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of the

FLORIDA
MUSEUM OF

NATURAL HISTORY

THE MAMMAL SPECIES OF
NORTH-CENTRAL VENEZUELA

John F. Eisenberg and John R. Polisar

Volume 42 No. 3, pp. 115-160 1999

E E' A-L***L 3 ir/Kil'* ~t / i &*#
UNIVERSITY OF FLORIDA GAINESVILLE



Numbers of the BULLETIN OF THE FLORIDA MUSEUM OF NATURAL HISTORY are published
at irregular- intervals. Volumes contain about 300 pages and are not necessarily completed in any one
calendar year.

JOHN F. EISENBERG, EDHOR
RICHARD FRANZ, Co-EDrroR

RHODA J. BRYANT, A*INWGINGEDROR

Communications concerning purchase or exchange of the publications and all manuscripts should be
addressed to: Managing Editor, Bulletin; Florida Museum of Natural History; University of Florida;
P. O. Box 117800, Gainesville FL 32611-7800; U.S.A

This journal is printed on recycled paper.

ISSN: 0071-6154 CODEN: BF 5BA5

Publication date: October 8, 1999 Price: $ 7.00



*r

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

background.



Plate 2. Giant anteater (Myrmecophaga tridactyla) along dry savanna/dry forest ecotone. Las Penitas, Hato Pifiero, Estado Cojedes.



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



Plate 4. Puma (Puma concolor) traveling along foot-transect. Cabo Caujaral, Hato Pihero.



C

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



THE MAMMALS OF NORTH-CENTRAL VENEZUELA

John F. Eisenbergl and John R. Polisa2

ABSTRACT

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 lianos A diverse array of habitats are
represented in this area: xeric scrub forests  montane rain forests, cloud forests  deciduous forests, and the
savannas ofthe lianos.

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 lianos 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. T#date the most successful
protection of 11anos flora and fauna seems to have been the result of private initiatives, on ranches such as
Hato Pificro and Hato Masaguaral. In the lianos, maintenance of large mammals such as white-lipped
peccaries (Tayassit pecar,1 jaguar (Panthera oncal 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

RESUMEN

Se han registrado 327 especies de mamiferos terrestres para Venezuela. En este trabajo analizamos la
na de los seis estados del nolte central y del distrito federal. Se incluyen por lo tanto, la costa del

Caribe, el ramal nortefio de la Cordillera de la Costa, los ramales surefios de la Cordillera de la Costa y los
Ilanos norteflos. Estan representadas una variedad de habitats: matorral des6rtico, bosques hOmedos
montanos, bosques de neblina, bosques deciduos y las sabanas de los Ilanos.

Esta regidn de Venezuela alberga 180 est,ecies de mamiferos que cuentan con especimenes
depositadas en algOn museo. La Cordillera de la Costa incluye sitios come el Parque Nacional de Guatop6
que alberga 95 especies, mientras que en los ilanos, en localidades que incluyen bosques de galeria se pueden
ilegar a registrar hasta 75 especies. Algunas especies de quir6pteros tienen ranges de distribucian
restringidos a un tipo de vegetaci6n especifico y/o a un rango de altitudes.

En esta regi6n muestras representativas de todos los hibitats importantes esun bajo alguna forma de
proteccidn. En la Cordillera de la Costa la proteccidn estA dada por parques nacionales. En los ilanos la
proteccitn mas efectiva de especies de flora y fauna ha sido debido a la iniciativa privada, por ejemplo en
ranchos como Hato Piaero y Hato Masaguaral. Es por eso que en los lianos pam la proteccidn de mamiferos

-'' . 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 Natu,al History, University of Florida, P. O. Box 117800.
Gainesvitle FL 32611-7800, U. S. A

Eisenber& J. F., and J. R. Polisar. 1999. The mammals ofnorth-central Venezuela. Bull. Florida Mus. NaL
Hist 42(3):115-160 + Plates 1-5.



