Biological Conservation 109 (2003) 297–310

www.elsevier.com/locate/biocon

Jaguars, pumas, their prey base, and cattle ranching: ecological

interpretations of a management problem
John Polisara,*, Ines Maxitb,1, Daniel Scognamillob,2, Laura Farrellc,
Melvin E. Sunquistb, John F. Eisenberga
a
Florida Museum of Natural History, University of Florida, PO Box 117800, Gainesville, FL 32611-7800, USA
b
Department of Wildlife Ecology and Conservation, University of Florida, PO Box 110430, Gainesville, FL 32611-0430, USA
c
Museum of Comparative Zoology, Harvard University, 26 Oxford Street, Cambridge, MA 02138, USA

Received 27 August 2001; received in revised form 29 December 2001; accepted 25 April 2002

Abstract
Jaguar and puma depredation on livestock may be influenced by (1) innate and learned behavior; (2) health and status of indi-
vidual cats; (3) division of space and resources among jaguar and puma; (4) cattle husbandry practices; and (5) abundance and
distribution of natural prey. Our study in Los Llanos of Venezuela aimed to establish how all these elements related to cattle being
lost to cat depredation. Prey distribution was influenced by forest composition, topographical characteristics, and degree of habitat
interspersion. The biomass of natural prey in the study area was adequate to support the resident large cats without a subsidy of
livestock. Selective rather than opportunistic hunting by the cats reinforced that conclusion. Puma were responsible for more
attacks on livestock than jaguar, frequently in maternity pastures in upland areas of relatively low prey availability. Management
recommendations are discussed that may be relevant to other savanna/forest mosaics of South America.
# 2002 Elsevier Science Ltd. All rights reserved.
Keywords: Jaguar; Puma; Prey; Predator; Cattle ranching; Venezuela

