Jaguar research in Venezuela
79
ANARTIA
Publicación del Museo de Biología de la Universidad del Zulia
ISSN 1315-642X (impresa) / ISSN 2665-0347 (digital)
Anartia, 34 (junio 2022): 79-95
e rich tradition of jaguar research and conservation
in Venezuela and its impact on human-jaguar coexistence
throughout the species’ range
La rica tradición de investigación y conservación del jaguar en Venezuela
y su impacto en la coexistencia entre humanos y jaguares
en el área de distribución de la especie
John Polisar1, Almira Hoogesteijn2, Lucy Perera-Romero3, María Fernanda Puerto-Carrillo4,5,
Emiliana Isasi-Catalá6, Włodzimierz Jędrzejewski4 & Rafael Hoogesteijn7
1Department of Enironment and Development, Zamorano Biodiversity Center, Zamorano University,
Tegucigalpa P.O. Box 93, Honduras.
2Centro de Inestigación y de Estudios Avanzados, Unidad Mérida, México (CINVESTAV).
3Caura Program – Wildlife Conservation Society 2009 – 2015.
Current: Mammal Spatial Ecology and Conservation Laboratory, Washington State University, WA, USA.
4Laboratorio de ecología y genética de poblaciones, Centro de Ecología, Instituto Venezolano de Inestigaciones Cientícas (IVIC).
5Departamento de Biología, Facultad Experimental de Ciencias, Universidad del Zulia, Venezuela.
6Wildlife Conservation Society, Perú.
7Panthera, 8 West 40th Street, 18th Floor, New York, NY 10018, USA.
Correspondence: raoogesteijn@gmail.com
(Received: 10-06-2022 / Accepted: 26-07-2022 / Online: 30-09-2022)
gesteijn 1986). Much of the work of the three researchers
focused on human-jaguar conict, a fair amount of that in
the state of Cojedes.
By the early 1990s, Venezuela was a leader in the young
eld of jaguar research and hosted the conference “Felinos
de Venezuela” in 1991 (FUDECI 1992), with specialists
from Switzerland, the United States, Mexico, Brazil, and
Venezuela. R. Hoogesteijn, A. Hoogesteijn and E. Mon-
dol published the rst study on cattle depredation by
jaguars (Hoogesteijn et al. 1993); this publication was
followed by Hoogesteijn and Mondol’s very comprehen-
sive and lavishly illustrated book (Hoogesteijn & Mon-
dol 1992) summarizing all the knowledge of jaguars at
the time, including information on Venezuelan natural
history richness. As Venezuelan researchers developed
further knowledge, they established uid contact with
colleagues from Brazil and Belize. Venezuela was the rst
country to experiment with jaguar translocations to solve
HISTORY
Some of the earliest studies on jaguars and human-
jaguar coexistence took place in Venezuela, including by
Ambassador (to Kenya) Edgardo Mondol, a Cornell
graduate (Animal Production and Zoology), professor at
the Central University of Venezuela, Director of several
Ministries, with links to the Smithsonian Institution, the
Species Survival Commission of the IUCN, and the Unit-
ed Nations Environmental Program (UNEP). Mondol
is considered the “father” of modern Mammalogy in Ven-
ezuela (Fig. 1). His mastery of all matters mammalian led
to the mentoring of many researchers, amongst them two
veterinarians, Rafael Hoogesteijn and Ernesto Boede. e
three carved a place for Venezuela as jaguar studies started
in the 1980s. eir chapter in the 1986 edited volume Cats
of the World is testimony to that, one of the earliest serious
publications on jaguars in the world (Mondol & Hoo-
J. Polisar et al.
80
the cat-cattle conict, although the results were not par-
ticularly encouraging (Fig. 2).