116 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL- 42(3)

grandes como Btiquiro Cachete Blanco (Tayassu pecan), el jaguar (Pantera onca) y el puma (Puma
concolor), se debe considerar una red integrda por grandes ranchos y parques que contengan habitats
pristinos sin presi6n o una presi6n muy baja de can

Introduction

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 (Sirenig 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 dornesticates) 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 km2),
Parque Nacional Henri Pittier (1078 km2) and Parque Nacional San Esteban (440
kmi). Protected areas in the interior of the coast range include Parque Nacional
Guatopo (1224 kme), Parque Nacional Macarao (150 km2) and Parque Nacional
Tirgua (910 kmD. A partially isolated inland range, the Sierra Aroa, contains
Parque Nacional Yurubi (236.7 knf). Elevations gradually decrease as these
mountains undulate into another, lower, biographic region, los 11anos. The
southern drainage from the mountains forms streams and rivers that meander south
through los 11anos, all eventually draining into major rivers, such as the Apure, and
ultimately exiting into the Atlantic via the Orinoco.

The Ilanos are usually divided into 11anos altos (above 100 m), and further
south, los tlanos 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 11anos 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 11anos. In some cases, enlightened management by private owners



EISENBERG & POLISAR: VENEZUELAN MAMMALS 117

72 W 64W

CARIBBEAN SEA

N
vene 1299 11'N

COASTAL RANG68
O

D 0/4 , 6*· .«
Maracal 

bo

Parque Nacional Guatopo
040 Ports uesa 2 -9

80 Gu nare
+---Hato Masaguaral

R\00

Bjo Apure

Hato PiAero

 40 Apure 6 TN

50 0 100 200 300km

Figure 1. The moist forest zones of'north-central Venemela 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 lianos 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 consitute an area of over 1,500 km: 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 Ilanos sites include Hato Masaguaral (3000 ha) in Estado
Guarico and Hato Pifiero (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 11anos. 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 Ilanos (Ewel and Madrig 1968). Falling entirely



118 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 42(3)

within a Holdridge Tropical Dry Forest classification, los 11anos 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 dfy season from December
through March, and April and November are transitional months. Low elevations,
high temperatures, clay soils, and impeded drainage in los 11anos 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 Magaguaral 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 land represents the efforts of Handley (1976) and Eisenberg
et al. (1979) with some ammendations. Some marginal bat species are not
included in Table 1 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.



EISENBERG & POLISAR: VENEZUELAN MAMMALS 119

72' 68' 64 60

12* %9 /GRENADA - 12'
Distrito Federal

Yaracuy TRill~ BTOBAGO
rabobo 

<2~ganta

, i
Guanco  i 1

8.

r
i GUYANA

COLOMBIA 7< -
r 1

f l

1. Hato Masaguaral ~-LVENE~ht-..r- 
4. -

4 2. R N. Guatopo
3. Hato fero
4. R N. Henry Pittier International Boundary
5. R N. El Avila - ~ State Boundary ~
6. R N. Macarao 1-$

 .9.2
7. R N. San Esteban
8. RN.Yunlbi --- =-' A .ueyu· '9. R N. Tirgua

50 0 50 100 km10. P N. Laguna De Tacangua ~~ BRAZIL
11.R N. Aguaro-Guariquito <

I

77 68' 64' 60 '

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 lianos, 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 woodiand savannas in the Neotropics (Voss and Emmons,
1997).

Trapping effort  microhabitats sampled, and genuine differences between the
"front" range and the "back" ranges of the north coast all contribute to difTerences
in species numbers recorded. The figure of 180 species for the total area of six



120 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 42(3)

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

Tan 82 (23

Didelphimorphia (10)
Didelphidae

Caluromys philander L g g
Chironectes mimmus Lg g
Didelphis marsupialis Lg Gm
Marmosopsfiscata Lg g
Marmosa munna Lg g
M. robinsoni X. g im
Micoureus demararae (cinerea) X. g g
Gracitinanus manca x o Perez Hernandez, 1989
Monodelphis brevicaudata X. g g
M. onnod o o Perez-Hernandez. 1989