1. Introduction unpredictable environments (Emlen, 1966; MacArthur

Jaguar (Panthera onca) and puma (Puma concolor)
depredation on livestock may be influenced by: (1)
innate and learned behavior; (2) health and status of
individual cats; (3) division of space and resources
among jaguar and puma; (4) cattle husbandry practices;
and (5) abundance and distribution of natural prey.
Predators select prey based on a cost–benefit analysis
(search time, handling costs, and energy gained in the
context of prey abundance) (Emlen, 1966; MacArthur
and Pianka, 1966) and vulnerability factors (Curio,
1976; Taylor, 1976; Temple, 1987). In productive envir-
onments, whether homogenous or heterogenous, pre-
dators can be expected to be more selective than in
* Corresponding author.
E-mail addresses: jjpolisar@aol.com (J. Polisar), sunquist@mail.
ifas.ufl.edu (M.E. Sunquist), lfarrell@oeb.harvard.edu (L. Farrell).
1
Present address: Louisiana Natural Heritage Program, Louisiana
Department of Wildlife and Fisheries, P.O. Box 98000, Baton Rogue,
LA 70898-9080, USA.
2
Present address: School of Renewable Natural Resources,
Louisiana State University AgCenter, Baton Rouge, LA 70803, USA.
0006-3207/02/$ - see front matter # 2002 Elsevier Science Ltd. All rights reserved.
PII: S0006-3207(02)00157-X
and Pianka, 1966). The value of a patch, in terms of
available prey, is usually reduced by predators, stimu-
lating them to search for alternative patches (Charnov,
1976). This predicts roaming among patches in all
instances except those where patch values are resilient.
These general postulates have to be able to absorb the
variation introduced by learned behaviors and indivi-
dual preferences. Among five intensively monitored
female mountain lions (Puma concolor) in Alberta, two
never killed bighorn sheep (Ovis canadensis), one killed
one sheep, one killed five, and one killed 17, in 1 year
killing 8.7% of an early-winter herd (26.1% of its
lambs) (Ross et al., 1997). All five cats were healthy, had
alternative prey available, and made varying use of
those alternatives. The learned ability to handle bighorn
sheep, normally more difficult to take than mule deer
(Odocoileus hemionus) reduced handling costs for one
puma. Risk of injury is a component of potential handling
costs (Sunquist and Sunquist, 1989). Preference for certain
natural or domestic prey may be transmitted from mother
to young (Hoogesteijn and Mondolfi, 1993; Mondolfi and
Hoogesteijn, 1986; Quigley and Crawshaw, 1992).
298 J. Polisar et al. / Biological Conservation 109 (2003) 297–310
Interactions among predators may influence choice of
prey. Seidensticker (1976) commented on the potential
effects of social dominance. In areas shared with tigers
(Panthera tigris) the behaviorally flexible social sub-
ordinate leopard (P. pardus) appeared to allow the
dominant tiger first choice of both habitats and prey
(Eisenberg and Lockhart, 1972; Seidensticker, 1976).
Spatial avoidance of a larger predator is likely to influ-
ence diet. Where there is a choice, leopards emphasize
smaller prey than tigers (Panthera tigris) (Karanth and
Sunquist 1995), but Karanth and Sunquist (2000) found
no evidence of spatial exclusion of leopards by tigers.
Overlap in carnivore diets may increase when a
resource is so abundant that there is little competition
(Colwell and Futumaya, 1971; Dailey et al., 1984; Spo-
wart and Thompson Hobbs, 1985). Environments fluc-
tuate; seasonally, annually, with patterns, even
erratically. It follows that levels of interspecific compe-
tition fluctuate. The present versions of jaguar and
puma coexistence are recent and perhaps still in flux
(Culver et al., 2000; Morgan and Seymour, 1997).
A high proportion of study animals have been killed
by ranchers and poachers in every jaguar study to date
(Sunquist, in press). Measures that reduce the frequency
of large cat depredation on livestock may go a long
ways towards maintaining cat populations. The con-
cerns of cattle ranchers are immediate and practical.
How can cattle losses to cats be reduced?
Roosevelt (1914) observed that ranches in Brazil that
possessed abundant native prey experienced fewer
jaguar problems. Shaw (1977) hypothesized that the
number of cattle taken by puma in Arizona was inversely
proportional to the size of the deer herd. Mondolfi and
Hoogesteijn (1986) and Hoogestein et al. (1993) hypothe-
sized a similar relationship for jaguar and puma in Vene-
zuela, where the cats exploit a more diverse prey base.
Cattle management offers some possibilities. In some
areas cattle have been so lightly managed that they
resemble wild prey (Hoogesteijn et al., 1993; Hooges-
teijn and Mondolfi, 1993; Mondolfi and Hoogesteijn,
1986; Quigley and Crawshaw, 1992; Schaller and
Crawshaw, 1980). Hoogesteijn et al. (1993) suggested
that losses could be reduced by: (1) excluding cattle
from forest; (2) maintaining adequate distance between
calving areas and forests; (3) moving calves out of
problem areas and replacing them with bulls; and 4)
maintaining adequate populations of wild prey.
In 1996, we initiated field work on a team project
designed to examine all the factors that could contribute
to cat-cattle conflicts: (1) ecology and behavior of jaguar
and puma; (2) abundance and distribution of natural
prey; and (3) cattle management practices. This paper
addresses the following questions: Could the natural prey
base in the study area support the cats or did they need a
subsidy from domestic livestock? What were the dominant
components of jaguar and puma diet? How did prey
selection relate to prey availability? What scenarios lead
to cat/cattle conflicts? What measures can be taken to
reduce those conflicts?
2. Methods
2.1. Study area
Hato Piñero is a working 80,000 ha cattle ranch/
wildlife preserve located between 8
400 and 9
00 N
and 68
00 and 68
18 W in the southeast corner of
the state of Cojedes in north-central Venezuela (Eisen-
berg and Polisar, 1999; Miller, 1992). The northern
boundary of Piñero lies among hills that rise to 396 m
above sea level (Farrell, 1999). The western boundary is
formed by the Cojedes and Portuguesa rivers, the south-
ern and eastern boundaries by the Chirgua and Pao rivers
(Fig. 1). Smaller streams (caños) run through this basin.
The lowest elevations are approximately 65 m above
sea level in the open savannas in the southern part of
the ranch. The landscape can be characterized as a
complex mosaic of interdigitated forests and open
areas with vegetation types based on interactions of
elevation, substrates, and hydrology. The ratio of
open to forested areas is roughly 50:50. Our 63,227
ha study area contained seasonally flooded lowland
savanna (39.06%), seasonally flooded semi-deciduous
forest (33.90%), dry hillside savannas with chaparral
(15.26%), dry hillside semi-deciduous forest (7.88%),
pastures in highlands that never flooded (2.86%),
evergreen forest (.08%) and mango groves (0.01%),
with the remainder developed for human habitation
and livestock maintenance.
The climate is strongly seasonal, with the majority of
the 1468.8 mm of precipitation falling during the wet
season between the beginning of May and the end of
November. The dry season, from December though
April is hotter. Relatively impermeable soils causes sur-
face water to accumulate starting in June and peaking in
July and August. The inundation is relatively shallow,
with greatest depths occurring in low savannas in the
southern portion of the ranch. Forests typically retain
pockets of dry land whereas the savannas in the south
flood completely for several months.
The majority of Piñero’s 14,000 head of cattle are Bos
indicus cebú races (nelore, brahma, guzerat, gir).
Approximately 420 horses, mules, and burros fulfill
working and breeding needs. A herd of approximately
150 water buffalo are maintained in the southern
savannas. Many cattle are moved from lowland pas-
tures to higher areas during the wet season. Cows can
forage in chest-deep water, but calf mortality is high in
flooded conditions. Nursing is hindered and the wet
muddy calves are vulnerable to biting insects. Artificial
insemination results in a pulse of calving from between
 J. Polisar et al. / Biological Conservation 109 (2003) 297–310
Fig. 1. Northwest and north-central Venezuela. Shaded areas show location of Hato Piñero study area in relation to locations where Smithsonian
research projects took place in the 1970s.
July and September. The maternity pastures where this
takes place are high, well-drained areas.
2.2. Prey abundance and distribution
Prey abundance, distribution, and population structure
were assessed using vehicle transects, linear foot transects,
point counts, night counts, camera trapping, capture–
mark–recapture methods, and opportunistic observations
(Buckland et al., 1993; Lancia et al., 1994; Seber, 1982).
2.3. Habitat characteristics
Patterns of forest vegetation were assessed using
35,000 m2 of quantitative sampling in a vertical profile
of Piñero (Polisar, 2000). Parameters describing phy-
siognomy were recorded at 100 m intervals along the 26
foot transects used for animal observations (Polisar,
2000). Forest types were classified using cluster analyses
(SPSS, 1999). Forest composition was tabulated for
different forest types, and number of species and percent
of individuals in trees, vines, and under story plants that
provided food for primary prey extracted for each type
(Polisar, 2000). A vegetation map used for areal esti-
mates of habitat types and felid habitat selection ana-
lyses using ArcView (1995) was based on Landsat
Thematic Mapper imagery, cluster analysis based forest
299
classifications, and ground-truthing with global posi-
tioning units.
2.4. Jaguar and puma prey consumption
Of five jaguars and six pumas captured, four jaguars
were radio-monitored for a total of 53 months and five
pumas were radio-monitored for a total of 68 months
(Farrell, 1999; Maxit, 2001; Scognamillo, 2001). Tele-
metry data, and tracks of jaguars and puma in areas not
used by collared animals, augmented by observations
and camera trapping images accrued during prey survey
efforts provided estimates of the study area’s cat popu-
lation. Jaguar and puma scats were collected opportu-
nistically on trails and roads, identified by the presence
of tracks or through mitochondial DNA analyses (Far-
rell et al. 2000), and prey species in scats were based
upon hair fibers, teeth, or scales (Farrell, 1999; Maxit,
2001; Scognamillo, 2001). Detailed notes were taken at
all cat kills, whether domestic livestock or free-ranging
natural prey (Farrell, 1999; Maxit, 2001; Scognamillo,
2001). Age/stage of natural prey kills was determined
using indices provided in Dimmick and Pelton (1994) and
Ojasti (1973) for white-tailed deer, collared peccary, and
capybara (Maxit, 2001; Scognamillo, 2001), and by cali-
brations of head-length to snout-vent-length and weight
obtained from spectacled caiman in Piñero (Polisar, 2000).
300 J. Polisar et al. / Biological Conservation 109 (2003) 297–310
The minimum annual requirement of killed prey neces-
sary to sustain Hato Piñero’s jaguar and puma population
was estimated by multiplying average weights of captured
animals by estimated numbers of cats (jaguars: 2.5 adult
males and 4 adult females, 2 subadults, and 3 cubs; pumas:
5 adult males, 7 adult females, 3 subadults, 5 cubs) using
consumption requirements of 34 g/day/kg of cat for
jaguars and 38.5 g/day/kg cat for pumas (extracted from
Emmons, 1987; Scognamillo, 2001). We used an estimate
of 70% consumption of killed prey based on estimates of
70–79% for pumas (Ackerman, et al. 1986; Hornocker,
1970) 75% for jaguars (Emmons, 1987) and 70% for tigers
(Sunquist, 1981). Average weights of captured adult male
and female jaguars respectively were 87.5 kg (n=2) and 52
kg (n=2). Average weights of adult male and female pumas
respectively were 51 kg (n=2) and 25.5 kg (n=4). Since
average jaguar weights (70 kg) were 1.88 times average
puma weights (37 kg) (Maxit, 2001), we presumed higher
metabolic rates and consumption needs per kg of body-
weight for puma (Emmons, 1987). Subadult consumption
requirements were considered equal to adult females of
respective species (Sunquist, 1981). Cub requirements were
estimated as 25% that amount (Sunquist, 1981), an estimate
that averaged size-classes from weaning to subadult.
2.5. Prey biomass estimates
Standing crop biomass of prey was derived from
abundance estimates and population structure data
based on group counts, night counts, transect-based
density estimates, and capture–mark–recapture sampling
(Polisar, 2000). Body weights were obtained in single or
repeated captures (Polisar, 2000) or in appropriate lit-
erature from studies completed nearby (Ayarzagüena,
1983; Brokx, 1972; Eisenberg et al., 1979; Linares, 1998;
Ojasti, 1973). Livestock biomass was estimated using fig-
ures provided by Ferdinando Corrales (Manager of Agro-
pecuaria San Francisco, father company of Hatos Piñero,
Paraima, Sembra, and slaughterhouse; Corrales, 1998).
Gross productivity was figured using our standing crop
biomass and population structure estimates, combined
with values for demographic parameters (average number
of litters per year/average number of young per litter/stage
specific survival rates) and growth rates that were either
local or realistic as possible (Bodmer et al., 1997; Brokx,
1972; Eisenberg et al., 1979; Hayne, 1984; Hellgren et al.,
1995; Kleiman et al., 1979; Ojasti, 1973; Ojeda and Keith,
1982; Smythe, 1978; Sowls, 1997; Teer, 1984).
3. Results
3.1. Prey biomass
Standing crop biomass of all major food species
(excluding livestock) was 374,489 kg, of which 149,988
(40%) was mammalian and 224,501 (60%) was reptilian
(Table 1). Annual minimum killing requirements for
resident jaguars and pumas were estimated at 10,100 kg
and 10,878 kg respectively, or 20,978 kg combined.
Minimum killing requirements for both cats combined
represented 5.6% of the standing crop: 2.7% for jaguar,
2.9% for puma. Including tapir, terrestrial tortoises,
and iguanas, potential food items not actually repre-
sented in the diet, standing crop was 470,825 kg
(Table 1).
The standing crop of cattle was around 4,656,000 kg
of which 160,000–384,000 kg were in the size class most
vulnerable to large cats (Table 2). Buffalo constituted
around 123,750 kg, and horse, mules, and burros pooled
118,300 kg (Table 2). Biomass estimates, for the entire
63,227 ha study area, are presented as kg/km2 in Table 3.
Percentages of the kg/km2 pooled along taxonomic
groups and the domestic (introduced)–wild (recent
native) dichotomy are presented in Table 4. Piñero bio-
mass estimates including and excluding domestic live-
stock are compared to other sites in the New and Old
World in Table 5. An estimate of annual gross pro-
ductivity of major native mammalian prey is presented
in Table 6. Cat killing needs represented a maximum of
29% of gross annual mammalian productivity (Table 6).
Average adult weights for deer, capybara, collared
peccary, and white-lipped peccary are 40, 46, 23, and 35
kg, respectively (Bodmer et al., 1997; Brokx, 1972;
Eisenberg et al., 1979; Ojasti, 1973; Sowls, 1997). Large
male deer may weigh slightly over 50 kg, as may large
capybara (Brokx, 1972; Ojasti, 1973). The largest cai-
man kill recorded was around 50 kg. The largest caiman
weighed was 75 kg. The largest anaconda weighed was
50 kg. In general, 50 kg was large prey in the region.
3.2. Jaguar and puma use of prey and habitats
Despite considerable overlap in prey species, jaguar
and puma diets were significantly different whether
examined with livestock included or excluded (Table 7)
(Maxit, 2001; Scognamillo, 2001). Jaguar showed a
preference for large-bodied prey, with puma taking
more medium-sized and smaller prey wild prey than
jaguar (Maxit, 2001; Scognamillo, 2001). Puma dietary
niche breadth exceeded that of jaguar (Maxit, 2001).
Jaguars preyed selectively upon capybaras and col-
lared peccaries, took white-tailed deer and caiman less
than expected, and killed white-lipped peccaries in pro-
portion to availability (Maxit, 2001; Scognamillo, 2001).
The most important wild prey in the jaguar diet in terms
of frequency of occurrence and biomass consumed were
collared peccary, capybara, and white-lipped peccary
(Table 7). Puma took most large prey in proportion to
availability, but selected for collared peccaries, and took
caiman less than expected according to availability
(Maxit, 2001; Scognamillo, 2001). The frequency of
 J. Polisar et al. / Biological Conservation 109 (2003) 297–310 301

