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Journal for Nature Conservation 31 (2016) 9–15

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Journal for Nature Conservation
journal homepage: www.elsevier.de/jnc

Puma density, habitat use and conflict with humans in the Argentine
Verónica A. Quiroga a,b,∗ , Andrew J. Noss c , Agustín Paviolo a,b , Gabriel I. Boaglio d ,
Mario S. Di Bitetti a,b,e
Instituto de Biología Subtropical (IBS), Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET) and Universidad Nacional de Misiones
(UNaM), Bertoni 85, Puerto Iguazú, Misiones CP 3370, Argentina
Asociación Civil Centro de Investigaciones del Bosque Atlántico (CeIBA), Bertoni 85, Puerto Iguazú, Misiones CP 3370, Argentina
Department of Geography, University of Florida, 1405 NW 38th St., Gainesville, FL, USA
Instituto de Diversidad y Ecología Animal (IDEA), Universidad Nacional de Córdoba (UNC), Av. Vélez Sarsfield 299, Córdoba Capital, Córdoba CP 5000,
Facultad de Ciencias Forestales, UNaM, Bertoni 124, Eldorado, Misiones CP 3380, Argentina

a r t i c l e

i n f o

Article history:
Received 27 September 2015
Received in revised form 22 February 2016
Accepted 23 February 2016

a b s t r a c t
The puma Puma concolor is the most widely distributed felid in the Americas. Although it utilizes humanmodified landscapes, its extensive territorial requirements, trophic needs, and real or perceived threats to
livestock render the puma susceptible to conflict with humans. Our objectives were to evaluate the population density, habitat use, and puma-human conflict in the Argentine Chaco. We conducted camera-trap
surveys and interviews over a three-year period, at three sites with different levels of legal protection and
with different ranch outpost and livestock densities: Copo National Park (1204 trap days, 24 stations, 17
interviews, national park, lowest ranch/livestock density), Aborigen Reserve (1993 trap days, 29 stations,
13 interviews, indigenous reserve, medium ranch/livestock density) and El Cantor (2129 trap days, 35
stations, 11 interviews, no protection, highest ranch/livestock density). Puma population density was
low (<1 individual/100 km2 ) and we found no significant differences in puma density across the three
sites. Occupancy models show a positive relationship between puma detectability and the distances from
vehicle roads. Legal protection status of the área does not positively affect puma density, probably due to
the large edge effect, and weak law enforcement capacity at Copo National Park. Low density of pumas
at the three sites could result primarily from retaliation killing of pumas by local ranchers in response
to predation on goats. Pumas in the Chaco require effectively managed protected areas, regulation of
wildlife hunting and livestock management practices to minimize depredation.
© 2016 Elsevier GmbH. All rights reserved.

1. Introduction
The puma Puma concolor is the most wide-ranging felid in the
Americas, from Canada in North America to Patagonia in South
America (Currier, 1983; Shaw, Beier, Culver, & Grigione, 2007).
The species is relatively tolerant to anthropogenic disturbance and
maintains populations in some human-modified landscapes (De

∗ Corresponding author at:, Instituto de Biología Subtropical (IBS), Consejo
Nacional de Investigaciones Científicas y Técnicas (CONICET) and Universidad
Nacional de Misiones (UNaM), Bertoni 85, Puerto Iguazú, Misiones CP 3370,
E-mail addresses: veroquiroga@gmail.com, veroquiroga@hotmail.com
(V.A. Quiroga), anoss@ufl.edu (A.J. Noss), paviolo4@gmail.com (A. Paviolo),
gabrielboaglio@yahoo.com.ar (G.I. Boaglio), dibitetti@yahoo.com.ar
(M.S. Di Bitetti).
1617-1381/© 2016 Elsevier GmbH. All rights reserved.

