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Nombre del archivo original: Paviolo et al 2008 Jaguar pop decline Oryx.pdf
Título: Jaguar Panthera onca population decline in the Upper Paraná Atlantic Forest of Argentina and Brazil
Autor: Agustín Paviolo, Carlos Daniel De Angelo, Yamil Edgardo Di Blanco, Mario Santiago Di Bitetti

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Jaguar Panthera onca population decline in the
Upper Parana´ Atlantic Forest of Argentina and Brazil
A g u s t I´ n P a v i o l o , C a r l o s D a n i e l D e A n g e l o , Y a m i l E d g a r d o D i B l a n c o and
Mario Santiago Di Bitetti
Abstract The Green Corridor of Argentina and Brazil is

the largest forest remnant of the Upper Parana´ Atlantic
Forest. The jaguar population of this region is highly
fragmented and reduced. To assess the status of the
subpopulation of jaguars of the Green Corridor we conducted four camera-trap surveys in three sites with
different levels of protection. At Urugua-ı´ (34 stations,
1,495 trap-days) we recorded one individual (minimum
density 5 0.12–0.33 per 100 km2). At Yabotı´ Biosphere
Reserve (42 stations, 1,871 trap-days) we recorded two
individuals (minimum density 5 0.11–0.25 per 100 km2).
At Iguazu´ National Park we conducted two surveys. In
2004 (39 stations, 1,839 trap-days) we recorded four adult
individuals, estimating a density of between 0.49 – 0.16
and 1.07 – 0.33 per 100 km2. In 2006, we increased the area
sampled (47 stations, 2,059 trap-days) and recorded 11
adult individuals, estimating a density of 0.93 – 0.2 to 1.74 –
0.34 per 100 km2. These density estimates are the lowest
recorded for the species. Estimates for Iguazu´ are between
2–7.5 times lower than those reported in the early 1990s.
This population decline probably results from the interaction of several factors, including lack of prey as a result
of poaching and persecution. We estimate that there is
currently a population of 25–53 adult jaguars in the Green
Corridor. In spite of having sufficient potential habitat
available this population is threatened and urgent conservation action is required.
Keywords Argentina, Atlantic Forest, camera trap, density

estimate, Green Corridor, jaguar, Panthera onca, poaching.
Introduction

T

he jaguar Panthera onca is the largest felid of the
Americas. During the 20th century its distribution was
reduced to almost half of its original range (Sanderson et al.,
2002). This reduction was particularly severe in Argentina
(Perovic, 2002) and the species now occurs in only three

AGUSTI´N PAVIOLO (Corresponding author), C ARLOS DANIEL DE ANGELO
and M ARIO SANTIAGO DI BITETTI National Research Council of Argentina–
Laboratorio de Investigaciones Ecolo´gicas de las Yungas, Universidad
Nacional de Tucuma´n and Centro de Investigaciones del Bosque Atla´ntico,
Yapeyu´, 23, CP 3370 Puerto Iguazu´, Misiones, Argentina. E-mail
paviolo4@arnet.com.ar
Y AMIL EDGARDO DI BLANCO Centro de Investigaciones del Bosque Atla´ntico,
Yapeyu´, 23, CP 3370 Puerto Iguazu´, Misiones, Argentina.
Received 4 April 2007. Revision requested 20 August 2007.
Accepted 3 December 2007.

