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Journal of Zoology. Print ISSN 0952-8369

Density, habitat use and activity patterns of ocelots
(Leopardus pardalis) in the Atlantic Forest of Misiones,
M. S. Di Bitetti, A. Paviolo & C. De Angelo
National Research Council of Argentina (CONICET) and Laboratorio de Investigaciones Ecologicas
de las Yungas (LIEY), Universidad Nacional de
´ Argentina

ocelot; density; habitat use; daily activity
patterns; moonlight activity.
Mario S. Di Bitetti, CONICET-LIEY, Yapeyu´
23, 3370 Puerto Iguazu´, Misiones,
Received 8 June 2005; accepted
6 December 2005

Camera-trap surveys were carried out at two different sites within the Atlantic
Forest of Misiones province, Argentina, to study the density, habitat use and
activity patterns of ocelots. At Urugua- ´ı Provincial Park, 17 different individuals
were captured (nine females, six males, two of unknown sex) during a 3-monthlong survey (34 camera stations, 1409 trap days). At Iguazu´ National Park,
34 different individuals were trapped (20 adult females, nine adult males, two subadult females and three of unknown sex) during the survey (39 sampling stations,
1631 trap days). Population density estimates ( SE) for Urugua- ´ı, in
an area of between 150 and 259 km2 (depending on the buffer used to estimate the area effectively sampled), range from 7.7 1.4 to 13.4 2.6 individuals 100 km 2, whereas at Iguazu´, in an area of between 275 and 428 km2,
a population density of between 12.8 2.7 and 20.0 4.2 individuals 100 km 2
was estimated. Minimum observed range estimates for individuals with
43 capture sites range from 3.19 to 37.09 km2 for four males and from 4.17 to
7.11 km2 for three females, but underestimate the true home range size. Ocelots
were captured more frequently along old roads than on new trails opened with
machetes. Ocelots were captured more frequently at night than during the day and
reduce their use of roads and trails during the week previous to and during full
moon nights, a behavior previously reported for Amazonian ocelots. Population
density estimates for ocelots in the Upper Parana´ Atlantic Forest ecoregion are
lower than those at other neotropical sites. The whole Green Corridor contains a
population of about 1280 individuals. This estimate should bring our attention to
the larger cats (pumas and jaguars) that live at lower population densities because
the future of their local populations is compromised if protected areas are not
urgently created and properly managed.

The ocelot Leopardus pardalis is a medium-sized (adult
weight range: 7–16 kg) neotropical cat with a distributional range from southern Texas in the USA to northern
Argentina (Emmons & Feer, 1997; Murray & Gardner,
1997). Ocelots are opportunistic predators that consume
any type of small and medium-size (usually less than 2 kg)
terrestrial prey available (Emmons, 1987; Murray & Gard´
ner, 1997; De Villa Meza, Martinez Meyer & Lopez
2001). The species was heavily hunted for its skin
and, as a result of this and of habitat loss, local populations
were declining until the 1980s, when the international trade
was banned. Currently, the ocelot is a widespread and sometimes locally common wild cat in the remaining neotropical
forests, but habitat loss remains a threat to populations.

Field studies on ocelot ecology and behavior have focused on their abundance, diet, activity patterns, home
range and habitat use (see the review in Murray & Gardner,
1997). Many of these studies have made use of radiotracking
as a tool to study ocelot activity patterns and habitat use.
Most recently, camera traps have become available as a tool
to study the behavior of cryptic animals, which, like ocelots,
can be identified by their spotted fur patterns (e.g. Karanth
& Nichols, 1998; Silver et al., 2004). Recently, Trolle & Ke´ry
(2003, 2005) and Maffei et al. (2005) used camera traps to
estimate ocelot densities in the Brazilian Pantanal and the
Bolivian dry forests, respectively. Camera-trap records can
also provide information on home range, daily and seasonal
activity patterns, and population dynamics. However, these
other aspects have been rarely explored in camera-trap
studies on felids, probably as a result of the small sampling

c 2006 The Authors. Journal compilation
c 2006 The Zoological Society of London
Journal of Zoology 270 (2006) 153–163


Density and habitat use of ocelots in NE Argentina



M. S. Di Bitetti, A. Paviolo and C. De Angelo


B r a z i l



P a r a g u a y



ne ido
sio orr
M nC



A r g e n t i n a




Figure 1 Location of the study sites.

effort or the low population density of the study animals,
which rendered few data available for statistical analyses
(but see Maffei et al., 2005).
Information on ocelot activity patterns, habitat use and
densities have just started to provide information on the
degree of variability of this cat species. For example, in the
Amazonian rainforest of Manu, Peru, ocelots live at high
densities and use small home ranges (Emmons, 1988). In the
southern USA, ocelots occupy larger home ranges, apparently a consequence of the low prey availability at the
margin of their distribution (Tewes, 1986). In the Amazon
rainforest of Peru and in the Venezuelan Llanos, ocelots are
a mostly nocturnal species (Ludlow & Sunquist, 1987;
Emmons, 1988; Sunquist, Sunquist & Daneke, 1989), but in
Belize, the Brazilian Pantanal and the Upper Parana´ Atlantic Forest ecoregion (Iguazu´ National Parks of Brazil and
Argentina) they seem to have a proportionally higher daytime activity (Crawshaw & Quigley, 1989; Konecny, 1989;
Crawshaw, 1995).
In this study, we used camera traps to study the activity
patterns, habitat use and absolute density of a population of
ocelots in the Upper Parana´ Atlantic Forest of Argentina.
This population has been previously studied by Crawshaw
(1995) using radiotelemetry, which allows for a comparison
of the results obtained by the two methods.

