The Mammal species of north-central Venezuela

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The Mammal species of north-central Venezuela
Eisenberg, John Frederick
Polisar, John R
Florida Museum of Natural History
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Gainesville, Fla.
Florida Museum of Natural History, University of Florida
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City of Gainesville ( local )
Florida Museum of Natural History ( local )
bibliography ( marcgt )
government publication (state, provincial, terriorial, dependent) ( marcgt )
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Includes bibliographical references (p. 153-160).
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Bulletin of the Florida Museum of Natural History, volume 42, number 3, pp. 115-160
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John F. Eisenberg and John R. Polisar.

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John F. Eisenberg and John R. Polisar

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Plate 1. Female jaguar (Panthera onca) near prestamo. La Candelaria, Hato Piflero, Estado Cojedes. Eye shine of young caiman (Caiman crocodilus) in

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PnM ra a along dr- a r f c eHo d
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Plate 2. Giant anteater (Myrmecophaga tridactyla) along dry savanna/dry forest ecotone. Las Penitas, Hato Pifiero, Estado Cojedes.

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Plate 3. White-lipped peccaries (Tayassu pecar exiting mud wallow. Dr season, Cao Caujaral, Hato Piro.

Plate 3. White-lipped peccaries (Tayassu pecan) exiting mud wallow. Dry season, Caiio Caujaral, Hato Pifiero.

Plate 4. Puma (Puma concolor) traveling along foot-transect. Cafio Caujaral, Hato Pifiero.



* ,-~.-

~! A~. A

Plate 5. Capybara (Hydrochaeris hydrochaeris) carcass killed by a jaguar. Rio Pao, Hato Piftero.


John F. EisenbergI and John R. Polisar2


The terrestrial mammal fauna of Venezuela includes 327 species. In this paper we examine the
mammal fauna from the six north-central states and the federal district. This area includes the Caribbean
coast, the north coastal ranges, the interior ranges, and the northern Ilanos. A diverse array of habitats are
represented in this area: xeric scrub forests, montane rain forests, cloud forests, deciduous forests, and the
savannas of the llanos.
This north-central portion of Venezuela contains 180 species which have been vouchered and
deposited in museums. The forested north coastal ranges include sites such as Guatopo National Park with
some 95 species, while Ilanos sites, when gallery forests are included, may have up to 75 species. Some bat
species have ranges restricted to specific vegetation complexes and/or altitudes.
Representative examples of all major habitat types in this area currently receive some protection.
National parks perform this role in the north coastal ranges and interior ranges. To-date the most successful
protection of Ilanos flora and fauna seems to have been the result of private initiatives, on ranches such as
Hato Pifiero and Hato Masaguaral. In the Ilanos, maintenance of large mammals such as white-lipped
peccaries (Tayassu pecari), jaguar (Panthera onca), and puma (Puma concolor) will depend on linked
networks of large ranches and parks in which habitats are intact and hunting levels low to nonexistent


Se han registrado 327 species de mamiferos terrestres para Venezuela. En este trabajo analizamos la
mastozoofauna de los seis estados del norte central y del distrito federal. Se incluyen por lo tanto, la costa del
Caribe, el ramal nortelo de la Cordillera de la Costa, los ramales surefios de la Cordillera de la Costa y los
llanos nortefios. EstAn representadas una variedad de habitats: matorral desdrtico, bosques hamedos
montanos, bosques de neblina, bosques deciduos y las sabanas de los Ilanos.
Esta region de Venezuela alberga 180 species de mamiferos que cuentan con especimenes
depositadas en algun museo. La Cordillera de la Costa incluye sitios como el Parque Nacional de Guatop6
que alberga 95 species, mientras que en los Ilanos, en localidades que incluyen bosques de galeria se pueden
Ilegar a registrar hasta 75 species. Algunas species de quir6pteros tienen rangos de distribuci6n
restringidos a un tipo de vegetaci6n especifico y/o a un rango de altitudes.
En esta region muestras representatives de todos los habitats importantes estAn bajo alguna forma de
protecci6n. En la Cordillera de la Costa la protecci6n esta dada por parques nacionales. En los Ilanos la
protecci6n mas efectiva de species de flora y fauna ha sido debido a la iniciativa privada, por ejemplo en
ranchos como Hato Pifiero y Hato Masaguaral. Es por eso que en los Ilanos para la protecci6n de mamiferos

The senior author is the Katharine Ordway Chair and Professor of Ecosystems Conservation, Florida Museum of Natural History,
University of Florida, P. O. Box 117800, Gainesville FL 32611-7800, U. S. A.
The junior author is a Graduate Research Assistant, Florida Museum of Natural History, University of Florida, P. O. Box 117800,
Gainesville FL 32611-7800, U. S. A.

Eisenberg, J. F., and J. R. Polisar. 1999. The mammals of north-central Venezuela. Bull. Florida Mus. Nat.
Hist. 42(3):115-160 + Plates 1-5.


grandes como Biquiro Cachete Blanco (Tayassu pecari), el jaguar (Pantera onca) y el puma (Puma
concolor), se debe considerar una red integrada por grandes ranchos y parques que contengan hAbitats
pristinos sin presi6n o una presi6n muy baja de caza.


As of 1998 roughly 337 species of mammals have been recorded within the
political boundaries of Venezuela and within its coastal waters. Excluding strictly
aquatic forms (Sirenia, Cetacea), there is a terrestrial fauna of some 327 species
(Linares, 1998). The order Chiroptera includes 154 species and Rodentia 84
species; thus, seventy-two percent of the species are accounted for by just these two
orders. This is a rich fauna by temperate-zone standards. Whitaker and Hamilton
(1998) listed (exclusive of exotics and feral domesticates) 106 species for the
twenty-seven states east of the Mississippi River in the U.S.A. Of this total, fifty-
four percent belong to the Chiroptera and Rodentia.
Eisenberg and Redford (1979) conducted a comparative analysis of the
Venezuelan mammal fauna across seven broad biogeographical regions. The
northern coast ranges contain a number of peaks over 2,000 m. Much of this
terrain is covered by evergreen forests, which are partially isolated from similar
forests in the Andes by lowlands which contain dry deciduous forest and
grasslands. Complex folding has resulted in a series of ranges roughly parallel
along an east-west axis. It is possible to refer to two major components: (1) the
coastal range abutting the sea and (2) a series of interior ranges separate from the
former by the valley of Caracas and the Lake Valencia valley (Fig. 1). Protected
areas in the front coastal range include Parque Nacional El Avila (852 kin2),
Parque Nacional Henri Pittier (1078 km2) and Parque Nacional San Esteban (440
km2). Protected areas in the interior of the coast range include Parque Nacional
Guatopo (1224 km2), Parque Nacional Macarao (150 km2) and Parque Nacional
Tirgua (910 km2). A partially isolated inland range, the Sierra Aroa, contains
Parque Nacional Yurubi (236.7 kmin2). Elevations gradually decrease as these
mountains undulate into another, lower, biographic region, los llanos. The
southern drainage from the mountains forms streams and rivers that meander south
through los llanos, all eventually draining into major rivers, such as the Apure, and
ultimately exiting into the Atlantic via the Orinoco.
The llanos are usually divided into llanos altos (above 100 m), and further
south, los llanos bajos. Both are characterized by a habitat mosaic of low,
seasonally flooded grasslands and gallery forests. The gallery forests provide a
"ribbon of life" for forest forms. In los Ilanos altos, these gallery forests are
augmented by large patches of semi-deciduous forest not associated with rivers.
These connected forest corridors and patches maintain mammalian diversity in a
savanna dominated landscape. Cattle ranching is the dominant land use
throughout los llanos. In some cases, enlightened management by private owners


Figure 1. The moist forest zones of north-central Venezuela are associated with the higher elevation of the
coastal ranges (Ewel and Madriz, 1968). Contour lines are at intervals of 500, 1000, 2000, and 3000 m.
The llanos lie to the east of the Andes and south of the coastal ranges. Black dots indicate major centers of
urbanization. The major study areas are shaded.