isolated areas in the north (Altrichter et al., 2006; Di Bitetti
et al., 2006a). The jaguar is considered a threatened species
in Argentina (Dı´az & Ojeda, 2000), and is categorized as
a National Natural Monument (Ley No. 25463, 2001). In
Brazil it is considered threatened (Reis et al., 2006) and it
has disappeared from most of the Atlantic Forest, where
only a few small and isolated populations remain (Leite
et al., 2002; Cullen et al., 2005).
Globally the Atlantic Forest of South America is one of
the most threatened rainforests. Distributed across northeast Argentina, south-west Brazil and eastern Paraguay, the
Upper Parana´ Atlantic Forest is the innermost region of the
Atlantic Forest complex (Di Bitetti et al., 2003) but is being
reduced and fragmented as a result of its conversion to
other land uses (Holz & Placci, 2003).
The largest remnant of the Upper Parana´ Atlantic
Forest, known as the Green Corridor (c. 10,000 km2, Fig. 1a)
lies in the Misiones Province of Argentina and neighbouring areas of Brazil. The Corridor holds the southernmost population of jaguars and has been identified as a high
priority area for the conservation of the species because
of its long-term conservation potential and its ecological
uniqueness (Sanderson et al., 2002; Marieb, 2006). This
jaguar population is the only one that has received that
category in Argentina and in the Atlantic Forests (Eizirik
et al., 2002; Sanderson et al., 2002).
Crawshaw (1995) provided relevant ecological information for this jaguar population, including home range size,
population density and diet, and identified important
threats. However, the decline in the rate of observed jaguar
signs and in the number of reported attacks on domestic
animals suggest that its population could have declined
since the 1990s (Crawshaw, 2002).
The goal of this study was to estimate jaguar population
densities widely across the Green Corridor and thus make
a robust estimate of population size, assess the species’
conservation status, and compare the population estimate
with earlier values from the region (Crawshaw, 1995; Cullen
et al., 2005) to assess population trends.
Study area
The Green Corridor is dominated by semi-deciduous forest
and has a humid subtropical climate with hot summers
(December–March) and winters with frosts (June–August).
Mean total annual precipitation is 1,700–2,200 mm, with no
marked dry season (Crespo, 1982).

ª 2008 Fauna & Flora International, Oryx, 42(4), 554–561

doi:10.1017/S0030605308000641

Printed in the United Kingdom

Jaguar decline in the Atlantic Forest

FIG. 1 (a) The Green Corridor of Argentina and Brazil and its main protected areas; the circles indicate the three study sites. The
rectangle on the inset illustrates the location of the main map at the border between Argentina and Brazil. (b) The Iguazu´ study site,
showing the minimum convex polygons that include all the camera-trap stations during the surveys of 2004 and 2006, and Crawshaw’s
(1995) study area.

ª 2008 Fauna & Flora International, Oryx, 42(4), 554–561

555

556

A. Paviolo et al.

We conducted the first camera-trap survey at Urugua-ı´
in 2003 in an area that comprises the Urugua-ı´ Private
Wildlife Reserve (32 km2), a portion of the Urugua-ı´ Provincial Park (840 km2) and a portion of Campo Los Palmitos
(300 km2), which belongs to a timber company and
contains old pine (Pinus taeda and Pinus ellioti) plantations
in a matrix of native forest (Fig. 1a). This forest was
selectively logged until the beginning of the 1990s but is
in a relatively good condition (Di Bitetti et al., 2006b). Lack
of resources precludes effective control of poaching, and the
area suffers a moderate to high hunting pressure (Table 1).
The second study site is Iguazu´, which we surveyed
twice, in 2004 and in 2006–2007. The first survey comprised
the central area of the Iguazu´ National Park of Argentina
(670 km2, Fig. 1). This is the best-protected area of the
region (Giraudo et al., 2003) and suffers a relatively low

TABLE 1 Area, law enforcement capacity and evidence of
poaching in the three surveyed areas (Iguazu´ was surveyed in
both 2004 and 2006; Fig. 1).
Iguazu´, Iguazu´,
Measure of hunting pressure Urugua-ı´ 2004 2006
Yabotı´
Law enforcement capacity
1,132
670
2,694
3,160
Area (km2)
Number of rangers
9
25
70
7
Number of vehicles
3
4
9
3
Rangers km-2
0.8
3.73
2.6
0.22
Vehicles km-2
0.27
0.6
0.33
0.1
Use of fire arms by
No
Yes
Yes
No
rangers
Evidence of hunting
Encounters with hunters
4
0
1
2
Encounters with dogs
4
0
0
0
Photographic records of
4
2
6
8
dogs or people
Hunting camp sites
6
0
0
.31
Artificial salt licks or
3
0
0
4
poaching platforms
Gunshots heard2
5
1
0
0
Hunting trails
7
0
0
03
Spent shotgun cartridges
1
0
0
2
2
0
0
04
Human tracks not
associated with logging
activities or patrols
by rangers
Camera trap stations
0
0
3
1
robbed or destroyed
1

This is an underestimate. We found three hunters’ camp sites but there
were many logging camps, employees of which usually hunt at the
weekends. On the two occasions we found spent shotgun cartridges it was
at logging camps.
2
Number of independent events, at each of which we may have heard . 1
shot.
3
At Yabotı´ we opened few trails and this reduced the likelihood of crossing
or detecting hunting trails.
4
Most private properties were under logging operations and we were not
able to distinguish tracks of workers from those of poachers.

