaxodontids.
The Hispaniolan “heptaxodontid” genus
Quemisia is similar in dental morphology to
capromyids, especially Plagiodontia. It is possible
to derive all of the conditions seen in
heptaxodontids from dental patterns found within
the Capromyidae. However, at this time we are
reluctant to consider the taxa grouped together as
heptaxodontids to be part of the capromyid
radiation. While that remains a working
hypothesis, it is equally possible that these forms
do not constitute a monophyletic family, and that
Quemisia may be distinct from other
heptaxodontids and more closely related to
capromyids than are the remaining heptaxodontids.
We currently believe that it is best to consider
the Heptaxodontidae to be an endemic Antillean
family. Heptaxodontids are often considered
closely related to the Chinchillidae based on similar
molariform laminar plates. However, as noted
above, we consider heptaxodontids to be closely
related to the Capromyidae. Whether Amblyrhiza
and Clidomys became extinct before or after
humans settlement in the West Indies is debatable.
Subfamily Clidomyinae Woods, 1989.
Biogeography of the West Indies, p. 753.
Molariform teeth generally without reentrant folds.
Molariform teeth with three complete laminae
except M3, which have four; each laminar plate is
separated by a wide band of cement which also
encases the outer surface of the entire tooth as a
thin sheath; molariform teeth hypsodont with open
roots (hypselodont) throughout life.
Clidomys Anthony, 1920. Bull Am. Mus. Nat.
Hist., 42:469.
TYPE SPECIES Clidomys osborni Anthony, 1920
SYNONYMS Alterodon Anthony, 1920;
Speoxenus Anthony, 1920; Spirodontomys
Anthony, 1920
COMMENT Characters of the genus include:
molariform teeth high crowned, rootless,
evergrowing and very oblique, maxillary toothrows

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strongly converge anteriorly with only about 3 mm
distance separating left and right P4 teeth, left and
right mandibular rami strongly united.
Clidomys osborni Anthony, 1920. Bull Am. Mus.
Nat. Hist., 42:469.
COMMON NAME Osborn’s key mouse
TYPE LOCALITY Jamaica, Balaclava,
Wallingford Eastate, Roadside Cave
DISTRIBUTION Jamaica
STATUS Extinct
SYNONYMS Clidomys cundalli (Anthony, 1920);
jamaicensis (Anthony, 1920); Alterodon major
(Anthony, 1920); parvus (Anthony, 1920)
COMMENT C. parvus is here synomized with C.
osborni based on Morgan and Wilkins (2003), who
noted the wide size range of C. osborni specimens
from Slue's Cave, Jamaica. They interpreted this
size range to indicate that C. osborni is the only
valid species of Clidomys present in Jamaica.
Considered Jamaica’s largest Quaternary mammal
until the discovery of a gigantic proximal right
femur from Sheep Pen (MacPhee and Flemming
2003). The possible affinities of the Sheep Pen
specimen is discussed in detail by MacPhee and
Flemming in their paper and they conclude that it is
most likely from a very large rodent, and that it is
somewhat similar to the femur of Amblyrhiza. The
specimen has not been described because there are
serious questions about its history. The specimen
was found by Ross MacPhee in a box labelled
“Sheep Pen” in the collections of the Florida
Museum of Natural History. Alterodon major is
considered a Clidomys (MacPhee et al. 1983;
MacPhee 1984; MacPhee and Flemming 2003).
The extinction of Clidomys probably occurred
before humans arrived in the West Indies since
dates for the associated cave materials where
Clidomys has been collected exceed 100,000 years
in age.
Subfamily Heptaxodontinae Anthony, 1917.
Bull Am. Mus. Nat. Hist., 37(4):183.