has resulted in wildlife refuges that rival national parks in efficacy. The
aggregated forests and savannas of Hatos Pifiero, Mata Clara, Socorro in Estado
Cojedes and Hato Corralito in Guarico constitute an area of over 1,500 km2. This is
larger than any one effective national park in our entire study area (Estados
Aragua, Carabobo, Cojedes, Guarico, Miranda, and Yaracuy, and the Distrito
Federal). The fauna retained (jaguars, tapirs, white-lipped pecccaries) reflect that.
Two well-studied llanos sites include Hato Masaguaral (3000 ha) in Estado
Guarico and Hato Piflero (84,000 ha) in Estado Cojedes.
Overall, habitat diversity is high. There are xeric habitats along the
Caribbean Coast, humid and dry forest types in the northern mountains, and
additional heterogeneity to the south in los llanos. Holdridge Life Zone
classifications adequately describe the forests of the northern part of the study area
(Wet Tropical, Dry Tropical, Very Dry Tropical, Very Wet Montane Tropical, Wet
Montane Tropical, and Dry Montane Tropical Forests), but fail to adequately
explain habitat variation in los llanos (Ewel and Madriz, 1968). Falling entirely


within a Holdridge Tropical Dry Forest classification, los llanos actually contain
diverse habitats: seasonally burnt chaparrales, dry deciduous forests on low hills,
semi-deciduous forests in valleys, palm forests along savanna edges, pockets of
evergreen forests, and a wide variety of savanna types. Small changes in
elevations, combined with variation in soils and hydrology, create a strong
horizontal heterogeneity. The entire study area experiences strong seasonality. A
wet season runs from May through October, the dry season from December
through March, and April and November are transitional months. Low elevations,
high temperatures, clay soils, and impeded drainage in los llanos result in
exceptionally strong seasonality: annual cycles of flooding and dessication there
greatly affect both flora and fauna.
The mammals of Venezuela were characterized in Handley (1976) and
summarized by Linares (1998). Original ecological research in Parque Nacional
Guatopo and Hato Masaguaral is included in Eisenberg (1979b). This paper
aggregates information regarding the terrestrial mammal fauna of the north-central
portion of Venezuela (Estados Aragua, Carabobo, Cojedas, Guarico, Miranda,
Yaracuy, and the Distrito Federal). This area was chosen because of the depth of
the studies that have taken place in this area, and its high diversity owing to a
juxtaposition of biogeographic regions. The area is unique for another reason.
Despite inclusion of several large cities, large scale industrial centers, and
widespread agricultural activities, this area retains much of its original fauna. We
document that situation and hope that this contribution helps perpetuate it (Fig. 2).
The mammal fauna of the area defined in the previous paragraphs includes
180 terrestrial species; 108 are Chiroptera and 27 belong to the Rodentia. The area
is comparable to about seventy percent of the state of Georgia, U. S. A. The fauna
is typical of the lowland Neotropics, although not as species-rich as comparable
areas in the western Amazon region of Peru and Ecuador. A "list" of 161 species
is included in Table 1 and represents the efforts of Handley (1976) and Eisenberg
et al. (1979) with some ammendations. Some marginal bat species are not
included in Table I but are listed in Table 2. Combining Tables 1 and 2 could
yield 180 species.
No one sampling effort in the major study areas discussed in Eisenberg et al.
(1979) included the full faunal complement. This occurred for a variety of reasons.
For example, let us consider Guatopo and vicinity. Handley (1976) collected 39
species of bats and 13 species of rodents. Eisenberg, O'Connell, and August (1979)
recorded 18 species of bats and 13 species of rodents. The lists were not identical,
reflecting different collecting efforts. When the lists are combined, the totals
would be 50 species of bats and 17 species of rodents. Combining all collection
efforts in Guatopo would yield about 95 species of mammals or about 55 percent of
the total estimated fauna for the defined 113,430 km2 six-state area; Guatopo
covers 1,224 km2.


Figure 2. The six states of north-central Venezuela plus the federal district. Study and protected areas and
parks are represented by the numbers.

Proceeding south, from the montane forests to the Ilanos, species richness
declines somewhat. Where gallery forests are present, species numbers remain
elevated, but the stochastic loss of species contributes to an uneven distribution.
For example, in the savanna woodlands and gallery forests of Hato Masaguaral,
seventy-four species of mammals were tabulated of which forty-two were bats and
twelve rodents. The areas surveyed included some 3000 ha. The level of species
richness is typical for woodland savannas in the Neotropics (Voss and Emmons,
Trapping effort, microhabitats sampled, and genuine differences between the
"front" range and the "back" ranges of the north coast all contribute to differences
in species numbers recorded. The figure of 180 species for the total area of six


Table 1. The Mammals of North-Central Venezuela (1976-1998).

Taxa A' B2 C3

Didelphimorphia (10)
Caluromys philander x, g g
Chironectes minimus x, g g
Didelphis marsupialis x, g g, m
Marmosops fuscata x, g g
Marmosa murina x, g g
M. robinsoni x, g g, m
Micoureus demararae cinereaa) x, g g
Gracilinanus marica x o Perez Hernandez, 1989
Monodelphis brevicaudata x, g g
M. orinoci o o Perez-Hernandez, 1989
Cryptotis meridensis o o Linares, 1998
Xenarthra (Edentata) (7)
Myrmeccphaga tridactyla o m
Tamandua tetradactyla x, g m, g
Bradypus variegatus x, g g
Cabassous unicinctus a g
Dasypus novemcinctus x, g g, m
D. sabanicola o o Wetzel, 1985
Priodontes maximus o o Wetzel, 1985
Primates (3)
Cebus olivaceous (nigrivittatus) x, g g. m
Alouatta seniculus x, g g, m
Ateles belzebuth o g Mondolfi and Eisenberg, 1979
Chiroptera (88)
Emballonuridae (7)
Peropteryx kappleri x, g o
P. macrotis x, g o
P. trinitatus x, g o
Rhynchonycteris naso x, g m
Saccopteryx bilineata x, g g, m
S. canescens x, g 0
S. leptura x, g g, m
Noctilionidae (2)
Noctilio leporinus x, g m
N. albiventris x, g m
Mormoopidae (4)
Pteronotus davyi x, g g
P. parnellii x, g g
P. personatus x, g o


Table 1 Continued.

Taxa Al B2 C3

P. suapurensis x, g o
Mormoops megalophylla x, g o
Phyllostomatidae (45)
Chrotopterus auritas x, g g
Lonchorchina aurita x, g g
Macrophyllum macrophyllum x, g g
Micronycteris brachyotis
M. megalotis x, g m
M. nicefori x, g g
M. minute x, g m
Mimon crenulatum x, g m
Phyllostomus discolor x, g g. m
P. elongatus x, g m
P. hastatus x, g g, m
Tonatia bidens x, g o
T. brasiliense x, g m
Tonatia sylvicola o o Linares, 1998
Trachops cirrhosis x, g m
Vampyum spectrum x, g o
Anoura caudifer x, g g
A. culturata x, g o
A. geoffroyi x,g o
Glassophaga longirostris x, g m
G. soricina x, g g, m
Carollia brevicauda x, g a
C. perspicillata x, g g, m
Sturnira erythromos x, g g
S. lilium x, g g. m
S. ludovici x, g g, m
Chiroderma salvini x g
C. villosam x, g o
Uroderma bilobatum x, g g
U. magnirostrum x, g g, m
Vampyressa pusilla x, g g
Vampyrodes caraccioli x g
Platyrrhinus helleri x, g g
P. umbratus x, g o
P. vittatus x o
Ametrida centurio x, g g
Artibeus cinereus x, g g, m
A. harti x
A. phaotis o g
A. jamaicensis x, g g, m
A. lituratus x,g g
Centurio senex o o Linares, 1986


Table 1 Continued.

Taxa Al B2 C3

Sphaeronycteris toxophyllum x g, m
Desmodus rotundus x, g m
Diphylla ecaudata. x o
Vespertilionidae (14)
Eptesicus brasiliensis x o
E. diminutus x o
E. furinalis x, g g, m
E. fuscus x o
Histiotus humbolti x o
Lasiurus blossevilli x g, m
(L. borealis)
L. cinereus x o
L. ega x,g m
Myotis albescens x, g m
M. keaysi x, g o
M. nigricans x, g m
Myotis simus o g
M. oxyotus x o
Rhogeessa tumida x, g g, m
Natalidae (1)
Natalus tumidirostis x o
Molossidae (15)
Eumops auripendulus x m
E. bonairiensis o m
E. dabbenei x m
E. glaucinus x m
E. hansae o m
Molossops greenhalli o m
M. paranus x o
M. temmincki o m
Molossus after x o
M. azteus x o
M. bondae x o
M. mollosus x, g m
M. sinaloae x o
Nyctinomops laticaudatus x o Linares, 1998
Tadarida brasiliensis o o Linares, 1998
Carnivora (17)
Urocyon cineroargenteus x o
Cerdocyon thous x g, m
Speothos venaticus a o Bisbal, 1989
Procycon cancrivorus x, g g, m
Potos flavus x, g g
Mustelafrenata o o Bisbal, 1989
Eira barbara o g, m
Galictis vittata o m


Table 1 Continued.

