Critical condition of the jaguar Panthera onca population in Corcovado National Park, Costa Rica

Abstract

The jaguar Pantheraonca is threatened throughout its range and categorized as Near Threatened on the IUCN Red List. To inform conservation of the jaguar population in Corcovado National Park, Costa Rica, population size was estimated using data from a 3-month camera trap study. Individuals were identified from their coat patterns. The resulting density estimate of 6.98 ± SD 2.36 individuals per 100 km2 was lower than expected. The sex ratio was 1.33 males per female, and the minimum home ranges of two males were 25.64 and 6.57 km2. Hunting pressure on jaguar and white-lipped peccaries Tayassupecari, the jaguar's main prey in the Park, may be responsible for the low jaguar density as space does not seem to be a limiting factor. The numbers of females may have been underestimated because of sampling bias and therefore the sex ratio obtained in this and similar studies must be interpreted cautiously. Better protection of the corridor that connects the Park with other protected areas is essential to guarantee long-term survival of the jaguar in Costa Rica.

Full text

Critical condition of the jaguar Panthera onca population in Corcovado
National Park, Costa Rica
Roberto Salom-Pe
´rez, Eduardo Carrillo, Joel C. Sa
´enz and Jose
´M. Mora
Abstract The jaguar Panthera onca is threatened
throughout its range and categorized as Near Threat-
ened on the IUCN Red List. To inform conservation of
the jaguar population in Corcovado National Park,
Costa Rica, population size was estimated using data
from a 3-month camera trap study. Individuals were
identified from their coat patterns. The resulting density
estimate of 6.98 ¡SD 2.36 individuals per 100 km
2
was
lower than expected. The sex ratio was 1.33 males per
female, and the minimum home ranges of two males
were 25.64 and 6.57 km
2
. Hunting pressure on jaguar
and white-lipped peccaries Tayassu pecari, the jaguar’s
main prey in the Park, may be responsible for the low
jaguar density as space does not seem to be a limiting
factor. The numbers of females may have been under-
estimated because of sampling bias and therefore the sex
ratio obtained in this and similar studies must be
interpreted cautiously. Better protection of the corridor
that connects the Park with other protected areas is
essential to guarantee long-term survival of the jaguar in
Costa Rica.
Keywords Abundance, camera traps, Corcovado
National Park, Costa Rica, jaguar, Panthera onca.
Introduction
Jaguar Panthera onca, categorized as Near Threatened on
the IUCN Red List (IUCN, 2006), are found throughout
the Americas, ranging from northern Mexico to south-
ern Argentina (Seymour, 1989; Sanderson et al., 2002b).
There has, however, been a decrease in the number of
their prey and increased fragmentation of their natural
habitat, and so-called problem jaguars accused of
preying on livestock have been killed (Swank & Teer,
1989; Emmons, 1990; Sa
´enz & Carrillo, 2002; Sanderson
et al., 2002b). Only 4% of the most important areas
for jaguar are currently being protected effectively
(Sanderson et al., 2002b) and Costa Rica is one of the
countries where, because of habitat loss and hunting,
the jaguar is most threatened (Swank & Teer, 1989;
Sanderson et al., 2002a). Although forested areas large
enough to support 500 or more jaguars may no longer
exist in Central America (Emmons, 1990; Ceballos et al.,
2002; Maffei et al., 2004) connections between popula-
tions living in distinct areas could help to guarantee the
survival of the species in the long term (Shaffer, 1989;
Swank & Teer, 1991).
Camera traps have been used to estimate populations
of tiger Panthera tigris in India (Karanth & Nichols, 2000;
Carbone et al., 2001) and are also now being used
with jaguar and other felids in the Neotropics (Tro
´lle &
Ke
´ry, 2003; Sarmiento, 2004; Silver et al., 2004). The
present study is one of the first investigations in
Central America to use this methodology for jaguar.
Our objective was to estimate the population size and
examine the conservation status of jaguar in Corcovado
National Park.