hunting frequency, restricted mainly to the park edges
(Table 1). In the second survey we expanded the sampled
area to cover most of Iguazu´ National Park, the whole San
Jorge Forest Reserve and the western area of the Brazilian
Iguaçu National Park of 1,850 km2 (Fig 1b). The surveyed
area overlaps extensively with Crawshaw’s (1995) study area
(Fig. 1b). The absence of a buffer zone and the extensive
edge of the western portion of the Brazilian Park makes
this area accessible to poachers (Crawshaw, 2002). The San
Jorge Forest Reserve (174 km2) is owned by a timber
company and is covered by native forest that was selectively
logged until 20 years ago (O. Lescano, pers. comm.).
The third study site is in the Yabotı´ Biosphere Reserve
(2,600 km2), which we surveyed in 2005, and comprises
private properties with forest that are being currently logged
(2,200 km2). It also comprises Esmeralda Provincial Park
(300 km2, logged until 1990). Poachers frequently arrive from
villages located outside the reserve and from the Brazilian
border. Anti-poaching control is insufficient and the area
suffers moderate to high hunting pressure (Table 1).
Methods
To estimate jaguar densities we used a standard cameratrapping protocol (Karanth, 1995; Karanth & Nichols, 1998)
that has been used to estimate jaguar densities (Wallace
et al., 2003; Maffei et al., 2004; Silver et al., 2004; Cullen
et al., 2005; Soisalo & Cavalcanti, 2006; Salom-Pe´rez et al.,
2007). The method makes use of capture-mark-recapture
closed population models to estimate animal densities (Otis
et al., 1978). Individuals are identified in photographs by
their distinctive spotted coat. From the capture-recapture
history of individuals we estimated population size
(Karanth, 1995).
At each study site we set 34–47 sampling stations
(Table 2), each of which consisted of a pair of camera
traps operating independently and facing each other. The
stations were located at regular intervals on trails and
rarely-used dirt roads. Prior to the full survey period we
conducted preliminary surveys of 4–7 months to identify
promising sites for placing the stations. Each full survey
lasted 96 days. During this period we set up camera traps in
half of the sampling stations for the first half of the survey
period (day 1–(45–50)), after which we moved the stations
to the remaining sampling stations for the rest of the study
period (day (45–50)–96).
In Urugua-ı´ and Yabotı´ we could not estimate the
abundance of jaguars by means of capture-recapture
population models because of the small number of individuals recorded. In these sites we provide an estimate of
a minimum number of individuals present in the area from
the photographs obtained by camera traps and the observation of jaguar tracks. To estimate jaguar abundance
at Iguazu´ we used the software CAPTURE (Rexstad &
ª 2008 Fauna & Flora International, Oryx, 42(4), 554–561

Jaguar decline in the Atlantic Forest
TABLE 2 Number of camera-trap stations, mean distance between stations, date of the full survey, and survey effort for full and for both
surveys combined, in the three surveyed areas (Iguazu´ was surveyed in both 2004 and 2006; Fig. 1).

Site
Urugua-ı´
Iguazu´, 2004
Iguazu´, 2006
Yabotı´

No. of
stations
34
39
47
42

Mean (– SE)
distance
between stations
(km)
1.25 – 0.46
2.08 – 0.41
2.58 – 0.60
2.43 – 0.81

Date of full
survey
Dec. 2003–Feb. 2004
Aug.–Nov. 2004
Oct. 2006–Jan. 2007
Sep.–Dec. 2005