Insectivora
Soricidae

Coprons mendensis o o Linares, 1998
Xenarthra (Edentata) (7)

Mynnecophagidae
Myrmeccphaga tridactyla o m
Tamandua tetradactyla X. g rn, g

Bradypodidae
Bradypus variegatus X. g g

Dasypodidae
Cabassous unicinctus o g
Dasypus novemcinctus Lg & m
D. sabonicola o o Wetzel, 1985
Priodontes marimus 0 0 Wetzel, 1985

Primates (3)
Cebidae

Cebus olivaceous (nigrivittatus) Lg gm
Alouatta seniculus X. g im
Ateles belzebuth o g Mondolfi and Eisenberg, 1979

Chiroptera (88)
Emballonuridae (7)

Peropteryx kappleri Lg 0
P. macrotis Lg 0
P. trinitatus Lg 0
Rhynchonycteris naso Lg m
Saccopteryx bilineata Lg Bm
S. canescens Lg 0
S. leptura Lg Bm

Noctilionidae (2)
Noctilio leporinus 14 g
N. albiventris Lg

Mormoopidae (4)
Pteronotus davyi Lg
P. parnellii Lg

E E

P. personatus X, g



EISENBERG & POLISAR: VENEZUELAN MAMMALS 121

Table 1 Continued.

Tan Al B~ C~

P. suapurensis X. g
Mormoops megalophylla Lg

Phyllostomatidae (45)
Phyllostomatinae

Chrotopterus auritas X. g
Lonchorchina aurita Lg
Macrophyllum macrophyllum X. g
Micronycteris brachyotis
M. megatons Lg
M. nicefori X. g

o 
o 

&0 00 00 
E 

w
 E 

E 
E 

O
E

O
E

O
 

H
O

O
E

M. minuta Lg
Mimon crenulatum X, g
Phyllostomus discolor Lg Bm
P. elongatus X. g
P. hastatus X. g Bm
Tonatia bidens Lg
T. brasiliense X. g
Tonatia sylvicola o Linares, 1998
Trachops cirrhosis X. g
Vampyum spectrum X. g

Glossophaginae
Anoura caudifer Lg
A. culturata Lg
A. geofroyi X, g
Glassophaga longirostris Lg
G. soricina X. g Bm

Carolliinae
Carollia brevicauda Lg 0
C. perspicillata Lg Bm

Sturnirinae
Sturnira erythromos Lg g
S. lilium Lg & m
S. ludovici X. g Gm

Stenodermatinae
Chiroderma salvini x g
C. villosam X, g 0
Uroderma bilobatum Lg g
U. magnirostrum Lg & m
Vampyressa pusitia Lg g
Vampyrodes caraccioli x g
Platyrrhinus hellen Lg g
P. umbratus Lg 0
P. vittatus
Ametrida centurio Lg g
Artibeus cinereus Lg Bm
A. harti x
A. phaous o g
A. jamaicensis Lg Bm
A. lituratus Lg g
Cenmrio senex 0 0 Linares, 1986



122 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 42(3)

Table 1 Continued.

Taxa 82 (3

Sphaeronycteris toxophyllum Bm
Desmodontinae

Desmodus rotundus Lg m
Diphylla ecaudata. x

Vespertilionidae (14)
Eptesicus brasiliensis x
E. diminutus x
E. furinalis Lg Bm
E. fuscus x
Histiotus humbom x
Lasiurus blossevilli Bm

(L. borealis)
L cinereus x
L ega 14 g
Myotis albeacens Lg
M. keaysi X. g

O 
0

0
 

0
0
 

O
S

E
O

E
 

H
O
 

o 
E

E
S

S
E

E
0

5
0

0
©

5
0

0
0

M. nigricans Lg
Myous simus o
M. 00'oms X
Rhogeessa tumida Lg Bm

Natalidae (1)
Natalus tumidirostis x

Molossidae (15)
Eumops auripendulus
E. bonairiensis
E. dabbenei
E glaucinus
E. hansae
Molossops greenhalli