Study site
The 425 km
2
Corcovado National Park on the Osa
Peninsula of the Pacific coast of Costa Rica borders the
Guaymı
´Indigenous Reserve and the Golfo Dulce Forest
Reserve. A portion of the latter forms a corridor that
connects the Park with Piedras Blancas National Park
and Golfito National Wildlife Refuge (Fig. 1). The
altitude of the Park is 0–745 m, annual maximum and
minimum temperatures for the nearest weather station
are 31.7 and 22.1uC, respectively, and mean precipitation
is 4,656.5 ¡SD 43.8 mm, one of the highest in Costa
Rica. The Park has a rich and diverse flora and fauna
and a relatively large number of endemic species
(Hartshorn, 1983; Soto, 1994; Naranjo, 1995). This and
other forests on the Osa Peninsula are the last of
the tropical rainforests on the Pacific side of Central
America (Hartshorn, 1983).
Roberto Salom-Pe
´rez (Corresponding author) P.O. Box 350-2300,
Curridabat, San Jose
´, Costa Rica. E-mail robersalom@yahoo.com
Eduardo Carrillo and Joel C. Sa
´enz Programa Regional en Manejo de Vida
Silvestre, Universidad Nacional, P.O. Box 1350-3000 Heredia, Costa Rica.
Jose
´M. Mora Escuela de Biologı´a, Universidad de Costa Rica, Costa Rica.
Received 27 September 2005. Revision requested 17 January 2006.
Accepted 7 June 2006.
Oryx Vol 41 No 1 January 2007
ß2007 FFI, Oryx,41(1), 51–56 doi:10.1017/S0030605307001615 Printed in the United Kingdom 51

Methods
A pilot project was carried out in Corcovado National
Park from August 2002 to January 2003 with five camera
trap stations, each with two cameras sensitive to heat
and motion (CamTrakker, Watkinsville, USA). The data
from this was not used to determine jaguar population
size but was used for other calculations (as explained
below). For the full study 12 trap stations, each also with
two cameras, were run from 19 January to 18 April 2003.
Because of robbery or failure of the cameras we were
only able to use the information from 11 stations. We
divided the trap stations into two blocks to be able to use
only the days in which all trap stations were active
continuously in each block. Block 1 consisted of traps
1–6 (data collected 21 January–22 February) and Block
2 of traps 7–11 (data collected 23 February–27 March).
Based on a previous radio telemetry study the
minimum home range for a female jaguar in the Park
is 12 km
2
(Carrillo, 2000) and therefore we placed at least
one trap station in every 12 km
2
circular area to ensure
that all individuals had a probability .0.0 of being
photographed. The two cameras at each trap were
focused at the same spot but not at each other, to avoid
flash interference, and were 0.5 m above the ground and
2–4 m from the centre of the trail. The cameras were
active 24 hours per day and the minimum interval
between photographic events was set to 25 minutes.
Trap stations were located in areas where signs of
felid or other mammal activity had been observed
(based on photos from the pilot project, faeces and
tracks). The cameras were checked c. every 15 days to
change film and ensure they were functioning correctly.
Every photograph obtained of a jaguar in a sampling
occasion is equivalent to one capture. Photos of the same
individual in successive sampling occasions were
considered recaptures. Individuals were identified by
their fur pattern, which is unique to each jaguar (Silver
et al., 2004).
The area covered by the study was calculated by
drawing a polygon, which we refer to as the perimeter
area, whose vertices were formed by the outermost trap
stations. A buffer area was added to the polygon in the
form of a band to determine the total area covered by
the study (Fig. 1). The width of this band was half of the
52
Fig. 1 Protected areas of the Osa Peninsula, Costa Rica, with perimeter and buffer areas (see text for details) and location of the camera-trap
stations (numbered) used to estimate the jaguar population within Corcovado National Park (in light grey).The inset shows the location of
the Osa Peninsula in Costa Rica.
R. Salom-Pe
´rez et al.
ß2007 FFI, Oryx,41(1), 51–56

mean maximum distance moved (MMDM) by indivi-
duals photographed more than once during the 3-month
study period (Wilson & Anderson, 1985; Karanth &
Nichols, 1998). To improve the calculation of MMDM
we also used the maximum distance moved by six
jaguars monitored in the Park with radio telemetry
during January–March of 1996–98 (E. Carrillo, unpubl.
data; Table 1).
To estimate the jaguar population and the average
probability of a capture per sampling occasion (P
ˆ)we
used the software CAPTURE (Otis et al., 1978; Rexstad &
Burnham, 1991; Karanth & Nichols, 1998). One of the
assumptions of CAPTURE is that the study population is
closed, i.e. individuals do not enter or leave the area. We
considered the study period of 3 months short enough to
be certain that this assumption held. Nevertheless,
CAPTURE also tests for closure. Every sampling occasion
was set to 3 days. Because of differences in mobility
between male and female and between adult and juvenile
jaguar it cannot be assumed that there is no variation in
the probability of capture of individuals. We therefore
used the heterogeneity estimator in CAPTURE. The
population estimate and the size of the study area were
used to calculate jaguar density.