Angelo, Paviolo, & Di Bitetti, 2011; Haines, 2006). However, similar
to other large carnivores, individual and population fitness is weakened as human disturbance reduces natural prey availability and
fragments suitable habitat, forcing pumas to increase their already
extensive territorial requirements and cross patches of unsuitable
habitat (Gittleman & Harvey, 1982; Ripple et al., 2014; Wilmers
et al., 2013). As a top predator, the puma also enters into conflicts
with livestock owners, and is persecuted because of its perceived or
actual predation of livestock (Kissling, Fernandez, & Paruelo, 2009;
Soto-Shoender & Giuliano, 2011). At the same time, the puma is
valued as a trophy, and direct hunting pressure is one of its principal conservation threats across its range (Carvalho & Pezzuti, 2010;
Mosa & Goytia, 2004; Paviolo, Di Blanco, De Angelo, & Di Bitetti,
The implementation of protected areas is one of the most
widespread strategies for the conservation of biodiversity. How-


V.A. Quiroga et al. / Journal for Nature Conservation 31 (2016) 9–15

ever, if the protected areas are not correctly designed and
implemented they may be ineffective in conserving populations of
large carnivores. Home ranges of resident carnivores often extend
beyond the edges of protected areas, where they suffer humaninduced mortality (Balme, Slotow, & Hunter, 2010; Woodroffe &
Ginsberg, 1998). This “edge effect” could drive important changes
in the abundance and population dynamics of carnivores inside
protected areas (Balme et al., 2010; Newby et al., 2013; Revilla,
Palomares, & Delibes, 2001). In addition, many countries lack
the resources to fully implement legally protected areas, thereby
diminishing biodiversity conservation outcomes. In areas where
poaching is widespread and law enforcement capacity is scarce,
puma densities can be significantly reduced (Paviolo et al., 2009).
Poaching or hunting not only can affect population numbers, but
also can have an important impact on carnivore behavior (Newby
et al., 2013; Ordiz, Støen, Delibes, & Swenson, 2011; Ordiz et al.,
2012; Paviolo et al., 2009; Valeix, Hemson, Loveridge, Mills, &
Macdonald, 2012). The “landscape of fear” phenomenon was originally described for herbivores that tried to avoid natural predators
by reducing their use of areas with high predation risk (Laundré,
Hernandez, & Anteldorf, 2001). This hypothesis was tested subsequently in other species and systems, confirming that carnivores
also change their activity patterns, movement patterns, and habitat use in high-risk areas (Ordiz et al., 2011, 2012; Paviolo et al.,
2009; Valeix et al., 2012).
The Chaco is the second largest eco-region of South America
after the Amazon, and the largest dry forest in the world (Morello
& Adámoli, 1974; Morello, Rodríguez, & Silva, 2009). Jaguars (Panthera onca) and pumas are the top predators in the region, and were
originally distributed throughout the Chaco. But the jaguar population has been reduced drastically by hunting and habitat loss,
virtually disappearing in the southern part of the eco-region, in
the Argentine Chaco (Quiroga, Boaglio, Noss, & Di Bitetti, 2014).
Albeit with a higher tolerance, we expect the puma to react in a
similar way as the jaguar to human disturbance (Currier, 1983; De
Angelo et al., 2011). The objective of this study was to evaluate
puma population density, habitat use and puma-human conflict in
the Argentine semi-arid Chaco at three sites across a gradient of
legal protection status and human disturbance.

2. Materials and methods
2.1. Study area
The semi-arid Chaco is located in the central part of the ecoregion and includes parts of Argentina, Paraguay and Bolivia. The
portion in Argentina covers approximately 270,000 km2 and is
characterized by extensive plains of dry forests dominated by red
quebracho (Schinopsis lorentzii), white quebracho (Aspidosperma
quebracho-blanco), palosanto (Bulnesia sarmientoi), and mistol (Ziziphus mistol). These canopy trees reach 18 to 20 m in height. The
dense shrub understory, from 1 to 10 m high, is dominated by
species of the Capparidaceae family and by thorny species like Acacia praecox and Celtis pallida (Tálamo & Caziani, 2003). The climate
is markedly seasonal, with a mean annual temperature of 24 ◦ C
and annual rainfall between 400 and 800 mm, concentrated from
October to April. Three rivers traverse the semiarid Argentine Chaco
from northwest to southeast, while the remainder of the region is
devoid of permanent water sources—giving the central portion of
region its name of “Impenetrable”—with the exception of artificial
ponds constructed for livestock (Caziani et al., 2003).
The region was colonized in the early 20th century by “criollos” who settled in isolated ranch outposts (permanent settlements
with 1–5 family houses, livestock corrals but otherwise unfenced,
and boreholes), 3–15 km apart from each other. The settlers estab-