Table 1
Natural prey abundance, distribution, and biomass

Species Habitat Habitat specific Prey abundance in 63,227 ha study area Total prey biomass in
density (km2) of prey study area (kg/63,227 ha)

OV All forest BSD & BS & BSV 1.0 234.73

High dry pasture PS 1.5 26.84
Low flooding pasture SI 8.3 1,973.28
All habitats pooled 2,234.85 78,890
TT All forest BSD & BS & BSV 7.5 1,760.5 35,835
TP All occupied habitats (primarily BSD) 167 5005
DA All forest BSD & BS & BSV 1.2 282 1072
SF Semi-deciduous forest BSD 0.8 151.21
Dry forest BS 1.7 77.32
Forests pooled 228.52 183
DN Dry forest BS 4.0 181.14
Semi-deciduous forest BSD 4.0 756.04
Forests pooled 937.19 3561
MT All forest 0.4 103.25 2788
HH All occupied habitats 547 22,654
GC All forest BSD & BS & BSV 95 22,300 96,336
CC All occupied aquatic habitats 15,408 167,827
FWT All occupied aquatic habitats 45,448.33 56,674

Species codes as follows: OV Odocoileus virginanus; TT Tayassu tajacu; TP Tayassu pecari; DA Dasyprocta agouti; SF Sylvilagus floridanus; DN
Dasypus novemcinctus; MT Myrmecophaga tridactyla; HH Hydrochaeris hydrochaeris; GC Geochelone carbonaria; CC Caiman crocodilus; FWT
freshwater turtles (Podocnemis voglii, Chelus fimbriatis, P. unifilis). Habitat codes as follows: BSD semi-deciduous forest; BS dry forest; BSV ever-
green forest; SI seasonally flooding savanna; and PS high dry pastures. Abundance based upon DISTANCE density estimates except where noted
(Polisar, 2000). TP and HH abundance estimates based upon counts in concentration areas, precluding density estimates. CC and FWT abundance
estimates based upon night counts adjusted for sighting fraction and mark-recaptures in concentrations that result from dry season low water levels.
Water surface area is in constant flux, thus no density estimates are presented (Polisar, 2000).

Table 2
Patterns of biomass among livestock at Hato Piñero

Sex/age class Crude Crude Biomass/class Biomass/class Biomass/class

numbers weight (kg) (Jul–Sep) (Oct–Dec) (Jan–Mar)

Bulls 200 750 150,000 150,000 150,000

Cows 7000 420 2,940,000 2,940,000 2,940,000
Newborns (0–3 months) 3200 50 160,000
Young calves (3–6 months) 3200 120 384,000
Calves
6–9 months 3200 180 576,000
10–24 months 3600 275 990,000 990,000 990,000
Total biomass (max and min) Minimum 4,240,000 4,464,000 Maximum 4,656,000
Biomass of class (most vulnerable to big cats) Minimum 160,000 Maximum 384,000

Sex and age specific numbers, weights, and biomass of cattle (Bos indicus and Bos taurus). Estimated cattle total is 14,000. The peak of parturition
is during July, August, and September. Stages most vulnerable to attacks by large cats are shown in italics. In addition to cattle there are: buffalo
(Bubalus bubalis) 123,750 kg (150825 kg); horses and mules 106,750 kg (305350 kg); young horses and mules 11,100 kg (111100 kg); and burros
450 kg (3150 kg). Total equidae biomass estimate is 118,300 kg, of which 11,100 would be colts.

capybara in puma scats was equal to that of deer,
although the large rodents were only one quarter as
abundant. The four most important wild prey items in
puma diet on bases of both frequency of occurrence and
biomass were collared peccary juveniles, deer, capybara,
and caiman (Table 7). While both jaguar and puma
showed a preference for collared peccaries, pumas were
mostly taking juveniles (Maxit, 2001; Scognamillo, 2001).
Capybara represented 71% of the sample of wild prey
killed by puma and 33% of the jaguar wild prey kill
sample. Spectacled caiman were 39% of the sample of
wild prey killed by jaguar (Table 7) (Maxit, 2001; Scog-
namillo, 2001). In terms of numbers of large prey avail-
able at any given time, the cumulative hunting pressure
upon capybara by big cats was high, and the numbers of
large reptiles consumed was less than expected.
Analyses of Hato Piñero’s books indicated that 13%
of total livestock losses in 10 years could be attributed to
large cats. These losses were highest during the peak cal-
ving months of August–October when on average 20
calves annually were killed by cats. Puma were respon-
sible for 86% of that damage (Maxit, 2001; Scognamillo,
302 J. Polisar et al. / Biological Conservation 109 (2003) 297–310

Table 3 2001). The results of our study support those trends

Conversions of biomass estimates for 63,227 ha study area into kg/ documented in the ranch ledgers (Table 7). Sixty-nine
km2 estimates
percent of attacked calves were between 1 and 30 days
Item kg/632.3 km2 kg/km2 of age (Scognamillo et al., in press). All five adult puma
home ranges included some of the high dry pastures
Native mammals
Capybara 22,654 35.83
used for calving during the wet season (1.7–23.9% of
Agouti 1072 1.69 the home ranges). For three pumas, use of the vegeta-
White-lipped peccary 5005 7.91 tion type exceeded within-home-range-availability by
Collared peccary 35,835 56.67 factors of 1.39–3.68 (Maxit, 2001; Scognamillo, 2001).
White-tailed deer 78,890 124.77 In contrast, that vegetation type was included in two of
Cottontail rabbit 183 0.29
Nine-banded armadillo 3561 5.63
four adult jaguar home ranges, and even then scarcely
Giant anteater 2788 4.41 so (0.0005–5.7% of those home ranges).
Subtotal native 149,988 237.21 There was no evidence of significant temporal segre-
mammals recorded in gation between the two cats (within 3-h intervals), and
jaguar and puma diet inter-specific spatial overlap was high on a coarse scale
Tapir 3000 4.74
(Scognamillo, 2001). On a fine scale, puma avoided the
Total native 152,988 241.95 immediate presence of jaguar (Scognamillo, 2001). Both
big cats intensively used the forest–savanna ecotone
Domestic ‘‘introduced’’ mammals (Scognamillo, 2001). Pumas were located significantly
Cattle 4,656,000 7363.59 more often within the first 500 m of large (> 300 ha)
Buffalo 123,750 195.71
Horses, mules, burros 118,300 187.09
forest patches than jaguars, which were located sig-
Domestic subtotal 4,898,050 7746.40 nificantly more often within the interior of such patches
(Maxit, 2001).
Total mammalian 5,051,038 7988.36

Reptiles
Iguana 669 1.06
4. Discussion
Caiman 167,827 264.95
Freshwater turtles 56,674 89.63 In Nepal, the Serengeti, and the Amazon, large pre-
Terrestrial tortoises 96,336 152.36 dators kill approximately 8–10% of the standing crop
Total reptilian 321,506 508.47 biomass of mammalian prey (Emmons, 1987; Schaller,
1972; Sunquist, 1981). In Hato Piñero that proportion
would require a minimum of 209,780 to 262,225 kg.
Table 4 With wild mammals and reptiles combined there was a
Percentage of total crude mammalian biomass expressed in kg/km2 minimum of 374,489 kg (Table 1), enough to support
represented by select groups 1.43–1.78 as many cats as were present. If potential
Group Pooled group Percent of total
dietary components apparently being bypassed were
mammalian biomass factored in, such as red-footed tortoises (Table 1) the
margin by which the estimated minimum is exceeded
Bovidae (introduced) becomes wider. In terms of standing crop prey biomass,
&
Cervidae (native)
the cats were killing approximately 4.46% of all poten-
& tial wild prey, 5.61% of the species actually being used,
Tayassuidae (native) Artiodactyla 96.99% and 13.99% of mammals alone.
This suggests that, in context of the entire study area,
Equidae (introduced) a subsidy from domestic livestock was not necessary to
&
Tapiridae (native) Perissodactyla 2.40%
sustain Hato Piñero’s large cats. Biomass ratios indicate
that wild mammals could provide approximately 57–
Agoutidae (native) 71% of the annual requirements, predicting some use of
& reptiles, which did occur (Table 7). It is fitting to assess
Hydrochaeridae (native) Rodentia 0.47% the proportion that mammals contribute before the
Myrmecophagidae Xenarthra 0.13%
reptiles. Many caiman and turtles were inaccessible to
cats, safe in the depths of their aquatic environments.
Bovidae (introduced) Artiodactyla The estimated annual needs of the cats constitute 29%
& of gross productivity of major native mammalian prey
Equidae (introduced) Perissodactyla 96.97% (Table 6). When considering sustainable consumption by
Native mammalian prey 3.03%
humans, Robinson and Redford (1991) and Robinson
and Bodmer (1999) suggest a maximum harvest of 20%
 J. Polisar et al. / Biological Conservation 109 (2003) 297–310 303