*
*
*
0
*
0
0
*
X
O
X

M.paranus
M. temmincki
Molossus ater
M. azteus
M. bondae
M. mollosus Lg
M. sinaloae x
Nyctinomops laticaudams x Linares, 1998
Tadarida brasiliensis o Unares, 199%

Carnivora (17)
Canidae

Urocyon cineroargenteus
Cerdocyon thous Bm
Speothos venaticus 0 0 Bisbal, 1989

Procyconidae
Procycon cancrivorus Lg Bm
Potos flagus Lg g

Mustelidae
Mustela#enata o o Bisbal, 1989
Eira barbara o Bm
Galictis vittata o m



EISENBERG & POLISAR: VENEZUELAN MAMMALS 123

Table l Continued.

Tan 82 (3

Conepatus semistriatus o Bm
Lontra longicaudif o o Bisbal, 1989
Preronura brasiliensis o o Bisbal, 1989

Felidae
Puma concolor o m Hato Pificro
Panthera onca o g Hato Pifiero
Herpailurus yagouaroundi Gm
Leopardus pardalis X. g Bm
Leopardus wied# 0 0 Bisbal, 1989
Leopardus tignna x o

Perissodactyla (1)
Tapiridae

Tapints terrestris o g
Artiodactyla (5)

Tayassuidae
Tayassu pecan o o Hato Piaero
Tayasm tajacu gm

Cervidae
Odocoileus virginiana Lg Gm
Mazama amencana x g
Mazama gouazoubira o o Bisbal 1991

Rodentia (27)
Sciuridae

Sciurus granatensis X, g Bm
Heteromyidae

Heteromys anomalus X, g Bm
Muridae

Sigmonotinae
Oryzomyini

Oecomys bicolor (seeted)
0. concolor (see text)
0. speciosus X. g
0. trinitatis X, g

E 
00 00 

.
 0 

E w
 

0

Oryzomys albigularis x
0. cap,to (see teL gt)
0. megocephalus (see text)
O. talamancae x
Microryzomys minutus x
Oligoryzomys fulve3cens X, g
Neacomys tenuipes X. g

nomasomyini
Rhipidomys venustus x
R. venezuelae 14 g & m

Phyllotini
Catomys hummelinki o o Linares, 1998

Ichthyomyini
Ichthromys pirrier, 0 0 Handley and Mondolfi, 1963

Akodontini
Akodon urichi Lg g

Zygodontornyini



124 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL„ 42(3)

Table 1 Continued.

Tan 82 (3

Zygodontomys brevicauda Lg
Sigmodontini

Sigmodon hispi<luz X, g
S. alstom x

E 
0
2
0
 

0 
M
 

no E

Holochilus sciureus x
Erethizontidae

Sphiggurus pru,nosus o CEBRG
Couendou prehensilis Lg Bm

Caviidae
Cavia aperea Lg 0

Hydrochacridae
Hydrochaeris hydrochaens o m

Agoutidae
Agouti paca Lg g

Dasyproctidae
Dasyprocta agouti (leporina) Xg Bm

Echimyidae
Echimys semiviliosus X Bm
Proechimys guairae Lg

Lagomorpha (2)
Lzporidae

Sylvilagus brasiliensis x
S. floridanus x

'x= occurs in one ormon of the six states of north-central Venezuela, g = collected at or near Guatopo; as documented in Handley (1976).
2 Found in g (Guatopo) and'or m (MasaguaraD as documented in Eisenbeig et al. (1979) or Sunquist, Sunquist, and Daneke (1989) (vouchers

of Chiroptm and Rodentia at the Museum, Texas Tech University, Lubbock).
3 Other vouchered localities either by site, museum reference. or Unares„ 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
Century.