Taxa Al B2 C3

Conepatus semistriatus
Lontra longicaudis
Pteronura brasiliensis
Puma concolor
Panthera onca
Herpailurus yagouaroundi
Leopardus pardalis
Leopardus wiedii
Leopardus tigrina
Perissodactyla (1)
Tapirus terrestris
Artiodactyla (5)
Tayassu pecari
Tayassu tajacu
Odocoileus virginiana
Mazama americana
Mazama gouazoubira
Rodentia (27)
Sciurus granatensis
Heteromys anomalus
Oecomys bicolor (see text)
0. concolor (see text)
0. speciosus
0. trinitatis
Oryzomys albigularis
0. capitol (see tex, gt)
0. megocephalus (see text)
0. talamancae
Microryzomys minutus
Neacomys tenuipes
Rhipidomys venustus
R. venezuelae
Calomys hummelinki
Ichthyomys pittieri
Akodon urichi

Bisbal, 1989
Bisbal, 1989

Hato Pifnero
Hato Pifero

Bisbal, 1989

o g

o 0
o g,m

Hato Pifiero

Bisbal, 1991

x 0
x,g g,m

o o Linares, 1998

o o Handley and Mondolfi, 1963

x,g g


Table 1 Continued.

Taxa Al B2 C3

Zygodontomys brevicauda x, g m
Sigmodon hispidus x, g o
S. alstoni x m
Holochilus sciureus x o
Sphiggurus pruinosus o a CEBRG
Couendou prehensilis x, g g, m
Cavia aperea x, g a
Hydrochaeris hydrochaeris o m
Agouti paca x, g g
Dasyprocta agouti (leporina) x, g g, m
Echimys semivillosus x g, m
Proechimys guairae x, g g
Lagomorpha (2)
Sylvilagus brasiliensis x g
S. floridanus x m

x = occurs in one or more of the six states of north-central Venezuela, g = collected at or near Guatopo; as documented in Handley (1976).
'Found in g (Guatopo) and/or m (Masaguaral) as documented in Eisenberg et al. (1979) or Sunquist, Sunquist, and Daneke (1989) (vouchers
of Chiroptera and Rodentia at the Museum, Texas Tech University, Lubbock).
Other vouchered localities either by site, museum reference, or Linares,, 1998.
Also see Tables 2 and 3 for additional Chiroptera records.

states comprising north-central Venezuela is probably still low. Taxonomic
revisions will inflate the value and new discoveries may await us, but this present
value of 180 species is probably representative of our knowledge of Neotropical
mammals from comparable areas in Latin America at the close of the 20th
Venezuelan mammalogists have been enormously productive in the last
twenty years (Aguilera, 1985; Fernandez-Badillo et al., 1988; Linares, 1998). A
new journal, Marmosana, was established in 1995, and specific references within
the text attest to their activity.
Throughout the text abbreviations for collections will be employed: AMNH,
American Museum of Natural History; USNM, National Museum of Natural
History; CEBRG, Coleci6n de la Estaci6n Biologica de Rancho Grande;
CMNHNS, Coleci6n Museo Historia Naturales, La Salle; CMCN, Coleci6n Museo



Ciencias Naturales, Caracas. The three Venezuelan collections contain more than
22,000 specimens (Aguilera, 1985).


The senior author would like to thank the following individuals for their support and kindness
through the years: Ambassador Edgardo Mondolfi, Sr. Tomnis Blohm and his wife Cecilia, Charles 0.
Handley, Jr., Sr. Rafael Garcia, and the late Juan Gomez-Nuflez. During the late 1990s Sr. Antonio Julio
Branger made the grounds of Hato Pinlero available as a study site, and his staff was extremely helpful.
During our work at Hato Pinero, Francisco Bisbal and his staff at Rancho Grande were supportive in many
ways. Rafael Hoogesteijn played critical roles in initiating the effort at Piflero. Francisco Bisbal was helpful
and supportive in many ways. Melvin E. Sunquist was a source of support and gladly aided at many phases
of the research project in Venezuela. Mel and his wife, Fiona, were part of the original team at Hato
Field research at Hato Pifiero was funded by The National Geographic Society, the Wildlife
Conservation Society, the British Embassy to Venezuela's Cooperation Fund, and the Lincoln Park Zoo
Scott Neotropic Fund. Critical assistance in the field was provided by: Juan Victor Meires, Orlando
Ramirez, Gilson Rivas Fuenmayor, Diego Giraldo, Sandra Melman, Emiliana Isase, Telva Carantona, and
Marcus Trepte. Important observations were contributed by Daniel Scognamillo, Ines Maxit, Laura Farrell,
Roy McBride, Rafael Hoogesteijn, Rocky McBride, Tibisay Escalona, Bruno Pampour, Rafael Ortiz, Dr.
Francisco Delascio, the llaneros of Piinero, and students and staff of Dr. Juhani Ojasti's wildlife management
class (UNELLEZ at Guanare).
The authors thank Melvin Sunquist and Charles A. Woods for helpful comments on the manuscript


The annotated list does not include the ubiquitous Homo sapiens. The
numerous feral populations it has introduced are also excluded: Felis catus, Canis
familiaris, Equus caballus, E. asinus, Sus scrofa, Capra hircus, Bos taurus,
Bubalus bubalis, Mus musculus, Rattus rattus, and R. norvegicus. Also excluded
are the strictly aquatic manatee (Trichechus manatus) and two species of
freshwater dolphins (Inia and Sotalia).


The marsupials of Venezuela have received monographic treatment by Perez-
Hernandez, Soriano, and Lew (1994).


The distribution of the didelphids in Venezuela was reviewed by Perez-
Hernandez (1989). O'Connell (1979) described the ecology of the common
marsupials in northern Venezuela. The absence of the widespread Philander and
Metachirus from north-central Venezuela is suggestive of the "insular" character of
the north coastal ranges and Ilanos.


Didelphis marsupialis
Common Opossum

The opossum is widespread in the neotropics below 1500 m. It is tolerant of
human disturbance. It prefers forested habitats and is most abundant in evergreen
forest although it exploits dry, deciduous forest that occurs near permanent water.
It can achieve densities of 50-100/km2 in optimum habitat. Home ranges overlap
and adult females have the smallest range at 16 ha; males can range over 120 ha
(Sunquist, Austad, and Sunquist, 1987).

Chironectes minimus
Water Opossum

In north-central Venezuela the species is confined to streams in the north
mountain ranges.
P. August trapped several individuals in Guatopo.

Marmosops fuscata

This species is more terrestrial than other mouse opossums. It prefers
premontane forests and is most abundant at about 1000 m elevation. O'Connell
(1979) studied it in Guatopo.

Marmosa murina

The species occurs in only a few locations in northern Venezuela. Handle
(1976) assigned specimens from Yaraguy to the species. It is strongly associated
with lowland, tropical evergreen forest. Perez-Hernandez et al. (1994) recorded
specimens from Cojedas and Aragua; Eisenberg et al. (1979) recorded specimens
in Guatopo.

Marmosa robinsoni

The common lowland mouse opossum in northern Venezuela is tolerant of
dry, deciduous forest. It was studied extensively at Hato Masaguaral (O'Connell,
1979, 1983).

Gracilinanus marica

This diminutive mouse opossum is found in the north coast range of
Venezuela. Most specimens are taken between 1500 and 2000 m in moist,
tropical, montane forest.


Micoureus demararae (Marmosa cinerea)

In northern Venezuela it is confined to moist, lowland tropical forest and is
widely present in the coast ranges. It has been studied in Guatopo by O'Connell

Monodelphis brevicaudata
Short-tailed opossum

In north-central Venezuela it is confined to the moist forests of the northern
and southern coastal ranges. It has been studied in Guatopo by O'Connell (1979).

Monodelphis orinoci

Reig et al. (1977) separated specimens of Monodelphis from the Ilanos into
the species orinoci while retaining brevicaudata for specimens taken in the coastal
ranges. Perez-Hernandez (1989) and Perez-Hernandez, Soriano, and Lew (1993)
retained the separation and plot locations in Cojedas, Guarico, and Bolivar. It is
apparently smaller in size than M. brevicaudata (see also Ventura, Perez-
Hernandez, and Lopez Fuster, 1988).

Caluromys philander
Wooly Opossum

This species is widespread in northern Venezuela but is strongly associated
with lowland, tropical evergreen forest and is highly arboreal. It was studied by
O'Connell (1979) in Guatopo.

Note: Linares (1998) notes records based on single specimens of
Gracilnanus agilis and G. emiliae from north-central Venezuela which were not
noted by Perez-Hernandez et al. (1994) nor Perez-Hernandez (1989). Further
study is necessary since these records are distant from defined centers of their
geographical distribution.



Long-nosed Armadillos

The long-nosed armadillos have been reviewed by Wetzel and Mondolfi
(1979 and Wetzel (1985). Redford (1985b) summarized feeding ecology.