lished extensive cattle ranching systems and today raise goats
(10–200 animals per outpost) and cows. Goats are enclosed in corrals only at night and by day range freely, susceptible to puma
predation. Wildlife hunting is widespread, both by local residents
and by hunters from nearby towns (Altrichter, 2006).
We compared puma population density, habitat use and pumahuman conflict across three sites that vary in legal protection status
as well as in ranch outpost and livestock density (Fig. 1). Within
each site, hunting and livestock pressures are associated with the
proximity of ranch outposts and roads, which provide access to the
forest by people and livestock.
1) Copo National Park (1180 km2 ): this site has the highest legal
protection level, with four park rangers (1 park ranger/295 km2 )
responsible for anti-poaching and other activities within the
park. Several small human settlements exist near the Park borders, with a ranch density of 0.4 outposts/100 km2 . Livestock
from neighboring ranch outposts enters the Park, but the livestock burden is relatively low in comparison to the other sites
(71% of camera trap stations with photos of cows). Lacking roads
and trails, the interior portion of Copo National Park is inaccessible to hunters. However, two roads constitute the northern
and eastern boundaries of the Park. We did not quantify the
frequency, but commonly observed hunters along these boundaries. The camera trap survey covered 367 km2 in the northern
portion of the Park, including both internal areas (i.e., inaccessible to poachers) and edges.
2) The Aborigen Reserve (2500 km2 ): although it is categorized
as an indigenous Reserve, no indigenous people reside inside
its boundaries. It is sparsely populated by criollos (0.8 outposts/100 km2 ), and has an intermediate livestock burden
relative to the other two survey sites (93% of camera trap stations
with photos of cows). The Reserve has no game rangers and no
anti-poaching activities. Three roads cross the Reserve and facilitate access by hunters. The camera trap survey covered 455 km2
in the center of the Reserve.
3) El Cantor (1966 km2 ): this site is not legally protected, and has no
game rangers nor anti-poaching activities. The resident criollos
hunt. The cattle stocking rate is the highest of the three sites
(100% of camera trap stations with photos of cows), and cattle
have been established the longest here. The density of ranches
is also the highest, at 1.3 outposts/100 km2 . Five roads cross the
area and facilitate access by hunters. The camera trap survey
covered 363 km2 in the center of El Cantor.
The percentage of stations with presence of cows varies significantly among sites (X-squared = 13.71, df = NA, p-value = 0.001),
and is lowest in Copo National Park. In the three sites the number
of goats is proportional to the number of cattle. However, goats
rarely move further than 2 km away from the ranch outposts, and
did not use the trails, therefore we rarely recorded them during
the camera-trap surveys. Nevertheless, they may attract pumas to
the posts and are occasionally preyed upon by pumas. In the case
of Copo National Park the goats rarely enter the park, but their
presence is conspicuous near its borders.
2.2. Camera-trap surveys
We undertook camera-trap surveys during three consecutive years (2008–2010), during a three-month period each year
(between June and November according to the site). At each site
we installed 24–35 camera-trap stations separated by an average of
3 km, using two cameras, 30 cm above the ground, facing each other
across foot trails or dirt roads in each station. We used a pool of analog 35 mm film cameras (Deer Cam® Model DC 300, Cam Trakker® ,
and Leaf River Trail Scan® Model C-1) at all three sites, supple-

V.A. Quiroga et al. / Journal for Nature Conservation 31 (2016) 9–15


Fig. 1. The three study sites in the Argentine semi-arid Chaco.

mented by digital (Cuddeback® Expert) cameras at El Cantor. In
each survey the camera models were interspersed across the survey area in order to avoid any biases that could result from differing
sensibility of the camera models. Camera-traps were active 24 h a
day, with a minimum one-minute delay between consecutive photos. Sensitivity was set at medium or high depending on camera
model and previous experience. Camera-traps were active for an
average of 58 days at each site (Table 1). We checked the cameras
weekly to change film, batteries and memory cards as needed.
We used the camera-trap photographs to confirm puma presence, to determine the number of individuals, and to estimate puma
density according to the captures and recaptures of the distinct
individuals. We used the same photos in combination with occupancy models (MacKenzie et al., 2002) to estimate the proportion
of the study area utilized by pumas, and the detectability of the
species relative to environmental variables indicating human disturbance. In the case of species with spotted pelts, like the jaguar,
individuals can be distinguished by spot patterns (Karanth, 1995).
In the case of pumas, individuals are more difficult to distinguish,
but can also be identified following established protocols (Kelly
et al., 2008; Paviolo et al., 2009; Rich et al., 2014). Two investigators worked independently to identify the individual pumas in the
photos and in turn to estimate population density for each site. They
considered only adult individuals, and used unique marks such as
the shape and color of the tail and its tip, how the tail is carried,
scars, and spots on the inside of the legs (Kelly et al., 2008; Rich
et al., 2014). We used the presence or absence of scrotum to define
the sex of the photographed individuals.