In the mid-late 1990s, a team from the University of
Florida – Gainesville (UF), consisting of John Eisenberg
(globally renowned mammalogist), Melvin Sunquist (a
tiger and ocelot expert), Rafael Hoogesteijn (a cattle vet-
erinarian nishing his MSc; Fig. 3) and John Polisar (a
PhD candidate with wide-ranging interests and a back-
ground that started with carnivores in the northern
Rocky Mountains; Fig. 4), initiated a project in Hato
Piñero, Cojedes state, an 80,000-ha cattle ranch and na-
ture preserve. e setting was a ooded savanna-forest
mosaic, working cattle ranch, with 14,000 heads of cattle
and 450 heads of riding stock, along with white-lipped
peccaries, tapirs, deer, capybaras, anacondas, caimans,
jaguars, and pumas.
e project encountered numerous obstacles and over-
came them for success (Fig. 5). One of the main products,
led by Polisar (Polisar et al. 2003), included a complete
(and laborious) accounting of prey distribution and bio-
mass on the ranch and remains one of the most intensive
and thorough studies of the origins of and solutions for
human-cat conict- until today. A companion paper led
by UF MSc students Daniel Scognamillo and Inez Maxit
was also published in 2003 (Scognamillo et al. 2003). It
presented a detailed examination of ecological coexistence
Figure 1. Dr Edgardo Mondol, considered the “father” of
modern mammalogy in Venezuela, holding the largest jaguar
skull measured in all the Americas (shot in 1945 by Paul Stem-
pel at Hato La Rubiera in the Guárico State Llanos). Photo: Ra-
fael Hoogesteijn (~1990).
through resource partitioning of jaguar and puma and
has been widely cited by jaguar researchers. e environ-
mental and management setting in the seasonally ooding
lowlands of the Orinoco Basin in many ways resembles the
Brazilian Pantanal, and a substantial portion of the early
stages of jaguar research started there. Hato Piñero was the
rst place in Latin America where specially designed elec-
trical fences (Fig. 6) were tried in the eld as a successful
deterrent for jaguar/puma/cattle depredation problems
(Scognamillo et al. 2003). Two jaguars (one male and one
female), one female puma, and two female ocelots were
captured and tted with radio collars for the rst time
in Venezuela for telemetry studies during the rst stage
of this project (Hoogesteijn et al. 1996). By the time the
project concluded, researchers had captured and studied
ve jaguars (3 females and two males), and six pumas (four
female and two males).
Another signicant development in the nineties was
the export of two Venezuelan jaguars as wild founders for
a “Llanos Jaguar Breeding Group” to the Jacksonville Zoo
in Florida, USA. Negotiations between the Fundación
Nacional de Parques Zoológicos y Anes of Venezuela
(FUNPZA), the Jacksonville Zoo in Florida, USA, and
the Wildlife Oce of the Venezuelan Ministry of the En-
vironment (Profauna) began by inscribing the Venezuelan
Jaguars of wild origin in the studbooks held by the Ameri-
can Association of Zoos and Aquariums.
Venezuelan negotiators included Edgardo Mondol,
Pedro Trebbau, Esmeralda Mujica, Ernesto Boede and
Israel Cañizales (all associated to FUNPZA). e Envi-
ronment Ministry had two representatives that worked
for Profauna (the section in charge of wildlife protection);
biologists Mirna uero and Salvador Boher. Rafael Hoo-
gesteijn, acted as the initial liaison between Venezuela and
the Jacksonville Zoo Director.
To start a “Jaguar Breeding Group” of jaguars from the
Llanos region, a US veterinarian was hosted in Venezuela
by Boede and Mujica, to collect genetic samples from cap-
tive jaguars of wild origin, kept in dierent collections. Af-
ter almost two years of paperwork and sanitary trials, all
the needed authorizations and CITES permits were issued
for a very popular male (Bruno, kept at the Cattle Ranch-
ers Association of Carabobo State) and a female (from a
local zoo). Both animals were deposited in the new facili-
ties of the “Orinoco Floodplain” exhibition at the Jackson-
ville Zoo. is early model of international cooperation
pioneered processes and procedures potentially relevant
throughout the jaguar range.
Francisco Bisbal (1953-2020), was the chair of the
Museo de la Estación Biológica de Rancho Grande
(EBRG) for many years, an institution attached to the
Jaguar research in Venezuela
81
Figure 2. Team that completed one successful Jaguar translocation in Venezuela, le to right: Omar Hernández (Fudeci), Jose Luis
Méndez-Arocha (Profauna-MARNR), Edgardo Mondol (mammalogist and associate researcher), Ernesto O. Boede (wildlife special-
ist veterinarian). Entre Ríos region, Caura River, Bolívar State, Venezuela (February 1993). Photo: Ernesto Boede.
