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De Angelo et al 2011 Oryx Monitoring of large carnivores .pdf



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Participatory networks for large-scale monitoring of
large carnivores: pumas and jaguars of the Upper
Parana´ Atlantic Forest
C a r l o s D e A n g e l o , A G U S T ´I N P a v i o l o , D a n i e l a R o d e , L a u r y C u l l e n J r
Denis Sana, Kaue Cachuba Abreu, Marina Xavier da Silva
Anne-Sophie Bertrand, Taiana Haag, Fernando Lima
A l c i d e s R i c i e r i R i n a l d i , S i x t o F E R N A´ N D E Z , F r e d y R A M ´I R E Z , M y r i a m V E L A´ Z Q U E Z
C r i s t i a n C o r i o , E s t e b a n H a s s o n and M a r i o S . D i B i t e t t i
Abstract Most large carnivores are secretive and threatened, and these characteristics pose problems for research
on, and monitoring of, these species across extensive areas.
Participatory monitoring, however, can be a useful tool for
obtaining long-term data across large areas. Pumas Puma
concolor and jaguars Panthera onca are the largest predators in the threatened Upper Parana´ Atlantic Forest. To
survey the presence of these two species we established
a participatory network of volunteers and a partnership
with researchers in the three countries that share the
Upper Parana´ Atlantic Forest (Argentina, Brazil and
Paraguay). We trained participants in simple methods of
collecting faeces and track imprints of large felids. Between

CARLOS DE A NGELO (Corresponding author), AGUSTI´N PAVIOLO and
M ARIO S. DI BITETTI National Research Council, Instituto de Biologı´a
Subtropical, Facultad de Ciencias Forestales, Universidad Nacional de Misiones, Puerto Iguazu´, Argentina, and Asociacio´n Civil Centro de Investigaciones del Bosque Atla´ntico, Yapeyu´ 23, CP 3370, Puerto Iguazu´, Misiones,
Argentina. E-mail biocda@gmail.com
DANIELA RODE Fundacio´n Vida Silvestre Argentina, Puerto Iguazu´, Misiones,
Argentina
LAURY CULLEN JR and FERNANDO LIMA Instituto de Pesquisas Ecolo´gicas,
Teodoro Sampaio, Brazil

2002 and 2008 . 100 volunteers helped with monitoring,
obtaining 1,633 records identified as pumas or jaguars across
c. 92,890 km2. We confirmed jaguar presence in a large section
of the Misiones Green Corridor in Argentina and in the largest
protected areas of Brazil and Paraguay. Pumas exhibited
a wider distribution, being recorded throughout Misiones
province in Argentina and in some areas of Brazil and
Paraguay where jaguars were not detected. Both species, and
especially jaguars, were detected mainly in the few remaining
medium and large forest fragments in this Forest. Although
these carnivores are often in conflict with local people, their
charisma and cultural significance makes them flagship species
that motivated the participation of volunteers and institutions.
Participatory monitoring allowed coverage of a vast area at
relatively low cost whilst enhancing collaborative management
policies among people and institutions from three countries.
Keywords Atlantic Forest, distribution, flagship species,

habitat loss, jaguar, large carnivore, participatory monitoring, puma
This paper contains supplementary material that can be
found online at http://journals.cambridge.org

DENIS SANA, Instituto Pro´-Carnı´voros, Atibaia, Brazil
KAUE CACHUBA ABREU Laboratorio de Biogeogra´fia da Universidade Federal
do Parana´, Goia´s, Brazil
M ARINA X AVIER DA SILVA Instituto Chico Mendes de Conservaça˜o da
Biodiversidade, Foz do Iguaçu, Brazil
ANNE-SOPHIE BERTRAND Rede Verde Conservation Network, Foz do Iguaçu,
Brazil
TAIANA HAAG Programa de Po´s-Graduaça˜o em Gene´tica e Biologia Molecular, Universidade Federal do Rio Grande do Sul, Porto Alegre, Brazil, and
Laborato´rio de Biologia Genoˆmica e Molecular, Faculdade de Biocieˆncias,
Pontifı´cia Universidade Cato´lica do Rio Grande do Sul, Porto Alegre, Brazil
ALCIDES RICIERI RINALDI Laborato´rio de Biodiversidade, Conservaça˜o e
Ecologia de Animais Silvestres, Universidade Federal do Parana´, Foz do
Iguaçu, Brazil
SIXTO FERNA´NDEZ, FREDY RAMI´REZ and MYRIAM VELA´ZQUEZ Fundacio´n
Moise´s Bertoni, Asuncio´n, Paraguay
CRISTIAN CORIO and ESTEBAN HASSON Laboratorio de Evolucio´n, Facultad
de Ciencias Exactas y Naturales, Universidad de Buenos Aires, Buenos Aires,
Argentina
Received 22 March 2010. Revision requested 17 May 2010.
Accepted 1 July 2010. First published online 31 May 2011.