2.3. Density estimations using capture-recapture models
To estimate puma population density, we applied both traditional (Karanth & Nichols, 2002), as well as spatially-explicit (Royle,
Karanth, Gopalaswamy, & Kumar, 2009), capture-recapture models. We used both approaches in order to be able to compare our
density estimates with previous studies elsewhere across the continent. Both models use the number of individuals identified and
the capture-recapture history at each camera-trap station. In the
traditional models, we first estimated puma abundance in the survey area using the software CAPTURE 2 (version <050810.1025>
by Jim Hines; http://www.mbr-pwrc.usgs.gov), and in turn estimated the effective survey area by applying a buffer around the
camera-trap polygon equivalent to the Mean Maximum Distance
Moved (MMDM) by all individuals captured in more than one station (Karanth, 1995; Karanth & Nichols, 2002; Kelly et al., 2008;
Maffei, Noss, Silver, & Kelly, 2011). Consistent with most other studies on large carnivores, we used the heterogeneity model (Mh ) to
estimate abundance, as this model best approximates the species’
behavior (Karanth, 1995), and we used the jackknife estimator,
which is more robust than other estimators (Silver et al., 2004).
Spatially-explicit capture-recapture (SECR) models estimate population density directly by using the capture-recapture history in
combination with the spatial distribution of the captures and recaptures (Noss et al., 2012; Reppucci, Gardner, & Lucherini, 2011; Royle
et al., 2009; Singh, Gopalaswamy, Royle, Kumar, & Karanth, 2010).
We used the software SPACECAP (version 1.0.1), which applies
a Bayesian modeling framework (Gopalaswamy et al., 2012). In

Table 1
Camera trap survey effort for pumas at three Argentine semi-arid Chaco sites.

Survey dates

N◦ of stations Trap-days (systematic survey) Total trap-days (including pilot) Mean distance among cameras (km ± SD)

El Cantor
1/Jul/2010 to 9/Sep/2010
23/Jun/2008 to 7/Sep/2008 29
Aborigen Reserve
Copo National Park 4/Sep/2009 to 19/Nov/2009 24



2.99 ± 0.32
3.04 ± 0.98
2.85 ± 0.65


V.A. Quiroga et al. / Journal for Nature Conservation 31 (2016) 9–15

accordance with these previous studies, for each area we ran
100,000 iterations. To define the extent of the state space for the
analysis we selected an area that included all the cameras plus a
buffer area of 50 km. That distance is approximately 10 times the
average radius of home ranges for four radio-collared male pumas
in the Paraguayan Chaco (the closest region for which such information is available, McBride, 2009). The probability of recording an
animal from outside this area during the survey period is very low
(Gopalaswamy et al., 2012). We applied a standard pixel area of
1 km2 for each potential activity center; this is a reasonable size to
ensure many potential activity centers per home range, as well as
reasonable computational times (Gopalaswamy et al., 2012). Camera trap data confirm that pumas do move large distances across the
study sites, therefore a pixel smaller than 1 km2 would be unnecessary. We based the augmentation values (150–300 individuals) on
the number of puma observations obtained in each area, adjusted
in accordance with the density plot for and N. We based the burnin period (20,000–35,000 iterations) on the values of the Geweke
test (with a z-score between −1.6 and +1.6.) (Gopalaswamy et al.,
2012). For each estimation, we ran the models 2–4 times in order
to make the appropriate adjustments for the final analysis.
2.4. Occupancy models
Occupancy models can be used to reveal what factors affect,
positively or negatively, site occupancy or proportion of sites used
( ), modeling the effects of these factors according to the species’
detection probability (p) (MacKenzie et al., 2002, 2006). In our analyses we estimate the proportion of sites used by pumas (with the
software PRESENCE version 4.4; Hines, 2006). We considered the
88 camera-trap stations across the three surveys to be distinct survey points, and we assumed no temporal changes in population
abundance among sites. During the three-year study period no
unusual climatic or environmental events occurred in the study
area. For each station we grouped the maximum number of 64 survey days in eigth capture occasions of eigth days each. If a station
was active fewer days overall, then we used the greatest multiple of
eigth possible, and eliminated the extra days. We ranked models by
their Akaike Information Criteria (AIC) weights. Instead of choosing
the single highest-ranked model, we estimated an average of the
highest-ranked models (those with AIC weights 10% or greater of
the weight of the highest-ranked model) (MacKenzie et al., 2006).
We evaluated the effects of three co-variables on the proportion of
sites used ( ) or detectability (p) of pumas:
(1) The distance (km, range 0.5–13.8) to the nearest ranch outpost.
We expected this distance to be positively related to and p,
because of direct puma hunting by ranch inhabitants, and/or
the combined disturbance effects of the ranch—including
human presence, dogs, cattle, goats, and vegetation modification (Altrichter, 2006).
(2) The distance (km, range 0–15) to the nearest vehicle road. We
expected that and p would vary positively with this distance,
because of the hunting pressure concentrated along vehicle
roads (e.g., Boas Goulart et al., 2008). For these two co-variables,
distance to vehicle road and distance to nearest ranch outpost,
we applied one-tailed statistical tests because our hypothesis
was directional.
(3) The type of trail/road (vehicle roads, abandoned roads and narrow footpaths) where the camera-trap station was installed.
There is a generalized preference of felids for using roads or
trails rather than bushwhacking (Cusack et al., 2015), and they
are known to prefer unpaved roads over narrow footpaths. We
therefore predicted this co-variable would affect p because the
pumas, when present, would more frequently use vehicle roads