Study sites
We carried out this study at two sites within the Green
Corridor of Misiones province, Argentina (Fig. 1). The area
belongs to the innermost ecoregion of the Atlantic Forest
complex, the Upper Parana´ Atlantic Forest ecoregion
(Di Bitetti, Placci & Dietz, 2003). The Green Corridor

contains the largest forest remnants in the Atlantic Forest
and still contains a complete native species assemblage,
including top predators like jaguars Panthera onca and
harpy eagles Harpia harpyja (Galindo-Leal & de Gusma˜o
Caˆmara, 2003). The area has a humid subtropical climate
with marked seasonality in day length and temperature
(Crespo, 1982; Di Bitetti & Janson, 2001). The mean annual
precipitation for the area varies between 1700 and 2200 mm,
with no marked dry season (Crespo, 1982; Brown & Zunino,
1990). There is strong seasonal variation in the production
of leaves, flowers, fleshy fruits and arthropods, which is
lowest in the winter months of May–August and reaches its
maximum between October and January (Placci, Arditi &
Cioteck, 1994; Di Bitetti, 2001; Di Bitetti & Janson, 2001).
The first study site, Urugua- ´ı (25158 0 S, 54106 0 W), comprised a large portion of the Urugua- ´ı Provincial Park
(84 000 ha), most of the Urugua- ´ı Private Reserve (3243 ha)
and a portion of a large private property, Campo Los
Palmitos (c. 26 000 ha), that belongs to a timber company
(Alto Parana´ Sociedad Anonima).
At Urugua- ´ı Private
Reserve and at Urugua- ´ı Provincial Park, the native forest
was exploited until the mid-1980s; however, the forest is
representative of a mature forest in a relatively good condition. At Campo Los Palmitos, pine Pinus elliotis plantations
are embedded in a matrix of native forest and comprise
20–25% of the area of the property that was surveyed. Of
the total area surveyed (150–259 km2, depending on the
buffer added), about half lay within the protected areas and
half within Campo Los Palmitos. Poaching of wild animals
(mainly pacas Agouti paca, peccaries Tayassu pecari and
Tayassu tajacu, brocket deer Mazama americana and Mazama nana, and tapirs Tapirus terrestris) still occurs in the
area, specially within the Urugua- ´ı Provincial Park, despite
efforts by park rangers, government authorities and private
property owners to control it. We conducted the study at
Urugua- ´ı between May 2003 and February 2004.
The second study site was located at Iguazu´ National Park,
Argentina (25140 0 S, 54130 0 W, 67 000 ha, Fig. 1). We conducted the study at Iguazu´ between late April 2004 and early
December 2004. This strictly protected area was created in
1934, and since then the native forest has not been exploited.
Its fauna and flora are representative of the original Upper
Parana´ Atlantic Forest communitites, and the park is protected by a well-trained corp of elite park rangers that make
illegal poaching of wildlife very uncommon.
At both sites there were few dirt roads available and they
only cover a fraction of the study area. To distribute evenly
and regularly the sampling stations in the study areas, we
opened 86 km of trails (1.5 m wide) with a machete, taking
care that the vegetation was cut very short to encourage the
use of the trails by the wild cats.
A previous study of ocelots and jaguars was conducted by
Peter Crawshaw (1995) at the Iguazu´ National Park of
Brazil (170 000 ha) and the Iguazu´ National Park of Argentina in an area contiguous to our study site in Iguazu´ (Fig. 1).
The two protected areas are separated only by the Iguazu´
River, so there is continuity in ocelot movements between
our study sites and that of Crawshaw’s study.

c 2006 The Authors. Journal compilation
c 2006 The Zoological Society of London
Journal of Zoology 270 (2006) 153–163

M. S. Di Bitetti, A. Paviolo and C. De Angelo

Study methods and data analysis
Ocelots are cryptic and mostly nocturnal, and traditional
methods used to estimate absolute densities of mammals
(e.g. line transect census) are inappropriate for studying
them. For these reasons, we used a new methodology based
on photographic records of individual ocelots obtained with
camera traps (see Karanth & Nichols, 1998; Moraes Tomas
& de Miranda, 2003). Camera traps have been recently used
with success to study population densities of wild cats: tigers
Panthera tigris (Karanth, 1995; Karanth & Nichols, 1998;
Carbone et al., 2001), jaguars P. onca (Wallace et al., 2003;
Maffei, Cue´llar & Noss, 2004; Silver et al., 2004) and ocelots
(Trolle & Ke´ry, 2003, 2005; Maffei et al., 2005). The method
to estimate densities of animals with camera traps is based
on traditional methods to estimate animal densities using
the capture–mark–recapture of individuals (Otis et al., 1978;
White et al., 1982), where a recapture consists in the
appearance of the same individual in subsequent photographic records (Karanth, 1995; Karanth & Nichols, 1998).
The photographs provide information on the date and hour
when the picture was taken and can be used to study the
daily or seasonal activity patterns of wild animals (Moraes
Tomas & de Miranda, 2003).
This study was originally designed to study jaguars,
pumas and ocelots in an effort to assess their population
status and trends. At Urugua- ´ı we deployed 34 sampling
stations at regular intervals of 1–2 km among the nearest
sites. At Iguazu´ we placed 39 sampling stations with a mean
distance of 2–3 km among nearest sites. After applying a
buffer equivalent to 1/2 of the mean maximum distance of
recaptures (MMDM) to each sampling station, no ‘holes’
were left within the sampling areas. This means that any
individual present in the study areas had a probability
greater than zero to be captured by at least one of the
sampling stations.
To explore the possibility that ocelots prefer certain forest
types, we categorized the forest cover and the understory in
the area surrounding each sampling station by walking 50 m
in each direction along the trail or road and by visually
inspecting the canopy and the understory. The canopy layer
was characterized as (1) very discontinuous canopy, (2) open
canopy layer (gaps present) and (3) closed (continuous)
canopy layer. The understory was categorized according to
the presence of bamboos (two native species of bamboo are
very invasive) as (1) no bamboo (or very little) present,
(2) presence of bamboos but no dominance and (3) closed
understory dominated by bamboos. Bamboos usually invade open canopy areas and we found a strong dependence
of our canopy and bamboo indexes estimates (w2 = 21.953,
P= 0.0002, d.f.= 4, n = 73 stations).
Each sampling station consisted of two camera traps
operating independently and facing each other at both sides
of the trails or roads that were not regularly used by
vehicles. This allowed us to obtain pictures of both flanks
of the animals and unambiguously identify them in a
subsequent capture. The equipment consisted of two CamTM
trakker (Camtrakker, Watkinsville, Georgia, USA), 17