Ferguson-Laguna (1984) studied dasypodid ecology in the savannas of northern

Dasypus novemcinctus
Nine-banded Armadillo

A highly adaptable species, it is found in most lowland habitats of north-
central Venezuela. No specific studies have been made in northern Venezuela but
it is well studied elsewhere (see McBee and Baker, 1982). It is widely hunted for
its meat and could be managed as a small game species. Density estimates range
from 5 to 30.4/km2. Individual home ranges have been measured at 3.45-13.8 ha
in the southern United States.

Dasypus sabanicola
Savanna Armadillo

This species exhibits a fragmented distribution in the Ilanos north of the
Orinoco, and is primarily associated with los llanos bajos. It was first described by
Mondolfi in 1968; the ecology was studied by Ferguson-Laguna (1984).

Cabassous unicinctus
Naked-tail Armadillo

The genus Cabassous was reviewed by Wetzel (1980). The fossorial habits of
naked-tail armadillos means they are seldom seen or collected. Records of
occurrence are scattered in the literature. A specimen was collected in Guatopo in
1967 and deposited with E. Mondolfi.

Priodontes maximus

This species has been hunted to extinction over much of its range in northern
Venezuela. It is still to be found in Apure but is probably extinct in Guarico
(Mondolfi pers. comm.).


Myrmecophaga tridactyla
Giant Anteater

Although the giant anteater is tolerant of a range of lowland tropical habitats,
it appears to be most abundant in the open woodlands of the Ilanos. It was studied
by Montgomery and Lubin (1977) at Masaguaral. Home ranges were large,


exceeding 25 km2. Redford (1985a) summarized its feeding ecology. Because it is
vulnerable to hunting, it is under threat.

Tamandua tetradactyla
Tamandua Anteater

This species is found in a variety of lowland habitats in northern Venezuela.
It seldom ranges above 500 m. It is abundant in dry, deciduous forest but also is
found in moist evergreen forests. The species has been studied by Montgomery
and Lubin (1977) at Hato Masaguaral. Home ranges of 375 ha were recorded.


Bradypus variegatus
Three-toed Sloth

This species is confined to forested areas, especially moist, evergreen forest.
It can extend to dry forest but always near permanent water. The sloth is common
in Guatopo but is absent from the llanos, except where there is extensive evergreen
forest near permanent water. The species has been well studied in Panama (see
Montgomery and Sunquist [1974, 1975, 1978] and Sunquist and Montgomery
[1973]). In appropriate habitats the three-toed sloth may reach high densities (3-
4/ha). The home range is quite small, usually less than 1.0 ha.



Cryptotis meridensis
Short-tailed Shrew

Linares (1998) notes two vouchered specimens from the north coastal range
in the state of Aragua.


The isolated nature of the north-central mountain ranges and associated
llanos is dramatically illustrated by the distribution of primates. The area covered
in this paper has just three primate species (see also Eisenberg, 1979a; Bodini and
Perez-Hernandez, 1987).


Cebus olivaceous
Wedge-capped Capachin Monkey

Handley (1976) and others have employed the name C. nigrivittatus, but see
Groves (1993). This species is tolerant of a range of lowland forest types but is
never found far from permanent water. It penetrates the llanos through forests
associated with major rivers. Within the llanos, it primarily uses gallery and
semi-deciduous forests, but will make forays into more open forest types and even
savannas at times. This species was intensively studied at Hato Masaguaral and to
a lesser extent at Hato Piflero. Foraging, diet, and social structure were studied by
Robinson (1981, 1984a, 1986, 1987); vocalization by Robinson (1984b and Norris
(1992); grouping tendencies by Ruiter (1986) and O'Brien and Robinson (1993);
development by Fragaszy (1990); all at Masaguaral. Miller (1996) has initiated a
long-term study at Hato Piflero.

Alouatta seniculus
Red Howler Monkey

One of the most adaptable species of New World primates, it enjoys a wide
distribution. It has colonized the llanos occupying open woodlands and tree
islands (matas) within the grasslands. This is one of the best studied species.
Braza (1980) conducted a major study at Hato El Frio, Apure. The species was
worked on for two decades at Hato Masaguaral (see Neville, 1972, Rudran, 1979,
Mack, 1979). Vocalizations were studied by Sekulic (1981, 1982b, 1982c) and
Sekulic and Chivers (1986); locomotion and anatomy by Schon-Ybarra (1984);
demography by Rumiz (1992); population genetics by Pope (1989, 1990, 1992,
1998); and finally a classic series of papers on demography and infancticide by
Crockett (1984, 1996), Crockett and Eisenberg (1987), Crockett and Pope (1984,
1988), Crockett and Rudran (1987a, b) and Crockett and Sekulic (1984).

Ateles belzebuth
Spider Monkey

The main distribution in Venezuela is south of the Orinoco and the
Maracaibo Basin. Mondolfi and Eisenberg (1979) discussed distributions in
northern Venezuela. Isolated populations still persist in Guatopo and Cupira. The
ecology has not been studied in the north, but Castellanos and Chanin (1996) have
studied this species in Bolivar. Mature forest with continuous canopy seems to be
the preferred habitat. It can occupy semi-deciduous tropical forest but evergreen
forest seems to be optimal.



Bisbal (1989) has reviewed the distribution and the status of the Carnivora of

Urocyon cinereoargenteus
Gray Fox
Apparently this species is confined to north-central Venezuela. There it
prefers tropical dry forest at elevations of 100-700 m (Bisbal, 1982). Vouchered
records from Cojedes come from the southern limits of the coast range. Mondolfi
and Bisbal (1985) analyzed the diet.

Cerdocyon thous
Crab-eating Fox
The most widespread canid in Venezuela, it is tolerant of a range of habitats
but is most abundant in dry deciduous forest, particularly along savanna/forest
ecotones. Brady (1979), Montgomery and Lubin (1978a), and Sunquist et al.
(1989) studied its behavior and ecology at Hato Masaguaral. Adult foxes live in
monogamous pairs and defend a home range of 45-100 ha. A crude density of
2.5/km2 was estimated for Masaguaral.

Speothos venaticus
Bush Dog
This species is still noted south of the Orinoco. Bisbal (1989) plotted
specimens from Carabobo, and Linares (1968) discussed the former presence of
specimens from northern Venezuela. Strahl et al. (1992) summarized previous
records, added accounts from south of the Orinoco, and discussed hunter's
observations in northern Venezuela in the vicinity of Parque Nacionales San
Esteban and Guatopo. This species is vulnerable to infection by canine distemper
and may be wiped out by contact with domestic dogs. Its status is unclear in
North-Central Venezuela.


Procyon cancrivorus
Crab-eating Raccoon
This species is common throughout Venezuela. It is associated with both
humid forest and seasonally wet forest. It has usually been taken at elevations
below 500 m and generally near water.



In north-central Venezuela this species is found in tropical humid forest and
premontane humid forest. It is absent over much of the Ilanos.


Mustela frenata
Long-tailed Weasel

In northern Venezuela it is confined to the north coastal ranges and is absent
from the llanos. It is abundant in the Andean states of Tachira and Merida.

Eira barbara

In northern Venezuela it is strongly associated with tropical humid forest but
extends into the llanos in gallery forests. Sunquist et al. (1989) studied the species
at Masaguaral and confirmed its dependency on forest cover. One female had a
home range of 9 km2 during the dry season.

Galictis vittata

This species is widely distributed in north-central Venezuela. It prefers
tropical dry forest at modest elevations. Sunquist et al. (1989) studied this species
at Masaguaral and found a female used a minimum area of 4.15 km2. The species
is mainly carnivorous (Bisbal, 1986).

Conepatus semistriatis
Hog-nosed Skunk

In Venezuela the species does not occur south of the Orinoco River. It is
strongly tied to dry deciduous forest but may also be found in moist forest. It was
studied by Sunquist et al. (1989) at Masaguaral, where one female had a home
range of 53 ha in the dry season. Density estimates for Masaguaral were 6-12/km2.

Lontra longicaudis

The species is found in rivers of the north coast range, but records are sparse.
The status is unknown for north-central Venezuela (Bisbal, 1989).


Pteronura brasiliensis
Giant Otter

Although the species occurs in the Orinoco River and Territorio Amazonas, it
has been exterminated over much of its range. There is one vouchered specimen
from southern Guarico (Bisbal, 1989).


The biology of the smaller cats is reviewed by Mondolfi (1986).

Leopardus pardalis

The ocelot is found in all the life zones of Venezuela at lower elevations. It
prefers forested habitats, but makes use of forest edge. Extending into the llanos
via gallery forests, it is common in all semi-deciduous forest types there. It was
studied by Sunquist and Ludlow at Masaguaral (Ludlow, 1986; Ludlow and
Sunquist, 1987; Sunquist et al., 1989) and at Hato Pifiero by Farrell (1998). Adult
males defend a territory that includes the territories of from one to three adult
females. Male home ranges of 6.2-9.4 km2 during the wet and dry seasons contrast
with female ranges of 1.4-2.6 km2. Densities of 0.25 individuals/km2 can be
achieved in gallery forest habitat.

