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Quiroga et al. 2014 Jaguar in argentinean Chaco Oryx .pdf

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Critical population status of the jaguar Panthera
onca in the Argentine Chaco: camera-trap surveys
suggest recent collapse and imminent
regional extinction
and M A R I O S A N T I A G O D I B I T E T T I

Abstract The population of jaguars Panthera onca in
the semi-arid Chaco Province is the least well-known in
Argentina. Its status in the region is described only from
interviews that confirmed its presence until 2003. To update
information on the distribution and population density
of this species we undertook three camera-trap surveys,
combined with searches for sign, at sites across latitudinal
and protection gradients, and 156 interviews with local
inhabitants across three larger areas. The camera-trap sites
were located in areas with the highest density of records in
the Argentine Chaco: Copo National Park (1,204 trap days,
24 stations, 344 km of transects), Aborigen Reserve (1,993
trap days, 30 stations, 251 km of transects) and El Cantor
(2,129 trap days, 35 stations, 297 km of transects). We did not
obtain any photographs of jaguars. We recorded very few
jaguar tracks, and only in the Aborigen Reserve (n 5 3) and
El Cantor (n 5 1). The map of distribution points confirmed
through interviews suggests that the jaguar range has not
changed significantly in the past 10 years; however, the
camera-trap and sign surveys suggest that densities are
extremely low. Before our study the Chaco population was
thought to be the largest in Argentina. This perception
was incorrect: the Chaco jaguar population is the most
threatened in the country. Systematic, intensive studies are
essential to provide the necessary information for decisionmaking for the management and conservation of threatened

BITETTI National Research Council of Argentina, Instituto de Biología
Subtropical—nodo Iguazú, Facultad de Ciencias Forestales, Universidad
Nacional de Misiones, Argentina, and Asociación Civil Centro de
Investigaciones del Bosque Atlántico, Puerto Iguazú, Misiones, Argentina.
GABRIEL IVÁN BOAGLIO Proyecto Elé, Dirección de Fauna Silvestre, Secretaría de
Ambiente y Desarrollo Sustentable, Argentina, and Asociación para la
Conservación y el Estudio de la Naturaleza, Córdoba Capital, Argentina
ANDREW JAY NOSS University of Florida, Gainesville, Florida, USA
*Also at: Asociación para la Conservación y el Estudio de la Naturaleza, Córdoba
Capital, Argentina
Received 7 February 2012. Revision requested 14 March 2012.
Accepted 11 June 2012.

© 2013 Fauna & Flora International, Oryx, Page 1 of 8


Keywords Argentina, Chaco, conservation, distribution,
interview, monitoring, Panthera onca, presence



he distribution of the jaguar Panthera onca in
Argentina has declined by . 90% and , 200 individuals survive in three isolated regions: the Atlantic Forest,
the Yungas and the Chaco forest (Di Bitetti et al., in press).
The jaguar is a National Natural Monument protected by
national and provincial laws, although these are rarely
enforced. The most recent national categorization based on
IUCN criteria considers the species Critically Endangered in
Argentina (Di Bitetti et al., in press). The Argentine jaguar
population is at the southern limit of the species’ range.
Therefore, it may be more vulnerable to human activities
and persist at lower densities than populations nearer to
the centre of the species’ range, enduring ecosystems with
marginal climatic conditions and low productivity, as do
jaguars at the northern limit of their range in the south-west
USA (Gaston et al., 1997; Laliberte & Ripple, 2004; Yackulic
et al., 2011). However, in Argentina the Chaco is not the
historical limit of the species’ range (it was formerly present
in northern Patagonia, 1,500 km further south), rather it
became so only after a considerable contraction of the
regional range caused by intensive land conversion
combined with heavy hunting pressure (Di Bitetti et al.,
in press).
The extreme isolation, aridity and high temperatures
of the Argentine Chaco have resulted in the Chaco jaguar
population being the least known and studied of the
country’s three populations. There have been only two
previous studies of jaguars in the region. One short cameratrap survey attempted to estimate population density in
Copo National Park (Denapole, 2007) but did not record
jaguars. The second, broader study, based on interviews
with inhabitants and the compilation of information on
jaguar records, evaluated jaguar distribution across the
whole Argentine Chaco (Altrichter et al., 2006). Better
knowledge of the situation of jaguars in the Argentine
Chaco is necessary because this region has been identified as