Introduction

M

ost species of large carnivores are threatened as
a result of their large territorial requirements, their
naturally low densities and direct persecution by humans
(Noss et al., 1996; Woodroffe & Ginsberg, 1998). Knowledge
of the presence of carnivores in human-dominated landscapes constitutes the basis for determining the conservation status of these species (Karanth & Nichols, 2002;
Sanderson et al., 2002). However, their secretive behaviour
and low densities make research on, and monitoring
of, large carnivores difficult (Karanth & Chellam, 2009).
Indirect evidence of animal presence (e.g. tracks and scats)
is a valuable resource for surveying the distribution of
secretive species (Wemmer et al., 1996; Karanth & Nichols,
2002), and sign and questionnaire surveys are commonly
used to determine the presence of large carnivores

ª 2011 Fauna & Flora International, Oryx, 45(4), 534–545

doi:10.1017/S0030605310000840

Monitoring of large carnivores

(McNab & Polisar, 2002; Altrichter et al., 2006; Carroll &
Miquelle, 2006). However, limited funds and personnel
often make it difficult to obtain this type of data across
extensive areas and over long periods of time (Karanth &
Nichols, 2002; Danielsen et al., 2005), and it is even more
challenging when the search involves different countries
with different cultures and languages.
Locally-based participatory monitoring programmes
have demonstrated potential for surveying large areas and
are particularly valuable in developing countries where the
involvement of local volunteers may prompt awareness and
management interventions (Danielsen et al., 2003, 2007). In
spite of relatively low cost and the possibility of covering
large areas, participatory monitoring networks require tools
for motivating the volunteers and maintaining their interest
(Bell et al., 2008). In developing countries, where resources
are often limited and the culture of volunteering is not
necessarily common or organized, simple, inexpensive and
motivating methodologies are required to guarantee the
success of participatory monitoring activities (Danielsen
et al., 2003, 2009; Bell et al., 2008). Charismatic species have
been important motivators for promoting local participation (Bowen-Jones & Entwistle, 2002; Gray & Kalpers, 2005;
Danielsen et al., 2009) and large carnivores are frequently
selected as flagship species in conservation programmes,
although mainly for fund-raising and educational purposes
(Dalerum et al., 2008; Karanth & Chellam, 2009). However,
as these species are often perceived as a threat to human
lives and livestock by local people, the utility of their image
as flagship species where local support is needed has been
disputed (Bowen-Jones & Entwistle, 2002).
The jaguar Panthera onca and puma Puma concolor are
the largest carnivores of the threatened Upper Parana´
Atlantic Forest on the tri-national border area formed by
Argentina, Brazil and Paraguay (Myers et al., 2000; Fig. 1).
The jaguar is a species of particular conservation concern in
this region. In Argentina it is both an Endangered species
(Dı´az & Ojeda, 2000) and a National Natural Monument
(Law #25,463), and it is categorized as Vulnerable in Brazil
(Chiarello et al., 2008) and as Critically Endangered in
Paraguay (SEAM, 2010). Because of its large territorial
demands and requirement for an adequate prey base, the
jaguar has been identified as an umbrella species to design
a biodiversity conservation landscape for the Upper Parana´
Atlantic Forest in a tri-national conservation initiative
(the Biodiversity Vision; Di Bitetti et al., 2003). In addition,
two areas in this part of the Atlantic Forest were
selected by a group of jaguar experts as significant areas
for the conservation of the species (Jaguar Conservation
Units; Sanderson et al., 2002). Therefore, studying and
monitoring jaguars in the Upper Parana´ Atlantic Forest
were considered priorities for the species and for ecoregional conservation strategies (Sanderson et al., 2002; Di
Bitetti et al., 2003).
ª 2011 Fauna & Flora International, Oryx, 45(4), 534–545

Here we discuss the use of locally based monitoring
programmes as a tool for a large-scale survey, using as
a case study a project in which large carnivores were used as
flagship species to promote participation. We present an
updated distribution map for pumas and jaguars in the
Upper Parana´ Atlantic Forest resulting from a collaborative
effort of local people, professionals and institutions in
Argentina, Brazil and Paraguay. We also assess the effect
of habitat loss on the distribution of pumas and jaguars in
this Forest and the advantages of local participation and
collaborative work for monitoring large carnivores at a regional scale.
Study area
We conducted this study in the core areas for long-term
conservation of the Upper Parana´ Atlantic Forest identified
by the Biodiversity Vision (Di Bitetti et al., 2003; Fig. 1). The
Upper Parana´ Atlantic Forest is the largest of the 15
ecoregions that comprise the South American Atlantic
Forest. It was formerly characterized by 470,000 km2 of
subtropical semi-deciduous rain forests but , 8% of its
original forest cover remains (Di Bitetti et al., 2003; Fig. 1).
This ecoregion contains high levels of biodiversity and
a diversity of human cultures (Galindo-Leal & de Gusma˜o
Caˆmara, 2003), with a high, unevenly distributed human
density and diverse economic activities (Jacobsen, 2003).
The Upper Parana´ Atlantic Forest is distributed in eastern
Paraguay, most of Misiones Province in Argentina and
portions of southern Brazil (Fig. 1). The largest area of this
forest is in Brazil but most large fragments (. 100 km2) are
in eastern Paraguay and the Misiones Green Corridor in
Argentina, a corridor connecting two large Brazilian
protected areas (Di Bitetti et al., 2003; Fig. 1).
Methods
We used two monitoring approaches to survey the puma
and jaguar in the Upper Parana´ Atlantic Forest. In Paraguay, Argentina and the two Brazilian protected areas
connected by the Misiones Green Corridor (Fig. 1) we
implemented a locally based participatory monitoring
network: the Jaguar Project Monitoring Network. Additionally, we used data gathered during studies of jaguars
and pumas by LC, DS, KCA and FL in the Upper Parana´–
Pontal do Paranapanema Region, the northern portion of
the Brazilian Upper Parana´ Atlantic Forest (Fig. 2).
The Jaguar Project Monitoring Network
In October 2002 we held a workshop in Puerto Iguazu´,
Argentina, integrating researchers and stakeholders from
the main institutions involved in the conservation and
management of the Upper Parana´ Atlantic Forest. In this