Figure 3. Rafael Hoogesteijn working a large herd of cattle in the Porto Jofre region of the Northern Brazilian Pantanal (2013).
Photo:Rafael Hoogesteijn.
J. Polisar et al.
82
Venezuelan Ministry of Environment. Francisco orga-
nized a number of collecting expeditions in many places
in Venezuela, being also an extraordinary museologist. In
the 1980s he published papers relevant to the Venezuelan
carnivores, including the jaguar.
During the 1990s, Almira Hoogesteijn (Venezuelan
Veterinarian; Fig. 7) began working in jaguar conservation
and research activities; she has since participated in many
important publications with her brother Rafael.
By the time large-scale jaguar conservation ignited,
with the January 1999 First Range Wide Priority Setting
Exercise conducted in Cocoyoc, Mexico; Venezuela was a
recognized force in jaguar conservation. Representations
at that conference associated with Venezuela included
Ernesto Boede, John Polisar, and Daniel Scognamillo.
Polisar, Hoogesteijn, and Boede had prepared the maps
for the workshop in late November 1998 in Valencia, Ven-
ezuela.
Hoogesteijn is currently a Ranch Manager and globally
renowned human-jaguar coexistence expert employed by
Panthera in the Pantanal of Brazil, advising other Panthera
teams and conict resolution projects, along with numer-
ous colleagues across a diverse range of Latin American
countries. Boede is now deceased, as is Mondol, and both
le a notable written legacy. Hoogesteijn is a prolic au-
thor and adviser on human-jaguar conict across the spe-
cies’ range.
e international educational institutions most tightly
involved with this genesis of applied research on jaguars
include the University of Florida and Cornell University.
From UF, this includes, Rafael Hoogesteijn, Laura Far-
rell, Inez Maxit (Fig. 8), John Polisar, Daniel Scognamillo
Figure 4. A recent photograph of John Polisar doing eldwork
in La Mosquitia, Honduras. Photo: John Polisar.
(Fig. 9), John Eisenberg, Melvin Sunquist, Jim Nich-
ols, and Wayne King. Peter Crawshaw was completing
his studies at UF as the Piñero project started. Edgardo
Mondol and Almira Hoogesteijn received their PhDs
from Cornell. John Polisar began his education in natu-
ral resources at Cornell. As the new millennium started,
Venezuela was a source of academically strong, ground-
breaking, and widely recognized contributions to jaguar
research and conservation.
VENEZUELA’S IMPORTANCE FOR JAGUAR
RESEARCH AND CONSERVATION 20032022
Venezuela is a physically beautiful country. It has sev-
eral biomes essential for jaguars and is of extreme strate-
gic importance for jaguar conservation on a range-wide
level (Fig. 10). Post-2003, jaguar research continued in
several distinct regions of Venezuela, including the Ama-
zon, the coastal range, the Llanos, and the Lake Mara-
caibo basin.
Emiliana Isasi-Catalá (who had worked as a volunteer
for Polisar in Hato Piñero) conducted her rst investiga-
tion on jaguars and pumas in Hato El Socorro (Cojedes
State), a large cattle ranch neighboring Hato Piñero. As
part of the objectives, this study evaluated habitat use and
diet of these big cats to determine possible conicts due to
predation on livestock and other domestic animals. As part
of her research interests, Emiliana has sought to improve
jaguar and puma population assessments using reliable es-
timation methods. erefore, she tested the identication
of individual jaguars and pumas based on morphometric
analysis of footprints, which later became part of her un-
dergraduate thesis. Additionally, she began the study of
these species using camera traps in protected areas such as
the Aguaro-Guariquito and Guatopo National Parks to
determine the eectiveness of this tool in population esti-
mates. Subsequently, she evaluated the presence of jaguars
in the Aguaro-Guariquito National Park, evaluating its
potential for jaguar conservation (Fig.11). She published
the results of that research, and now works on jaguars
and their prey in Peru. Her study area in Venezuela was
extremely rugged, which helped protect it since several
of Venezuela’s larger cities are nearby. Emiliana’s doctoral
research resulted in the rst formal abundance estimate
for jaguars in Venezuela. She went on to study, at a larger
landscape scale, the threats to connectivity and the tools to
maintain that connection, among which were traditional
cacao and coee agroforestry practices, and establishing
and maintaining good rapport and working relationships
with local communities, which can be eective stewards
of the land. reats include forest conversion for mono-
Jaguar research in Venezuela
83
Figure 6. First Latin-American eld tested electrical fences that successfully controlled jaguar/puma predation problems in a cattle
maternity paddock at Hato Piñero. Photo: Daniel Scognamillo.