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



EISENBERG & POLISAR: VENEZUELAN MAMMALS 125

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

ACKNOWLEDGEMENTS

The senior author would like to thank the following individuals for their support and kindness
through the years: Ambassador Edgardo Mondolfi, Sr. Tomas Blohm and his wife Cecilia, Charles O.
Handley, Jr., Sr. Rafael Garcia, and the late Juan Gomez-Nufter During the late 19905 Sr. Antonio Julio
Branger made the grounds of Hato Piaero available as a study site, and his stair was extremely helpful.
During our work at Hato Pifiero, Francisco Bisbal and his staff at Rancho Grande were supportive in many
ways. Rafael Hoogesteijn played critical roles in initiating the effort at Piaero. Francisco Bisbal was helpful
and supportive in many ways Melvin E. Sunquist was a source ofsupport 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
Maraguaral

Field research at Hato Pifiero was funded by The National Geographic Society, the Wildlife
Conseivation 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 Giraido, Sandra Melman, Emiliana Isase, Telva Carantona, and
Marcus Trepte. Important observations were contributed by Daniel Scognamillo, Ines Maxit Laura Famll,
Roy McBride, Rafael Hoogesteijn, Rocky McBride, Tibisay Escalona  Bruno Pampour, Rafael Ortiz, Dr.
Francisco Delascio, the Ilaneros ofPifiero, 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

ANNOTATED LIST OF TERRESTRIAL SPECIES

The annotated list does not include the ubiquitous Homo sapiens. The
numerous feral populations it has introduced are also excluded: Felis can,s, 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 Unia and Sotalia).

ORDER POLYPROTODONTA

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

FAMILY DIDELPHIDAE

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.



126 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 42(3)

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. Handley
(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 (0'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.



EISENBERG & POLISAR: VENEZUELAN MAMMALS 127

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
(1979).

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 at. (1977) separated specimens of Monodelphis from the tlanos 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 phitander
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 nortli-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.

ORDER XENARTHRA (EDENTATA)

FAMILY DASYPODIDAE

GENUS DASYPUS
Long-nosed Armadillos

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



128 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 42(3)

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

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/kmz. 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, 11anos north of the
Orinoco, and is primarily associated with los 11anos bajos. It was first described by
Mondolfi in 1968; the ecology was studied by Ferguson-Laguna (1984).

Cabassous unicinctus
Naked-tail Armadillo

The genus Cahassous 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 noghern
Venezuela. It is still to be found in Apure but is probably extinct in Guarico
(Mondolfi pers. comm.).

FAMILY MYRMECOPHAGIDAE

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 11anos. It was studied
by Montgomery and Lubin (1977) at Masaguaral. Home ranges were large,



EISENBERG & POLISAR: VENEZUELAN MAMMALS 129

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.

FAMILY BRADYPODIDAE

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 lianos, except where there is extensive evergreen
forest near permanent water. The species has been well studied in Panama (see
Montgomery and Sunquist [1974, 1975, 19781 and Sunquist and Montgomery
[19731). 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.

ORDER INSECTIVORA

FAMILY SORICIDAE

Cryptotis meridensis
Short-tailed Shrew

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

ORDER PRIMATES

The isolated nature of the north-central mountain ranges and associated
11anos 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).



130 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL- 42(3)

FAMILY CEBIDAE
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 lianos through forests
associated with major rivers. Within the 11anos, 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 Pifiero. Foraging, diet, and social structure were studied by
Robinson (1981, 19842, 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 Pifiero.

Alouatta seniculus
Red Howler Monkey

One of the most adaptable species of New World primates, it enjoys a wide
distribution. It has colonized the Ilanos 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 Sch6n-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.



EISENBERG & POLISAR: VENEZUELAN MAMMALS 131

ORDER CARNIVORA

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

FAMI~ CANTDAE
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/kn12 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.

FAMILY PROCYONIDAE

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.



132 BULLETIN FLORIDA MUSEUMNATURAL HISTORY VOL. 42(3)

Potos jlavus
Kinkajou

In north-central Venezuela this species is found in tropical humid forest and
premontane humid forest. It is absent over much of the 11anos.