535

536

C. De Angelo et al.

FIG. 1 Main protected areas and forest
remnants in the core area of the Upper
Parana´ Atlantic Forest ecoregion, in which
puma Puma concolor and jaguar Panthera
onca presence was surveyed. The rectangle
on the inset indicates the location of the
main map in South America.

workshop we defined the methodology and identified
potential participants for constructing a network of volunteers to monitor the presence of pumas and jaguars.
Between mid 2002 and 2008 we conducted 70 field training
workshops on sampling techniques and data collection for
people living or working in areas where large carnivores are
potentially present. Initially most of the participants were
park rangers and field biologists (both men and women)
and members of local governmental and non-governmental
institutions, who supported this initiative and provided
data from protected areas and their surroundings. However, as each volunteer encouraged the integration of more
people into the network, we included farmers, ranchers,
forestry workers, army patrols and students, whose joint
efforts allowed us to obtain information from many regions
both within and beyond protected and forested areas.
We trained the participants to search for and collect
track imprints and faeces of large carnivores following
simple instructions: ‘to collect tracks wider than 6 cm with
the impression of the heel pad and four toes, and faecal
samples . 2 cm diameter with prey content (hair, bones,

hoofs)’. The main objective was to ensure that collaborators
would collect every potential presence sign of pumas or
jaguars, without the need for species-specific sign recognition in the field. Additionally, we designed a simple and rapid
methodology of data collection to allow participants to
incorporate the monitoring as a routine activity in their
normal work without demanding extra costs and time. We
provided them with a kit containing instructions and
supplies for making plaster moulds of tracks and to collect
and store faecal samples (Appendix 1). The kit also contained
easy-to-complete cards to record data about the samples
(date, collector name, site, environment). For faeces sampling, volunteers used disposable gloves and labelled paper
bags. Faecal samples were dried by the volunteers inside the
paper bags and stored with silica gel (Amato et al., 2006).
Volunteers also recorded sightings and livestock depredation reports following pre-designed questionnaires. A coordinator (CDA) periodically visited or contacted the
volunteers to compile data and provide extra kit supplies.
All participants worked for free, with the only motivation being that of collaboration for the conservation and
ª 2011 Fauna & Flora International, Oryx, 45(4), 534–545

Monitoring of large carnivores

by Haag et al. (2009; Appendix 3). We classified sightings and
livestock depredation events by pumas or jaguars according
to their reliability (high: direct detection by a reliable
observer or with physical evidence; medium: indirect detection obtained by a reliable informer but without physical
evidence; low: not able to establish reliability with confidence). We did not consider records of low reliability as
presence data, and we only considered records with medium
reliability if they came from regions with physical evidence
of species’ presence recorded in nearby areas (, 10 km). To
these data we added presence data collected by research
projects (e.g. photographs from camera traps; Paviolo et al.,
2008, 2009) and information from governmental agencies
(e.g. denunciations compiled and verified by the Ministry of
Ecology of Misiones or the National Park Administration of
Argentina).
Collaborative research in the Upper Parana´–Pontal do
Paranapanema Region

FIG. 2 Distribution of the data records (n 5 2,666) collected by
research groups in the Upper Parana´–Pontal do Paranapanema
Region (UPPR) and by the Jaguar Project Monitoring Network
(JPMN). Dashed lines indicate the surveyed areas estimated
using the density of records (99% kernel density estimator; see
text for details).

management of both carnivore species. To help maintain
motivation we prepared and distributed bulletins to inform
participants about the progress of the project and provide
practical information (e.g. tips for searching for presence
signs). We also produced and distributed complementary
materials, such as a track identification book (De Angelo
et al., 2008), to help improve the quality of data collected.
Additionally, we organized a tri-national workshop once
per year (Appendix 2) to discuss progress and future plans
with volunteers.
Although data collection was by volunteers, data selection, species identification and final analysis were by
professionals, using accurate techniques and conservative
criteria to avoid potential false positives of species’ presence.
To identify tracks we followed the protocol established by
De Angelo et al. (2010), using multiple measurements combined with a discriminant function analysis (Appendix 3).
We extracted a piece of the best-preserved faecal samples for
DNA identification of species following methods developed
ª 2011 Fauna & Flora International, Oryx, 45(4), 534–545

Since the late 1990s various research groups have been
gathering information on large predators in the Upper
Parana´–Pontal do Paranapanema Region (Crawshaw, 2006).
The records of jaguar and puma presence collected between
2002 and 2008 by these research groups include tracks,
camera-trap photographs, records of individuals captured or
killed and radio-tracking data (Cullen et al., 2005; Cullen,
2006; Abreu et al., 2009; Pro´-Carnı´voros, 2009; IPEˆ, 2010).
Data interpretation
We determined the geographical coordinates of each record
using the information provided by collectors, local maps
and satellite images. Less accurate records, such as locations
of low certainty, degraded faecal samples, older records and
material that we could not accurately identify as being from
pumas or jaguars were only used to estimate the size of the
surveyed area. We used the kernel density estimator tool
(Bayer, 2004) to estimate the area covered by our survey.
Kernel density estimation is a non-parametric way of
estimating the probability density function of a random
variable, incorporating information about both spatial
distribution and density, and is typically used in homerange studies (Worton, 1989).
To construct distribution maps of each species we used
all presence points obtained between July 2002 and July
2008 that were confidently identified as puma or jaguar and
were precisely located. We examined the locations of both
species in relation to native forest cover (using forest cover
estimated by supervised classification of Landsat Satellite
images from 2004 by De Angelo, 2009, where native marshland habitat was included as native forest) and protected
areas (using a compilation of protected areas maps for the
Upper Parana´ Atlantic Forest region from De Angelo, 2009).