Table 5
Mammalian biomass in New and Old World study areas

Sites Comments kg/km2

New World
Hato Piñero, llanos, Venezuela Not including small mammals Including livestock 7988
Hato Piñero, llanos, Venezuela Not including small mammals Excluding livestock 242
Hato Masaguaral, llanos, Venezuela All nonvolant mammals Including livestock 8315
Barro Colorado Island, Panama All nonvolant mammals 2115
Guatopo, Coastal Range, Venezuela All nonvolant mammals 1001
Urucu, Brazilian Terra Firma Amazon All nonvolant mammals 891
Acurizal, Pantanal, Brazil Most nonvolant mammals 380

Old World
Wilpattu, Sri Lanka Ungulates only 766
Kanha, India Primarily ungulates 1708
Nagarahole, India Wild & domestic ungulates, primates 15,094
Serengeti Unit, Tanzania Primarily ungulates 4222
Manyara, Tanzania Primarily ungulates 7785
Ngorongoro Crater, Tanzania Primarily ungulates 10,363

Sources are as follows: Hato Masaguaral, Barro Colorado, Guatopo (Eisenberg, 1980); Urucu (Peres, 1999); Pantanal (Schaller, 1983) Wilpattu
(Eisenberg and Lockhart, 1972; McKay and Eisenberg, 1974; Eisenberg and Seidensticker, 1976); Kanha (Schaller, 1967; adapted by Eisenberg and
Seidensticker, 1976); Nagarahole (Karanth and Sunquist, 1992) Serengeti, Manyara, and Ngorogoro Crater (Schaller, 1972).

Table 6
Gross productivity (of major mammalian prey) at Hato Piñero

Species Abundance Standing crop (kg) Annual addition of biomass Annual gross
per 100 animals (kg) Productivity (kg)

Capybara 547 20,315–22,654 2295.00 12,554

White-lipped peccary 167 5005 1615.60 2698
White-tailed deer 2235 78,890 1292.50 28,887
Collared peccary 1760 35,835 1660.00 29,224

Total 73,363

Presented as increments of biomass added per year. Subtractions (adult mortality and otherwise) not included. Estimated annual kill needed to
support resident cats (20,978 kg) is 29% of gross productivity estimate above.

of production for long-lived species, and 40% for short-
lived species. Last reproduction occurred at over 10
years of age in long-lived species and between 5 and 10
years of age in short-lived species. White-tailed deer are
short to medium lived species, with emphasis on short.
Few deer live over 10 years and life expectancy in the
wild is frequently less than 3 years (Brokx, 1972; Win-
ston, 1991). There were no capybara over 5 years of age
in a harvested population in Apure, Venezuela (Lord
and Lord, 1988), although Robinson and Redford
(1986) estimated age at last reproduction as 9 years
using Ojasti (1973). Kleiman et al. (1979) commented on
the high reproductive potential of capybara. Using last
age of reproduction estimates of 13 years for collared
and white-lipped peccaries, Robinson and Redford
(1991) classify them as long-lived species. Hellgren et al.
(1995) found fecundity rates in collared peccaries over 7
years of age were less than animals between 3 and 7
years of age, roughly equal to animals 2–3 years of age,
and greater than animals between 1 and 2 years of age.
Some wild females in Texas and Arizona did exceed 10
years in age (Hellgren et al., 1995; Sowls, 1997). Though
classified as a long-lived species, Robinson and Redford
(1991) commented on the high productivity of peccaries
and Robinson and Bodmer (1999) consider a harvest
< 40% of production sustainable for both species. The
addition of caiman and freshwater turtle production
would elevate the total prey production estimate for
Hato Piñero considerably. Production estimates support
the assertion that, when the entire study area is con-
sidered, resident cats did not require a livestock subsidy.
An efficient predator will accept all potential prey
encountered when food is scarce or unpredictable, and
exercise greater selectivity when food is common and
adequate productive patches known (Emlen, 1966;
MacArthur and Pianka, 1966; Sunquist and Sunquist,
1989). Thus, diet breadth, in the context of diversity of
potential prey, can reflect relative scarcity or abundance
of prey. Analyses of jaguar scats from the Peruvian
Amazon yielded 40 prey taxa (n=25) (Emmons, 1987).
Rabinowitz and Nottingham (1986) recovered 17 taxa
from 228 scats in Belize. Analyses from the Chaco of
304 J. Polisar et al. / Biological Conservation 109 (2003) 297–310

Table 7
Numbers of items and proportions of different prey species in jaguar and puma scats and kills at Hato Pinero (data from Scognamillo, 2001)

Prey size Prey species Jaguar Puma

Scats n (%) Kills n (%) Scats n (%) Kills n (%)

Large size prey ( >15 kg) Tayassu tajacu 11 (26) 5 (16) 2 (5) 1 (2)
Tayassu pecari 5 (12) 0 (0) 1 (2) 0 (0)
Hydrochaeris hydrochaeris 9 (21) 6 (20) 4 (10) 15 (30)
Odocoileus virginianus (adult) 2 (5) 0 (0) 4 (10) 4 (8)
Myrmecophaga tridactyla 4 (10) 0 (0) 0 (0) 0 (0)
Caiman crocodilus 0 (0) 4 (13) 0 (0) 0 (0)

Livestock 3 (7) 10 (33) 10 (24) 29 (58)

Subtotals 34 (81) 25 (83) 21 (50) 49 (98)

Medium size prey (1–15 kg) Tayassu tajacu (juvenile) 1 (2) 0 (0) 5 (12) 0 (0)
Odocoileus virginianus (juvenile) 0 (0) 0 (0) 0 (0) 1 (2)
Procyon cancrivorus 2 (5) 0 (0) 0 (0) 0 (0)
Sylvilagus floridanus 1 (2) 0 (0) 3 (7) 0 (0)
Dasyprocta agouti 0 (0) 2 (7) 1 (2) 0 (0)
Dasypus novemcinctus 0 (0) 0 (0) 1 (2) 0 (0)
Caiman crocodilus 3 (7) 1 (3) 4 (10) 0 (0)
Podocnemis voglii 0 (0) 2 (7) 0 (0) 0 (0)

Subtotals 7 (17) 5 (17) 14 (33) 1 (2)

Small prey ( <1 kg) Small rodents or marsupials 0 (0) 0 (0) 7 (17) 0 (0)
Unidentified birds 1 (2) 0 (0) 0 (0) 0 (0)

Subtotals 1 (2) 0 (0) 7 (17) 0 (0)