Full survey
effort (trap
days)
1,495
1,839
2,059
1,871

Total survey
effort* (trap
days)
2,611
2,942
2,287
2,676

*Includes effort of the preliminary and full surveys

Burnham, 1991), which provides population estimates using
various models (Otis et al., 1978; White et al., 1982).
We report the results of model Mh using a Jackknife
estimator. Mh assumes heterogeneity among individuals
in their capture probabilities and is the most appropriate
model because of the unequal access to sampling stations
by different individuals (Karanth & Nichols, 2002). We
divided the survey period into 16 trapping occasions of
6 consecutive days each with the aim of having a capture
probability higher than 0.1, as recommended by Otis et al.
(1978) and White et al. (1982). Finally, we performed the
closure test provided by CAPTURE to test the closed
population assumption.
To estimate density it is necessary to calculate the area
effectively sampled. This is usually accomplished by applying
to each sampling station a buffer equivalent to ½ of the mean
maximum distance of recaptures (MMDM) for the individuals recorded at more than one station (Silver et al., 2004).
However, in some situations using ½ MMDM as the buffer
could inflate density estimates (Trolle & Ke´ry, 2003; Soisalo
& Cavalcanti, 2006; Maffei & Noss, 2007) and, if radiotelemetry data exist, the best buffer should be the radius of
the estimate of mean home range (Soisalo & Cavalcanti,
2006). We provide three density estimates using different
buffers to estimate the effectively sampled areas: (1) the
radius of the mean adult home range (n 5 3 individuals)
from Crawshaw (1995); (2) ½ MMDM; (3) MMDM. Because
of the small number of jaguars photographed we calculated
MMDM as the mean maximum distance moved by all the
individuals with recaptures at more than one station, pooling
the data from the four surveys (n 5 8). For individuals that
were captured (and recaptured at . 1 station) in the two
surveys at Iguazu´ (n 5 2), we averaged their maximum
distance moved (MDM) for both surveys, thus contributing
only one value each to the estimate of MMDM. After
applying the buffer to the sampling stations we subtracted,
from the total area thus obtained, the portions of unsuitable
jaguar habitat (e.g. cities, annual crops, airports) to estimate
the effectively surveyed area. We used the geographical
information system ArcView v. 3.2 (ESRI, Redlands, USA)
to estimate MMDM values and surveyed areas.
ª 2008 Fauna & Flora International, Oryx, 42(4), 554–561

We used an ANOVA, and post hoc comparisons among
samples using the Tukey-Kramer test, to compare our
estimates of jaguar density with others for the same region
(Iguazu´: Crawshaw, 1995; Morro do Diabo State Park: Cullen
et al., 2005) and with other published estimates from across
the jaguar’s range (Ceballos et al., 2002; Nun˜ez et al., 2002;
Maffei et al., 2004; Silver et al., 2004; Soisalo & Cavalcanti,
2006; Salom-Pe´rez et al., 2007). For these comparisons we
used our estimates obtained using ½ MMDM as the buffer.
For Iguazu´ we used only one density value (the mean of the
two surveys). After checking for deviations from normality
we performed the ANOVA with ln(density).
To estimate the population of adult jaguars in the Green
Corridor we first appraised the area of potential jaguar
habitat by using detailed jaguar presence information
obtained in 4 years of intensive large-scale census of jaguar
presence (C.D. De Angelo, unpubl. data), and then digitized
forest and plantation cover that surrounded jaguar presence points from 2004 Landsat satellite images using
ArcView. We categorized this potential area into three
habitat categories: (1) not suitable (jaguars absent), (2) well
protected, and (3) poorly protected. Finally, we extrapolated our maximum and minimum density estimates for
areas with low protection (mean of Yabotı´ and Urugua-ı´)
and high protection (mean of the Iguazu´ surveys) to the
available area in categories 2 and 3 of habitat quality to
estimate the number of jaguars.

Results
We photographed 13 different adult jaguars during the four
surveys, eight at more than one station. The maximum (25.08
km) and minimum (2.35 km) distances of recapture were for
an adult male and female, respectively, at Iguazu´. The MMDM
was 11.33 – SE 2.7 km (n 5 8), which is not statistically
different from the mean diameter of the home ranges of the
adult jaguars estimated by Crawshaw (1995; 8.55 – SE 1.45 km,
n 5 3; ANOVA F 5 0.36, P 5 0.563).
Urugua-ı´ We recorded only one individual, an adult male,
captured twice (at two sampling stations; MDM 5 9.44 km)