537

538

C. De Angelo et al.

Results
Between 2002 and 2008 we trained c. 320 people to collect data
on the presence of pumas and jaguars in the Jaguar Project
Monitoring Network, representing 40 institutions (governmental, non-governmental and private) and individuals (farmers, students and others). Although most of the volunteers were
park rangers (30% including governmental and private protected areas), many collaborators were personnel or owners of
private properties/companies (14%, mainly timber and forestry
companies), NGO members (12%), students (11%), researchers
(10%, including biologists, anthropologists and forestry engineers), local government staff (8%), army or border security
patrols (5%), tourism guides (5%) and farmers (4%). Volunteer
participation in the network was dynamic and not all people
were active throughout the 6 years but at least 100 volunteers
participated for the entire monitoring period. This collaborative
effort resulted in 2,667 records (Fig. 2): 33.2% from researchers
working in the Upper Parana´–Pontal do Paranapanema Region and the remainder from the Jaguar Project Monitoring
Network. The distribution of all records covered an area of
92,890 km2 (99% kernel, h 5 13,000 m; Fig. 2) but most of the
data were concentrated in 54,181 km2 (95% kernel, h 5 13,000
m). The surveyed area included 86% of the protected areas of
the Upper Parana´ Atlantic Forest (Appendix 4) and covered
66.2% of the remaining forest, including 68% of fragments 100–
1,000 km2 in size and seven of the eight largest forest fragments
. 1,000 km2 (considering the Green Corridor divided by main
roads into four large forest fragments; Fig. 2). Of the total data
61.2% of records were classified as reliable evidence of the
presence of pumas or jaguars (Table 1).
Records of pumas were more abundant than those of the
jaguar, independently of the type of record (mean 5 2.25 – SE
0.63 times more abundant; Table 1), with the exception of
radio-tracking records that were obtained from an unbalanced number of animals (one puma versus 10 jaguars
monitored; Cullen, 2006). The puma was present in most
of the study area except for those areas with high human
disturbance (Fig. 3). The jaguar, in contrast, was concentrated
mainly in the Green Corridor of Argentina–Brazil and in the

largest forest fragments in Brazil and Paraguay (Fig. 3). In
Brazil we recorded jaguars only in or near the largest protected
areas (Fig. 3, Appendix 4): Morro do Diabo State Park,
Ivinhema State Park, Ilha Grande National Park, Perobas
Biological Reserve, Iguaçu National Park and Turvo State Park.
In Paraguay the largest protected areas also contained most of
the country’s jaguar records (Mbaracayu´ Natural Reserve,
Morombı´ Private Reserve, Limoy and Itabo´ Biological Refuges
and the southernmost record in Paraguay, in San Rafael
Reserve Area for National Park, S 26 38913.70 W 55 39942.10;
Fig. 3, Appendix 4). We recorded the southernmost jaguars in
the entire Upper Parana´ Atlantic Forest in Turvo State Park in
Brazil (S 27 10914.60 W 53 5194.90) and in Mocona´ Provincial
Park in Argentina (S27 9933.40 W 53 53928.70). Puma records
extend further south in both Argentina and Brazil (Fig. 3).
We obtained most puma (83%) and jaguar (83%) records
in areas covered by native forest or native marshland
habitat, and we confirmed puma presence in more forest
fragments than jaguars (Fig. 4). Both species occurred in all
the larger fragments (. 100 km2) but , 2% of surveyed
forest fragments , 10 km2 in area had evidence of the
species. We detected pumas in a higher proportion of small
and medium sized fragments than jaguars (Fig. 4) and the
total area covered by fragments with confirmed puma
presence (19,266 km2) was larger than the total area with
confirmed presence of jaguars (16,585 km2). For both
species these areas are , 25% of the total area surveyed.
We detected jaguars in seven large forest fragments (i.e.
. 100 km2) outside the Jaguar Conservation Units defined
for this region (Sanderson et al., 2002): Salto Encantado and
Cun˜a Piru´ provincial parks in Argentina, Morro do Diabo
State Park in Brazil and five fragments in Paraguay (Fig. 3).