Density and habitat use of ocelots in NE Argentina

Leaf Rivers Trail Scan Model C-1 (Vibra Shine, Taylorsville, MS, USA) and 30 TrailMACs 35 mm Standard Game
(Trail Sense Engineering, LLC, Middletown, DE, USA)
scouting cameras.
Because of the paucity of camera traps, at both sites we
subdivided the study area into two time periods, and half of
the stations were operating during each period. Each survey
consisted of two 42-day-long sampling periods separated by
3 (Urugua- ´ı) or 4 (Iguazu´) days during which we shifted the
cameras between the camera sites assigned to the first period
to those assigned to the second. Thus, the surveys lasted
88–89 days. The sampling periods at both sites corresponded
to three consecutive lunar cycles. Assignation of cameras to
each period was carried out so that the cameras assigned to
each period cover the same general area, not as in other
studies where the sampled area was subdivided into two or
more adjacent areas and each portion was surveyed at
sequential time periods (e.g. at Chiquibul, Belize and Madidi, Bolivia jaguar’s studies; Silver et al., 2004; Trolle & Ke´ry,
2005). For this reason most individuals with 42 captures
(15 out of 21) were caught during both sampling periods.
A few days were lost at four stations because camera
traps ran out of batteries or film, creating temporary holes in
the grid of short duration. Camera stations were checked
once every 5–8 days, and if a camera trap had a malfunction
it was replaced by a replacement one to avoid camera
stations working with only one camera trap. Sampling effort
during the survey was 1409 trap days at Urugua- ´ı (19 days
were lost in total at two stations) and 1631 trap days at
Iguazu´ (7 days were lost at two stations).
To estimate the abundance of ocelots using the program
CAPTURE (see below) we clumped both study periods such
that we treated the sample as coming from a 42-day-long
survey consisting of 34 sampling stations at Urugua- ´ı and
39 sampling stations at Iguazu´ (e.g. day 1 of both study
periods was considered as day 1 of the survey, day 2 of both
study periods as day 2 of the survey, etc.). To increase the
individual probability of capture and to make it 40.10 per
trapping occasion, as recommended by Otis et al. (1978) and
White et al. (1982), we pooled three successive days as one
trapping occasion (e.g. days 1–3= first trapping occasion,
days 4–6= second trapping occasion, etc.). Thus, the trapping history of each individual consisted of a string of 14
trapping occasions. For each trapping occasion an individual could get a 1 or a 0, where a 1 indicates that the animal
was captured at any of the sampling stations during that
trapping occasion and a 0 if it was not captured.
Prior to the survey at each site, we conducted a preliminary survey. The preliminary survey started on April 2003 at
Urugua- ´ı and the systematic survey was conducted between
24 November 2003 and 18 February 2004. At Iguazu´ the
survey was conducted between 6 August 2004 and 1 November 2004, with a preliminary survey that started in late
April 2004 and a post-survey period that extended into early
December 2004. During the preliminary and post-survey
periods, sampling effort was not kept constant but the
information from these periods was used only for estimation
of the daily activity patterns.

c 2006 The Authors. Journal compilation
c 2006 The Zoological Society of London
Journal of Zoology 270 (2006) 153–163