FAMILY MUSTELIDAE

Mustela frenata
Long-tailed Weasel

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

Eira barbara
Tayra

In northern Venezuela it is strongly associated with tropical humid forest but
extends into the 11anos 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.

Gatictis vittata
Grison

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 km: 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 diy season. Density estimates for Masaguaral were 6-12/kmt

Lontra longicaudis
Otter

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).



EISENBERG & POLISAR: VENEZUELAN MAMMALS 133

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).

FAMILY FELIDAE

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

Leopardus pardalis
Ocelot

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 11anos
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 kmi 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
Jaguarundi

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



134 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL- 42(3)

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

Puma concolor
Puma

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), widle-tailed deer (Odocoileus
virginianus), and collared peccary (Tayassu tajacu) were among the top prey items
(Farrell, 1999).

Panthera onca
Jaguar

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 Ilanos 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). Pifiero is connected to other ranches containing
intact habitat and adequate natural prey. It is this network of connected forests that
maintain jaguars in los 11anos 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.

ORDER PERISSODACTYLA ,

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



EISENBERG & POLISAR: VENEZUELAN MAMMALS 135

associated with tropical evergreen forests in much of its range, in the tlanos 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.

ORDER ARTIODACTYLA
FAMILY TAYASSUIDAE

Tayassu pecari
White-tipped 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 paits 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 11anos 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 offirearms, 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/km: In the dry,
deciduous forests of the Ilanos at the Hato Masaguaral densities of 8.5/km2 were
estimated. Food habits were studied at Hato Pifiero by Bamto 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



136 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 42(3)

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 Pifiero more frequently encountered groups of 11-15 animals. T.
tajacu is a preferred prey of both pumas and jaguars.

FAMILY CERVIDAE

Mazama americana
Red Brocket

Present over much of lowland Venezuela, absent over most of the 11anos,
penetrating only the western 11anos 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 tlanos. 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 11anos. Brokx (19724 b, 1984) and Brokx and Anderssen (1970)
pioneered the study of O. virginianus in no~'them Venezuela. Stowe (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.



EISENBERG & POLISAR: VENEZUELAN MAMMALS 137

ORDER RODENTIA
SUBORDER SCIUROGNATHI
FAMILY SCIURIDAE
Squirrels

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.

FAMILY HETEROMYIDAE

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.

FAMILY MURIDAE
Rats and Mice

SUBFAMILY SIGMODONTINAE
New World Rats and Mice

Note: The names Oryzomys capito, 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.

TRIBE ORYZOMYINI
GENUS OECOMYS
Arboreal Rice Rats

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



138 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 42(3)

Oecomys bicolor

Handley (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

Handley (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 jlavicans

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
Masaguaral.

Oecomys trinitatus

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

GENUS ORYZOMYS

Oryzomys albigularis

Nominal forms of O. a/bigu/aris 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
Venezuela.



EISENBERG & POLISAR: VENEZUELAN MAMMALS 139

Oryzomys capito

Handley (1976) employed capito as the species name for the terrestrial rice
rats inhabiting the lower altitudes in northern Venezuela. At the time 0. capito
included many subspecies following Hershkovitz (1944). Current thinking has
resulted in the break-up of capito into some full species. Redefinitions of species
initiated by Musser and Williams (1984) established that O. talamancae be
recognized in northern Venezuela and was reinforced by Perez-Zapata et al.
(1986). Oozomys 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 O. capito and suggest O. megacephalus be
employed as the senior name for the remaining forms previously included in 0.
capito.

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 O. talamancae. O'Connell (1981, 1989) studied it at Guatopo.

GENUS MICRORYZOMYS
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.

GENUS OLIGORYZOMYS
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 dfy, deciduous forests.



140 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 42(3)

GENUS NEACOMYS
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.

TRIBE THOMASOMYINI

GENUS RHIPIDOMYS
Climbing Rats

Handley (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 atoreal 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.