Discussion
Partnership for regional surveys of large carnivores
Participatory monitoring and collaboration among scientists
allowed us to obtain and compile data on the distribution of

TABLE 1 Confirmed records of puma Puma concolor and jaguar Panthera onca in the Upper Parana´ Atlantic Forest (Fig. 1) between July
2002 and July 2008.
Upper Parana´–Pontal do
Paranapanema Region records

Jaguar Project Monitoring Network records
Species
Puma
Jaguar

Tracks1
236
150

Scats1
25
13

Sightings
50
33

Conflicts
with cattle
19
6

Others2
136
83

Various3
168
48

Telemetry locations4
18
650

Total5
634 (651)
333 (982)

1

Details of track and faecal sample identification in Haag et al. (2009), De Angelo et al. (2010) and Appendix 3
Includes skin and tissue samples, camera-trap photographs and poached animals
3
Data from various surveys by research groups in the Upper Parana´–Pontal do Paranapanema Region, in the northern Upper Parana´ Atlantic Forest. It
includes captured/removed individuals, poached individuals, tracks, sightings, camera-trap photographs and conflicts with cattle.
4
Telemetry locations were obtained from one puma and 10 radio-collared jaguars (details in Cullen, 2006).
5
Total without considering radio-tracking locations (total including radio-tracking locations in parentheses).
2

ª 2011 Fauna & Flora International, Oryx, 45(4), 534–545

Monitoring of large carnivores

FIG. 3 (a) Puma and (b) jaguar distribution in the Upper Parana´ Atlantic Forest as determined by collaborative and participatory
monitoring between July 2002 and July 2008. Jaguar Conservation Units (Sanderson et al., 2002): (A) Green Corridor, (B) Upper
Parana´–Pontal do Paranapanema Region. Note the numerous areas with jaguar records detected outside the Jaguar Conservation Units,
particularly in Paraguay. Horizontal black arrows indicate the area of the Paraguayan Upper Parana´ Atlantic Forest with large forest
fragments that are poorly surveyed and where there is a high probability of the presence of both species. See caption to Fig. 2 for details
of delimitation of the surveyed areas in the UPPR (Upper Parana´–Pontal do Paranapanema Region) and JPMN (Jaguar Project
Monitoring Network).

two secretive species, the puma and jaguar, in most of the
remnants of the Upper Parana´ Atlantic Forest ecoregion. This
survey had two important characteristics. One was the
interaction between research groups from different countries,
combining their local knowledge to understand patterns
occurring at a regional scale. Partnership between researchers

FIG. 4 Percentage of surveyed forest fragments with confirmed
puma and jaguar presence, by fragment size categories (in
parentheses: number of fragments with presence/total number
of fragments in category included in the surveyed area).
ª 2011 Fauna & Flora International, Oryx, 45(4), 534–545

has produced useful results for jaguars and other species in
other parts of the jaguar’s distribution (Grigione et al., 2009)
and at a continental scale (Sanderson et al., 2002).
The second characteristic was the participatory monitoring carried out by a wide network of locally based teams.
Danielsen et al. (2009) suggested a typology for monitoring
programmes according to the degree of local participation.
Following this typology the Jaguar Project Monitoring
Network could be classified as ‘collaborative monitoring
with external data interpretation’ because local people
participated not only in data collection but also in the design
of the monitoring, whereas data analyses and decision
making were by professionals. Danielsen et al. (2009)
defined eight characteristics of the benefits and demands
of participatory monitoring, and Table 2 summarizes these
for the Jaguar Project Monitoring Network.
The implementation of the Jaguar Project Monitoring
Network faced not only the challenges associated with the
survey of a large area by people from different countries
and cultures but also communication difficulties (e.g.
isolated areas, difficult access by vehicle and no telephones).
Nevertheless, it yielded a large amount of data and involved
many local people, with no costs for participants and low

539

540

Cost to local
stakeholders
None or low
Participants
selected who worked
or lived in or close
to potential puma/
jaguar habitat to
avoid cost of
transportation
# 20 minutes
required to collect
samples
Simple & costless
methods allowed
volunteers to
incorporate survey
in their routines
without extra costs
in time & materials
(e.g. park rangers
collected tracks &
faecal samples
during their patrols)

Cost to others
(outsiders)
Total external
investment in 6
years of monitoring
of most intensively
surveyed area was
,USD 0.01 ha-1
year-1 & c. USD 124
per accurately
identified puma/
jaguar record
Local institution
involvement helped
reduce external
costs by providing
continuous support
for mobility &
communication
with & among
volunteers

Requirement for
local expertise
All participants had
opportunity to be
trained, so no
previous expertise
required

Requirement for
external expertise
To train volunteer
teams & for network
coordination but
mainly in data
processing

However, volunteers
with previous
expertise of
fieldwork normally
obtained more &
better information

Improved accuracy
& precision of data

Participants’ local
knowledge favoured
success in finding
new records

Catalysed transfer of
information to
national &
international
monitoring schemes
Increased costs &
time needed to
report results (i.e.
feedback required to
keep people
interested)

ª 2011 Fauna & Flora International, Oryx, 45(4), 534–545

External expertise
other than field
biologists was
important for
communication &
conservation
activities (e.g.
communication
campaign)

Accuracy &
precision
Records were
accurately identified
to species
(Appendix 3) &
precisely located by
professionals
Accuracy was useful
for distribution
mapping &
knowledge of
habitat use but large
area sampled &
difficulty of
quantifying
sampling effort
hindered
comparisons of
relative abundance
between species,
areas & over time

Promptness of
decision making
Facilitated prompt
& coordinated
actions among
individuals &
institutions for
mitigation of
human–felid
conflicts*
Many volunteers
participated in
educational
activities (Campan˜a
Yaguarete´, 2007)
Promoted
development of
action plans for
jaguars, integrating
institutions from
three countries
Participants had
direct involvement
in action plans for
large felids (e.g.
Chalukian, 2006),
which will result in
greater diffusion &
local acceptance of
implementation

Potential for
enhancing local
stakeholder capacity
Permanent contact
with coordinator,
seven informative
bulletins, four trinational workshops
& a track
identification guide
(De Angelo et al.,
2008) are examples
of tools used for
enhancing
stakeholders’
capacity
Fostered
communication
among volunteers
about felid
conservation
Improved response
of local managers to
potential human–
felid conflicts*