Density and habitat use of ocelots in NE Argentina

M. S. Di Bitetti, A. Paviolo and C. De Angelo

To estimate the abundance of animals in the study area,
we used the program CAPTURE (Rexstad & Burnham,
1991). CAPTURE provides population estimates using
different models (for details on these models, see Otis et al.,
1978; White et al., 1982). CAPTURE compares all possible
models among themselves and indicates which of the models
best fits the data. The model Mh was the model that best
fitted the capture–recapture history of individuals at Iguazu´
and ranked second after the null model M0 at Urugua- ´ı,
with a high value of 0.95 (models are ranked by CAPTURE
ranging from 0 to 1, with 1 indicating best fit). Model
Mh assumes heterogeneity among individuals in their capture probabilities, and is the most logically correct because
of the expected behavioral differences among individuals
and their unequal access to camera traps (see Karanth &
Nichols, 1998, 2002). We here report the results of model
Mh using a jackknife estimator.
To estimate the density (and SE) of animals, we followed
the methodology described by Silver et al. (2004) and Maffei
et al. (2005), based on that of Karanth & Nichols (1998) and
Karanth & Nichols (2002). CAPTURE gives an estimate of
the number of individuals present in the study area (and its
SE and 95% confidence limits). To estimate the absolute
density of animals, we need to estimate the area effectively
sampled and its SE. There is no complete agreement among
researchers on how to estimate the actual sampling area.
This has been traditionally estimated by applying a buffer
zone to each sampling station (or to the minimum convex
polygon comprising all sampling stations) equivalent to
1/2 of the MMDM for all individuals observed at two or
more stations (e.g. Karanth & Nichols, 1998, 2002; Maffei
et al., 2004, 2005). MMDM is used as an approximation of
the home range diameter. However, Trolle & Ke´ry (2005)
and the papers presented at a recent workshop on camera
trapping held at the 2005 SCB meeting (Brasilia, Brazil)
suggest that 1/2 of MMDM is an underestimate of the actual
buffer, thus increasing density estimates. The actual value
should lie between 1/2 MMDM and MMDM, and where
radiotelemetry data are available, the best estimate for the
buffer should be the radius of the mean home range estimate
(Trolle & Ke´ry, 2005). We thus provide three different
density estimates for each study site using the following
as the buffer to estimate the actual area sampled:
(1) 1/2 MMDM, (2) MMDM and (3) the mean radius of
the mean home range estimates (from Crawshaw, 1995).
To estimate distances between cameras where ocelots
were caught and areas of home ranges, we used the program ArcView 3.2. The location of trails and sampling

stations was obtained with a GPS receiver (Garmins eTreck
Venture; Garmin International Inc., Olathe, KS, USA).
Individuals were sometimes photographed by only one of
the two cameras operating at a sampling station. The event
of capturing an individual, whether it was photographed by
the two camera traps or only by one of them, was considered
a record of that animal. On rare occasions, an individual
was captured more than once in a sampling station during a
short period of time (o1 h), and to avoid pseudoreplication
we only considered the first capture of that animal as a
record. In some of the analyses (e.g. ocelot activity in
relation to moonlight), we used only the records obtained
during the survey period to keep the sampling effort constant. In other analyses (when no sampling bias was possible
as a result of trapping effort, e.g. comparison of male and
female activity patterns), we used all the ocelot records
obtained during the study, including both the preliminary
and post-survey periods.
Following Batschelet (1981), we used circular statistical
analyses for temporal data that follow a cycle. We used the
Rayleigh test (Batschelet, 1981) to test whether ocelot
captures were randomly or uniformly distributed along a
lunar cycle. We used Kuiper’s test to test whether the daily
frequency distributions of captures of two different samples
(e.g. males vs. females) have the same distribution (Batschelet, 1981). We performed the non-circular statistical tests
with the program JMPs (version 3.2). All statistical tests are
two tailed and with an a level of 0.05 for statistical

Ocelot abundance and population
At Urugua- ´ı we recorded 17 individuals (Table 1) from an
estimated population of 20 3.43 individuals in the sampled
area (Table 2). At Iguazu´ we captured twice that number of
individuals from an estimated adult population of
55 11.02 individuals.
Females outnumbered males at both sites (Table 1). The
M:F sex ratio of our ocelot population (both sites combined,
15 males, 29 females) is different from the sex ratio found at
the Kaa-Iya del Gran Chaco National Park of Bolivia
(Maffei et al., 2005; four study sites combined, 79 males,
68 females, likelihood ratio test of independence, w2 = 5.31,
P = 0.021).

Table 1 Total number of ocelot Leopardus pardalis records and number of individuals captured according to their sex during the survey period
at Iguazu´ and Urugua-´ı
Study site

Trap days







Sex ratiob










These individuals were the immature daughters of two of the adult females, as indicated by their joint presence in photographic records and
overlap in their observed ranges.
Adult sex ratio (M/F).


c 2006 The Authors. Journal compilation
c 2006 The Zoological Society of London
Journal of Zoology 270 (2006) 153–163

M. S. Di Bitetti, A. Paviolo and C. De Angelo

Density and habitat use of ocelots in NE Argentina

Table 2 Population estimates provided by CAPTURE for the two study sites using model Mh (jackknife) and three different effective areas
sampled and density estimates for each site (depending on the method used to estimate the buffer)



( SE)




20 3.43




55 11.02


Method used to
estimate buffer
and density

MMDMd or
radius of HRe
( SE)(km)

Areaf ( SE)

( SE)

1/2 MMDM
1/2 MMDM

3.96 0.63
3.96 0.63
2.63 0.28
3.95 0.64
3.95 0.64
2.63 0.28

150 13.6
259 17.9
184 4.8
275 18.9
428 23.6
327 8.8

13.36 2.60
7.71 1.43
10.88 1.89
19.99 4.23
12.84 2.67
16.80 3.40


Individual estimated capture probability (per trapping occasion) for model Mh.
Population (number of individuals present in the study area) estimate ( SE).
Approximate 95% confidence interval for population estimate.
Mean maximum distance of recapture (MMDM) ( SE) for individuals captured at two or more sampling stations. Sample sizes are n = 7
individuals for Urugua-´ı and n = 15 individuals for Iguazu´.
Home range estimates based on Crawshaw (1995) using the minimum convex polygon method and not including subadults.
Estimate of the area sampled ( SE) in km2 after applying a buffer.
Density estimate (individuals 100 km 2).