Figure 5. e Piñero Project encountered numerous obstacles to overcome yet generated considerable pioneering contributions. Here
one of the principal researchers Laura Farrell, is inspecting one of the pig-baited jaguars-traps, with contracted professional trapper, Roy
McBride (March 1996). Photo: Rafael Hoogesteijn.
J. Polisar et al.
84
Figure 8. Inez Maxit and assistant Victor Juan Meires, with a tranquilized jaguar of the Piñero Project. Photo: Daniel Scognamillo.
Figure 7. Almira Hoogesteijn presents predation mitigation methods in a workshop for livestock owners in Autlán de Navarro, Jalisco,
México (2014). Photo: Almira Hoogesteijn.
Jaguar research in Venezuela
85
Figure 10. Prime jaguar habitat in Hato El Socorro, near El Baúl, Cojedes State, a mix of swamps, forested savannas, gallery forests, and
rocky hills that oer jaguars, excellent habitat and abundant prey-base. Photo: Rafael Hoogesteijn.
Figure 9. Local assistant together with Daniel Scognamillo in the process of radio-collaring a tranquilized jaguar at hato Piñero. Photo:
Daniel Scognamillo.
J. Polisar et al.
86
cultures. Emiliana formed a new project in Serranía El
Bachiller, an area contiguous with Guatopo National Park,
dedicated to community work to conserve this rugged and
beautiful region and the jaguars it harbors. Emiliana also
assisted Lucy Perera-Romero in her studies in the Upper
Caura.
Aer acquiring a remarkable experience of many
years researching on carnivore populations in the forests
of the Białowieza region on the Poland-Belarus border,
Włodzimierz Jędrzejewski came to Venezuela about a de-
cade ago to work for the Venezuelan Institute for Scientic
Research (IVIC– Instituto Venezolano de Investigaciones
Cientícas) based near Caracas. His extensive research
has yielded an important publication record on jaguars,
including a range-wide analysis, intensive camera trap-
ping studies principally in the Cojedes State but also al-
most in the entire country (with the help of Ernesto Boede
and several others (Fig. 12). More recently, Jędrzejewski
has published detailed research on jaguar populations
range-wide and has generated manuscripts on the status,
distribution, and connectivity of jaguar sub-populations
throughout South America. Jędrzejewski’s 2016 paper on
population density and structure in Hato Piñero (a decade
aer Polisar et al. 2003 and Scognamillo et al. 2003) re-
vealed high densities (7.67 jaguars/100km²). It represents
one of the few before-aer assessments with a larger than a
10-year interval of jaguar status on a working cattle ranch.
It provides strong evidence that coexistence is possible.
e population of jaguars in and around the large
ranches of Cojedes (Fig. 13) comprises their jaguar con-
servation unit (JCU), and there are also jaguars in the
Delta of the Orinoco River, as pointed out by Almira
Hoogesteijn et al. (2016). Venezuela and Colombia share
borders and biomes and have joint populations which do
not recognize political boundaries. Whether it is Llanos,
Amazonas, Caribbean slope, or lower slopes of the Andes
– these two countries share ecological characteristics and
management challenges. Venezuelan research relates to
Colombian realities. Colombia’s adjacency to Venezuela is
extremely important for range-wide jaguar connectivity in
the northern part of the continent.
To the west, where there are jaguars around the Mara-
caibo Lake region, Maria Fernanda Puerto-Carrillo, stud-
ies them and recently received an international “Future
for Nature” prize for her outstanding work; she founded
the Sebraba’s Project (Sebraba is the jaguar’s name in the
Bari indigenous language). Her research takes place in the
lowlands adjacent to the southern part of the Lake Ma-
racaibo basin, in the hottest and rainiest climate in the
country, where approximately 90% of the original for-
est has been lost. is region shares its fauna more with
northeastern Colombia than the remainder of Venezuela.
e region is essential for large-scale jaguar connectivity.
reats include habitat fragmentation and loss, reduced
natural prey, and direct killing, with consumption, sport,
and livestock losses being the drivers. Although these for-
Figure 11. Running/reviewing the camera-trap grid of Emiliana Isasi Catalá in Guatopo National Park, from le to right: Ernesto
Boede, Rafael Hoogesteijn, Emiliana Isasi-Catalá, and assistant José Infante (~2010). Photo: Ernesto Boede.