557

558

A. Paviolo et al.

during the full survey and once during the preliminary
survey. Jaguar tracks were found only three times and
probably belonged to the photographed jaguar, because they
were similar in size. Considering the low occurrence of tracks
and the low number of photographic captures (Table 3),
it is possible that the animal photographed could have been
the only resident jaguar in the surveyed area. The effectively
sampled area varied widely (299.01–823.63 km2) depending
on the buffer used. We estimate a minimum density of
0.12–0.33 per 100 km2 at Urugua-ı´ (Table 2).
Iguazu´ During the first full survey we obtained 10 photographs of four different adults (three females, one male).
We also photographed two other jaguars that were not
included in the analyses because they were captured outside
the full survey period of 96 days. CAPTURE indicate that
this population is not different from a closed one (z 5 -0.942,
P 5 0.173) and estimated a population of 5 – SE 1.41 adults.
The effectively sampled area was 467.65–1,023.78 km2,
depending on the buffer used (Table 3). We estimated an
adult jaguar density of between 0.49 – SE 0.16 and 1.07 – SE
0.33 per 100 km2 (Table 3). During the second survey
we obtained 28 photographs of 11 different adult individuals (six females, four males, one unsexed individual),
four of which were photographed during the 2004 survey.
Two of the females were photographed with large cubs
(c. 1 year old). CAPTURE indicated there was no violation of the closed population assumption (z 5 -1.392, P 5
0.082) and estimated an adult population of 14 – SE 2.45.
The estimated sampled area was 804.88–1,499.52 km2,

depending on the buffer. We estimated an adult jaguar
density of between 0.93 – SE 0.2 and 1.74 – SE 0.34 per
100 km2 (Table 3).
Yabotı´ We photographed only one male, recorded once and
three times during the preliminary and full surveys, respectively. We recorded large tracks (right front track 12.1
cm long 3 13.1 cm wide), most probably belonging to the
photographed male, and medium size tracks (right front
track 9.1 cm long 3 10.6 cm wide) probably from an adult
female or a subadult male. Thus, the surveyed area was
occupied by at least two jaguars, giving a minimum
estimate of 0.11–0.25 per 100 km2 (Table 2).
There are statistically significant differences between the
estimated jaguar densities at the three study sites and those
obtained in the same area . 10 years before or in other
portions of the same region (n 5 2) and with other jaguar
studies (n 5 10; ANOVA F2,12 5 17.49, P 5 0.0003;
Fig. 2). Tukey-Kramer comparisons among samples indicated that the jaguar densities of the Green Corridor
(mean 5 0.545 – SE 0.31 per 100 km2) are lower than those
of other sites outside the Upper Parana´ Atlantic Forest
(5.243 – SE 0.77 per 100 km2) and those previously obtained in the Green Corridor or in other fragments of the
Upper Parana´ Atlantic Forest (2.96 – SE 0.74 per 100 km2).
The latter two estimates are not statistically different.
We calculated an area of 9,234 km2 of habitat with
potential jaguar presence in the Green Corridor. The extrapolation of the minimum and maximum jaguar density
estimates to the available areas of high and low protection

TABLE 3 Number of jaguar photo-captures per 1,000 trap-days, number of adult jaguars recorded, population estimate, buffer applied for
calculations (see text for further details), area surveyed, and density estimates for each of the four full camera-trap surveys.

Site
Urugua-ı´

Jaguar
captures
per 1,000
trap-days1
1.34

No. of
adult
jaguars
recorded
1

Population
estimate
– SE
12

Iguazu´, 2004

5

4

5 – 1.417

Iguazu´, 2006

14

11

14 – 2.457

Yabotı´

1.63

2

22

Buffer
applied
(km)
4.283
5.675
11.336
4.283
5.675
11.336
4.283
5.675
11.336
4.283
5.675
11.336

Surveyed
area
(km2)
299.01
367.69
823.63
467.65
576.61
1,023.78
804.88
958.16
1,499.52
807.94
1,000.67
1,762.62

Density
estimate
– SE
(100 km-2)
0.334
0.274
0.124
1.07 – 0.33
0.87 – 0.3
0.49 – 0.16
1.74 – 0.34
1.46 – 0.34
0.93 – 0.2
0.254
0.24
0.114

1

. 1 h had to pass for consecutive photographs of a jaguar to be considered independent records
Number of jaguars recorded in the study area during the survey
3
½ of the radius of the mean adult home range estimates (n 5 3) from Crawshaw (1995)
4
Minimum density estimate
5
½ of the mean maximum distance of recapture (½ MMDM) for all the individuals recaptured at . 1 sampling station during the four surveys
6
Mean maximum distance of recapture (MMDM) for all individuals recaptured at . 1 sampling station during the four surveys
7
Abundance estimate obtained with CAPTURE (Rexstad & Burnham, 1991) using model Mh
2