Despite differences in the mean distance among nearest
camera stations between sites, there was no difference in the
MMDM (and SE) between Urugua- ´ı (3.96 0.63 km, n = 7)
and Iguazu´ (3.95 0.64 km, n = 15) (Table 2; ANOVA
based on the ln of MDM, F1,20 = 0.087, P= 0.77). Thus,
we applied basically the same buffer (either 1/2 MMDM,
MMDM or the radius of the mean home range estimated by
radiotelemetry) at both sites. The mean radius of the home
range estimates based on a radiotelemetry study (Crawshaw,
1995) was larger than the mean 1/2 MMDM (ANOVA
using the ln of MDM or radius of HR, F1,33 = 4.05,
P= 0.05).
The density estimate ( SE) for the Urugua- ´ı study area
using model Mh (jackknife) range from 7.71 1.43 to
13.36 2.6 individuals 100 km 2, depending on the buffer applied to the camera stations (Table 2). The density
estimates for the adult population of ocelots at Iguazu´ were
higher, ranging from 12.84 2.67 to 19.99 4.23 individuals 100 km 2. The density estimates using as a
buffer the radius of the home range estimates from Crawshaw (1995) lay within the estimates we obtained using
1/2 MMDM and MMDM as a buffer (Table 2).
The density estimate of 14 individuals 100 km 2 obtained by Crawshaw (1995) for Iguazu´ using radiotelemetry
lay within the range of our density estimates. The density
estimates for ocelots in the Upper Parana´ Atlantic Forest
are generally lower (but not statistically) than estimates
from other sites, even when using our larger density estimates based on 1/2 MMDM (Fig. 2; Wilcoxon two-sample
test, S= 10.5, Z= 1.693, P= 0.090).

Spatial patterns of habitat use
We performed a three-way factorial ANOVA to test for the
effect of (1) study site (Iguazu´ vs. Urugua- ´ı), (2) effect of a
road versus a trail and (3) degree of protection of the area
(whether the sampling station was located within a pro-

Density estimates (indiv. 100 km−2)











Other sites
Study area
Figure 2 Density estimates for ocelots Leopardus pardalis at the
Upper Parana´ Atlantic Forest (UPAF) and other neotropical study
sites. Data sources: 1, Trolle & Ke´ry (2005); 2–5, 8, Maffei et al.
(2005); 6, Ludlow & Sunquist (1987); Sunquist et al. (1989); 7, Trolle &
Ke´ry (2003); 9, Crawshaw & Quigley, in Crawshaw (1995);
10, Emmons (1988); 11, 13, this study; 12, Crawshaw (1995).

tected area or in an area under forest exploitation) on ocelot
capture rates at each camera station (number of captures
during the survey/number of days the camera station was
active during the survey). The whole model test explained
almost 2/3 of the variation (r2 = 0.61, F3,69 = 35.49,
Po0.0001). Capture rates at Iguazu´ were not higher than
at the Urugua- ´ı study site (F1,69 = 1.36, P= 0.25). Capture
rates were higher on camera stations located on roads than
on trails (F1,69 = 77.69, Po0.0001) and were higher in the
protected areas than in areas with pine plantations
(F1,69 = 23.26, Po0.0001). At both study sites we found no
effect of the forest cover and the abundance of bamboos on
the ocelot capture rate, either alone or after controlling for
the effect of other variables in factorial analyses.

c 2006 The Authors. Journal compilation
c 2006 The Zoological Society of London
Journal of Zoology 270 (2006) 153–163


Density and habitat use of ocelots in NE Argentina



Relative frequency of captures


Home range estimate (km2)

M. S. Di Bitetti, A. Paviolo and C. De Angelo



Camera traps

Figure 3 Home range estimates ( SE) for ocelots Leopardus pardalis.
First two bars, radiotelemetry estimates for males and females using
the minimum convex polygon method (from Crawshaw, 1995,
subadults not included); last two bars, our camera-trap minimum
observed range estimates for the males and females recorded at
more than three different camera stations at our study sites.

Males (n= 87)
Females (n =138)


13 14 15 16 17 18 19 20 21 22 23 0 1 2 3 4 5 6 7 8 9 10 11 12

Figure 4 Daily activity patterns of male and female ocelots Leopardus
pardalis at Iguazu´ National Park for the period May–December 2004.


For those individuals observed at four or more sampling
stations at both sites (three females and four males), we
estimated a minimum observed range using the minimum
convex polygon method. The mean ( SEM) home range size
for females was 6.01 0.92 km2 (range 4.18–7.11 km2). The
mean ( SEM) home range size for males was
13.41 8.00 km2 (range 3.19–37.09 km2). A two-way ANOVA using home range size estimates as the dependent
variable and sex and study method [our camera-trap minimum observed range estimates vs. those obtained by Crawshaw (1995, data from his table 4.2, excluding subadults)
using radio tracking and the minimum convex polygon
method] as the independent variables indicates that males
have larger territories than females (F1,18 = 5.574,
P= 0.0297) and that our home range estimates are lower
than those obtained using radiotracking data (F1,18 = 5.897,
P= 0.0259; Fig. 3).