Leopardus wiedii
Margay Cat

In north-central Venezuela this cat has been taken in the north coastal ranges.
It is strongly tied to humid, closed canopy forests, no doubt reflecting its strong
arboreal tendencies (Bisbal, 1989).

Leopardus tigrinus
Little Spotted Cat

Very few specimen records exist for this cat in Venezuela. It is known from
the premontane forests of Aragua, Miranda, and the Districto Federal. (See also
Bisbal, 1989.)

Herpailurus yagouroundi

A frequently seen cat due to its diurnal habits, it is strongly associated with
tropical dry forest and is commonly encountered at the interface of grasslands and


forests. Although most abundant in the lowlands, it can occur up to 900 m
elevation. The jaguarundi is rarely found south of the Orinoco (Bisbal, 1989).

Puma concolor

Records from north-central Venezuela are scattered. This cat has suffered
greatly from persecution by humans, but is still found in virtually all national parks
and better protected larger cattle ranches in North-Central Venezuela. It has been
taken in Guatopo, and is an occasional visitor at Masaguaral. Pumas used nearly
all habitat types in Hato Pifiero, Cojedes, slightly overlapping the habitats preferred
by jaguars. Capybara (Hydrochaeris hydrochaeris), white-tailed deer (Odocoileus
virginianus), and collared peccary (Tayassu tajacu) were among the top prey items
(Farrell, 1999).

Panthera onca

The status of the jaguar in Venezuela has been discussed by Mondolfi and
Hoogesteijn (1986) ; Hoogesteijn and Mondolfi (1993); and Hoogesteijn,
Hoogesteijn, and Mondolfi (1993). The jaguar was widespread in Venezuela, but
persecution has confined it to the larger national parks or large tracts of ranch
lands. We found jaguar in Guatopo and recorded occasional visits at Masaguaral.
Jaguars occur at low densities in most of the national parks in the northern states.
More suitable habitat and greater numbers occur in los llanos altos, where large
connected patches of semi-deciduous forest and gallery forest create suitable areas
for residency that typically include several cattle ranches. This was species was the
subject of an intensive study at Hato Pifiero, where abundant habitat remains and
persecution is rare (Farrell, 1999). Pifilero is connected to other ranches containing
intact habitat and adequate natural prey. It is this network of connected forests that
maintain jaguars in los llanos altos. Both species of peccaries, capybaras, and
spectacled caiman (Caiman crocodilus) are among the most important prey items.
Though clearly fond of forest cover, when secure from hunting, jaguars will use
savanna edges, small savannas, and dirt roads.


Tapirus terrestris

Once widely distributed in northern Venezuela, this species has become
locally extinct in many areas through hunting and land development. In north-
central Venezuela it still occurs in the larger national parks and some of the larger
cattle ranches. Tapirs occurred at a crude density of 0.6/km2 in Guatopo. Though


associated with tropical evergreen forests in much of its range, in the Ilanos it uses
gallery forest and semi-deciduous forest. During the dry season, it may reside in
quite dry habitats, but will travel regularly to water holes. Food habits and
preferred habitat use patterns were studied by Salas (1996) and Salas and Fuller
(1996) in southern Venezuela.


Tayassu pecari
White-lipped Peccary

Although widely distributed south of the Orinoco River and present in
western Apure, this species is rarer north of the Orinoco. Groups occur along the
more heavily forested parts of the coastal range. Hernandez et al. (1995) described
behaviors and Barreto et al. (1997) described diet in Hato Pifiero (Barreto and
Hernandez, 1988). Hato Pifiero is part of the range of two large groups (60 and
115 individuals) and there are reliable reports from other parts of Cojedes and
extreme western Guarico. More tied to deep forest than Tayassu tajacu, T. pecari
persists in los llanos altos where long strips of relatively moist forest along active
waterways or old river beds, combine with large patches of semi-deciduous forest.
Despite a clear preference for forest and a strong tie to water in the dry season,
these groups will use savanna edges and small savannas particularly when the
introduced mango fruits (Mangifera indica) are in season. Predictable visitation
sites at water during the dry season, combined with a fearlessness maladaptive for
the age of firearms, makes this species very vulnerable to overhunting.

Tayassu tajacu
Collared Peccary

Grubb (1993) recommended the generic name Pecari for this species. It was
found over much of lowland Venezuela, has been locally extirpated in many areas,
yet is more resilient than its larger cousin T. pecari, due to: 1) smaller, more
widely dispersed groups that occupy a larger variety of habitats; and 2) its greater
caution in response to humans, and fragmentation of groups when in flight.
Robinson and Eisenberg (1985) discussed seasonal variation in group size, and
Castellanos (1983) analyzed the ecology of the species with the use of
radiotelemetry. In Guatopo populations had a crude density of 1.9/km2. In the dry,
deciduous forests of the llanos at the Hato Masaguaral densities of 8.5/km2 were
estimated. Food habits were studied at Hato Pifiero by Barreto et al. (1997). A
greater proportion of Pithecellobium saman (a moisture loving tree) seeds were
found in the T. pecari diet. Forest edge seeds were slightly more common in T.
tajacu stomachs, forest interior seeds more common in T. pecari stomachs (Barreto


et al., 1997). Still on a preliminary basis, with a small sample, this resource
partitioning on a dietary basis was equivocal. Responses to human hunting
pressure now affect these ecological relationships and T. pecari may in part be
where it is now because it can persist there. Although T. pecari groups are larger,
and on a year-round basis make use of a larger net area, T. tajacu groups are more
generalists, taking advantage of more habitat types, and specifically, more dry
ecotones than their larger cousins. Although groups as large as 30-40 may occur,
observers at Hato Piflero more frequently encountered groups of 11-15 animals. T.
tajacu is a preferred prey of both pumas and jaguars.


Mazama americana
Red Brocket

Present over much of lowland Venezuela, absent over most of the llanos,
penetrating only the western llanos altos in wide gallery forests. It seems to prefer
moist evergreen forest and is seldom found far from permanent water. Bisbal
(1991) reviewed the distribution. It is a browser, but feeds seasonally on fruit
(Bisbal, 1994). The species is abundant in Guatopo and Avila.

Mazama gouazoubira
Brown Brocket

This species has a disjunct distribution in northern Venezuela (Bisbal, 1991).
It is more tolerant of drier sites and deciduous forests than is M. americana. It has
been collected in deciduous forests in the northern coastal range in Estados Aragua
and Carabobo, and occupies thorn forests in Falcon and Yaracuy.

Odocoileus virginianus
White-tailed Deer

This species is found over much of lowland Venezuela and is tolerant of a
wide range of habitat types. Highest densities occur in the forested savannas
found over much of the llanos. It will use the fringes of tropical evergreen forest
where it can co-occur with M. americana. Densities of 2/km2 are an average value
for much of the llanos. Brokx (1972a, b, 1984) and Brokx and Anderssen (1970)
pioneered the study of 0. virginianus in northern Venezuela. Stiiwe (1985)
compared the biology of Venezuelan and Virginian (U.S.) populations. 0.
virginianus is both important to pumas as a prey item, and over-hunted virtually
everywhere it is not rigorously protected.



Sciurus granatensis
Neotropical Red Squirrel

This species is abundant in north-central Venezuela but is confined to
forested tracts. It is able to exploit dry, deciduous forest and thereby extends into
the Ilanos. O'Connell (1981, 1982) has studied the species at Masaguaral and
Guatopo; Heaney and Thorington (1978) studied it in Panama. Nitikiman (1985)
reviewed the species.


Heteromys anomalus
Forest Spiny Pocket Mouse

This mouse is widely distributed in northern Venezuela but is most abundant
in moist evergreen forests, although it extends into dry zone forests near permanent
water. O'Connell (1981) studied the species at Masaguaral and Guatopo and Rood
(1963) and Valdez et al. (1984) at Rancho Grande.

Rats and Mice

New World Rats and Mice

Note: The names Oryzomys capitol, 0. bicolor, and 0. concolor were employed
by Handley (1976), Eisenberg, O'Connell, and August (1979), O'Connell (1981,
1982), and August (1981, 1983, 1984). The taxonomic revisions by Musser and
Williams (1985) and Musser, Carleton, Brothers, and Gardner (1998)
supplemented by Aguilera et al. (1995) and Perez-Zapata et al. (1986) have led to
the usage of new binomials throughout the following discussions of the species.

Arboreal Rice Rats

This group was raised from the level of subgenus to full genus (see Musser
and Carleton, 1993).