Capacity to inform
national &
international
monitoring schemes
Jaguar data were
included in Zeller
(2007)
Felid records
incorporated in
IUCN Neotropical
felids’ database
(CSG–IUCN, 2005)
Data are being used
to update
Argentinian
mammal Red List
Presence records of
white-lipped
peccary Tayassu
pecari & tapir
Tapirus terrestris
were requested of
volunteers &
incorporated in
continental surveys
by the Wildlife
Conservation
Society & IUCN
specialists groups

Local NGOs used
results to define
important areas of
forest to protect
under Misiones
Province territorial
plan
*Some examples of combined management interventions with the involvement of different parts of the Jaguar Project Monitoring Network, as communicated in the local, national and international press, can be
found in Territorio Digital (2004, 2008) Mullen (2006), La Nacio´n (2007) and Misiones Online (2009)

C. De Angelo et al.

TABLE 2 Evaluation of the Jaguar Project Monitoring Network according to the main characteristics of monitoring schemes suggested by Danielsen et al. (2009).

Monitoring of large carnivores

costs for external institutions (Table 2). Considering Danielsen et al.’s (2005) summary, external cost investment in
the Jaguar Project Monitoring Network (, USD 0.01 ha-1
year-1) is among the lowest costs estimated for similar
participatory monitoring programmes (USD 0.01–0.13 ha-1
year-1) and much lower than the cost estimated for professional surveys (c. USD 3.6 ha-1 year-1; Danielsen et al.,
2009; Table 2). Although professional surveys (e.g. with
camera traps) may allow more detailed information to be
obtained (e.g. density estimates), their application at broader
scales may not only be more expensive but also practically
infeasible or unsustainable for long periods of time. Locally
based surveys have the advantage of being able to maintain
permanent long-term monitoring in many areas simultaneously, which is the best survey technique for obtaining
data in areas where large carnivores live at extremely low
densities (Karanth & Nichols, 2002).
The low cost associated with the Jaguar Project Monitoring
Network was not only the result of a simple affordable
methodology but also of the commitment of participants
and the support received from many local governmental and
non-governmental institutions that allowed the monitoring to
be incorporated in the routine activities of their personnel.
Volunteers showed great interest and enthusiasm for seeking
evidence of large predators, and the subsequent recognition for
their work from the coordinator and other volunteers promoted increased efforts. Enthusiasm, credit and desire for
learning have been recognized as important drivers for
maintaining interest amongst volunteers (Danielsen et al.,
2007; Bell et al., 2008). However, the powerful image of the
species monitored was essential to enthuse and involve local
people and institutions in the Network, and both species, but
mainly the jaguar, were important motivators for people and
institutions, and also contributed to the growth of the Network
(e.g. people who spontaneously offered to join the Network
because of their interest in jaguars). This demonstrated that
with adequate motivation and methods, volunteer work is
possible in developing countries even though volunteering is
not as culturally common as in developed countries (Danielsen
et al., 2003, 2009; Bell et al., 2008).
Comparable experiences of motivation were reported by
Poulsen & Luanglath (2005) in participatory biodiversity
monitoring in Laos and by park rangers monitoring gorilla
Gorilla beringei beringei populations in Rwanda, Uganda
and the Democratic Republic of Congo (Gray & Kalpers,
2005). As in the Jaguar Project Monitoring Network,
gorillas were used as flagship species to help maintain
participants’ interest and motivation. Through the Network
we showed that, despite their conflicting image for cattle
ranchers and some local people (Conforti & Azevedo,
2003), large carnivores may have special value as flagship
species when they have positive associations for the selected
focal audience (Bowen-Jones & Entwistle, 2002; Dalerum
et al., 2008). Although most participants in the Network
ª 2011 Fauna & Flora International, Oryx, 45(4), 534–545

had a positive relationship with jaguars some ranchers, who
may be expected to have a negative relationship with the
species, joined the network because of their concern for the
conservation of felids. Although we did not put much effort
into the recruitment of ranchers as volunteers, we believe
that rancher-based local monitoring with adequate advice
from professionals (i.e. agronomists, sociologists) and
government involvement may prove a useful tool not only
in monitoring predators but also in reducing human–
predator conflicts.
In spite of low costs and strong local support, the Jaguar
Project Monitoring Network demanded large and permanent coordination efforts (e.g. personal contact with volunteers and communication activities). Because the
coordinator (CDA) could not commit himself to this
endeavour for . 5 years and because funds were not
secured for the long-term (the project was conceived and
funded mostly by NGOs), the continuity of the coordination of the Network and the monitoring was not ensured,
and finished in 2008. Discontinuous funding is a common
problem for sustaining monitoring over long periods of
time (Brashares & Sam, 2005; Poulsen & Luanglath, 2005).
The involvement of local institutions is essential to guarantee monitoring continuity over time without permanent
external funds, enhancing the important role that involvement of local institutions has for participatory monitoring
(Danielsen et al., 2005; Gray & Kalpers, 2005; Poulsen &
Luanglath, 2005).
Similar surveys have frequently left identification of
records in charge of local people (e.g. using interviews;
McNab & Polisar, 2002; Altrichter et al., 2006). However,
jaguars have a conspicuous cultural value in local communities in the Upper Parana´ Atlantic Forest (Conforti &
Azevedo, 2003) and their strong image and the similarity of
their signs with those of pumas may cause a bias towards
overrating jaguar presence and to misidentify the signs of
pumas as belonging to jaguars, as occurs with other large
carnivores (Lynam, 2002). The use of physical evidence and
precise identification methods reduces this problem but
demands more professional involvement. However, professional participation may not only increase accuracy and
precision but also result in a wider use of the data gathered
(Table 2).
There are two other important aspects of local participation in monitoring biodiversity: its implications for
promptness of decision making and its potential for
enhancing local stakeholder capacity (Danielsen et al.,
2009). The Jaguar Project Monitoring Network had significant outcomes in both aspects (Table 2) but, most
importantly, it promoted collaborative work in conservation and management of large felids at a regional scale
through the involvement and training of people from
different institutions, professions and countries. Since the
creation of the Network complaints of potential conflicts