V. A. Quiroga et al.

FIG. 1 The study area in the
Argentine semi-arid Chaco,
with the three extensive areas
surveyed using interviews
during 2004–2010, minimum
convex polygons of the three
camera trap surveys (2008–
2010), and the locations of 35
jaguar records obtained during
the interviews. The rectangle
on the inset indicates the
location of the main map in

a Jaguar Conservation Unit (Sanderson et al., 2002) and is
important for connectivity with other areas where jaguars
persist, such as the Bolivian and Paraguayan Chaco as well
as the Argentine and Bolivian Yungas forests (Rabinowitz &
Zeller, 2010; Rumiz et al., 2011).
For species that are difficult to detect because of their
low population density or behaviour, studies in other
regions have used presence records to evaluate the status of
the population (Perovic et al., 2003; De Angelo et al., 2011;
Karanth et al., 2011a; Pereira Munari et al., 2011; Salvador
et al., 2011). Such information allows investigators to
develop a baseline for a species’ distribution and to identify
favourable research sites, particularly when information
is scarce or non-existent. However, presence data and
distribution maps alone are not sufficient to determine the
conservation status of threatened species (Can & Togan,
2009; Karanth et al., 2011b). Altrichter et al. (2006) provided
information on the range of the jaguars in the region up to
2003 but they focused on interviewing local residents, and
did not have information about whether reports represented
a few wandering male jaguars or a reproducing population.
We evaluated the density of the jaguar in areas of various
categories of legal protection and intensities of hunting and
ranching in the Argentine Chaco using the first large-scale
camera-trap survey in the region. We also evaluated
whether presence data reflect the conservation status and
current range of the species in the region, and whether stable
populations persist at each site.

Study area
The Gran Chaco is the second-most extensive forest in the
Americas and it is the largest subtropical dry forest (Morello
& Adámoli, 1974; Morello et al., 2009). Sixty percent of
the Chaco (675,000 km2) is in Argentina and c. 270,000 km2
are semi-arid Chaco forests. The Argentine semi-arid
Chaco comprises dry forest plains with a marked seasonal
climate, a median annual temperature of 24 °C and annual
precipitation of 400–800 mm, falling mostly in October–
April. The area was colonized early in the 20th century by
non-indigenous people who settled on isolated ranches. The
region also has many indigenous settlements. The characteristics of the region support an extensive cattle ranching
system, combined with heavy hunting of wildlife both by
local inhabitants and by hunters from nearby towns
(Baxendale & Buzai, 2009).
We compared three sites across a north-south gradient
(Fig. 1) and across a gradient of conditions related to
hunting pressure, livestock burden and legal protection
status (Table 1).
Copo National Park is the site with the highest degree of
protection within the study area. Although there are a few
small human settlements near the Park, only one person
lives inside the surveyed portion of the Park and the
livestock burden is the lowest of any of the surveyed sites.
Lacking roads and trails, the interior of Copo National Park
is inaccessible and therefore hunting pressure is very low.
© 2013 Fauna & Flora International, Oryx, 1–8








2.85 ± 0.65





3.04 ± 0.98



2.99 ± 0.32

© 2013 Fauna & Flora International, Oryx, 1–8

Pilot survey
in Aborigen

Copo National
Park (south)

Aborigen Reserve

1 July–9
23 June–7
4 September–19
17 September–3
October 2007
El Cantor (north)


Survey period

Cameratrap days


Mean distance
cameras± SD



convex polygon
covered by
camera traps
Total camera-trap
days (including
pilot and
main surveys)
No. of

TABLE 1 Survey effort and characterization of the three survey sites (Fig. 1), 2007–2010.

surveys for
sign (km)

No. of
in and
around site

Livestock burden
(no. of photos of
cattle per 100

The jaguar in the Argentine Chaco

Aborigen Reserve is not a protected area but because it is
categorized as an indigenous reserve it is sparsely populated
and has an intermediate livestock burden relative to the
other two survey sites. Three roads cross the reserve and
facilitate access by hunters.
El Cantor is not legally protected but is considered a
candidate for the creation of a protected area (Brown et al.,
2010). The site is currently inhabited and people hunt
wildlife there. The livestock burden is the highest of the
three study sites.