To determine the sex ratio, activity patterns, and the
minimum home range of jaguar we used data from this
3-month study, the pilot project, and a parallel study of
sea turtle predation by jaguar carried out within the
limits of the study area (September 2002–June 2003;
Salom-Pe
´rez, 2005). Minimum home ranges were esti-
mated as the minimum convex polygon. To calculate the
perimeter and buffer areas we used the geographical
information system ArcView v. 3.2 (ESRI, Redlands,
California). Distances between trap stations were
obtained using a global positioning system.
Results
Average distance between consecutive trap stations was
2.75 ¡SD 0.67 km (range 1.10 – 3.64 km), i.e. there was
no area .10.41 km
2
without a camera (mean of one
camera per 7.82 km
2
; Fig. 1). In a total of 363 trap nights
four individual jaguars were photographed (Table 2).
The total number of captures (including recaptures)
used for the population estimate was seven, and P
ˆwas
0.11. The closure test of CAPTURE indicated that the
population was closed (z52.13, P 50.98) and the
population estimate was 6.0 ¡SE 1.96 individuals (95%
confidence interval 5–14). All four jaguars photographed
were recaptured during the 3-month study period
(Table 2). The MMDM used to calculate the buffer area
was 3.48 ¡SD 0.47 km. The total area (perimeter area of
29.46 km
2
+buffer area of 56.55 km
2
) studied was 86.02
53
Table 1 Maximum distance moved (MDM) by 10 jaguars in Corcovado National Park. These distances were used in the calculation of the
total area covered by the study (see text for further details).
Jaguar MDM (km) Gender Source
jm1 3.17 Male This study
jm2 5.21 Male This study
jm3 3.81 Male This study
jh1 0 Female This study
rjm1 3.1 Male Radio telemetry (E. Carrillo, unpubl. data)
rjm2 5.2 Male Radio telemetry (E. Carrillo, unpubl. data)
rjh1 2.8 Female Radio telemetry (E. Carrillo, unpubl. data)
rjh2 3.7 Female Radio telemetry (E Carrillo, unpubl. data)
rjh3 3.3 Female Radio telemetry (E. Carrillo, unpubl. data)
rjh4 4.5 Female Radio telemetry (E. Carrillo, unpubl. data)
Table 2 Camera-trap capture history of jaguars in Corcovado
National Park, 2002–2003.
Jaguar
1
Date Time Trap station
2
Jm1 8/9/2002 0.35 Pilot study 5
Jm1 3/10/2002 23.38 Pilot study 4
Jm1 23/2/2003 7.49 10
Jm1 25/3/2003 4.20 10
Jm1 6/4/2003 17.42 8
Jm2 13/9/2002 22.38 Pilot study 1
Jm2 26/1/2003 8.14 4
Jm2 4/2/2003 2.52 1
Jm2 14/2/2003 23.42 4
Jm2 18/4/2003 4.24 4
Jm2 18/4/2003 5.31 3
Jm3 14/3/2003 1.57 7
Jm3 9/4/2003 15.07 10
Jm4 22/8/2002 19.55 Pilot study 3
Jh1 8/3/2003 10.58 8
Jh1 18/4/2003 9.19 8
Jh2 14–16/10/2002 Turtle study 3
Jh2 10/11/2002 21.44 Turtle study 5
Jh3 2/9/2002 Turtle study 2
1
Only the records in italics were used for the population estimate
(see text for further details).
2
In this study (see numbered locations in Fig. 1) and in an earlier
pilot study and a study of jaguar predation of turtles (see text for
details)
Jaguar in Corcovado, Costa Rica
ß2007 FFI, Oryx,41(1), 51–56

¡SD 7.75 km
2
(Fig. 1), giving a jaguar density of 6.98 ¡
SD 2.36 individuals per 100 km
2
.