Oecomys bicolor

Handle (1976) employed bicolor as the species name for the second smaller
species of Oecomys found in northern Venezuela, the name 0. speciosus is now
recognized for the lowland dry deciduous forest forms (see below).

Oecomys concolor

Handle (1976) used this name for the larger species of Oecomys found in
northern Venezuela. The elevation of 0. trinitatus to a full species suggests this
name be applied to the northern Venezuelan forms (see below), while 0. concolor
be retained for forms south of the Orinoco (Linares, 1998).

Oecomys flavicans

Linares (1998) lists this species as a shorter-tailed form that, in a macro-
habitat sense, may co-occur with 0. trinitatis. Its relationship with trinitatis and
0. concolor remains to be determined.

Oecomys speciosus

This arboreal rice rat is found in the palm and open woodlands of northern
Venezuela (Musser and Carleton, 1993). O'Connell (1981) studied it at

Oecomys trinitatus

This arboreal rice rat is found in the evergreen forest habitats of northern
Venezuela. O'Connell (1981, 1989) studied it at Guatopo.


Oryzomys albigularis

Nominal forms of 0. albigularis were studied by Aguilera, Perez-Zapata, and
Matino (1995) in northern Venezuela. A diploid number (2n) of 66 was recorded
for all samples, but the number of autosomal arms (FN) varied: 90,92,104. They
postulated the variations represent an allospecies. 0. caracolus Thomas, 1914,
(2n=66, FN=90) is assigned to populations from the north coastal range of


Oryzomys capitol

Handley (1976) employed capitol as the species name for the terrestrial rice
rats inhabiting the lower altitudes in northern Venezuela. At the time 0. capitol
included many subspecies following Hershkovitz (1944). Current thinking has
resulted in the break-up of capitol into some full species. Redefinitions of species
initiated by Musser and Williams (1984) established that 0. talamancae be
recognized in northern Venezuela and was reinforced by Perez-Zapata et al.
(1986). Oryzomys yunganensis may be the more appropriate name for the non-
talamancae forms in northern Venezuela and south of the Orinoco, but more
research is necessary.
Musser et al. (1998) have revised 0. capitol and suggest 0. megacephalus be
employed as the senior name for the remaining forms previously included in 0.

Oryzomys talamancae

Perez-Zapata et al. (1986) investigated the 0. capito-like complex in northern
Venezuela. The low chromosome numbered form (2n=34) was provisionally
assigned to 0. talamancae. O'Connell (1981, 1989) studied it at Guatopo.

Microryzomys minutus
Pygmy Rice Rat

This species occurs in high montane areas of northern Venezuela at
elevations of 1150-3810 m; it is essentially terrestrial in the cloud forests. Handley
(1976) has taken specimens at Parque de Avila. Aagaard (1982) studied the
species at Merida.

Oligoryzomys fulvescens

This species is widespread in north-central Venezuela, ranging to 2000 m
elevation but most abundant at < 1500 m. It is tolerant of disturbed habitats and is
found in agricultural areas. Moist sites are preferred but it also is found in
evergreen and dry, deciduous forests.


Neacomys tenuipes
Spiny Mouse

The spiny mouse is abundant in the mountains of north-central Venezuela
from 404 m to 1655 m elevation. It is associated strongly with moist forest.
O'Connell (1981, 1989) studied the species in Guatopo.


Climbing Rats

Handle (1976) recognized seven species from Venezuela. Pending a
revision of the genus his designations are followed.

Rhipidomys couesi

Linares (1998) records this species from the north coastal range, although it
is associated strongly with the Andes.

Rhipidomys venzuelae

The common arboreal rat is found in the lowlands. The species is reasonably
abundant below 1500 m and occupies evergreen and deciduous tropical forests.
O'Connell (1981, 1989) studied it at Masaguaral and Guatopo.

Rhipidomys venustus

This higher altitude species is taken from 1160 m to 3160 m (Handley, 1976). In
north-central Venezuela it occurred in cloud forest and tropical evergreen forests in
the extreme northern ranges of Districto Federal and Miranda.


Calomys hummelincki

Although noted in the Falcon arid zone and scattered locations in Venezuela
(Eisenberg, 1989), Linares (1998) records two specimens from the state of Guarico.



Ichthyomys pittieri
Fishing Rat

The fishing rat was described by Handley and Mondolfi (1963) from Aragua
and subsequently studied by Voss, Siva, and Valdes (1982). This is a high altitude
form found near streams.


Akodon urichi

The only species of Akodon in north-central Venezuela, it is found in moist
sites usually in tropical evergreen forests above 500 m (Handley, 1976). It also is
found in north and south coastal ranges. O'Connell (1981, 1989) studied it at


Zygodontomys brevicauda
Cane Rat

This common lowland, terrestrial rat is found in the grasslands of Venezuela,
usually near streams or moist areas. Density can reach very high numbers during
the breeding season. Numbers can fluctuate from 260 to 7400/km2. O'Connell
(1981, 1989) and Vivas (1986) studied it at Masaguaral. Aguilera (1985) studied
growth and reproduction in captivity. Zygodontomys microtinus is a frequently
used synonym.


Sigmodon hispidus
Cotton Rat

The cotton rat occurs in northwestern Venezuela and is associated with moist
grassy sites in the lowlands. It is tolerant of human activities. It invades
agricultural fields and pastures. S. alstoni replaces it in the Ilanos.


Sigmodon (Sigmomys) alstoni

This rat is found in grassland sites, subject to seasonal inundation. Tolerant
of human activities, it occupies pasture and agricultural fields. O'Connell (1981,
1989) and Vivas (1986) studied it at Masaguaral and vicinity.

Holochilus sciureus
Marsh Rat

The name sciureus is usually applied to the central Venezuelan forms
(Musser and Carleton, 1993), but Handley (1976) employed H. brasiliensis.
Aguilera et al. (1993) separate populations of Holochilus from northern Venezuela
with a 2n chromosome number of 46 as H. venezuelae. They contrast this value
with samples they assigned to H. brasiliensis from the Amazon area. The problem
needs to be resolved. Marsh rats, as the name implies, are associated with lowland
wet sites. They can become a serious agricultural pest.



Prehensile-tailed Porcupines

The hairy dwarf porcupines were separated in their own genus Sphiggurus by
Husson (1978; Woods, 1993). Other authors include them in Coendou (Emmons
and Feer, 1997).

Coendou prehensilis

The common lowland, prehensile-tailed porcupine is usually found in tropical
evergreen forest below 1000 m. It can extend to lowland dry forest in the vicinity
of permanent water. Montgomery and Lubin (1978) studied it at Masaguaral.
Roberts, Brand, and Maliniak (1985) recorded reproduction and growth of this
species in captivity.

Sphiggurus vestitus pruinosus

Woods (1993) included pruinosus as a subspecies of vestitus. Handley (1976)
treated priunosus as a full species.
Concepci6n and Molinari (1991) examined a small, long-haired porcupine
from 3.5 km southwest of Zea in the state of Merida. The chromosome numbers


were 2n=42 and FN=76. These numbers are identical with Erethizon but different
from Coendou rothschildi. They advocate the separation of the genus Sphiggurus
from Coendou, a view opposed by Handley and Pine (1992).
I (JFE) examined a specimen at CEBRG with the following data on the label:
1 male caught at Rancho Grande, Aragua; Venezuela. Measurements 490-210-51-
23 wt 900 g. The tail is very short (42% of HB); the dorsal hair is long with the
underlying spines evident on the lower back; the spines are white-tipped.
This taxon is rare in collections and appears to be confined to montane
evergreen forests.


Cavia aperea
Guinea Pig

Handley (1976) listed the forms from Venezuela as C. porcellus. This name
is usually applied to feral domestic or domestic forms (Woods, 1993). Ojasti
(1964) reviewed the problem of the origins of guinea pigs in Venezuela. Whether
they are escaped domestics or represent relictual populations remains to be
determined. The distribution is fragmented. Handley (1976) took specimens from


Hydrochaeris hydrochaeris

This species is potentially abundant throughout the lowland tropics but is
always associated with wet sites. Its highest abundance is reached in the wet llanos
(llanos bajos). Ojasti (1973) demonstrated that it can be harvested as a renewable
resource. Its behavior has been studied by Ojasti (1986) and Azacarate (1980).
The current distribution of capybara is not only dictated by suitable mixes of
water, forage, and cover, but also by the exceptionally intense hunting pressure it
receives annually for semana santa. Though large groups occur on a number of
well-defended and managed private ranches, the situation "outside" is more grim.
An important prey item for both jaguar and puma.