Temporal patterns of habitat use
Both at Iguazu´ and at Urugua- ´ı, ocelots were captured more
frequently at night than during the day. Males seem to have
two peaks of activity during the night, one before midnight
and another just before sunrise (Fig. 4). However, males and
females did not differ in their daily activity patterns (Kuiper’s test grouping time of capture into 96 intervals
of 15 min each, each interval = 3.751, k = 1734, n1 = 87,
n2 = 138, P40.10).
Both at Urugua- ´ı and at Iguazu´, the frequency of ocelot
captures was not uniform or random during the lunar cycle
but increased during the week prior to and during the new
moon lunar phase, a period when light is at a minimum on
the forest floor (Fig. 5; data from Iguazu´ and Urugua- ´ı
combined, Rayleigh test, r = 0.176, Z =6.013, Po0.01,
n= 195). In a circular distribution of 29 days (the lunar
cycle lasts 29 or 30 days, but we omitted day 30 to make the

Frequency of captures


Day of lunar cycle (starting at full moon)


Figure 5 The frequency of ocelot Leopardus pardalis captures varies
along the lunar cycle. Six complete lunar cycles, three from each
study site.

three cycles at each site equal in length), the concentration
parameter is located 3–4 days before new moon. Ocelots
tend to be more active during the day and start their activity
later on days with a full moon, but the frequency distribution of daily captures is not statistically different from the
one observed on days with a new moon (Kuiper’s test,
96 intervals of 15 min each, each interval= 3.751, k = 2154,
n1 = 114, n2 = 147, P40.10; Fig. 6).

Density estimates
Density estimates for ocelots at Iguazu´ and Urugua- ´ı (near
the southern edge of the ocelot’s range in South America)
are low compared with those of other neotropical sites.
Tewes (1986) noted that home range size of ocelots in Texas
is larger than those of populations living at lower latitudes,
which is consistent with the general pattern found in other

c 2006 The Authors. Journal compilation
c 2006 The Zoological Society of London
Journal of Zoology 270 (2006) 153–163

M. S. Di Bitetti, A. Paviolo and C. De Angelo

Relative frequency of captures


New moon (n =147)
Full moon (n =114)


131415161718192021222324 1 2 3 4 5 6 7 8 9 101112

Hour (starting at 12:00 h noon)
Figure 6 Daily activity patterns of ocelots Leopardus pardalis on new
moon and full moon nights. New moon days are days 1–6 and 23–29
or 30 of the lunar cycle that starts on the new moon. Full moon days
are days 9–20 of the same cycle. Transitional days (days 7–8 and
21–22) were omitted.

carnivores (Gompper & Gittleman, 2001). Almost by definition home range size and density are negatively correlated in
within-species comparisons of territorial carnivores (e.g. Karanth & Chundawat, 2002), and both are correlated to prey
density (e.g. tiger densities, Karanth & Nichols, 1998;
carnivore densities, Carbone & Gittleman, 2002; home
range size, Herfindal et al., 2005). Thus, our lower ocelot
density estimates and the larger home ranges of North
American ocelots are consistent with these general trends
of lower ocelot densities and larger home ranges at both
extremes of their distribution. However, there is a large
variation in density estimates among study sites even within
regions (e.g. Pantanal, Trolle & Ke´ry, 2005). Maffei et al.
(2005) found no relationship between rainfall and ocelot
densities in a comparison of several study sites across the
neotropics. This suggests that rainfall alone may not be a
good indicator of ocelot prey availability. We are still
lacking reliable estimates of prey abundance at the sites
where ocelots have been studied to make further conclusions
on the relationship between ocelot territory size, population
density and food availability.
From a top-down perspective, ocelot density may also be
affected by the abundance of their predators. Even though
jaguars have suffered a recent population decline at our
study sites, their lower numbers may have been compensated by an increase in the abundance of pumas (M. S. Di
Bitetti, unpubl. data). Crawshaw’s (1995) ocelot density
estimate, although based on a different methodology, is very
similar to ours, which suggests that ocelot density have
remained stable within the last 15 years in the Iguazu´ area.

Habitat use
Ocelots are present in all the different forest communities at
Iguazu´ National Park and at Urugua- ´ı, including mosaic
habitats where the native forest has been partially replaced

Density and habitat use of ocelots in NE Argentina

by pine plantations. The distribution of ocelots throughout
their range in the neotropics coincides with the structurally
closed habitats of tropical and subtropical forests and dense
thorn shrubs (Brown, 1989; Shindle, 1995; Emmons & Feer,
1997; Murray & Gardner, 1997; Lopez
Brown &
Gallo Reynoso, 2003). At a finer scale, the canopy or the
understory structural characteristics of the locations where
the camera stations were deployed did not affect ocelot
capture rate.
Several authors have noted the preference of big cats
for walking on roads or trails instead of through the forest
(e.g. Emmons, 1988). Carbone et al. (2001) noticed that
camera-trap capture rates of tigers are much higher than
those expected under a simple random walk model, indicating that researchers place their camera traps at places where
tiger captures are maximized. At Iguazu´ ocelots have a
much higher capture rate on old dirt roads than on trails
opened with a machete. A similar pattern was found for
jaguars in the Kaa-Iya National Park of Bolivia (Maffei
et al., 2004) and ocelots in Pantanal (Trolle & Ke´ry, 2005).
This difference may be the result of old roads being traditional animal routes or even becoming landmarks that
animals frequently use to demarcate their territories
(e.g. jaguars, Rabinowitz & Nottingham, 1986).
In the areas with pine plantations capture rates were low
compared with the protected areas, even though most
camera stations in the first were located on old roads rather
than on trails. We do not find a logical explanation for this
reduction in capture rates in areas with pine plantations
other than a lower ocelot abundance in this portion of the
study site, probably as a result of natural habitat loss.
Females outnumbered males by about two to one in our
study population. Our adult sex ratio is correlated with the
larger home range sizes of males (Fig. 3), which are usually
two to four times larger in male than in female ocelots
(Tewes, 1986; Ludlow & Sunquist, 1987; Emmons, 1988;
Konecny, 1989; Sunquist et al., 1989). That adult males have
larger territories than females is the usual pattern for
solitary territorial cats (Sunquist & Sunquist, 2002), with
few exceptions reported for some populations in the literature (e.g. Canadian lynxes Lynx canadensis; see the review in
Poole, 2003). This pattern reflects the common social system
of solitary cats, where there is intra-sexual territoriality:
males and females evict any same-sex adult individual from
their own territories (Kitchener, 1991; Poole, 2003). The
difference between the female-biased sex ratio of our study
population and the more even (or male biased) sex ratio
reported by Maffei et al. (2005) for the ocelot populations of
the dry forests of Bolivia is suggestive of differences in the
social system of ocelots between sites.