In the pilot project and the parallel study of jaguar
predation on turtles two additional female (jh2 and jh3)
and one juvenile male (jm4) jaguar were photographed
(Table 2). The total of seven jaguars therefore gave a
1.33 male/female sex ratio. Of the 17 photographs in
which the time could be clearly determined, 11 (64.7%)
were during the night (18.00–6.00) and the others
(35.3%) during the day (6.00–18.00; Table 2). Mean home
range could only be determined for the two jaguars,
both males (jm1 and jm2), that were photographed
at .two trap stations: 25.64 km
2
(jm1, 6 captures) and
6.57 km
2
(jm2, 6 captures).
Discussion
Although we only photographed four jaguars during
the 3 months of the study we know that there were at
least three more jaguars present (Table 2). In Bolivia
Wallace et al. (2003) and Maffei et al. (2004) also captured
different individuals while performing consecutive
studies in the same area within ,1 year. It is possible
that several jaguars share an area but at different
times. Presumably they detect other individuals by
the presence of faeces or other sign and move to other
parts of their home ranges (Rabinowitz & Nottingham,
1986). The occurrence of a localized resource (marine
turtles) that are relatively easy to predate could explain
this convergence of home ranges, at least for jaguars
captured near the beach.
The density of jaguar calculated in this study (6.98 ¡
SD 2.36 per 100 km
2
) is higher than that reported in a
number of other localities in South America and Mexico
(0.45–5.23 per 100 km
2
; Schaller & Crawshaw, 1980;
Quigley & Schaller, 1988; Aranda, 1991; Nu
´n
˜ez et al.,
2002; Wallace et al., 2003; Maffei et al., 2004; Silver et al.,
2004), but similar to that determined from a 3-year radio
telemetry study of jaguar in Calakmul, Mexico (Ceballos
et al., 2002), and similar to or less than that determined
from camera-trap studies (Silver et al., 2004) in Cerro
Cortado, Bolivia (5.11 ¡2.10 per 100 km
2
) and in two
sites in Belize (8.80 ¡2.25 per 100 km
2
and 7.48 ¡2.74
per 100 km
2
). The jaguar density in Corcovado National
Park was, however, lower than we expected. It was
previously thought that, because of the abundance of
prey and high habitat quality, the Park had the highest
density of jaguar in Costa Rica, and that the density was
greater than that found in Belize (E. Carrillo, pers. obs.).
As our study only encompassed 20% of the Park it is
possible that we underestimated density. However, the
similarity of the maximum distance moved by jaguars
monitored with radio telemetry (E. Carrillo, unpubl.
data) and in this camera-trap study suggest that the
movement patterns reported here, and hence our
estimate of density, are relatively accurate.
As an individual jaguar’s range is usually not
exclusive (e.g. male jaguar jm1 shared his home range
with male jm3 and female jh4) it is likely that Corcovado
National Park could accommodate a greater density
than that estimated (Schaller & Crawshaw, 1980;
Quigley & Schaller, 1988; Nu
´n
˜ez et al., 2002; Maffei
et al., 2004). As space does not appear to be a limiting
factor it is possible that food supply is limiting jaguar
density in the Park. This is supported by the fact that in
the past few years hunting has caused a considerable
decrease in the numbers of white-lipped peccaries
Tayassu pecari, the primary prey of the jaguar in the
Park (E. Carrillo, unpubl. data; Chinchilla, 1994; Carrillo,
2000). Jaguar activity depends mainly on the prey being
hunted (Carrillo, 2000) and as c. 65% of the jaguar
photographs were taken during the night this suggests
they were probably searching for alternative, nocturnal
prey such as marine turtles.
The limitations of our data do not allow us to
confidently extrapolate our density estimate to the
whole of Corcovado National Park. However, if jaguar
occur at a similar density throughout the Park the total
population would be c. 30 individuals, and even if the
Park could support the maximum density reported for
the species in Central America (8.80 ¡2.25 per 100 km
2
;
Silver et al., 2004) it would contain no more than 50
jaguars. If this population is reproductively isolated its
survival is threatened as it probably does not contain
sufficient individuals for a minimum viable population
(Eizirik et al., 2002). However, jaguars killed in areas
surrounding the Park and individuals photographed in
the corridor that connects the Park with Piedras Blancas
National Park and Golfito National Wildlife Refuge
(E. Carrillo, unpubl. data), are evidence that there is
probably movement of jaguars between the Park and
other protected areas. This indicates the importance of
upgrading the management level of Golfo Dulce Forest
Reserve, or at least a part of it, because a National Park
in Costa Rica provides better protection to large and
medium size mammals than that given by a Forest
Reserve (Carrillo et al., 2000).