541

542

C. De Angelo et al.

(e.g. felid sightings in populated areas, cattle killed) are
communicated not only to local authorities but also to
Network members, favouring the intervention of many
institutions. As a result, a protocol of action is being drawn
up and will be implemented through the bi-national action
plan for jaguars in the Argentina–Brazil Green Corridor.
This plan incorporates not only this protocol but also the
information and ideas collected in the various different
meetings in which the Network volunteers participated
(Table 2). Additionally, the information gathered proved
useful for other regional conservation initiatives: an action
plan for jaguars in Paraguay is in the initial stages and, in
the Upper Parana´–Pontal do Paranapanema Region, a jaguar action plan is being developed (Table 2).
Jaguars and pumas in the Upper Parana´ Atlantic Forest
Although many human pressures can affect the persistence
of species, habitat destruction has particularly harmful
effects on large carnivores that require extensive territories
(Karanth & Chellam, 2009). Pumas and jaguars in the
Upper Parana´ Atlantic Forest have been seriously affected
by forest loss and fragmentation. A clear indication of this
is that these species are almost exclusively associated with
medium and large fragments of native forest in a region
where most of the landscape has been transformed to
anthropogenic land uses (Figs 3 & 4).
The puma still occupies most of its continental distribution but has disappeared or became rare in those areas
with the highest human pressures (Sunquist & Sunquist,
2002). In the Upper Parana´ Atlantic Forest pumas are
present in a significant proportion of the forest remnants,
including many areas where the jaguar was not recorded
(Fig. 3). However, fragments with puma presence represent
, 25% of the surveyed area and we did not find evidence of
pumas in areas with intensive agriculture or with high
human presence (Figs 3 & 4). Therefore, although pumas
have apparently suffered less range contraction than
jaguars, habitat loss and fragmentation along with the
impacts of intense logging and poaching have resulted in
the decrease of puma populations in the Upper Parana´
Atlantic Forest (Paviolo et al., 2009).
The continental distribution of the jaguar has contracted
severely (Sanderson et al., 2002; Zeller, 2007). In Brazil,
where c. 50% of the continental range of the jaguar persists,
habitat loss is the most important cause of jaguar decline
(Toˆrres et al., 2008). In the Brazilian Atlantic Forest habitat
destruction has been high (de Gusma˜o Caˆmara, 2003), and
Mazzolli (2009) described a south-to-north reduction of
jaguar distribution in coastal Atlantic Forest caused by
habitat fragmentation and poaching of jaguars. In the
Upper Parana´ Atlantic Forest of Brazil the main forest
fragments are within protected areas, and it is only in these
areas where the jaguar persists (Fig. 3, Appendix 4). Habitat

loss is probably the main reason for jaguar range contraction
in the Brazilian Upper Parana´ Atlantic Forest but poaching
of jaguars and their prey is also affecting jaguars in this
region (Conforti & Azevedo, 2003; Cullen et al., 2005; Abreu
et al., 2009). Reducing these threats is essential for jaguar
survival (Cullen et al., 2005) but habitat recovery and
connectivity are necessary to increase dispersal between
populations and to reduce the genetic loss that fragmentation is producing in this region (Haag et al., 2010).
In Paraguay we also obtained most jaguar records from
within protected areas that harbour the most extensive
forest fragments. The largest jaguar population in eastern
Paraguay is probably located in Mbaracayu´ Nature Reserve
and surrounding areas (Fig. 3). However, important forest
remnants not surveyed by the Jaguar Project Monitoring
Network persist in Canindeyu´, Amambay and San Pedro
departments, where we obtained informal data of puma
and jaguar presence (Fig. 3). These areas could be important for maintenance of the connectivity among jaguar
populations of the Atlantic Forest and chacoan and cerrado
jaguar populations in western Paraguay. Although 30 years
ago the Paraguayan Upper Parana´ Atlantic Forest was
a vast continuous forest it has suffered recent and rapid
conversion (Huang et al., 2007). Many of the areas with
jaguar presence in Paraguay correspond with recently
reduced and isolated fragments, and many of these isolated
and small jaguar populations will probably disappear in the
short-term. Although other pressures exist, habitat loss and
fragmentation are also the main threats to the jaguar in
eastern Paraguay.
In the Argentinian Upper Parana´ Atlantic Forest jaguar
records were concentrated in the Green Corridor (Fig. 4)
where a large and continuous corridor of forest remains.
Argentina has had the highest national rate of jaguar range
contraction (Di Bitetti et al., 2006), occurring in a south-tonorth pattern. The last records of jaguars in Corrientes
Province (south of Misiones province) date from the 1970s
(Giraudo & Povedano, 2003) and the latest reliable records
we obtained for southern Misiones date from the early
1990s. Since the mid 1990s the southernmost jaguar records
in Misiones are in the central and eastern parts of the
province (Giraudo & Povedano, 2003) where jaguars were
detected by the Jaguar Project Monitoring Network (Fig. 3).
These areas, and Turvo State Park in Brazil, are currently
the southernmost limit of jaguar distribution (Sanderson
et al., 2002; Zeller, 2007). Although habitat loss is affecting
jaguar range in the Argentinian Upper Parana´ Atlantic
Forest, many areas with large forest fragments outside the
Green Corridor harbour pumas but not jaguars (Fig. 3),
suggesting that other factors have caused the disappearance
of jaguars in these areas. A vast extent of the Argentinian
Forest has been heavily logged and there is high poaching
pressure on the jaguar and its prey, probably the main
causes of a recent decline of jaguars in the Green Corridor
ª 2011 Fauna & Flora International, Oryx, 45(4), 534–545