ª 2008 Fauna & Flora International, Oryx, 42(4), 554–561

Jaguar decline in the Atlantic Forest

FIG. 2 Estimates of jaguar density in the Upper Parana´ Atlantic
Forest (UPAF) from this (13, 14, 15, 16) and other studies
(8, Iguazu´: Crawshaw, 1995; 11, Morro do Diabo State Park: Cullen
et al., 2005), and from other regions (1, 2, 6, 7, 9, Silver et al., 2004;
3, Salom-Pe´rez et al., 2007; 4, Ceballos et al., 2002; 5, Soisalo &
Cavalcanti, 2006; 10, Maffei et al., 2004; 12, Nun˜ez et al., 2002).

results in an estimate of 25–53 adult jaguars in the Green
Corridor.
Discussion
Jaguars currently occur at low densities in the Green Corridor
of Argentina and Brazil. Our estimates of density are the
lowest obtained for the species across its range, and lower
than those obtained by Crawshaw (1995) in the same area and
by Cullen et al. (2005) in the northern part of the same
region. We believe we have documented a decline that has
taken place since the early 1990s. At Iguazu´ our estimates are
2–7.5 times lower than the 3.7 jaguars per 100 km2 calculated
by Crawshaw in 1995 (Fig. 2). The diminution of jaguar signs
(Crawshaw, 2002) and fewer recorded attacks by jaguar on
domestic animals during the last few years (K. Schiaffino,
pers. comm.) also suggest that jaguars are now less abundant.
Our low density estimates in comparison to other studies
could have resulted from methodological differences or
sampling artefacts but ancillary evidence suggests our estimates are reliable. Firstly, our sampling effort was similar or
greater than that of other studies in terms of the number of
stations and trap-days, and our camera-trap surveys at Yabotı´
and Iguazu´ are probably the largest ever conducted for jaguars
in terms of the area sampled (A. Noss, pers. comm.). Secondly,
although Crawshaw (1995) used radio-telemetry other studies
estimated similar jaguar densities using both radio-telemetry
and camera trapping in the same region (Cullen et al., 2005),
and our ocelot density estimates for Iguazu´ using camera traps
are not different from Crawshaw’s (1995) estimate using radiotelemetry (Di Bitetti et al., 2006b, 2008a).
The slightly higher jaguar density recorded in the 2006
survey at Iguazu´ may be related to the inclusion of San
Jorge Forest Reserve (c. 25% of the sampled area), where six
of the eleven adults photographed were recorded. This, and
ª 2008 Fauna & Flora International, Oryx, 42(4), 554–561

the fact that San Jorge is located in a strategic area serving
as a connection between Iguazu´ National Park and Urugua-ı´
Provincial Park, (Fig. 1b) suggest that this is an important
area for jaguar conservation.
It was previously assumed that Crawshaw’s density
estimates for Iguazu´ were representative of the whole Green
Corridor (Eizirik et al., 2002). For Urugua-ı´ and Yabotı´ we
only have conservative density estimates and there is no
previous information to ascertain whether there has been
a population decline in these areas as well. However, the
fact that we photographed only one jaguar at each site,
despite the long duration of the surveys and the large areas
studied, the difficulty of finding jaguar tracks, and the low
jaguar recording rate compared to similar studies (Table 1;
Maffei et al., 2004) indicate that jaguar densities are
currently extremely low in these areas.
Low availability of prey could have affected this population. A good prey base is essential for the maintenance
of healthy jaguar populations (Hoogesteijn & Mondolfi,
1992; Crawshaw, 1995). In Misiones poaching is common
and can reduce prey availability (Paviolo, 2002). Our
camera trap records of collared peccaries Pecari tajacu,
tapirs Tapirus terrestris and red brocket deers Mazama
americana were also fewer in Urugua-ı´ and Yabotı´ (with
high hunting pressure) than in Iguazu´ (Di Bitetti et al.
2008b; A. Paviolo et al., unpubl. data). Densities of other
large carnivores such as pumas and ocelots are also lower
at Yabotı´ and Urugua-ı´ (Di Bitetti et al., 2006b, 2008a;
A. Paviolo et al., unpubl. data), suggesting they are probably
affected by similar factors.
In the northern part of the Green Corridor (including
Iguazu´ and Urugua-ı´), white-lipped peccaries Tayassu
pecari have declined. In the early 1990s they were abundant
and comprised the most important prey item in the jaguar’s
diet (Crawshaw, 1995). This species has not been recorded
since 2000 in the Iguaçu´ National Park of Brazil (A. Ricieri,
pers. comm.). We did not record it in our survey at Urugua´ı and it was rarely recorded during the two surveys at Iguazu´
National Park, and in unusually small herds (A. Paviolo et al.,
unpubl. data). The scarcity of this major prey species could
have affected jaguars in this area but does not explain the
low density of jaguars at Yabotı´, where we observed and
recorded white-lipped peccaries in large herds.
Even though jaguars are protected by law in Argentina
and Brazil they are regularly killed. Jaguar poaching was an
important source of mortality in the Iguaçu National Park
(Crawshaw, 1995), and jaguars are also killed because they
occasionally prey on domestic animals (Schiaffino et al.,
2002) and because people consider them dangerous
(Conforti & Azevedo, 2003). Between 1995 and 2002 at least
70 jaguars were killed in areas neighbouring Iguaçu´ National
Park (Crawshaw, 2002), and during the last 10 years at least
47 were killed in Northern Misiones (A. Paviolo, unpubl.
data).