Paraguay yielded 23 taxa (n=106) (Taber et al., 1997).
Fifty jaguar scats from the dry forests of Jalisco, Mex-
ico, yielded seven prey species (Nuñez et al. 2000).
Based on 42 scats, jaguar diets at Piñero include
approximately 10 taxa (Table 7).
Comparing diet breadth among studies has several
confounding factors. Innate prey diversity among the
study areas varies. Sample sizes were not equal among
studies. Nuñez et al. (2000) estimated 35–50 scats as the
minimum to adequately document diet. Anderson
(1983) suggested a sample of 90–100 scats was necessary
to calculate food habits of pumas within 10% of actual
use patterns. Emmons (1987) obtained diverse taxa with
only 25 jaguar scats. The number at which the asymp-
tote is obtained is likely to vary among sites. In Piñero
rank importance of prey items varied little between
subsamples of 30 scats and the entire samples of 42 scats
for both cats (Maxit, 2001).
The diet of jaguars in Piñero appears to be more spe-
cialized than the diet in Peruvian and Belizean rain-
forest sites (Emmons, 1987; Rabinowitz and
Nottingham, 1986). Foraging theory predicts that items
will be added to the diet only when the energy gained
outweighs the costs invested (Emlen, 1966; MacArthur
and Pianka, 1966). In Belize, 54% of jaguar scats con-
tained armadillos (Rabinowitz and Nottingham, 1986).
In Piñero, armadillos outnumber capybara (Table 1).
The search time required to obtain armadillos (relatively
dispersed in forest habitats) probably equals or exceeds
the search time for capybara (relatively concentrated in
somewhat predictable habitats, rarely > 500 m from
water; Ojasti 1973). Handling costs to capture armadillo
may equal handling costs for capybara. Capybara
weights are ten times as much as armadillos, which in
Piñero, are practically ignored (Table 7). In the Per-
uvian Amazon, terrestrial tortoises tied with collared
peccary as the numerically most frequent items in the
diet (Emmons, 1987, 1989). In Piñero, where terrestrial
tortoises were an order of magnitude more abundant
than the larger mammalian prey (Table 1), they were
virtually ignored. The large cats in Piñero have ade-
quate natural prey to make choices, another indication
that the natural prey base is adequate.
The prey base is adequate when the entire Piñero
study area is considered, but it is far from uniformly
distributed (Fig. 2). Piñero has lower prey diversity than
rain forests, but its high horizontal heterogeneity results
in patches where prey production is high. Cats move
through and between those patches. Some semi-decid-
uous forests have seasonal concentrations of white-lip-
ped peccaries. Others have resident groups of collared
peccaries. Prey abundance is high in lowland forest-
savanna mixes and in well-watered small savannas sur-
rounded by forest. The latter contain capybara, caiman,
turtles, and deer, and collared peccaries often use the
adjacent forest edge.
 J. Polisar et al. / Biological Conservation 109 (2003) 297–310 305

Fig. 2. Comparative encounter rates (observations/km) among 26 linear transects. Sign (which does not directly measure group size) and visual
observations independent of group size are pooled. All taxa recorded are included in the figure. This includes preferred prey, less important prey,
and some taxa that would rarely be prey. BSD is semi-deciduous forest (Polisar, 2000).