We worked at two different scales to generate two types
of information: fine-scale data were generated by camera
traps, line transects and track plots to estimate presence,
number of individuals and density, based on captures and
recaptures; and coarse-scale presence–absence data were
generated through interviews, to determine the species’
regional distribution. In the camera-trap surveys we
established 25–35 camera stations, each with two camera
traps facing each other across foot trails or low-transit dirt
roads (Table 1). The camera pairs provided simultaneous
photographs of both sides of individual animals, facilitating
the identification of individuals and determination of
abundance (Karanth & Nichols, 2002; Maffei et al., 2002;
Paviolo et al., 2008). To generate presence and relative
abundance data we walked transects daily to record jaguar,
puma and prey data through direct observations and
through signs such as tracks, claw marks on trees and
scats (Carrillo et al., 2000; Silveira et al., 2003). To ensure
correct identification of jaguar and puma tracks we
photographed them with a size reference and applied
automated software for identifying tracks using keys and
discriminant models (De Angelo et al., 2010). The number
of km surveyed at each site (Table 1) was determined by the
number of footpaths and abandoned roads available, and
We conducted shorter preliminary surveys at the three
sites to familiarize ourselves with the area and adjust our
methodologies (Table 2). In the Aborigen Reserve we
conducted two preliminary surveys, using 75–90 track plots,
to generate presence data. We cleared 1 × 1 m square track
plots 50 m apart, sifting soil to create a better surface for
detecting tracks, along footpaths that were 2 km long and at
least 6 km apart. We checked track plots once per day,
identifying and counting any tracks and then clearing the
plots again, for 4–6 days.
During the coarse-scale survey in 2004–2010 we
conducted informal interviews (Marker et al., 2003; Figel
et al., 2011) over a vast area, to generate jaguar presence–
absence data and to compare our results with the only
previous information for the species in the region



V. A. Quiroga et al.
TABLE 2 Methods and survey periods across the three principal sites.
Preliminary survey

Complete survey






El Cantor

Walking trails for sign,
Track-plots, walking
trails for sign, interviews
Camera traps

June 2010

Camera traps, walking trails
for sign, interviews
Camera traps, walking trails
for sign, interviews

June–September 2010

Camera traps, walking trails
for sign, interviews

September–November 2009

Aborigen Reserve

Copo National Park Walking trails for sign,

June & October
July 2009

June–September 2008

TABLE 3 Jaguar records for the three principal survey sites, by methodology.

Camera trap

Tracks from transect surveys & track plots

Interview records (2004–2010)

El Cantor
Aborigen Reserve
Copo National Park


1 (+2 records during pilot survey in 2006)

Yes (but no records post 2008)

(Altrichter et al., 2006). We interviewed 156 people,
including local residents, rural teachers, park guards and
other researchers in the region. These interviews were
conducted opportunistically over a total of 20 field trips,
both during jaguar-specific field research and during
research on other species. Twelve field trips of 15–45 days
each were focused specifically on interviews. Interviews were
undertaken in three regions (Fig. 1) that included the
camera-trap sites and the region identified by Altrichter
et al. (2006) as having the highest probability of jaguar
presence (Fig. 1). In the interviews we asked about (1) current
and historical jaguar presence in the area, (2) the number
of years since the last encounter (direct sighting or tracks)
with the species, (3) conflicts between jaguars and livestock,
(4) the number of jaguars killed recently or historically in
the area and the motives for the killings, and (5) skins or
other remains of jaguars killed. Finally, we asked about the
interviewee’s general knowledge of the jaguar, to determine
whether they knew the species.
The interviews, rather than using structured questions,
were part of a general, informal conversation in which we
solicited information about jaguars as far as cordial
exchange with the interviewee permitted. The types of
questions and the way we posed them varied according to
the interviewee and the situation. We were previously
acquainted with most of the local inhabitants and they
trusted us sufficiently to provide sensitive information,
particularly about jaguars hunted or conflicts with the
species. With respect to jaguar tracks reported by local
people, without having photographs or plaster casts we
could not confirm identifications using the discriminant
analysis that we applied to the track data that we collected.