Activity patterns
Several authors have previously noted that ocelots are more
active at night than during the day, as was the case in our
study, but the degree of nocturnality varies from study to
study. Emmons (1988) noted that males and females have a
bimodal activity pattern at night, with a peak of activity

c 2006 The Authors. Journal compilation
c 2006 The Zoological Society of London
Journal of Zoology 270 (2006) 153–163


Density and habitat use of ocelots in NE Argentina

before midnight and another a few hours later. At Iguazu´
males seem to have a bimodal pattern similar to the one
described by Emmons, but females do not. However, the
activity patterns of males and females at Iguazu´ are not
statistically different.
Some of the differences observed in activity patterns
among sites could result from the different methodologies
used to study ocelots. Crawshaw (1995), using radiotelemetry, found smaller differences in activity levels between night
(41% of the readings) and day (34% of the readings) for
ocelots at Iguazu´ National Park. Our camera-trap data
show a pattern much more similar to that found for ocelots
in Manu by Emmons (1988) or to that described for the
Venezuelan Llanos by Ludlow & Sunquist (1987). The
difference between Crawshaw’s study and ours could reflect
the fact that animals could be active but not moving during
the day (giving a signal that indicates ‘activity’ to the
receiver) or that animals may walk relatively less frequently
on trails and roads during the day than at night.
Emmons et al. (1989) reported that ocelots use trails less
frequently during the week previous to full moon and during
peak full moon, a behavior that the authors interpreted as
an avoidance of open areas when there is more light in the
forest floor. Maffei et al. (2005) did not find any relationship
between the frequency of use of trails and roads and the
lunar cycle. Our results coincide with the results of Emmons
et al. (1989): ocelots at Iguazu´ also use the roads and trails
more frequently on new moon nights than on full moon.
Emmons et al. (1989) suggested that ocelots avoid open
areas when there is more light on the forest floor because
they may become more visible and thus less successful
hunting small prey or more vulnerable to the larger felids
that prey on them. This change in behavior associated with
the lunar phases is not the consequence of ocelots becoming
less active with full moon, but probably a general preference
for those portions of their habitat with lower light levels (see
Shindle, 1995). Emmons et al. (1989), using radiotelemetry
data, showed that ocelot activity levels did not change with
the lunar phases, only their preference to walk on trails. The
ocelot main prey in Manu, the spiny rats in the genus
Proechimys, become more active and visible on trails on
new moon nights, probably because they can more easily
avoid predation attempts from ocelots (Emmons
et al., 1989). We do not have information on the activity
pattern of ocelots’ small prey species at Iguazu´, but some of
the most important large and medium-size nocturnal prey
items in the ocelot’s diet (opossums Didelphis sp., armadillos
Dasypus novemcinctus, Brazilian rabbit Sylvilagus brasiliensis; Crawshaw, 1995) show no change in activity or they may
even increase their capture rates on roads and trails on full
moon nights (M. S. Di Bitetti et al., unpubl. data), showing a
pattern opposite to that observed for spiny rats in Manu.

Camera traps as a methodology to study the
natural history of wild animals
Camera trapping is becoming a standard methodology to
study wild animal populations (Karanth & Nichols, 1998;

M. S. Di Bitetti, A. Paviolo and C. De Angelo

Carbone et al., 2001; Lynam, 2002; Karanth, Nichols &
Cullen, 2003; Moraes Tomas & de Miranda, 2003; Silver
et al., 2004). Camera traps can be used to study aspects of
the natural history of wild animals other than abundance or
density. For example, they can provide important information on the seasonal and daily activity patterns (Moraes
Tomas & de Miranda, 2003; Maffei et al. 2005; this study).
Camera traps may also provide information on reproductive
and social behavior of animals if sampling effort is large
enough or if the study animals are relatively abundant.
During our survey we gathered few data on the reproductive
conditions of females. We believe that with long-term longitudinal studies it should be possible to get a good picture of
the reproductive patterns of ocelots.
Camera trapping is probably not the best methodology to
get good estimates of home range size and ranging patterns
of wild animals, unless sampling effort is very high and
sampling stations are very dense or they are constantly
shifted to new places. Our minimum observed ranges for
ocelots are clearly an underestimate of actual home range
sizes, because they are based on very few data points.
Estimates by Crawshaw (1995) are more accurate than ours,
and indicate that home ranges of ocelots in the Upper
Parana´ Atlantic Forest are much larger than those of ocelots
at other sites, which correlates with the low-density estimates of ocelots in our study site. However, at least in some
places (the dry forests of eastern Bolivia), minimum observed range estimates obtained using camera traps are
similar to home range estimates obtained using radiotelemetry (A. Noss et al., pers. comm.). Data from other
sites where radiotelemetry and camera trapping are conducted simultaneously are needed to understand, among
other things, (1) the relationship between home range
size and the size of the minimum convex polygons derived
from camera trapping, (2) what is the best buffer to apply to
the camera stations to estimate the effectively sampled area
(1/2 MMDM, MMDM or some value in between)
and (3) if there is a relationship between MMDM and the
distance between camera stations in the grids (not found
in our study but suggested by recent camera-trap
studies; M. Kelly, pers. comm.). For example, we found
that the mean radius of Crawshaw’s (1995) home range
estimates is larger than the mean 1/2 MMDM, despite
the latter being used as a proxy for the first one when
estimating densities with camera-trapping data (but see
Trolle & Ke´ry, 2005).