Jaguar research in Venezuela
87
ests are listed as Near reatened in the Red Book of Ter-
restrial Ecosystems of Venezuela, there are two protected
areas: Parque Nacional Ciénagas de Juan Manuel and
Reserva de Fauna Silestre Ciénagas de Juan Manuel de
Aguas Blancas y Aguas Negras, both totaling 3,400km² of
continuous forest, a good size even for the wide-ranging
jaguar (Fig. 14). Maria Fernanda studied diet, through
scats, and estimated population density, using a 600km²
camera trap polygon and spatially explicit models for an
estimate of 3.37 jaguars/100km². In 2021, she started two
new research projects on jaguars, one in the north of the
Amazon state with the Piaroa indigenous community and
another in areas near San Carlos, Cojedes State, where she
works with local producers, addressing the feline-human
conict.
Areas of great importance for jaguars include Venezu-
ela’s connections to the southwest, south, and southeast,
where the country has (in Apure, Amazonas, Delta Ama-
curo, and Bolivar states) physical and ecological connec-
tions with the Colombian, Brazilian, and Guyanese Ama-
zon-Guiana Shield greater Amazon region and comprises
one of the nine countries in the largest Jaguar Conserva-
tion Unit in the world.
e Wildlife Conservation Society supported Lucy
Perera-Romero’s research in the Upper Caura area (Gui-
ana Shield) from 2010 to 2015. Perera-Romero’s inves-
tigation was conducted 200 miles into the remote Gui-
ana Shield, upstream of river rapids and waterfalls, with
complete collaboration of the indigenous inhabitants
of ethnic Orinoco Basin groups (Fig. 15). ese studies
comprehensively focused on prey/game species, subsis-
tence hunting patterns, and jaguar abundance and distri-
bution (Fig. 16). Lucy’s studies included three separate
camera trapping campaigns across various cultural and
disturbance gradients (a relative term given the rela-
tively pristine characteristics of this area). e sampling
areas ranged from 235km² to 331km². Despite generat-
ing abundant data on how subsistence hunting aected
the natural prey base for jaguars/game species for hu-
mans, only one sample yielded a density estimate (~ 2.3
jaguars/100km²). e indigenous communities noted the
eect their hunting had on game and ecological equilib-
rium and designated “protection zones” related to each
community, where no hunting should occur. Despite the
documented low-scale threat that small-scale indigenous
subsistence hunting can represent, the indigenous are
vested in the environment that they inhabit. e threats
from deforestation and water contamination due to un-
controlled mining, the cultural erosion that the mining
camps bring, and, in particular, the high demands for
bush meat to serve hungry miners – represent a more se-
vere and, in some ways, existential threat for these spec-
tacular forests and their inhabitants, in an area that is
truly among one of the world’s natural wonders.
Lucy is currently a student at Washington State Uni-
versity, nishing up massive camera trapping research con-
ducted in the Maya Biosphere Reserve in Guatemala in the
Department of Petén.
Perera-Romero’s research in Venezuela’s Upper Caura
has been partially published, with some papers yet pend-
ing. Her abundant and deeply signicant data from Ven-
ezuela’s Upper Caura can be found in the 2022 Amazon
Cam Trap data paper. Her study and the studies by María
Figure 13. Gertrudis Gamarra (†), a great collaborator and
eld guide in Hato Piñero, Cojedes State. Photo: Włodzimierz
Jędrzejewski.
Figure 12. Ernesto Boede (le) and Włodzimierz Jędrzejewski
(right) camera-trapping at Hato El Frío in 2012, Apure State,
Llanos of Venezuela. Photo: Włodzimierz Jędrzejewski.
J. Polisar et al.
88
Figure 15. Lucy Perera-Romero camera-trapping in the upper Caura river region, a very remote and dicult access area, in which she
was widely helped by the local indigenous communities. Photo: Lucy Perera-Romero.