Nevertheless we collected these data because local knowledge is particularly important in complementing other data
when monitoring rare and elusive species, recognizing that
there are higher degrees of error when comparing with other
types of data. To avoid biases we considered data to be valid
only if provided by people whom we knew were knowledgeable about the species and could clearly describe the
typical characteristics that facilitate discrimination between
jaguar and puma tracks, or provide clear descriptions of how
the prey was killed and consumed. If a record was doubtful
but we could confirm the information, we did so before
including the record in our sample. Data from unreliable
informants and which we could not corroborate were
discarded. Thus our records from informants were collected
in a similar manner to those reported by Altrichter et al.
(2006), with which we compare our results.

We did not obtain camera-trap records of jaguars at any of
the sites. Only in the Aborigen Reserve and in El Cantor did
we ourselves observe jaguar tracks. Three of the track
records were in the Aborigen Reserve, one in each track plot
survey (June and October 2006) and the other during the
2008 camera-trap survey at this site. The other track record
was at El Cantor during 2010 (Table 3). Both records from
2006 were identified as jaguar tracks with a probability
. 93%. The 2008 and 2010 track records were identified as
jaguar tracks with a probability . 99%. In each case the
tracks were from a single individual recorded only once
during the field survey.
© 2013 Fauna & Flora International, Oryx, 1–8

The jaguar in the Argentine Chaco

Of 156 people interviewed, only 20% reported jaguar data
for 2004–2010, and 80% said they had not seen jaguars or
signs for many years or that jaguars no longer existed in the
region. A total of 31 independent records were obtained
through interviews. Together with our four track records,
we obtained 35 independent records of jaguar presence
(Fig. 1). The majority of the records since 2007 were limited
to the northern and eastern parts of the study area (Fig. 1),
despite a similar survey effort during the same period in
the south. Of the 35 records, 13 were direct observations of
the species, with confirmation of at least one female with a
cub (central Formosa Province, 2005), a juvenile (Aborigen
Reserve, 2008) and a further two solitary females (one in
Loro Hablador Provincial Park in 2004 and the other near
the Aborigen Reserve, both in Chaco Province). Of the three
records of livestock predation, one was in 2004 near the
Loro Hablador Provincial Park and the other two were in
Formosa during 2010, one at a ranch in the Bañados de la
Estrella and the other at a ranch near Posta Zalazar, 4 km
and , 10 km from the Paraguayan border, respectively. Two
records are of jaguars killed in 2007: one in Palmar Largo,
Formosa (40 km north of El Cantor) and the other north of
the Aborigen Reserve at El Recreo ranch, Chaco Province,
along the Bermejito River. The remaining 17 records were
tracks: four we confirmed ourselves and 13 were reported by

Of the 79,000 km2 that Altrichter et al. (2006) described as
the distribution of the jaguar in the Argentine Chaco up to
2003, we surveyed c. 60% during 2004–2010. Our range map
and the overall pattern of jaguar presence records are
broadly similar to the findings of Altrichter et al. (2006). We
could therefore conclude that the jaguar population has not
suffered any important change between the two studies.
However, distribution maps based on such records may
indicate the existence of transients or survivors rather than
a stable population if only wandering males occur in the
area, for example (Boydston & López González, 2005;
Hoogesteijn & Hoogesteijn, 2005; Conde et al., 2010).
Presence data alone do not facilitate a precise evaluation of
population status, particularly when generated only from
interviews. For this reason we emphasize the importance
of specific and systematic studies to confirm the status of
particular populations, especially when data are required
to make management decisions for the conservation of
threatened species (Karanth et al., 2011b). Currently, general
and wide-ranging studies (Altrichter et al., 2006; De Angelo
et al., 2011) or rapid biological inventories (Torres & Jayat,
2010) are commonly employed to provide baseline information on the conservation status of particular species or
for decision-making on management measures for
© 2013 Fauna & Flora International, Oryx, 1–8