The juxtaposition of contrasting productive habitats,
which also created productive edges, seemed key in
defining desirable jaguar home ranges. In this way, the
jaguar may be similar to the tiger, whose prey is most
abundant where ‘‘grasslands and forests form a mosaic
and the interdigitation of many different vegetation
types supports a rich ungulate community’’ (Sunquist et
al., 1999). The ungulate community of the llanos is
hardly equal to that of southeast Asia. It is neither cer-
vid nor bovid rich. Larger caviomorph rodents fill some
niches occupied by cervids and bovids in the Old World
Tropics (Eisenberg and McKay, 1974). Nonetheless, the
patterns of prey production across landscapes bear
similarities.
The areas that had low prey abundance were large
open savannas in the far south of the study area, and
the high dry pastures, set in hills, that were used for
calving (Fig. 2). These pastures were bordered by dry
forest, a habitat more frequently occupied by puma
than jaguar (Maxit, 2001). Pumas residing in the vicinity
of these maternity pastures had pockets of productivity
within their ranges, but the immediate vicinity of the
pastures had low native prey abundance and diversity.
In managing the calving season successfully by moving
cattle to higher ground, ranching operations may coin-
cidentally reduce some potential problems with jaguars
while increasing the potential for problems with pumas.
Prey-poor patches become rich, their wealth in calves.
Jaguar preyed selectively upon collared peccaries and
capybara. In puma scats, capybara remains were as fre-
quent as deer, although the latter were 4.1 times as
abundant. Capybara constituted 71% of the recorded
kills of wild prey by puma. Why might capybara be a
preferred prey? The profit margin must be high. Capy-
baras are as large as deer, and larger then peccary, yet
appear to have less flight capability and weaker defen-
ses.
In Piñero, jaguars make rounds, following a rough
circuit, as they check on productive patches. The mar-
ginal value that Charnov (1976) predicted occurs
because prey in patches become wary and/or flee in the
presence of a predator, particularly after a herd member
has fallen victim (Brown et al., 1999). Capybara possess
a behavioral constraint that imposes relatively tight site
fidelity. They rarely occur more than 500 m from water
(Ojasti, 1973). Activity centers are rarely more than 300
m from water (Herrera and MacDonald, 1989). As a
consequence, home ranges are very small. In Apure,
capybara ranges measured 6–16 ha (Herrera and Mac-
Donald, 1989), which would be approximately 20% of
the area of collared peccary home ranges in Guarico
(Castellanos, 1982). Capybara densities are exceptionally
306 J. Polisar et al. / Biological Conservation 109 (2003) 297–310
high locally (and exceptionally low away from water).
Although amphibious like the caiman, capybara spend
far more time in terrestrial habitats, including forest.
The patch they occupy may be slow to lose its value.
Since capybara need to reconvene at water, restoration
of the value of the patch they occupy may be more
rapid. A cat might decide to visit more frequently, or
even stay a while.
Handling costs of prey procurement include the phy-
sical hazards of capture. Adult caiman, when struggling,
represent a risk to dentition. Collared peccary canines
approach those of a jaguar in size, and are sharper.
Both peccary species rely on groups for vigilance and
defense. In Hato Piñero average jaguar weights were 1.9
times that of puma, a factor reflected in the jaguar’s
heavier use of large bodied risky prey. Young calves
weigh 30–50 kg, as much as large natural prey, but lack
many of the defenses that natural prey possess (Maxit,
2001; Scognamillo, 2001). Although cows do rally to the
defense of their calves, the short-term cost of a calf is
likely to seem low to a predator.
A maternity pasture, in which cows and calves are
fenced becomes a patch whose value may never become
marginal. Theory would predict no travel from such a
patch. Unfortunately some pumas follow the prediction.
Although the majority of cats do not make a habit of
preying on such situations, some do. It pays them high
profits with low costs, until their demise.
In Piñero, the frequency of cattle depredation was
inversely related to availability and vulnerability of
natural prey and directly related to availability and
vulnerability of livestock. There was some coincidence
in this. Young calves were not often pastured in the
prey-rich well-watered small forest-lined savannas at
low elevations. Cattle were virtually absent from some
of the most prey rich areas in high-stature semi-decid-
uous forest due to a lack of suitable forage in those
areas.
Tables 3 and 4 are revealing. The biomass of the
native artiodactyls (Cervidae 124.77 kg/km2 and Tayas-
suidae 64.59 kg/km2) is roughly equal to that of the
introduced Perissodactyls (Equidae 187.09 kg/km2). As
a result, the domestic mammalian biomass (introduced
Bovidae and Equidae) is roughly equal to the total
artiodactyl biomass (Bovidae, Cervidae, Tayassuidae),
both being around 97% (Table 4). Roughly 3% of the
mammalian biomass is large native prey (Table 4). The
mammalian biomass of the llanos is high, approaching
the richest sites of Africa and Asia, and exceeding many
productive sites of the Old World Tropics (Table 5;
Eisenberg, 1980; Eisenberg and Lockhart, 1972; Eisen-
berg and Seidensticker, 1976; Karanth and Sunquist,
1992; Mckay and Eisenberg, 1974; Peres, 1999; Schaller,
1967, 1972, 1983). This biomass is in human-facilitated
ecological replacements of the grazers that went extinct
in the Pleistocene. Approximately 10,000–11,000 years
ago the remaining ancient South American ungulates
(Toxodontia, Litopterna, and Glyptodonts) and
roughly half of the Pliocene’s northern immigrants
(Proscidea, Perissiodactyla, and Artiodactla) went
extinct (Cartelle, 1999; MacFadden and Shockey, 1997;
Martin, 1967; Webb, 1978). Bovidae never occurred in
South America. Although capybara are grazers, they
are small relative to the recent mega-grazers. At present,
there are 21 species of ungulates in tropical America
(Ojasti, 1983). Most are at least partially dependent
upon forest. In proportion to continent areas, by Afri-
can standards, there would be 55 ungulates in South
America (Ojasti, 1983). The savannas of South America
were, in some respects, empty when the Spanish arrived,
carrying Old World grazers. Feral on the landscape, the
bovids and equids multiplied. Managed by humans
(immunizations, predator control, forage improve-
ments), their biomass climbed even higher (Crosby,
1986).
Before the extinction, the New World tropics had
large herbivores, and an associated assemblage of large
predators. In the Pleistocene, there were lions (Panthera
atrox), jaguars (Panthera onca), pumas (Puma concolor)
and sabertooth cats (Smilodon fatalis) in Florida (Mor-
gan and Seymour, 1997). A community of a large
sabertooth cat (Homotherium serum), a smaller lighter
sabertooth cat (Smilodon gracilis) and a cheetah-like cat
(Miracinoyx inexpectatus) approximately mirrored the
size class distribution of the present community of lion,
leopard, and cheetah in Africa (Morgan and Seymour,
1997). The communities of large mammals in South
America are recent. Individual members may be
ancient, some more then others, but the communities
are very recent. The native herbivore biomass in the
llanos is miniscule compared to Africa. With large gra-
zers reintroduced, the biomass surpasses famous grazing
grounds in Africa (Table 5). The vast majority of that
biomass is in cattle. It is surprising that the problem of
livestock depredation by jaguar and puma is not even
more severe.
The perspective that fossils provide does little to
assuage the concerns of ranchers. At present, some
puma and fewer jaguars make decisions based on sound
energetics that fail to factor in lethal rancher’s respon-
ses. Cattle ranchers lose when cats take livestock. Ulti-
mately, the cats lose too. Humans introduced bovids
into the receptive environment of the New World tro-
pics. In doing so, we created this dilemma, and it thus
rests upon us to help the cats make wise decisions.
4.1. Recommendations for ranchers
Observations made by ranchers and researchers
working in Venezuela and Brazil have resulted in sug-
gestions for promoting the coexistence of cats and cattle
(Hoogesteijn et al., 1993, in press; Hoogesteijn and
 J. Polisar et al. / Biological Conservation 109 (2003) 297–310
Mondolfi, 1993; Quigley and Crawshaw, 1992; Scogna-
millo et al., in press). These include:
1. Protect all principal prey of the large cats by
preventing poaching;
2. Avoid commercial harvests of capybara and cai-
man. If harvests are conducted, exert strict con-
trol, particularly with capybara;
3. When feasible, impede the ability of cattle to
enter forest;
4. Concentrate calving seasons via artificial insemi-
nation. A shorter calving season facilitates con-
trol;
5. When possible, locate maternity pastures at a
distance from cover that cats may prefer;
6. Explore the application of electric fence around
maternity pastures, as developed by Scognamillo
et al. (in press);
7. If practical, move calves from pastures with
chronic depredation problems and replace with
older animals, over 1–2 years of age;
8. Move all cattle out of lowland flooding areas
before waters rise to avoid isolation and crowd-
ing in forest islands amidst flooded savannas;
9. Where possible, stock low flooding savannas
with water buffalo (less vulnerable);
10. Keep good clear records of losses from all causes
to facilitate planning and decision making; and
11. Do not clear all forests.
In our study area, large inter-connected cattle ranches
owned by ranchers that appreciated wildlife values
allowed large mammals such as tapir, white-lipped
peccary, and jaguar to persist in an agricultural mosaic.
These ranches constituted partially protected areas,
enforced more effectively than some national parks. The
natural prey base in Hato Piñero was judged to be ade-
quate. Cats were not forced to rely on domestic live-