The subfamily includes all of the giant hutias with
obliquely orientated reentrant folds on the
molarform teeth: Amblyrhiza = 35, Quemisia = 55,
Elasmodontomys = 45. They probably chewed
obliquely (= plagiognathously). The group is
convergent with the capromyid subfamily
Plagiodontinae in many dental and cranial
characters. Heptaxodontines can be distinguished
from clidomyines by the lack of complete dental
laminae and the presence of closed roots on the
molariform teeth. Heptaxodontines are more
similar to capromyids than are clidomyines and are
restricted to the northeastern Lesser Antilles,
Puerto Rico and to Hispaniola near the probable
origin of capromyids in the West Indies. As noted
by MacPhee and Flemming (2003), the
interrelationships of heptaxodontines, clidomyines
and the genus Quemisia remains unresolved, as
well as is the relationship of these forms to
capromyids.
Amblyrhiza Cope, 1868. Proc. Am. Philos. Soc., p.
313.
TYPE SPECIES Amblyrhiza inundata Cope, 1868.
SYNONYM Loxomylus Cope, 1869.
COMMENT Characters of the genus include: large
body mass, head, trunk and hind limb large relative
to elongated and gracile forelimbs, bimodal
distribution of incisor metrics (suggesting the
possibility of sexual dimorphism).
Amblyrhiza inundata Cope, 1868. Proc. Am.
Philos. Soc., p. 313.
COMMON NAME Blunt-toothed giant hutia
TYPE LOCALITY West Indies, Anguilla, Bat
Cave
DISTRIBUTION Anguilla, St. Martin
STATUS Extinct
SYNONYMS latidens (Cope, 1870); longidens
(Cope, 1871); quadrans (Cope, 1871)
COMMENT The world’s largest island rodent with
an estimated mass of 50 to 200 kg and it is
considered to be a hypervariable species
(Biknevicius et al. 1993; McFarlane et al. 1998). It
likely became extinct prior to humans arrival in the
West Indies. There is a nice artist’s reconstruction
of a pair of Amblyrhiza on the south shore of
Anguilla with St. Martin in the distance in
Biknevicius et al. (1993).
Elasmodontomys Anthony, 1916. Ann. N.Y. Acad.
Sci., 27:199.
TYPE SPECIES Elasmodontomys obliquus
Anthony, 1916
SYNONYM Heptaxodon Anthony, 1917
COMMENT Characters of the genus include:
moderate body mass (about the size of a paca), high
crowned rootless molariform teeth, molars with
five to six alternating enamel bands (laminar
plates), skull flattend and resembling Myocastor in
size and shape. The short digital phalanges suggest
that it was highly terrestrial.
Elasmodontomys obliquus Anthony, 1916. Ann.
N.Y. Acad. Sci., 27:199.
COMMON NAME Plate-toothed giant hutia,
Puerto Rican giant hutia
TYPE LOCALITY Utuado, Puerto Rico
DISTRIBUTION Known only from cave deposits
in Puerto Rico
STATUS Extinct
SYNONYM Heptaxodonbidens Anthony, 1917.
COMMENT H. bidens was an immature E.
obliquus. The remains of this species are abundant,

TERRESTRIAL MAMMALS OF THE WEST INDIES
414
widely distributed and well preserved in Puerto
Rico, and it is very likely that it survived into
Amerindian times (Woods 1996). There are
excellent descriptions and illustrations of
Elasmodontomys in Anthony (1918).
Tainotherium Turvey, Grady and Rye, 2006. J.
Zool. (London), 270: 586.
TYPE SPECIES Tainotherium valei Turvey, Grady
and Rye, 2006
COMMENT The description of this taxon is based
on an incomplete right femur of a single specimen.
Characters of the genus include: large-bodied
caviomorph rodent, enlarged femoral head, femoral
neck angled proximally at 50°; short, rectangular
greater trochanter not extending past the femoral
head or expanding laterally beyond the femoral
shaft, and with raised trochanteral crest; no ridges
running from greater trochanter on anterior surface;
large lesser trochanter not connected by
intertrochanteric crest; third trochanter
underveloped; diaphysis anteroposteriorly
flattened, and with distal half bowed posteriorly by
20°. The resemblance of this form with
Elasmodontomys is marked and some traits
regarded as unique to Tainotherium could be due to
taphonomy, disease, or other factors. Some
features fall within the well-documented range of
variation for Elasmodontomys obliquus (P. Bover,
personal communication). The characters look
insufficient for the designation of a new genus, but
until a more comprehensive analysis is completed,
we retain it as a valid taxon.