conservation. Although such studies are useful for generating information quickly or over large areas, they should
never replace intensive focused studies. Because it is
superficial and imprecise, coarse-scale information does
not clearly reflect the status of a species (Silveira et al., 2003;
Karanth et al., 2011b).
Prior to our study it was presumed that the Chaco
population was one of the largest in Argentina because of
the vast area of the region, the high percentage of native
forest cover, the low human population density and
the previous range maps based on presence information
(Altrichter et al., 2006; Di Bitetti et al., in press). This
perception was incorrect: the population is the most
threatened in the country. Our survey was one of the most
extensive jaguar studies conducted to date (Maffei et al.,
2004, 2011a,b; Paviolo et al., 2008; De la Torre & Medellín,
2011). However, the camera-trap capture rate for jaguars in
similar studies in the Bolivian Chaco (Maffei et al., 2004),
and even in regions with much lower jaguar density, such as
Argentina’s Atlantic Forest (Paviolo et al., 2008), was 2–20
and 1.34–14 photographs per 1,000 trap nights, respectively,
in contrast with no records in 5,326 camera-trap nights in
our study. Jaguars may live at lower densities in the
Argentine Chaco compared to more humid areas, assuming
lower resource availability in this arid region. However,
given that the Bolivian Chaco has similar rainfall and
environmental conditions to the sites surveyed in the
Argentine Chaco, the lack of camera-trap photographs
indicates an extremely low jaguar density and perhaps
extinction at some sites. Probable causes are the arid
environment, lower prey availability, historically high
hunting pressure and the intrinsic vulnerability of living at
the boundary of the species’ current range (Gaston et al.,
1997; Yackulic et al., 2011).
A relatively reliable indicator of the presence of the
jaguar is the frequency of hunted animals reported in
interviews, particularly in areas where jaguars are systematically eliminated by local people, as is the case in the
Chaco. Comparing the frequency of each type of presence
record in our study with the frequencies reported by
Altrichter et al. (2006) for the period 1993–2003, the hunting
records are those that diminish the most between the
two periods (χ2 5 6.05, df 5 1, P , 0.02). Hunted animals
represent only 2% of our records from interviews, compared
with 23% of records a decade ago. In the Argentine Chaco
the jaguar population has declined as a result of hunting,
such that well-conserved forest sites no longer maintain
jaguars (Altrichter et al., 2006). Hunting of jaguars and
other large mammals has been part of the lifestyle of Chaco
residents since the region was colonized (Altrichter, 2006)
and the absence of jaguars is directly related to the age of
ranch settlements (Altrichter et al., 2006). This implies that
jaguar density should be higher within protected areas
relative to less well-protected areas (Paviolo et al., 2008).



V. A. Quiroga et al.

During 2002–2010 there were notable environmental
changes in the Chaco (Torrella & Adámoli, 2006; Gasparri
& Grau, 2009). Jaguar distribution may also be affected by
large-scale habitat loss and the fragmentation of remaining
forest patches into small parks and reserves with little or no
connectivity (Matteucci, 2009; De Angelo et al., 2011).
Studies of threatened large felids suggest a general
pattern of higher probability of presence and higher
abundance in relatively large areas that are well conserved,
effectively managed and interconnected (Maffei et al., 2004;
Novack et al., 2005; Datta et al., 2008; Paviolo et al., 2008;
Rabinowitz & Zeller, 2010; Yackulic et al., 2011). However,
the results of the survey conducted in Copo National Park in
2009 suggest that the jaguar population is extremely small
or that the species may recently have gone extinct in this
protected area. Given that Copo National Park is the
southernmost limit of the species’ distribution in the Chaco,
of the three survey sites it could be the area most vulnerable
to extinction (Brown, 1984; Gaston et al., 1997; Laliberte &
Ripple, 2004). Copo is surrounded by ever more intensive
productive activities and is becoming isolated by forest
clearing for farming and ranching from the neighbouring
Loro Hablador Provincial Park, which had served as a
corridor connecting Copo with forested areas to the north
(Matteucci, 2009). A corridor among protected areas in the
region would reduce the isolation of Copo and potentially
facilitate recolonization of the Chaco. For this reason
the conservation and management of Copo are regional
The advance of the agricultural frontier in the Argentine
Chaco means not only habitat loss and fragmentation but
also a more extensive network of roads, and thus facilitates
access by hunters (Morello et al., 2006). This has probably
put greater pressure on the jaguar, as has occurred elsewhere
(Maffei et al., 2004). The status of the jaguar in the
Paraguayan and Bolivian Chaco is notably different. Jaguar
photographs have been obtained with much lower cameratrap effort and direct and indirect observations are common
(Maffei et al., 2004; McBride, 2009). The marked difference
between countries in this respect could be a result of their
historical differences in land use. The Argentine Chaco has a
longer history of colonization by descendants of Europeans,
who maintained a strong hunting tradition (Altrichter,
2006; Morello et al., 2006). In contrast, the Paraguayan and
Bolivian Chaco still contain vast areas that are uninhabited
and relatively inaccessible (Taber et al., 1997; Maffei et al.,
2004; Rumiz et al., 2011). The long-term presence of cattle
in the Argentine Chaco has probably also affected
jaguars, because ranchers hunt jaguars in response to real
or perceived conflicts with livestock (Hoogesteijn &
Hoogesteijn, 2005; Arispe et al., 2009).
Nevertheless, of the total area of Chaco forest that is
potential jaguar habitat, our camera-trap surveys covered
, 1.5%. Other sites in the region must be surveyed as a