As in other camera-trap studies of jaguar a greater
number of males were captured than females (Wallace
et al., 2003; Silver et al., 2004). The number of females
could, however, have been underestimated because
females have smaller territories and move less than the
males, and therefore have fewer opportunities to be
captured. In addition, females are known to be more
timid than males and are more likely to avoid walking
on man-made trails; six of the 12 trap stations were
located on man-made trails. Two females were photo-
graphed on the beach preying on turtles (jh2 and jh3, on
54 R. Salom-Pe
´rez et al.
ß2007 FFI, Oryx,41(1), 51–56

two and one occasions, respectively) and one was
photographed on a game trail (jh1, on two occasions),
but none of these individuals were photographed on
man-made trails.
The use of camera traps allows the calculation of more
accurate population estimates of felids and other
animals than earlier methods based on indirect signs
(Mondolfi & Hoogesteijn, 1991; Smallwood & Fitzhugh,
1993; Cutler & Swann, 1999; Grigione et al., 1999; Silveira
et al., 2003; Tro
´lle & Ke
´ry, 2003). Radio telemetry has
also been used to obtain data on home range size and
densities but, in addition to being expensive, has
problems related to topography, forest cover, data
collection and animal health (because of capture and
sedation; Rabinowitz & Nottingham, 1986; Mondolfi
& Hoogesteijn, 1991; Carrillo et al., 2000). However,
cameras are expensive (USD 90–400 per camera, plus the
price of the film and developing), must be checked
frequently, may be stolen, and may malfunction,
especially in extremely humid conditions (R. Salom-
Pe
´rez, pers. obs.; Maffei et al., 2004; Silver et al., 2004).
Our results emphasize the need to safeguard corridors
that connect protected areas in Costa Rica. Such
protection could include the possibility of modifying
the management categories of such prioritized areas. In
addition, more rigorous control of hunting and an
increase in environmental education in the communi-
ties neighbouring the protected areas are required.
Currently, RS-P, EC and others from various state
universities and the National Institute of Biodiversity
are carrying out investigations to facilitate delineation of
the corridor between Corcovado and Piedras National
Parks. Other governmental and non-governmental
organizations are resolving matters regarding land
possession, providing environmental education, and
supporting sustainable development programmes for
the area.
Acknowledgments
We thank Scott Silver for his guidance and valuable
comments. In addition we thank Daniela Araya, Juan de
Dios Valdez, Alexander Go
´mez, Alfonso Chamorro,
Carolina Orta, Nereyda Estrada, Marcela Ferna
´ndez,
Rebeca Chaverri, Martin Zeilbauer, Coral Pacheco,
Eduardo Toral, Franklin Castan
˜eda, Charles Foerester,
Rausel Sarmiento and the 14th promotion of PRMVS-
UNA for their invaluable help and companionship in
the field, the staff at Corcovado National Park for their
support and for sharing their facilities, Jose Pablo
Carvajal for his assistance with the maps, and Meg
Harper for translating this document. We thank the
Wildlife Conservation Society, Corcovado National
Park, the University of Costa Rica and the National
University of Costa Rica for their support, and Gerardo
Uman
˜a, Marcella Kelly and an anonymous reviewer for
their comments.
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56
Biographical sketches
This study was a part of Roberto Salom-Pe
´rez’s MSc thesis at
the University of Costa Rica, which included a population
estimate for ocelots and investigation of the predator/prey
relationship between jaguars and marine turtles. This
research was also part of the Jaguar Conservation Program
(JCP) of the Wildlife Conservation Society, which began in
1999 with the objective of reversing jaguar decline through-
out its range. Eduardo Carrillo is a professor at the Wildlife
and Management Institute (IMVS) of the National University
in Costa Rica and the coordinator for the JCP of Mesoamerica.
As part of the JCP Roberto Salom-Pe
´rez and Eduardo Carrillo
are now carrying out a camera-trap based investigation to
help delineate the corridor in Golfo Dulce Forest Reserve,
using spider monkey Ateles geoffroyi and large felids as key
species. Jose
´M. Mora is a professor at the University of Costa
Rica and has carried out research focused on mammals and
reptiles. Joel C. Sa
´enz is the director of IMVS and carries out
research on habitat fragmentation.
R. Salom-Pe
´rez et al.
ß2007 FFI, Oryx,41(1), 51–56