559

560

A. Paviolo et al.

The conversion of forests into other land uses is an
ongoing process that reduces the availability of adequate
habitat for jaguars, fragments forest and facilitates poaching in areas that were previously relatively inaccessible. The
decline of the jaguar population of the Green Corridor,
estimated to number several hundreds c. 15 years ago
(Eizirik et al., 2002) but now comprising , 60 adult animals,
may have resulted from the interaction of these factors.
Because this population is at the southernmost limit of the
species’ range its conservation has special relevance. Its
disappearance would be one of the final steps towards the
extinction of the species in Argentina, where other jaguar
populations are also threatened (Altrichter et al., 2006;
Di Bitetti et al., 2006a).
However, the population of jaguars in the Upper Parana´
Atlantic Forest is still that with the greatest potential for
long-term persistence in the Atlantic Forest, despite its
relatively small size. To mitigate the impact of the main
threats and allow the jaguar population to recover we
recommend five actions: (1) Halt the killing of jaguars and
their prey, by reinforcing anti-poaching measures both
inside and outside protected areas. (2) Implement the
existing protected areas and create new ones in strategic
locations to improve connectivity between forest areas, thus
conserving important habitat and limiting access for
poachers. (3) Reduce conflict between cattle ranchers and
jaguars by compensating ranchers for losses but also
demanding better management practices and penalizing
those who kill jaguar. (4) Change human perceptions of the
danger from jaguars and inform the public about the
species’ conservation status using campaigns and environmental education programmes (an example of which is
Campan˜a Yaguarete´, 2007, an education campaign recently
launched by local NGOs and governmental agencies). (5)
Coordinate joint conservation efforts between government
institutions in Argentina and Brazil and NGOs (Paviolo
et al., 2006). We are now working in partnership with these
institutions to develop a recovery plan for the jaguar
population of the Green Corridor (Chalukian et al., 2006).
Acknowledgements
We are grateful to all the volunteers and park rangers who
helped us with field work. We acknowledge the support and
permits provided by the Ministry of Ecology, Natural
Resources and Tourism of Misiones province (MERNRT)
and the National Parks Administration of Argentina. We
are grateful to Fundacio´n Vida Silvestre Argentina and
the property owners for their support and permission to
conduct this work. Financial support was provided by
CONICET, Fundacio´n Vida Silvestre Argentina, WWF–
USA, WWF–International, WWF–Switzerland, Lincoln
Park Zoo, Fundacio´n Antorchas, the Wildlife Conservation
Society, Idea Wild, the Rufford Foundation, MERNRT, and

the Eden Project through a grant from the Darwin
Initiative. We also thank Lı´a Montti, Andy Noss, Cristina
de De Angelo, Peter Crawshaw and an anonymous reviewer for help and comments on the manuscript.
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Biographical sketches
A G U S T ´I N P A V I O L O has been working on research related to jaguar
conservation in the Atlantic Forest since 2001. This research is focused
on the factors that affect the abundance of jaguar, puma and their
prey, and their conservation in the Atlantic Forest. C A R L O S D A N I E L
D E A N G E L O conducts research on jaguar distribution and population
genetics in relation to Atlantic Forest landscape fragmentation.
Y A M I L E D G A R D O D I B L A N C O is participating in various studies
on the ecology and conservation of mammals in the north-east of
Argentina. M A R I O S A N T I A G O D I B I T E T T I has been carrying out
research in the Atlantic Forest for the last 15 years, focusing on the
behaviour, ecology and conservation of primates and felids.

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