Figure 14. María Fernanda Puerto (IVIC), Raúl González (LUZ) and Orlando Gómez (INPARUES) checking data collected from
camera traps in Catatumbo River, Ciénagas de Juan Manuel National Park, Zulia State, 2018. Photo: Pedro L. Bermúdez.
Jaguar research in Venezuela
89
Fernanda Puerto-Carrillo, Emiliana Isasi-Catalá, and
Włodzimierz Jędrzejewski, have taken place thirty-plus
years aer the ground-breaking work of Edgardo Mon-
dol, Rafael Hoogesteijn, Ernesto Boede, and twenty
years aer the outcome of John Polisar, Daniel Scog-
namillo, and Inez Maxit – and are a solid testimony to
Venezuela’s rich history and continued active role in jag-
uar research.
Last but not least, it is worth mentioning that some
paleontological researches in Venezuela have rendered
information about the fossil record and paleodistribution
of the Jaguar in the region. e reports include Panthera
onca in the Late Pleistocene sites of Muaco, and possibly
in Taima-Taima, both in the Falcon state (Bocquentin
1979, Aguilera 2006, Chávez-Aponte & Carrillo-Briceño
2012, Carrillo-Briceño 2015), and the El Breal de Orocu-
al ORS20, in the Monagas state (Solórzano et al. 2015,
Ruiz-Ramoni 2016). Some remains assigned to P. onca
have been also reported from a cave named “La Cueva del
Cañón de Sorotamia”, in the Socuy River basin, Zulia state
(Rincón 2006); however, the geological age of the site is
not well dened, and this could be Pleistocene and Holo-
cene (see Ruiz-Ramoni 2016).
SUMMARY
Rafael Hoogesteijn, now stationed in Brazil, still pub-
lishes prolically. Polisar took what he learned in the Lla-
nos of Venezuela and applied it range-wide. Inez E. Maxit
is the Study Abroad Coordinator at Stephen F. Austin
State University in Texas, and Daniel G. Scognamillo
is an Associate Professor for Research in the Feline Re-
search Program in the Caesar Kleberg Wildlife Research
Institute-Texas A&M University-Kingsville. Isasi-Catalá,
Jędrzejewski, Perera-Romero, and Puerto-Carrillo have
skillfully carried the tradition forward with signicant new
contributions in Zulia State, in the Llanos, in the Guiana
Shield-Amazon, along with the coastal range, throughout
South America and range-wide. Almira Hoogesteijn con-
tinues working with ranchers in Mexico.
It is no exaggeration to say that work with Venezuelan
origins has played an essential role in jaguar research and
human-jaguar coexistence throughout the range of the
species. Because Venezuela has a rich history in jaguar re-
search and conservation and has played such an important
role in human-jaguar coexistence range-wide, it would oc-
cupy a welcome & strategic position as a member of the
Figure 16. Jaguar in upper Caura, Bolivar State, Venezuela. is area borders the Amazon and is part of the >2,000,000 km2 largest
Jaguar Conservation Unit worldwide. Photo: Lucy Perera-Romero.
J. Polisar et al.
90
2030 Jaguar Conservation Initiative. Venezuela is one of
the seventeen most biodiverse countries in the world. e
jaguar as an ‘umbrella species” can indicate that Venezuela,
apart from being the cradle of anti-depredation strategies
development in cattle ranching, is preserving its globally
signicant natural heritage and the cradle of anti-depreda-
tion strategies development in cattle ranching. e Jaguar
2030 Roadmap coincides with the Sustainable Develop-
ment Goals, the Convention for Biodiversity’s Global Bio-
diversity Framework, and the United Nations Decade of
Restoration.
Below (see appendix), we list publications focused on
Venezuela, of range-wide signicance, with the content of
Venezuelan origins, or Venezuelan co-authors, organized
alphabetically by author-year.
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rieur de Muaco, Etat de Falcón, Venezuela. Vol. 1-2. Palento-
logie des vertebres et paleontology humaine, L’ Université
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Ruiz-Ramoni, D. 2016. Paleobiología y ecología evolutiva de los
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Solórzano, A., A. D. Rincón & H. G. McDonald. 2015. A new
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tory Museum of Los Angeles County 42: 125–150.