TRIBE PHYLLOTINI

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.



EISENBERG & POLISAR: VENEZUELAN MAMMALS 141

TRIBE ICHTHYOMYINI

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.

TRIBE AKODONTINI

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
Guatopo.

TRIBE ZYOGDONTOMYINI

Zygodontomys brevicauda
Cane Rat

This common lowland, terrestrial rat is found in the grasslan(Is 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.

TRIBE SIGMODONTINI

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 tlanos.



142 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 42(3)

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. brasihensis.
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.

SUBORDER HISTRICOGNATHI

FAMILY ERETHIZONTIDAE
Porcupines

GENUS COENDOU
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 Mnwguaral.
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



EISENBERG & POLISAR: VENEZUELAN MAMMALS 143

were 2n=42 and FN=76. These numbers are identical with Erethizon but different
from Coendou rothschiMi. 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.

FAMILY CAVIIDAE
Cavies
-

Cavia aperea
Guinea Pig

Handley (1976) listed the forms from Venezuela as C porceUus. 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
Carabobo.

FAMILY HYDROCHAERIDAE
Capybara

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 11anos
(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.



144 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 42(3)

FAMILY DASYPROCTIDAE
Agouti

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.

FAMILY AGOUTIDAE

Agouti paca
Paca

This species can be common in tropical evergreen forests near permanent
water. It extends into the lianos 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.

FAMILY ECHIMYIDAE

Echimys semivillosus
Arboreal Spiny Rat

Makalata didelphoides (Echimys armatus) has been described from Trinidad
and the adjacent coast of Venezuela. Echimys semivil/osus is known from the
gallery forests of 11anos 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 (198lb) 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.).



EISENBERG & POLISAR: VENEZUELAN MAMMALS 145

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.

ORDER LAGOMORPHA
FAMILY LEPORIDAE
Hares and Rabbits

GENUS SYL#'7LAGUS
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.

Sylvilagusjioridanus
Cottontail

This species is common in lowland habitats, often quite dry such as thorn
forests, savannas, and range lands. It replaces S brasiliensis in the Ilanos. 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.

ORDER CHIROPTERA

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



146 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 42(3)

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

Emballonuridae (8) A. geofroyi
Rhynchonycteris naso A. latidens
Saccopteryx bilineata Choeroniscus godmani
S. leptura
S. canescens Carollinae (2)
Peropteryx macrotia Carollia brevicauda
P. kappleri C. perspicillata
P. trinitatus
Diclidurus albus

Sturnirinae (3)
Noctilionidae (2) Sturnira lucovici

Noctilio albiventris S. lilium
N. leporinus S. egthromos

Mormoopidae (5) Stenodermatinae (19)
Mormoops megalophylla Uroderma bilobatum
Pteronotus davyi U. magnirostrum
P. gymnonotus Platyrrhinus (Vampyrops) dorsalix
P. parnetti P. helleri
P. personatus P. umbratus

P. vittatits
Phyllostomidae (56) Vampyrodes caraccioli

Phyllostominae (21 ) Vampyressa pusilla
Micronycleris brachyons Chiroderma salvini
M. hirsuta C. trinitatum
M. megalotis C. villosum
M. microtis Artibeus harti
M. minuta A. cinereus
M. nicefori A. lituratus
M. schmidtorum A. jamaicensis
M. sylvestris Ametrida centurio
Tonatia bidens (T. saurophila) Sphaeronycteris toxophyllum
T. brasiliense g. venezuelae) Centurio senex
T. silvicola
Lonchorhina aurita Desmodontinae (3)
Macrophyllum macrophyllum Desmolus rotundus
Mimon cremilatum Diphylla ecaudata
Trachops cirrhosus Diaemus youngi
Phyllostomus discolor
P. elongatus Natalidae ( 1 )
P. hastatus Natalus stramineus
Phylloderma stenops
Chrotoprerus auntus Thyropteridae (1)
Vampyrum spectrum Thyroptera discifera

Glossophaginae (8) Vespertilionidae (15)

Glossophaga soricina Histiotus humboldti

G. longirostris Lasiurus cinereus
Leptonycteris curaswe L blossevilli (borealis)
Anoura caudifer L. ega
A. culturata Rhogoessa tumida



EISENBERG & POLISAR: VENEZUELAN MAMMALS 147

Table 2 Continued.