Dasyprocta aguti (D. leporina)

The specific name for agoutis in north-central Venezuela remains in dispute;
D. leporina is frequently used. Ojasti (1964) made an attempt at a preliminary
revision. Woods (1993) offers the latest thinking on the problem. Emmons and
Feer (1997) have attempted to standardize the nomenclature.
This species is associated with moist areas in lowland forests and can extend
into dry, deciduous forests. It is an important small game species but can become
locally extinct through hunting. Smythe (1978) has offered the most thorough
study of ecology and behavior of the closely related D. punctata in Panama.


Agouti paca

This species can be common in tropical evergreen forests near permanent
water. It extends into the llanos via gallery forests. Hunted for its meat, which is
the most prized in the neotropics, it has been driven to local extinction in many
areas. Smythe (1995) and associates have pioneered in the study of the
domestication process for this species as a means of supplying meat to the specialty
food markets.


Echimys semivillosus
Arboreal Spiny Rat

Makalata didelphoides (Echimys armatus) has been described from Trinidad
and the adjacent coast of Venezuela. Echimys semivillosus is known from the
gallery forests of llanos in north-central Venezuela, but specimens of Echimys from
the coastal ranges need to be checked for their species identity. Echimys are
present in Guatopo but were not collected. Echimys semivillosus were studied by
August (1981b) at Masaguaral; they were taken in gallery forest. Sunquist et al.
(1989) found Echimys was preyed upon by Eira barbara. The whole group is
being revised by Emmons (pers. comm.).



Proechimys guairae

Handley (1976) listed the northern Venezuelan specimens as P. semispinosus,
but see Gardner (1989). Reig et al. (1976, 1980) examined the karyotypic diversity
of spiny rats and proposed a revision of the group. Aguillera and Conti (1994)
discussed the chromosomal speciation exhibited by Proechimys in northern
Venezuela. Within the area discussed in this report the name P. guairae, a super
species, is applied. The 2n number varies: 44, 46, 48, 50, 52 with an FN ranging
from 72 to 74. This species is abundant in lowland to mid-elevation evergreen
forests and gallery forests. O'Connell (1981, 1989) studied it at Guatopo. Linares
(1998) lists this form as P. trinitatis. Its common occurrence in ocelot scats in
Hato Pifiero attest to its residency in the large tracts of semi-deciduous forest there.

Hares and Rabbits

Cottontail Rabbits and Allies

Sylvilagus brasiliensis
Brazilian Cottontail

The Brazilian cottontail has a fragmented distribution in north-central
Venezuela. It prefers moist habitats, usually in clearings or on the edge of
evergreen tropical forests. It can range to elevations of 1500 m. The species is
present in Guatopo.

Sylvilagus floridanus

This species is common in lowland habitats, often quite dry such as thorn
forests, savannas, and range lands. It replaces S. brasiliensis in the llanos. It is
abundant at Hato Masaguaral and Hato Pifiero. Ojeda-Castillo and Keith (1982)
present data on the breeding biology of this species from Guarico. Though
widespread in Hato Pifiero, frequencies were highest in well-drained dry habitats.


Numerous taxonomic and distribution studies of bats have been carried out in
Venezuela. Handley (1976) offered excellent data on habitat preferences and
altitudinal distributions. Eisenberg (1989, pp. 424-425) analyzed some trends in


Table 2. Species of Chiroptera with records in north-central Venezuela according to Linares (1987, 1998).

Emballonuridae (8)
Rhynchonycteris naso
Saccopteryx bilineata
S. leptura
S. canescens
Peropteryx macrotis
P. kappleri
P. trinitatus
Diclidurus albus

Noctilionidae (2)
Noctilio albiventris
N. leporinus

Mormoopidae (5)
Mormoops megalophylla
Pteronotus davyi
P. gymnonotus
P. parnelli
P. personatus

Phyllostomidae (56)
Phyllostominac (21)
Micronycteris brachyotis
M. hirsuta
M. megalotis
M. microtis
M. minute
M. nicefori
M. schmidtorum
M. sylvestris
Tonatia bidens (T. saurophila)
T. brasiliense (T. venezuelae)
T. silvicola
Lonchorhina aurita
Macrophyllum macrophyllum
Mimon crenulatum
Trachops cirrhosus
Phyllostomus discolor
P. elongatus
P. hastatus
Phylloderma stenops
Chrotopterus auritus
Vampyrum spectrum

Glossophaginae (8)
Glossophaga soricina
G. longirostris
Leptonycteris curasoae
Anoura caudifer
A. culturata

A. geoffroyi
A. latidens
Choeroniscus godmani

Carollinae (2)
Carollia brevicauda
C. perspicillata

Sturnirinae (3)
Sturnira lucovici
S. lilium
S. erythromos

Stenodermatinae (19)
Uroderma bilobatum
U. magnirostrum
Platyrrhinus (Vampyrops) dorsalix
P. helleri
P. umbratus
P. vittatus
Vampyrodes caraccioli
Vampyressa pusilla
Chiroderma salvini
C. trinitatum
C. villosum
Artibeus harti
A. cinereus
A. lituratus
A. jamaicensis
Ametrida centurio
Sphaeronycteris toxophyllum
Centurio senex

Desmodontinae (3)
Desmolus rotundus
Diphylla ecaudata
Diaemus young

Natalidae (1)
Natalus stramineus

Thyropteridae (1)
Thyroptera discifera

Vespertilionidae (15)
Histiotus humboldti
Lasiurus cinereus
L. blossevilli (borealis)
L ega
Rhogoessa tumida


Table 2 Continued.

R. minutilla
Myotis albescens
M. nigricans
M. oxyotus
M. keaysi
Eptesicus brasiliensis
E. diminutus
E. furinalis
E. fuscus
E. montosus (E. andinus)

Molossidae (20)
Molossops greenhalli
M. paranus
M. planirostris
M. temmincki

Eumops auripendulus
E. bonariensis
E. dabbenei
E. glaucinus
E. hansae
E. perotis
Nyctinomops aurispinosa
N. laticaudata
Molossus ater
M. bondae
M. coibensis
M. molossus
M. pretiosus
M. sinaloae
Promops centralis
Tadarida brasiliensis

A total of 108 species.

altitudinal distributions and concluded that the highest species richness for bats lies
below 700 m elevation. Some species are adapted for higher elevations, including
Lasiurus cinereus, Histiotus montanus, and Eptesicus fuscus among others. Some
species of Sturnira are also well adapted to moderate elevations. Ochoa-G. (1980)
provided a useful analysis of bat altitudinal distributions by means of a transect
over the north coastal range passing from the Valencia depression through Rancho
Grande to the Caribbean coast.
Linares (1987, 1998) provided a fine summary of the distribution and ecology
of Venezuelan bats. Ibafilez (1981) offers a comprehensive monograph on the
ecology and biology of bat communities in the llanos of Apure.
During our work in Guatopo and at Masaguaral, August (1979, 1981, 1985)
pursued research on Artibeus jamaicensis, and August and Baker (1982) made
important collections resulting in a paper on reproductive ecology for some key
Table 2 offers a possible list of bat species that could occur in north-central
Venezuela deriving from the range maps of Linares (1987, 1998). This list can be
compared with our vouchered records in Table 1. Table 3 lists the subset of bats
with restricted ranges within north-central Venezuela.




Taxonomic studies of mammals have helped to clarify the species
composition within the ecological communities to be found in northern Venezuela.
The bibliography concluding the volume prepared by Aguilera (1985) summarizes
those efforts, while Linares (1998) provides a recent synthesis. Work on the
mammals during the last forty years involved the efforts of investigators not only
within Venezuela but also from the United States, Canada, Spain, Colombia, and
many other nationalities. Ecological studies take a great deal of time and, in a
manner of speaking, are still in their infancy. Studies of small mammal ecology in
the Cordillera Oriental in the Andes have important implications for
understanding the fauna of the north coastal ranges for Venezuela. Only a handful
of studies have been performed, but are well worth noting (Aagaard, 1982; Pafaur
and Pasqual, 1985; Lopez-Arevalo, Montenegro-Diaz, and Cadena, 1993). The
ecological studies within the six state areas of this paper are amply cited within the
text, but O'Connell (1979, 1983, 1989), Ibafiez (1981), August (1983, 1984), and
Vivas (1986) deserve special mention as community studies. The efforts of Brokx
(1972b, 1984) with deer and Ojasti (1973) with capybara focus on species of great
economic importance.
Considering the six state areas (plus the Distrito Federal) of north-central
Venezuela, several patterns of mammalian distributions are noteworthy. Endemic
species of the llanos (Monodelphis orinoci and Dasypus sabanicola) are rare,
perhaps attesting to its recent configuration. Western Apure contains many
Amazonian elements and the Falcon Peninsula with a long history of aridity
contains some endemics (Marmosa xerophila). The Maracaibo Basin shows
affinities with northern Colombia and Central America (Eisenberg and Redford,
1979). At the present time, the north coastal ranges of Venezuela contain species
that are also broadly distributed in the Andes Mountains of Colombia to the west
and beyond. These include, among the marsupials, Gracilinanus marica and
Marmosops fuscatus. Among the Chiroptera, Eptesicus fuscus, Lasiurus cinereus,
and Histiotus humboldti are typical of the higher elevations. Microryzomys
minutus, Ichthyomys pittieri, Oryzomys talamancae, 0. albigularis, and
Sphiggurus vestitus represent the montane rodents. As an "island" of tropical
evergreen forest the coastal ranges support Bradypus infuscatus, Ateles belzebuth,
Felis (Leopardus) tigrina, and Potosflavus.
Certain wide ranging genera of mammals found in the northern Neotropics
are conspicuously absent from north-central Venezuela. These include: Primates--
Aotus, Saimiri, and Saguinus; Carnivora--Nasua and Bassaricyon; Marsupials--
Philander and Metachirus. The vast expanses of the Ilanos and the Andes to the
west must serve as a filter barrier (Eisenberg and Redford, 1979).
Haffer (1974) first proposed that the north coastal ranges of Venezuela might
serve as a refugial area for birds; Morton (1979) reinforced his ideas. In short,


Table 3. Species of Chiroptera having restricted ranges in north-central Venezuela (Linares, 1998).