APPENDIX
Scientic bibliography of the jaguar in Venezuela. e
following is a list of articles and books derived from the re-
search activities referred to in this paper. It does not intend
to be an exhaustive bibliographic list of the works related
to the presence of the Jaguar in Venezuela. It excludes a
number of popular articles, historical and travellers’ ac-
counts, ethnographic and etnozoological studies and most
references to Jaguar hunting activities in the country.
Antunes, A.C., A. Montanarin, D.M. Gräbin, E.C. dos Santos
Monteiro, F.F. de Pinho, G.C. Alvarenga, J. Auhumada, R.
Wallace, E.E. Ramalho, A.P.A. Barnett, A. Bager, A.M.C.
Lopes, A. Keuroghlian, A. Giroux, A.M. Herrera, A.P. de
Almeida Correa, A.Y. Meiga, A.T. de Almeida Jácomo, A. de
Barros Barban, A. Antunes, A.G. de Almeida Coelho, A.R.
Camilo, A.V. Nunes, A.C. dos Santos Marocio Gomes, A.C.
da Silva Zanzini, A.B. Castro, A.L.J. Desbiez, A. Figueire-
do, B. de oisy, B/ Gauzens, B.T. Oliveira, C.A. de Lima,
C.A. Peres, C.C. Durigan, C.R. Brocardo, C. A. de Rosa,
C. Zárate-Castañeda, C.M. Monteza-Moreno, C. Car-
nicer, C.T. Trinca, D.J. Polli, D. da Silva Ferraz, D.F. Lane,
D. Gomes de Rocha, D.C. Barcelos, D. Auz, D.C.P. Rosa,
D.A. Silva, D.V. Silvério, D.P. Eaton, E. Nakano-Oliveira,
E. Venticinque, E. Carhalvo Jr., E.N. Mendoca, E.M. Vieira,
E. Isasi-Catalá, E. Fischer, E.P. Castro, E.G. Oliveira, F.R.
de Melo, F. de Lima Muniz, F. Rohe, F. Beggiato Baccaro,
F. Michalski, F.P. Paim, F. Santos, F. Anaguano, F.B.L. Pal-
meira, F.da Silva Reis, F.H. Aguiar-Silva, G. de Avila Batista,
G. Zapata-Ríos, G. Forero-Medina, G. De Souza Ferreira
Neto, G.B.Alves, G. Ayala, G.H.P. Pedersoli, H.R. El Bizri,
H. Alves de Prado, H.B. Mozerle, H.C.M. Costa, I.J.Lima,
J. Palacios, J. de Resende Assis, J.P. Boubli, J.P. Metzgar, J.V.
Teixeira, J.M.D. Miranda, J. Polisar, J. Salvador, K. Borges-
Almeida, K. Didier, K. D. de Lima Pereira, K. Torralvo, K.
Gajapersad, L. Silveira, L. U. Maioli, L.Maracahipes-Santos,
L. Valenzuela, L. Benavalli, L. Fletcher, L.N. Paolucci, L.P.
Zanzini, L.C. da Silva, L.C. R. Rodrigues, M. Benchimol,
M.A. Oliveira, M. Lima, M.B. da Silva, M.A. dos Santos Jr.,
Jaguar research in Venezuela
91
M. Viscarra, M. Cohn-Ha, M.I. Abrahams, M.A. Benedet-
ti, M. Marmontal, M.R. Hirt, N.M. Tôrres, O.F. Cruz Jr., P.
Alvarez-Loayza, P. Jansen, P. Ribeiro Prist, P. Monteiro Bran-
do, P.B. Perônico, R. do Nascimiento Leite, R.M. Rabelo, R.
Sollman, R. Beltrão-Mendes, R.A.F. Ferreira, R. Coutinho,
R. da Costa Oliveira, R. Ilha, R.R. Hilário, R.A. Prudente
Pires, R. Sampaio, R. da Silva Moreira, R. Botero-Arias, R.V.
Martinez, R. A. de Albuquerque Nóbrega, R.F. Fadini, R.G.
Morato, R.L. Carneiro, R.P.S. Almeida, R.M. Ramos, R.
Schaub, R. Dornas, R. Cueva, S. Rolim, S. Laurindo, S. Es-
pinosa, T.N. Fernandes, T.M. Sanaiotti, T.H.G, Alvim, T.T.
Dornas, T.E.N. Piña, V’L.C. Andrade, W.T.V. Santiago, W.
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