Tainotherium valei Turvey, Grady and Rye, 2006.
J. Zool. (London), 270: 586
COMMON NAME Puerto Rican large hutia
TYPE LOCALITY Cueva de la Vaca, south of
Barahona, Morovis Municipality, Puerto Rico;
N18°20´43.3412´´, W66°27´05.5851´´
DISTRIBUTION Known only from the type
locality
STATUS Extinct
COMMENT Possibly a morph of Elasmodontomys
obliquus. See our comments on the genus above.
Quemisia Miller, 1929. Smithson. Misc. Coll.,
81(9):22.
TYPE SPECIES Quemisia gravis Miller, 1929.
COMMENT The genus is very similar in dental
morphology to capromyids. Among the three
genera assigned to the Heptaxodontinae, Quemisia
remains the most problematic one (Ray 1965).
Characters of the genus (based on very limited
material) include: massive dentary with broad
cheekteeth obliquely orientated in alveoli. The
molar occlusal surfaces have long reentrant folds
(not arranged in laminar plates). The mandibular
symphysis is strong and broad, but the dentaries are
not fused.
Quemisia gravis Miller, 1929. Smithson. Misc.
Coll., 81(9):22.
COMMON NAME Twisted-toothed giant hutia,
quemi
TYPE LOCALITY Haiti, Département de
l’Artibonite, 6 km east of St. Michel de l’Atalaye
DISTRIBUTION Hispaniola
STATUS Extinct
COMMENT The remains of this large and robust
rodent are surprisingly uncommon in cave deposits
in Hispaniola. This is probably because it was far
too large to be preyed upon by barn owls, which
were responsible for accumulating many small and
medium sized rodents and insectivores in cave
deposits. Even though Miller (1929b) considered it
to be a close match for the “quemi” of historical
lore, the most recent dated association is 3,000
yBP. It is not clear if it survived into Amerindian
times (Woods 1996). We believe that the “quemi”
could have been a large robust Plagiodontia, which
it would have closely resembled. Ray (1965)
speculated that there might have been a close
relationship between Quemisia and Plagiodontia.
Family incertae sedis
Subfamily ?Heptaxodontinae
The description of this taxon is based on a single
fragmented left mandible. MacPhee and Flemming
(2003: 35-38) carefully considered the cases for
assigning Xaymaca to each of the most likely
clades (echimyids, capromyids, clidomyines, or
heptaxodontines). They tentatively assigned
Xaymaca to the Heptaxodontinae, although they,
and we, feel that this designation is tentative. They
are quite sure that it is not a capromyid or
clidomyine, but are less sure that it is not an
echimyid. It is a fascinating conundrum.
Xaymaca MacPhee and Flemming, 2003. Am.
Mus. Novitates 3422: 1.
TYPE SPECIES Xaymaca fulvopulvis MacPhee
and Flemming, 2003
COMMENT Known only from a single left
hemimandible with a well-worn premolar.
Extensive efforts to locate additional materials at
the site were unsuccessful. Characters of the genus
include: molar alveoli are smooth walled with no
internal divisions, suggesting that the cheekteeth
are cylindriform (as in Amblyrhiza and
Elasmodontomys); both rami of the dentaries are
not fused (unlike Amblyrhiza and Clidomys).
Xaymaca fulvopulvis MacPhee and Flemming,
2003. Am. Mus. Novitates 3422: 3.
COMMON NAME Diminute Jamaican hutia

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TYPE LOCALITY Brown Dust Passage of Drum
Cave, near Entrance #3. Drum Cave, part of the
“upper” sequence of the ~10 km Jackson’s Bay
cave system, is located on the south slope of
Portland Ridge near Jackson’s Bay, in the
southernmost part of Clarendon Parish, Jamaica.
Entrance #3 is located at approximately
17044’05’’N, 77013’15’’W.
DISTRIBUTION Known only from the type
locality
STATUS Extinct
COMMENT From a late Pleistocene deposit.
Despite their small size it is tentatively placed
within the heptoxodontine group (MacPhee and
Flemming 2003).