Table 2
Comparison of 95% confidence intervals for density estimates (individuals per
100 km2 ) of pumas using traditional and spatially-explicit capture-recapture models, at three Argentine semi-arid Chaco sites.

Traditional model

capture-recapture model

Investigator 1 Investigator 2 Investigator 1 Investigator 2
El Cantor
Aborigen Reserve 0.51–0.97
Copo National Park 0.31–0.84




or abandoned roads than footpaths (e.g., Di Bitetti, Paviolo, & De
Angelo, 2006; Di Bitetti, De Angelo, Paviolo, & Di Blanco, 2008).
To build the models we tested the influence of the three variables at the same time on either or p.
Wild prey availability was not considered as an influential
covariate in our analysis since the most important puma’s prey
(e.g., Mazama gouazoubira, Sylvilagus brasiliensis, Dolichotus salinicola, Tolypeutes matacus, among others) are relatively abundant in
the study sites (Quiroga, V. A., 2013). Thus, we do not consider prey
availability to be an important and limiting factor for pumas in the
study areas, and therefore was not included in the analysis.
2.5. Interviews
During the systematic camera-trap surveys we conducted 41
informal semi-structured interviews with rural residents of the
three surveyed areas: 17 in Copo National Park, 13 in Aborigen
Reserve and 11 in El Cantor. All the people interviewed were adult
male and had livestock of their own. During the interviews we
asked about poaching of pumas and conflicts due to predation on
livestock during the previous year, potential causes of conflicts,
and solutions to solve the conflicts. The main goals of the interviews were to generate information about the frequency of puma
poaching in the different areas and help us understand the level of
puma-human conflict.
3. Results
3.1. Puma population density
We identified, from the camera trap photos, between 10 and
12 pumas at Copo National Park, between 7 and 8 at the Aborigen
Reserve, and 9 at El Cantor. We recorded adult males and females
as well as kittens and/or juveniles at all three sites. We recorded
between 4 and 5 females, between 5 and 6 males and 1 unsexed
individual at Copo National Park; 3 females and between 4 and
5 males at the Aborigen Reserve; and between 4 and 5 females
and between 3 and 5 males at El Cantor. Although the two investigators did not coincide in the number of individuals identified at
the two first sites, the density estimations using traditional models (and MMDM) as well as SECR models did not vary significantly
between the two investigators (95% confidential intervals overlapped). Also, the densities estimates were not statistically different
(95% confidential intervals overlapped) between the site with legal
protection (Copo National Park) and the sites without legal protection (Table 2).
3.2. Occupancy models
Using the model in which all sites had the same probability of
species presence, and in which the detection probability was constant across time and sites [ (.); p(.)], the proportion of sites used
by pumas ( ± SE) across the entire study area was 0.75 ± 0.07,