12 minutilla Eumops auripendulus
Myotia albescens E. bonariensis
M. nigricans E. dabbenei
M. oxyotus E. glaucinus
M. keaysi E. hansae
Eptesicus brasiliensia E. perous
E. diminutus Nyctinomops aurispinosa
E. jitrinalis N. laticaudata
E. fiscus Molosms ater
E. montoms (E andinus) M. bondae

M. coibensis
Molossidae (20) M. molosms

Molossops greenhalli M. pretioms
M. paranus M. sinaloae
M. planirostris Promops centralis
M. temmincki Tadari<la brasitiensis

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 ouiers. 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. Ibaflez (1981) offers a comprehensive monograph on the
ecology and biology of bat communities in the 11anos 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
species.

Table 2 offers a possible list of bat species that could occur in nonh-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.



148 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 42(3)

DISCUSSION

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 \lanos (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 histoiy 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. Microgzomys
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 Potos flavus.

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 A*tachirus. The vast expanses of the tlanos 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,



EISENBERG & POLISAR: VENEZUELAN MAMMALS 149

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

I . Confined to the arid crugtal strip Platyrrhinus helleri
P. dorsalis 04)

Glossophaginae P. vittata 04)
Leptonyctens curasoae Vampyrodes cariccioli
Choeronycteris godmani Vampyressa pusilla

Chiroderma tinitatiam (N)H. Confined to the coastal mountains Diphylla ecaudataor foothills
Thyropteridae

Emballonuidae Thyroptera disciferaM
Peropteryx kapplert Vespertilionidae
Diclidurus albus (N) Histiotis humboldti (N)
Mormoops megalophylla Lasiurus cinereus en
Pteronotus gymnonotus (5) Eptesicus h,scus 04)

E. montosus
Phyllostomidae

Micronycteris hirsuta (N) - Molossidae
M. microtis (N) Nyctinomops aurispinosa 04)
M. schmidtorum N. laticaudata (N)
Tonatia bidens Tadarida brasiliensis
Lonchirhina aurita Molossus bondae
Chrotopterus aurita M. pretiosus
Anoura geoffroyi 04) M. sinaloae
A. caudifer
Sturnira erythrymos

'N= found in the north coastal (front) ranges.
S = found m 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
advance.

The term "Hanoi' is broadly applied to the woodland savannas and
grasslands of central Venezuela and adjacent Colombia. The area exhibits great
heterogeneity. Under appropriate conditions permanent water courses support
multistratal tropical f6rest 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 tile Ilanos fauna is derived from the adjacent forest fauna
although there are a few species endemic to the open grasslan(Is. Mares and his



150 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 42(3)

2 e

80
4

5

6

7

Figure 3. Biogeographic regions of Venezuela. Circles refer to collection stations as given in the

publication by Handley (1976). 1 = Maracaibo Basin; 2 = Falcon add zone; 3 = North coast rangse; 4 =

Andean region; 5 = Llanos; 6 = Guyana Highlan(Is; 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).



EISENBERG & POLISAR: VENEZUELAN MAMMALS 151

associates (Lacher, Mares, and Alho, 1989) have described an analagous
phenomenon for the seasonally dry forests of Caatinga and Cerrado ofBrazil.

The seasonally dry 11anos 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 wellbe gone. Ateles belzebuth, Tayussa
pecari, Puma concolor, and Panthera onca are uireatened.

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 11anos 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 knf, 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 (iaguars, 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



152 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 42(3)

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
recover.

CONCLUDING REMARKS

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 nodh-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.



EISENBERG & POLISAR: VENEZUELAN MAMMALS 153

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