stock. Outside of these protected ranches prey can be
presumed to be less abundant. Where Piñero lay adja-
cent to unprotected land, capybaras were less abundant
than habitats could support. Wary behavior and noc-
turnal habits suggested intense poaching pressure.
Within Piñero, wildlife protection was compromised
where a small river that ran through the property (by
law a public thoroughfare). Capybara were nearly era-
dicated and caiman far fewer there than in better areas.
Camps and cleaned carcasses were evidence of poaching
activity. Piñero’s cats had the choice to not take more
domestic prey because (in most areas) vigilance had
maintained the natural prey.
In Piñero, progressive cattle management had already
reduced the potential severity of depredation. (Quigley and
Crawshaw, 1992) described how in lightly managed ran-
ches of the Brazilian Pantanal cattle were left in low-lying
307
areas during the wet season. Cattle and large cats were
confined to slightly elevated forest islands where the
cattle fell prey to jaguar. Piñero’s seasonal cattle drives
between lowlands and uplands were geared towards
increasing calf survivorship and maximizing forage
opportunities. Coincidentally they probably reduced the
frequency of jaguar attacks, but may have elevated the
likelihood of attacks on calves by puma in upland pas-
tures. From an economic perspective, calves lost to
pumas in uplands must have been less than would have
been lost by leaving herds in flooded savannas. The
persistence of cattle losses, albeit at a relatively low
level, in this well-managed setting is testimony that
depredation can be reduced but probably never fully
eliminated.
In the absence of regular herd inspections and written
records, calf losses due to injuries, diseases, poor nutri-
tion, or poisonous bites may be mistakenly attributed to
cats. Good records clarify the actual proportion of los-
ses that are due to cats, reducing the tendency to make
the cats scapegoats for poor management (Hoogesteijn
et al., 1993).
The recommendation that forests not be cleared may
be equivocal. Unrestrained forest clearing might alle-
viate conflicts between ranchers and wildlife altogether,
albeit eliminating all wildlife values. Our findings imply
that puma depredation may be less limited by proximity
to forest than jaguar depredation. It is clear from this
study that forests are a reservoir of prey. Some forests
are more productive than others (Fig. 2). In Piñero, the
highest depredation rates occurred in areas where prey
abundance and diversity was relatively low. If a ranch
has cats and wishes to accommodate them, the more
prey-rich forest it has, the less a temptation livestock
will be. Any measures that keep cattle from the forest
interior are likely to reduce attacks by jaguar. One of
the few documented cases of jaguar attacks on cattle
during our study took place when a young cow strayed
into dense riparian forest.
Motivated ranchers might consider locating some
excavated water retention ponds specifically for prey.
This would elevate prey numbers, direct dry-season
spatial distribution of prey, and thus potentially focus
cats’ activities in space. Savannas boasting abundant
ponds and bordered by forest were one of Piñero’s
richer habitat types in terms of prey (Fig. 2). In these
habitats, capybara and caiman frequented the water
bodies, deer traversed the ecotone, visiting the ponds to
drink, the adjacent forests supported peccary, and
attacks on cattle were scarce.
Acknowledgements
Funding was provided by The National Geographic
Society, the Wildlife Conservation Society, the British
308 J. Polisar et al. / Biological Conservation 109 (2003) 297–310
Embassy to Venezuela’s Cooperation Fund, the Cat
Specialist Group of the Species Survival Commission
(IUCN-World Conservation Union), and the Lincoln
Park Zoo Scott Neotropic Fund, and Katharine B.
Ordway Chair endowment funds via Dr. J.F. Eisenberg.
Faunal aspects of the study were assisted in the field by
Victor Juan Meires, Orlando Ramirez, Gilson Rivas
Fuenmayor, Diego Giraldo, Sandra Melman, Emiliana
Isase, Telva Carantona, and Marcus Trepte. Rafael
Ortiz and Dr. Francisco Delascio worked on floral
aspects. Rafael Hoogesteijn, Roy McBride, Rocky
McBride, Tibisay Escalona, Bruno Pampour, the lla-
neros of Piñero, and students and staff of Dr. Juhani
Ojasti’s wildlife management class (UNELLEZ at Gua-
nare) all contributed greatly. Francisco Bisbal per-
formed some scat analyses. His staff provided logistical
support. Edgardo Mondolfi served as liaison. Don
Antonio Julio Branger provided an excellent study area,
and facilitated. Matt Burgess, Kyle Harris, Stephen
Taranto, and Richard Owens condensed data. Maria
Fernanda Zermoglia, and Rosanna Rivero refined
maps. F. Wayne King, Jim Nichols, and Fred Thomp-
son reviewed the dissertation from which this paper
originated (Polisar, 2000).
References
Ackerman, B.B., Lindzey, F.G., Hemker, T.P., 1986. Predictive ener-
getics model for cougars. In: Miller, S.D., Everett, D.D. (Eds.), Cats
of the World: Biology, Conservation, and Management. The
National Wildlife Federation, Washington, D.C, pp. 333–352.
Anderson, A.E., 1983, A critical review of literature on puma. Color-
ado Division of Wildlife Special Report 54.
Anon., 1995. ArcView. Environmental Systems Research Institute,
Redlands, CA.
Ayarzagüena, J.S., 1983. Ecologia del caiman de anteojos o baba
(Caiman crocodilus) en los llanos de Apure (Venezuela). Doñana 10,
1–136.
Bodmer, R., Aquino, R., Puertas, P., Reyes, C., Fang, T., Gottdenker,
N., 1997. Manejo y uso sustentable de pecarı́es en la Amazonı́a
Peruana. Unión Internacional para le Conservación de la Natur-
aleza y los Recursos Naturales, Quito.
Brokx, P.A.J., 1972. A Study of the Biology of Venezuelan White-
tailed Deer (Odocoileus virginianus gymnotis Wiegmann, 1833), with
a Hypothesis on the Origin of the South American Cervids. PhD
dissertation, University of Waterloo, Waterloo.
Brown, J.S., Laundré, J.W., Gurung, M., 1999. The ecology of fear:
optimal foraging, game theory, and trophic interactions. Journal of
Mammalogy 80, 385–399.
Buckland, S.T., Anderson, D.R., Burnham, K.P., Laake, J.L., 1993.
Distance Sampling: Estimating Abundance of Biological Popula-
tions. Chapman and Hall, London.
Cartelle, C., 1999. Pleistocene mammals of the Cerrado and Caatinga
of Brazil. In: Eisenberg, J.F., Redford, K.H. (Eds.), Mammals of
the Neotropics: The Central Neotropics. The University of Chicago
Press, Chicago, pp. 27–46.
Castellanos, A.H.G., 1982. Patrones de movimiento y uso de habitat
del baquiro de collar Tayassu tajacu en los llanos centrales de Vene-
zuela. Tesis de Licenciatura, Universidad Central de Venezuela.
Charnov, E.L., 1976. Optimal foraging, the marginal value theorem.
Theoretical Population Biology 9, 129–136.
Colwell, R.K., Futumaya, D.J., 1971. On the measurement of niche
breadth and overlap. Ecology 52, 567–577.
Corrales, F., 1998. Libros de Ganaderia de Hato Pinero. Agropecuaria
San Franciso, Valencia, Venezuela.
Crosby, A.W., 1986. Ecological imperialism: the biological expansion
of Europe, 900–1900. Cambridge University Press, Cambridge.
Culver, M., Johnson, W.E., Pecon-Slattery, J., O’Brien, S.J., 2000.
Genomic ancestry of the American puma (Puma concolor). Journal
of Heredity 91, 186–197.
Curio, E., 1976. Ethology of Predation. Springer-Verlag, Berlin.
Dailey, T.V., Thompson Hobbs, N., Woodard, T.N., 1984. Experi-
mental comparisons of diet selection by mountain goats and moun-
tain sheep in Colorado. Journal of Wildlife Management 48, 799–
806.
Dimmick, R.W., Pelton, M.R., 1994. Criteria of sex and age. In:
Bookhout, T.A. (Ed.), Research and Management Techniques for
Wildlife and Habitats. The Wildlife Society, Bethesda, pp. 169–214.
Eisenberg, J.F., 1980. The density and biomass of tropical mammals.
In: Soulé, M.A., Wilcox, B.A. (Eds.), Conservation Biology: an
Evolutionary-ecological Perspective. Sinauer Associates, Sunder-
land, pp. 35–55.
Eisenberg, J.F., Lockhart, M., 1972. An ecological reconnaissance of
Wilpattu National Park, Ceylon. Smithsonian Contributions to
Zoology 101, 1–118.
Eisenberg, J.F., McKay, G.M., 1974. Comparison of ungulate adap-
tations in the New World and Old World tropical forests with spe-
cial reference to Ceylon and the rainforests of Central America. In:
Geist, V., Walther, F. (Eds.), The Behavior of Ungulates and its
Relation to Management. IUCN, Morges, Switzerland, pp. 585–
602.
Eisenberg, J.F., O’Connell, M.A., August, P.V., 1979. Density, pro-
ductivity, and distribution of mammals in two Venezuelan habitats.
In: Eisenberg, J.F. (Ed.), Vertebrate Ecology in the Northern Neo-
tropics. Smithsonian Institution Press, Washington, DC, pp. 187–
207.
Eisenberg, J.F., Polisar, J.R., 1999. The mammal species of north-
central Venezuela. Bulletin of the Florida Museum of Natural His-
tory 42, 115–160.
Eisenberg, J.F., Seidensticker, J., 1976. Ungulates in Southern Asia: a
consideration of biomass estimates for selected habitats. Biological
Conservation 10, 293–305.
Emlen, J.M., 1966. The role of time and energy in food preference.
The American Naturalist 100, 611–617.
Emmons, L.H., 1987. Comparative feeding ecology of felids in a neo-
tropical rain forest. Behavioral Ecology and Sociobiology 20, 271–
283.
Emmons, L.H., 1989. Jaguar predation on Chelonians. Journal of
Herpetology 23, 311–314.
Farrell, L., 1999. The Ecology of the Puma and the Jaguar in the
Venezuelan Llanos. MS thesis University of Florida, Gainesville.
Farrell, L., Roman, J., Sunquist, M.E., 2000. Dietary separation of
sympatric carnivores identified by molecular analysis of scats.
Molecular Ecology 9, 1583–1590.
Hayne, D.H., 1984. Population dynamics and analysis. In: Halls, L.K.
(Ed.), White-tailed Deer: Ecology and Management. Stackpole
Books, Harrisburg, pp. 203–210.
Hellgren, E.C., Synatzske, D.R., Oldenburg, P.W., Guthery, F.S.,
1995. Demography of a collared peccary population in south Texas.
Journal of Wildlife Management 59, 153–163.
Herrera, E.A., MacDonald, D.W., 1989. Resource utilization and ter-
ritoriality in group-living capybara (Hydrochoerus hydrochaeris).