V.A. Quiroga et al. / Journal for Nature Conservation 31 (2016) 9–15
Table 3
Occupancy models for puma in the Argentine Chaco ranked by their Akaike Information Criteria (AIC), considering only models with AIC weights >10% of the highest
ranked model.
(dis ranch, dis road), p(dis ranch, dis road)
(dis ranch, dis road), p(trail, dis ranch, dis road)
(.), p(dis road)
(.), p(trail, dis road)
(dis ranch, dis road), p(dis road)
(dis ranch, dis road), p(trail, dis road)
(.), p(dis road, dis ranch)
(dis ranch), p(trail, dis road)
(dis ranch), p(dis road)
(.), p(trail, dis ranch, dis road)
(dis ranch), p(dis ranch, dis road)
(dis ranch), p(trail, dis ranch, dis road)


AIC AIC wgt no.Par.





Table 4
Parameter estimates of the average model. The weighted averaged model is estimated using the highest-ranked model and models with AIC weights 10% or greater
of the weight of the highest-ranked one.
Beta model averaging

Site use ( )
Intercept estimate
Distance to the nearest ranch
Distance to the nearest road −1.93
Detectability (p)
Intercept estimate
Distance to the nearest ranch −0.18
Distance to the nearest road
Type of trail


95% confidence








with a 95% confidence interval of 0.59–0.86. The average of the
highest-ranked models indicates that the only parameter for which
the estimated 95% confidence interval did not include 0 was the ˇ
for detection probability (p) of pumas as a function of the distance
to the nearest road (Tables 3 and 4).
3.3. Puma-livestock conflict and puma retaliation killing
In Copo National Park all 17 interviewees reported puma predation on goats, and 8 (47% of the interviewees) admitted they
had killed pumas in the past year. Of 13 people interviewed at the
Aborigen Reserve, only one said he had had no puma predation on
livestock in recent years, though he knew pumas had been attacking
goats on his neighbor’s ranch. The other 12 interviewees confirmed
heavy losses by pumas, and at least 7 people (58% of the interviewees) had killed a puma in the past year. Similarly, in El Cantor all
11 interviewees reported puma predation on goats, and at least 5
of them (42% of the interviewees) had tried to kill pumas in the
past year, but reportedly without success. At all three sites, 100%
of interviewees affirmed that predation on goats was frequently by
female pumas teaching their young to hunt, and that conflicts were
more common in the dry season (May–October). All interviewees
also affirmed that the predation problem can be greatly reduced or
even eliminated by using well-trained shepherd dogs to tend the
goat herds and by enclosing the goats in corrals at night. However,
the interviewees explained that dogs are useless if not appropriately trained. The few people in the region who know how to train
shepherd dogs are old women who have inherited this skill from
their mothers or grandmothers. As for night-time corrals, 85% of the
interviewees said they lack the time or help to build and maintain
them, in addition to all their domestic and ranch duties.