Considerations on the conservation of
ocelots and other felids in the Upper Parana´
Atlantic Forest
Ocelots are probably the most abundant of the wild cats in
most of the neotropical forests (Emmons & Feer, 1997).
There are very few assessments of the populations of other
small cats in this region to corroborate this assertion, but
ocelot densities are clearly higher than those of pumas and
jaguars across most of their shared range (Emmons, 1988;
Maffei et al., 2004). What can our ocelot density estimates

c 2006 The Authors. Journal compilation
c 2006 The Zoological Society of London
Journal of Zoology 270 (2006) 153–163

M. S. Di Bitetti, A. Paviolo and C. De Angelo

tell us about the population status and conservation needs
of wild cats in the Upper Parana´ Atlantic Forest?
With a mean population density of 13.60 individuals 100 km 2, we can extrapolate numbers to the portion of the Green Corridor of Misiones and the contiguous
areas of Brazil that still contain continuous native forest
cover (about 9400 km2). This is the only large (41000 km2)
continuous forest fragment of the Upper Parana´ Atlantic
Forest ecoregion remaining on earth. Assuming densities on
our grids were representative of the entire area, we estimate
a population of about 1280 ocelots for this area (range
724–1879). The Green Corridor is being continuously encroached by poor people practising slash-and-burn agriculture or by timber companies seeking land for large-scale
pine plantations (Chebez & Hilgert, 2003; Holz & Placci,
2003). If what is left of the Green Corridor within the next
few decades is just the area strictly protected at the present
time, the available habitat for ocelots will be reduced to less
than 5000 km2, and only a portion of this area (3501.65 km2)
constitute a continuous block of strictly protected areas
(Iguazu´ National Park of Brazil, Iguazu´ National Park of
Argentina and Urugua- ´ı Provincial Park, plus other small
contiguous protected areas; see Giraudo et al., 2003). Thus,
if conservation measures are not implemented soon
(e.g. consolidation of the Yabot ´ı Biosphere Reserve as a
protected area and the creation of new protected areas
within the Green Corridor), this area will sustain, at most,
a population of between 500 and 700 adult ocelots in the
near future.
Recent comparative analyses of the extinction risk of
terrestrial vertebrate populations (Reed et al., 2003) indicate
that only populations above 5000 individuals are safe from
extinction (less than 99% probability) in the future
(40 generations). Our ocelot population estimate for the
Green Corridor is below this number. Even though ocelots
are not at a high risk of local extinction in the near future,
their population numbers and the high rate of habitat destruction should make us pay more attention to the population
status of sympatric felids that are less known than the ocelot
(the margay Leopardus wiedii, the oncilla Leopardus tigrinus
and the jaguarundi Herpailurus yaguarondi) or that live at
much lower population densities (the jaguar and the puma).
The area needed to protect a viable population of ocelots in
the Upper Parana´ Atlantic Forest ecoregion is of the same
magnitude of a protected area designed to protect jaguars in
other areas (Maffei et al., 2004). The ocelot is relatively easy to
study (to capture in live traps or camera traps) compared with
the other wild cats, makes a good population model for other
neotropical felids and may help guide conservation efforts
(e.g. habitat restoration) in this region.

We are grateful to all the students and park rangers who
helped us with the field activities, especially Esteban Pizzio,
Ricardo Melzew, Ilaria Agostini, Ingrid Holzmann, Jenny
Glikman, Fernanda Fabbio, Santiago Escobar, Jessica Pra-

Density and habitat use of ocelots in NE Argentina

da, Georgina Lanza, Yamil Di Blanco, Carolina Ferrari and
´ Vida Silvestre
Peonia Britos. We acknowledge Fundacion
Argentina, and especially their park manager Andre´s Johnson, for support and permission to conduct this study at the
Urugua- ´ı Biological Station. We are thankful to the National Parks Administration of Argentina and Karina
Schiaffino for permission to conduct fieldwork at Iguazu´
National Park and for the use of the Park’s Field Station
(CIES). We are also thankful to the Ministry of Ecology,
Natural Resources and Tourism of Misiones province for
permission to conduct fieldwork at Urugua- ´ı Provincial
Park and to Alto Parana´ S. A. for permission to work in
their property. Andrew Noss, Leonardo Maffei and Peter
Crawshaw made available unpublished data. Financial sup´
port for this project was provided by CONICET, Fundacion
Vida Silvestre Argentina, World Wildlife Fund-USA,
´ AnWWF-International, Lincoln Park Zoo, Fundacion
torchas, Wildlife Conservation Society and Idea Wild.
Isabella Levi kindly helped us scan photographs. We thank
Kathleen Conforti and Catherine Grippo (WCS) and Charlie Janson (SUNY at Stony Brook) for providing assistance
in finding bibliography. Andrew Noss, Marcella Kelly and
an anonymous reviewer made useful comments on this
manuscript. We are thankful to Tuto, Horacio and Mart ´ın
from Laboratorio Rolando for their invaluable help with
film processing.

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c 2006 The Authors. Journal compilation
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