Journal of Animal Ecology 58, 667–679.
Hoogesteijn, R., Boede, E.O., Mondolfi, E., in. Observaciones sobre la
depredacion de jaguares sobre bovinos en Venezuela y los pro-
gramas de control gubernamentales. In: Medellı́n, R.A., Chetkie-
wicz, C., Rabinowitz, A., Redford, K.H., Robinson, J.G.,
Sanderson, E., Taber, A. (Eds.), Los Jaguares en el Nuevo Milenio.
 J. Polisar et al. / Biological Conservation 109 (2003) 297–310
Un Análisis de su Estado, Detección de Prioridades y Recomenda-
ciones Para la Conservación del Jaguar en América. Instituto de
Ecologı́a, Universidad Nacional Autonoma de México/Wildlife
Conservation Society, México, D.F.
Hoogesteijn, R., Hoogesteijn, A., Mondolfi, X., 1993. Jaguar preda-
tion and conservation: cattle mortality caused by felines on three
ranches in the Venezuelan Llanos. In: Dunstone, N., Gorman, R.L.
(Eds.), Mammals as Predators. Zoological Society, London, pp.
391–407.
Hoogesteijn, R., Mondolfi, E., 1993. The Jaguar. Armitano Publish-
ers, Caracas.
Hornocker, M.G., 1970. An analysis of mountain lion predation upon
mule deer and elk in the Idaho primitive area. Wildlife Monographs
21, 1–39.
Karanth, K.U., Sunquist, M., 2000. Behavioral correlates of predation
by tiger (Panthera tigris), leopard (Panthera pardus) and dhole
(Cuon alpinus) in Nagarahole, India. Journal of Zoology London
250, 255–265.
Karanth, K.U., Sunquist, M.E., 1992. Population structure, density
and biomass of large herbivores in the tropical forests of Nagara-
hole, India. Journal of Tropical Ecology 8, 21–35.
Karanth, K.U., Sunquist, M.E., 1995. Prey selection by tiger, leopard,
and dhole in tropical forests. Journal of Animal Ecology 64, 439–
450.
Kleiman, D.G., Eisenberg, J.F., Maliniak, E., 1979. Reproductive
parameters and productivity of caviomorph rodents. In: Eisenberg,
J.F. (Ed.), Vertebrate Ecology in the Northern Neotropics. Smith-
sonian Institution Press, Washington, DC, pp. 173–183.
Lancia, R.A., Nichols, J.D., Pollock, K.H., 1994. Estimating the
number of animals in wildlife populations. In: Bookhout, T.A.
(Ed.), Research and Management Techniques for Wildlife and
Habitats. The Wildlife Society, Bethesda, pp. 215–253.
Linares, O., 1998. Mamı́feros de Venezuela. Sociedad Con-
servacionista Audubon de Venezuela, Caracas.
Lord, R.D., Lord, V.R., 1988. Cross checking censuses and a model of
the annual cycle of mortality and reproduction in Capybaras
(Hydrochaeris hydrochaeris). Studies on Neotropical Fauna and the
Environment 23, 213–224.
MacArthur, R.H., Pianka, E.R., 1966. On optimal use of a patchy
environment. The American Naturalist 100, 603–609.
MacFadden, B.J., Shockey, B.J., 1997. Ancient feeding ecology and
niche differentiation of Pleistocene mammalian herbivores from
Tarija, Bolivia: morphological and isotopic evidence. Paleobiology
23, 77–100.
Martin, P.S., 1967. Pleistocene overkill. In: Martin, P.S., Wright, H.E.
(Eds.), Pleistocene Extinctions: The Search for a Cause. Yale Uni-
versity Press, New Haven, pp. 75–120.
Maxit, I.E. 2001. Prey Use by Sympatric Puma and Jaguar in the
Venezuelan Llanos. MS thesis, University of Florida, Gainesville.
Mckay, G.M., Eisenberg, J.F., 1974. Movement patterns and habitat
utilization of ungulates in Ceylon. In: Geist, V., Walther, F. (Eds.),
The Behavior of Ungulates and its Relation to Management. IUCN
Publication, Morges, Switzerland, pp. 708–721.
Miller, L.E., 1992. Socioecology of the Wedge-capped Capuchin
Monkey (Cebus olivaceous). PhD dissertation, University of Cali-
fornia, Davis.
Mondolfi, E., Hoogesteijn, R., 1986. Notes on the biology and status
of the jaguar in Venezuela. In: Miller, S.D., Everett, D.D. (Eds.),
Cats of the World: Biology, Conservation, and Management.
National Wildlife Federation, Washington, D.C, pp. 125–146.
Morgan, G.S., Seymour, K.L., 1997. Fossil history of the panther
(Puma concolor) and the cheetah-like cat (Miracinoyx inexpectatus).
Bulletin of the Florida Museum of Natural History 40, 177–219.
Nuñez, R., Miller, B., Lindzey, F., 2000. Food habits of jaguars and
pumas in Jalisco, Mexico. Journal of Zoology 252, 373–379.
Ojasti, J., 1973. Estudio biólogico del chigüire o capibara. Fondo
Nacional de Investigaciones Agropecuarias, Caracas.
309
Ojasti, J., 1983. Ungulates and large rodents of South America. In:
Bourlière, F. (Ed.), Ecosystems of the World. Vol. 3. Tropical
Savannas. Elsevier Scientific Publishing Company, Amsterdam, pp.
427–439.
Ojeda, M.M., Keith, L.M., 1982. Sex and age composition and
breeding biology of cottontail rabbit populations in Venezuela.
Biotropica 14, 99–107.
Peres, C.A., 1999. The structure of nonvolant mammal communities in
different Amazonian forest types. In: Eisenberg, J.F., Redford, K.H.
(Eds.), Mammals of the Neotropics: The Central Neotropics. The
University of Chicago Press, Chicago, pp. 564–581.
Polisar, J., 2000. Jaguars, Pumas, their Prey Base, and Cattle Ranch-
ing; Ecological Perspectives of a Management Issue. PhD disserta-
tion, University of Florida, Gainesville.
Quigley, H., Crawshaw, P., 1992. A conservation plan for the jaguar
Panthera onca in the Pantanal region of Brazil. Biological Con-
servation 61, 149–157.
Rabinowitz, A.R., Nottingham Jr., B.G., 1986. Ecology and behavior
of the jaguar (Panthera onca) in Belize, Central America. Journal of
Zoology 210, 149–159.
Robinson, J.G., Bodmer, R.E., 1999. Towards wildlife management in
tropical forests. Journal of Wildlife Management 63, 1–13.
Robinson, J.G., Redford, K.H., 1986. Intrinsic rate of natural increase
in Neotropical forest mammals: relationship to phylogeny and diet.
Oecologia 68, 516–520.
Robinson, J.G., Redford, K.H., 1991. Sustainable harvest of Neo-
tropical forest mammals. In: Robinson, J.G., Redford, K.H. (Eds.),
Neotropical Wildlife Use and Conservation. The University of Chi-
cago Press, Chicago, pp. 415–429.
Roosevelt, T., 1914. Through the Brazilian Wilderness. Charles Scrib-
ner’s Presses, New York.
Ross, P.I., Jalkotzy, M.G., Festa-Bianchet, M., 1997. Cougar preda-
tion on bighorn sheep in southwestern Alberta during winter.
Canadian Journal of Zoology 74, 771–775.
Schaller, G.B., 1967. The Deer and the Tiger. The University of Chi-
cago Press, Chicago.
Schaller, G.B., 1972. The Serengeti Lion: a Study of Predator–prey
Relations. The University of Chicago Press, Chicago.
Schaller, G.B., 1983. Mammals and their biomass on a Brazilian
ranch. Arquivos de Zoologia 31, 1–36.
Schaller, G.B., Crawshaw, P., 1980. Movement patterns of jaguar.
Biotropica 12, 161–168.
Scognamillo, D., Maxit, I., Sunquist, M., Farrell, L., in. Ecologia del
jaguar y el problema de la depredacion sobre ganado en Hato
Piñero, Venezuela. In: Medellı́n, R.A., Chetkiewicz, C., Rabinowitz,
A., Redford, K.H., Robinson, J.G., Sanderson, E., Taber, A. (Eds.),
Los jaguares en el nuevo milenio. Un análisis de su estado, detec-
ción de prioridades y recomendaciones para la conservación del
jaguar en América. pressInstituto de Ecologı́a, Universidad Nacio-
nal Autonoma de México/Wildlife Conservation Society, México,
D.F.
Scognamillo, D.G., 2001. Ecological Separation Between Jaguar and
Puma in a Mosaic Landscape in the Venezuelan Llanos. MS thesis,
University of Florida, Gainesville.
Seber, G.A.F., 1982. The Estimation of Animal Abundance and
Related Parameters. Charles Griffin and Company, London.
Seidensticker, J., 1976. On the ecological separation between tigers
and leopards. Biotropica 8, 225–234.
Shaw, H., 1977. Impact of mountain lion on mule deer and cattle in
northwestern Arizona. Pages. In: Phillips, R.L., Jonkel, C. (Eds.),
Proceedings of the 1975 Predator Symposium. Montana Forest and
Conservation Experimental Station, Missoula, pp. 17–31.
Smythe, N., 1978. The natural history of the Central American agouti
(Dasyprocta punctata). Smithsonian Contributions to Zoology 257,
1–52.
Sowls, L.K., 1997. Javelina and Other Peccaries: Their Biology, Man-
agement, and Use. Texas A & M University Press, College Station.
310 J. Polisar et al. / Biological Conservation 109 (2003) 297–310
Spowart, R.A., Thompson Hobbs, N., 1985. Effect of fire on diet
overlap between mule deer and mountain sheep. Journal of Wildlife
Management 49, 942–946.
SPSS, 1999. SPSS Base 9.0 User’s guide. SPSS Inc, Chicago.
Sunquist, M.E., 1981. The social organization of tigers (Panthera
tigris) in Royal Chitawan National Park, Nepal. Smithsonian Con-
tributions to Zoology 336, 1–98.
Sunquist, M.E., in. History of jaguar research in the Americas. In:
Medellı́n, R.A., Chetkiewicz, C., Rabinowitz, A., Redford, K.H.,
Robinson, J.G., Sanderson, E., Taber, A. (Eds.), Los Jaguares en el
Nuevo Milenio. Un Análisis de su Estado, Detección de Prioridades
y Recomendaciones Para la Conservación del Jaguar en América.
Instituto de Ecologı́a, Universidad Nacional Autonoma de México/
Wildlife Conservation Society, México, D.F.
Sunquist, M.E., Karanth, K.U., Sunquist, F.C., 1999. Ecology, beha-
vior and resilience of the tiger and its conservation needs. In: Sei-
densticker, J., Christie, S., Jackson, P. (Eds.), Riding the Tiger:
Tiger Conservation in Human-dominated Landscapes. Cambridge
University Press, Cambridge, pp. 5–18.
Sunquist, M.E., Sunquist, F.C., 1989. Ecological constraints on predation
by large felids. In: Gittleman, J.L. (Ed.), Carnivore Behavior, Ecology,
and Evolution. Cornell University Press, Ithaca, pp. 283–301.
Taber, A.B., Novaro, A.J., Neris, N., Colman, F.H., 1997. The food
habits of sympatric jaguar and puma in the Paraguayan Chaco.
Biotropica 29, 204–213.
Taylor, R.J., 1976. Value of clumping to prey and the evolutionary
response of ambush predators. American Naturalist 110, 13–29.
Teer, J.G., 1984. Lessons from the Llanos basin, Texas. In: Halls, L.K.
(Ed.), White-tailed Deer: Ecology and Management. Stackpole
Books, Harrisburg, pp. 261–290.
Temple, S.A., 1987. Do predators always capture substandard indivi-
duals disproportionately from prey populations? Ecology 68, 669–
674.
Webb, S.D., 1978. A history of savanna vertebrates in the New World.
Part II: South America and the great interchange. Annual Review of
Ecology and Systematics 9, 393–426.
Winston, P.M., 1991. Odocoileus virginianus. Mammalian Species 388,
1–13.