4. Discussion
Puma density estimates were very low across the three sites,
and our surveys could not demonstrate a statistically significant
effect of legal protection status on density. It is important to note,
however, that the camera trap survey in Copo National Park may
be capturing important edge effects because it covered only the
northern site of the Park, where roads constitute two of the boundaries of the protected area. In addition, people living in the 10 ranch
outposts neighboring the Park frequently kill pumas in response to
attacks on livestock, as occurs within the other two survey areas.
Individual pumas utilize extensive territories (Beier, 1993; Shaw
et al., 2007; Smallwood, 1997), thus perhaps even those individuals
whose territories encompass the interior of the Copo National Park
move in and out of the park, thereby exposing themselves to the
same threats (for instance, road-based hunters) facing pumas at the
other two sites. Human-induced mortality at the edges of protected
areas can reduce the densities of carnivores, and the negative effect
may extend well inside protected areas (Balme et al., 2010; Davis,
Kelly, & Stauffer, 2011; Revilla et al., 2001; Woodroffe & Ginsberg,
1998). Our study suggests this might be occurring in Copo National
Park. On the other hand, is important to note that the small sample size (only three areas) of our study may also be influencing the
pattern we detected, and therefore the results should be carefully
Other studies undertaken across a variety of eco-regions and
conservation contexts reveal no consistent relationship between
pumas and human disturbance (Davis et al., 2011; De Angelo et al.,
2011; Foster, Harmsen, & Doncaster, 2010; Negrões et al., 2010;
Payán Garrido, 2009; Sollmann et al., 2012). We found only one
variable associated with human disturbance, distance to roads, that
affects puma detection probability (p) in the Argentine Chaco. We
expected that pumas would avoid ranch outposts, because pumas
are frequently killed in retaliation for livestock losses, but our data
did not support this hypothesis. Although the ranches are dangerous places to be avoided, perhaps pumas are drawn to the abundant
easy prey (goats) that concentrate around ranch outposts. Other
large carnivores face similar opposing pressures (Ordiz et al., 2011,
2012; Valeix et al., 2012). In Botswana, lions do not avoid cattle
posts, but use these areas at times when humans are least active,
to minimize the chance of encounters (Valeix et al., 2012). Similarly, brown bears in Scandinavia do not shift their ranges when
the hunting season starts, but change their movement patterns to
avoid encounters with people (Ordiz et al., 2012). We also expected
that pumas would avoid roads as risky places that facilitate access
for hunters from nearby cities and towns (Mosa & Goytia, 2004). On
the other hand, unpaved roads also make walking easier for pumas,
compared to traversing forest or brush, and big cats at many sites
seem to prefer such roads (Blake & Mosquera, 2014; Cusack et al.,
2015; Di Bitetti, Paviolo, & De Angelo, 2014; Harmsen, Foster, Silver,
Ostro, & Doncaster, 2009; Rabinowitz & Nottingham, 1986). In this
case our data do support our hypothesis, thus pumas’ wariness of
humans must outweigh the benefit of walking on roads at our study
sites, either because the risk from humans is especially high in the
Argentine Chaco, or the abundant cattle open numerous trails that
pumas also use, or a combination of the two.
According to the interviewees, puma predation on goats is
very common, especially during the dry season (May–October).
Although they know management measures that can reduce predation (shepherd dogs and night-time enclosures), time and resource
limitations prevent livestock owners from adopting them. As a
result, retaliation killing of pumas because of predation on livestock in criollo ranch outposts is very high, and is probably one
of the main factors causing low puma population density in this
region, as elsewhere across the puma’s range (Conforti & Cascelli de
Azevedo, 2003; Inskip & Zimmermann, 2009; Kissling et al., 2009;


V.A. Quiroga et al. / Journal for Nature Conservation 31 (2016) 9–15

Mazzolli, Graipel, & Dunstone, 2002; Novack, Main, Sunquist, &
Labisky, 2005; Perovic, 2003).
In some well-protected regions puma densities are much
higher: 1.55–2.89 pumas/100 km2 in Iguazú in the Atlantic Forest of Argentina, 7.99/100 km2 in Ravelo in the Bolivian Chaco,
and 3.42/100 km2 in Belize. The low puma densities in the Argentine Chaco are similar to other areas of high hunting pressure in
the Americas: 0.31–0.74/100 km2 in Yabotí in the Atlantic Forest of Argentina and 0.36/100 km2 in Cerro in the Bolivian Chaco
(Kelly et al., 2008; Mazzolli, 2010; Negrões et al., 2010; Noss et al.,
2012; Paviolo et al., 2009; Soria-Díaz, Monroy-Vilchis, RodríguezSoto, Zarco-González, & Urios, 2010). Although we did not find an
effect on puma density of legal protection status across our sites,
we consider that protected areas are essential to puma conservation. However, existing protected areas such as Copo National
Park must be managed more effectively, with sufficient staff and
resources to implement essential conservation measures. These
measures must include wildlife hunting regulation and a reduction of puma-human conflicts in the neighboring properties by
improved livestock management practices, especially for goats.
These management and conservation measures must also be
implemented on communal and indigenous lands such as the
Aborigen Reserve, as well as on private lands. Furthermore, zoning plans exist to conserve native forests and effective biological
corridors where puma and human activities can coexist, allowing pumas to move safely among protected areas. These zoning
plans must be implemented and must include the creation of new
effectively managed protected areas. These recommendations echo
those urgently required to protect jaguars in the region (Quiroga
et al., 2014); both pumas and jaguars (and several others through
trophic cascades) will only survive in the Argentine Chaco if these
recommendations are adopted.
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, 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, Peter
Feinsinger, Yamil Di Blanco, Paula Cruz, Carlos De Angelo and Susan
Walker for their support and advice at different stages of this
project. We particularly thank Yamil Di Blanco for his collaboration
in identifying individual pumas.
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  • Autor: Verónica A. Quiroga
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