I. Confined to the arid coastal strip

Leptonycteris curasoae
Choeronycteris godmani

II. Confined to the coastal mountains
or foothills

Peropteryx kappleri
Diclidurus albus (N)
Mormoops megalophylla
Pteronotus gymnonotus (S)

Micronycteris hirsuta (N)
M. microtis (N)
M schmidtorum
Tonatia bidens
Lonchirhina aurita
Chrotopterus aurita
Anoura geoffroyi (N)
A. caudifer
Sturnira erythrymos

Platyrrhinus helleri
P. dorsalis (N)
P. vittata (N)
Vamipyrodes cariccioli
Vampyressa pusilla
Chiroderma tinitatiam (N)
Diphylla ecaudata

Thyroptera discifera (N)
Histiotis humboldti (N)
Lasiurus cinereus (N)
Eptesicus fuscus (N)
E. montosus

Nyctinomops aurispinosa (N)
N. laticaudata (N)
Tadarida brasiliensis
Molossus bondae
M. pretiosus
M. sinaloae

* N = found in the north coastal (front) ranges.
S = found in the southern (back) ranges.
No symbol = found in both forested ranges.

with respect to the avifauna, the north coastal ranges exhibit characteristics of an
island. Taxa typical of the Andes are represented and basal taxa of Amazonas are
also present. To some extent Eisenberg and Redford (1979) have discussed this
problem with respect to the mammals. Clearly the students of the avifauna were in
The term "llanos" is broadly applied to the woodland savannas and
grasslands of central Venezuela and adjacent Colombia. The area exhibits great
heterogeneity. Under appropriate conditions pennanent water courses support
multistratal tropical forest and have an associated but often depauperate forest
fauna. Morton (1979) has analyzed the problem for the avifauna. August (1981)
demonstrated the role of the gallery forests in supporting complex mammal
communities. Almost all the llanos fauna is derived from the adjacent forest fauna
although there are a few species endemic to the open grasslands. Mares and his


Figure 3. Biogeographic regions of Venezuela. Circles refer to collection stations as given in the
publication by Handley (1976). 1 = Maracaibo Basin; 2 = Falcon arid zone; 3 = North coast range; 4 =
Andean region; 5 = Llanos; 6 = Guyana Highlands; 7 = Amazon Lowlands. Boundaries of the regions
are approximate, future research may define zone of intergradation. The region of the delta is excluded
from consideration as is Isla Margarita and Trinidad. From Eisenberg and Redford (1979).


associates (Lacher, Mares, and Alho, 1989) have described an analagous
phenomenon for the seasonally dry forests of Caatinga and Cerrado of Brazil.
The seasonally dry llanos and dry forests surround the north coastal ranges
and to the west the dry forest band joins the arid Falcon Peninsula and the arid
areas at the north end of the Maracaibo Basin. The eastern range of the Andes
blocks simple emigration of the terrestrial fauna to north-central Venezuela from
the Maracaibo region. At the present time the largest block of lowland, moist
forest north of the Orinoco River and east of the Andes is in southwestern Apure.
Some species of mammals known to have occurred in northern Venezuela
have been reduced to the point of local extinctions. Speothos venaticus, Pteronura
brasiliensis, and Priodontes maximus may well be gone. Ateles belzebuth, Tayussa
pecari, Puma concolor, and Panthera onca are threatened.
The national parks of north-central Venezuela are illustrated in Figure 2. El
Avila, Henri Pittier (Rancho Grande), and San Esteban protect parts of the north
coastal ranges. Macarao, Guatopo, Yurubi and Tirgua protect the interior ranges.
La Laguna de Tacarigua protects a piece of coastal habitat in Estado Miranda.
Aguaro-Guaraquito is a large park (4,000 km2) covering primarily private land in
the Ilanos of Guarico. Of the 113,430 km2 study area, a total of 9,282 km2 (8.2%)
lies within National Park Boundaries. However, the area covered by parks that
effectively protects larger fauna may be less. If we subtract an area of 4,200 km2 in
which we suspect that parks currently may not effectively protect all fauna, and add
1,500 km2 of private ranches where wildlife is protected, we arrive at a rough
estimate that 6,582 km2, or 5.8% of the total surface area of North-Central
Venezuela that functions as wildlife refuges.
North-central Venezuela has much to be proud about in terms of
mammalian conservation. Large natural areas abut modern urban areas. Large
ranches protect and manage fauna responsibly. There is a rich literature.
However, at the time of this publication, there exists cause for concern. The
noteworthy parks near Venezuela's largest cities (El Avila/Caracas; Henry
Pittier/Maracay; San Esteban/Puerto Cabello & Valencia) are already partly
compromised by their proximity to human developments. These parks face a
future of increasing isolation due to human expansion unless a conscious and
specific effort is made to prevent that. There will always be some problems of
poaching of "game" species (Silva and Strahl, 1996) in the parks. A very active,
well-funded, and continuous combined program of education and enforcement can
keep that in check. Large-bodied species (jaguars, white-lipped peccaries, pumas)
are native to the lower elevations of the coast range, but are presumed eliminated
from many areas and scarce in others. They may yet completely disappear, unless
their habitats are aggressively defended. Large cattle ranches can and do play a
role in the preservation of wildlife (Hoogesteijn and Chapman, 1998). The public
service that enlightened ranch management provides should not go unrecognized,
nor, perhaps, in some cases, unrewarded. Effective conservation requires serious
commitment. The long-term maintenance of the fauna described here will require


nothing less than a coordinated commitment, at the national level, and at every
other level of Venezuelan society, even in the face of economic unpredictability
and hardship. Once lost, these mammalian communities will be difficult to


The six-state area plus the Federal District may be considered a mega-
landscape. The north coastal ranges (both front range and south range) have
served as an island of forested habitat surrounded by lower elevational species that
wax and wane in terms of forest cover. The extreme north range exhibits Andean
connections. Voss and Emmons (1996) urge us to continue collecting until
complete lists of organisms can be compiled. A worthy aim, but the area we have
defined is a habitat mosaic with a long history of non-human disturbance. If the
effects of Homo sapiens beginning about 10-15 thousand years ago are added in,
the situation becomes even more complex. European agronomy was introduced in
the 16th and 17th centuries, and human influences together with their agronomy
led to profound changes. The deforestation of the last 50 years (Hamilton et al,
1976) has been dramatic.
We have tried to characterize the mammalian fauna of north-central
Venezuela recognizing that the sampling has occurred on fragments that are
increasingly fragmented. What is the mammal list for Parque Nacional Guatopo?
It depends. Handley (1976) published data from a survey of the southern portion.
Eisenberg, O'Connell, and August (1979) published a list that derived from a
longer, more intensive effort, yet with little collecting of Chiroptera. Only by
analyzing multiple data sets do we approach a true evaluation of biological
diversity. Yet some species (e.g. Panthera onca, Puma concolor) were better
studied at alternative sites. We are left with comments in the introduction;
combined lists suggest at least 95 species of mammals are to be found in Guatopo.
The inventories for Parque Nacional Henri Pittier (Rancho Grande), El Avila, Hato
Masaguaral, Hato El Pifiero also remain open-ended.
With increasing habitat fragmentation in most parts of the globe, it is all too
likely that most parks will lose more species than they gain, thus it may be more
useful to characterize a bio-region than to restrict to an "insular" survey area of 15
km2, 30 km2, etc. in hopes that one straggler will be caught, vouchered for, and
catalogued. With this said, we hope the list for the six states serves as some sort of
benchmark for 1999.



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