matter of urgency, to determine whether the jaguar is now
rare across the whole semi-arid Chaco. The creation of
corridors linking existing protected areas and the effective
management of corridors and protected areas so that they
provide refuge and protection for the species are high
priorities. Recent conservation efforts include the proposal
for a new National Park in La Fidelidad (2,500 km2). At only
20 km east of the Aboriginal Reserve (Fig. 1), there are
jaguars present in the forest of La Felidad, but the status of
this population has not yet been evaluated (Rumiz et al.,
2011). The establishment of this Park would provide jaguars
with large and well-conserved core areas, which could
maintain populations. The protection of the few remaining
well-conserved forests that connect the Argentine Chaco
with the Yungas in Argentina is also important. Such
corridors, to the Yungas as well as to the Paraguayan Chaco,
ensure connectivity of jaguars in the Argentine Chaco with
the southernmost populations of the species (Sanderson
et al., 2002; Rabinowitz & Zeller, 2010; Rumiz et al., 2011).
We recommend that the government of Argentina takes
measures to conserve jaguar habitat in the Chaco and
protect the species and its prey from illegal hunting. The
environmental conditions of the Argentine Chaco remain
suitable for maintenance of a viable jaguar population if
illegal hunting can be controlled and deforestation halted.
We thank the volunteers who helped with field work;
the Agency of Wildlife, Parks and Ecology, Chaco
Province; the Ministry of Production and Environment,
Formosa Province; Copo National Park, the National
Parks Administration; the National University of Córdoba;
the National Wildlife Service, National Environment
and Sustainable Development Secretariat. Financial and
logistical support was provided by CONICET-Argentina,
Jaguar Conservation Program-WCS, Rufford Small Grants
Foundation, Cleveland Metroparks Zoo and Cleveland
Zoological Society, Mohamed bin Zayed Species
Conservation Fund, Idea Wild, Elé Project, Stephen
F. Austin State University, Yaguareté Project Misiones and
Tapir Project Salta. We thank Pablo Perovic, Ricardo
Banchs, Gustavo Porini, Adrián Díaz, Daniel Scognamillo,
Andrés Ravelo, Agustín Paviolo, Peter Feinsinger and Susan
Walker for their support and advice at different stages of
this project.
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Biographical sketches
V E R Ó N I C A A N D R E A Q U I R O G A has conducted ecological and
conservation research on large mammals in Argentina’s semiarid Chaco for the past 10 years. She participates in a range of
conservation activities for species and ecosystems in Argentina.
G A B R I E L I V Á N B O A G L I O has conducted ecological and conservation
research on large mammals in Argentina’s semi-arid Chaco for the
past 11 years. He also studies bird and mammal populations in
the Cordillera region of western Argentina. A N D R E W J A Y N O S S
has worked with indigenous peoples in the Bolivian Chaco and
Ecuadorian lowlands for the past 15 years, on territorial management,
conservation of threatened species, including jaguars, and sustainable
use of natural resources. 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 subtropical forests and grasslands of
northern Argentina for the past 20 years, focusing on the behaviour,
ecology and conservation of mammals, particularly primates and

© 2013 Fauna & Flora International, Oryx, 1–8

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