Monitoring of large carnivores

(Paviolo et al., 2008). Although maintenance of forest cover
is essential, reducing other human threats is critical for
survival of the jaguar in this area (Paviolo et al., 2008).
The monitoring programme allowed us to detect both
species in areas where there was no information available or
where they were considered rare or locally extinct. The data
have contributed not only to a more complete understanding of the effect of habitat loss on both species (e.g. through
habitat suitability modelling, De Angelo et al., 2011, and
genetic studies, Haag et al., 2010) but is also helping in
conservation and management actions (Table 2). An
updated and more detailed definition of the Jaguar Conservation Units (Fig. 3) will help to improve local, regional
and continental conservation plans (Rabinowitz & Zeller,
2010). A re-evaluation of the conservation landscape
designed by Di Bitetti et al. (2003) for the Upper Parana´
Atlantic Forest is in progress, and this is another important
outcome for which the data collected in this participatory
survey were utilized.
Acknowledgements
We are very grateful to all the volunteers, collaborators and
institutions who participated in the Jaguar Project Monitoring Network, particularly to C. Boiero, P. Crawshaw Jr,
Y. Di Blanco, E. Eizirik, A. Garcı´a, M. Go´mez, E. Krauczuk,
M. Jaramillo, R. Melzew, E. Pizzio, G. Placci, M. Rinas,
A. Rodrigues, K. Schiaffino, S. Welcz, the National Parks
Administration (Argentina), the Ministry of Ecology of
Misiones Province (Argentina), Itaipu Binacional (Paraguay/Brazil), Pro-Cosara (Paraguay), Instituto de Derecho
y Economı´a Ambiental (Paraguay), and our hosting institutions. Financial support was provided by the National
Research Council of Argentina (CONICET), Fundacio´n
Vida Silvestre Argentina, WWF–USA, WWF–International, WWF–Switzerland, WWF Education for Nature
Program, Species Action Fund (WWF–US) and Lincoln
Park Zoo.
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Appendices
The appendices for this article are available online at http://
journals.cambridge.org
Biographical sketches
The authors work on various aspects of the ecology, genetics, conservation and management of biodiversity in Argentina, Brazil and
Paraguay. As scientists, managers or conservationists the authors
belong to a variety of governmental and non-governmental institutions but share a common interest in biodiversity conservation, and
are particularly concerned about the conservation of pumas and
jaguars as keystone and umbrella species in the Atlantic Forest. In this
study they combined their efforts and expertise, taking advantage of
their diverse capabilities to assess the population status of large felids
from a regional perspective and undertake coordinated conservation
actions.

545

Participatory networks for large-scale monitoring of
large carnivores: pumas and jaguars of the Upper
Parana´ Atlantic Forest
C a r l o s D e A n g e l o , A G U S T ´I N P A V I O L O , D A N I E L A R O D E , L A U R Y C U L L E N J R
Denis Sana, Kaue Cachuba Abreu, Marina Xavier da Silva
ANNE-SOPHIE BERTRAND, TAIANA HAAG, FERNANDO LIMA, ALCIDES RICIERI RINALDI
S I X T O F E R N A´ N D E Z , F R E D Y R A M ´I R E Z , M Y R I A M V E L A´ Z Q U E Z , C R I S T I A N C O R I O
E S T E B A N H A S S O N and M a r i o S . D i B i t e t t i
Appendix 1 Examples of (a) instructions and (b) cards included in the collection kits of the collaborators. Easy-to-fill in cards (b) were
prepared for the information associated with each track or faecal sample. The sighting register card (c) was for more experienced volunteers
only.

ª 2011 Fauna & Flora International, Oryx, Page 1 of 4

doi:10.1017/S0030605310000840

2

C. De Angelo et al.
Appendix 2 Participants of the Jaguar Project Monitoring Network in the annual tri-national workshop held in Eldorado, Misiones
Province, Argentina in May 2007. (Photograph: C. De Angelo).

Appendix 3 Details of track and faecal identification methods
We washed plaster moulds of tracks and then photographed them with a digital camera, including a metric
rule in the photograph for measurements. All tracks were
then digitized using the spline tool in AutoCAD 2004
(AutoDesk Inc., San Rafael, USA) and scaled using the
reference metric rule. We followed the protocol established
by De Angelo et al. (2010) to identify tracks to species using
multiple measurements combined in a discriminant function analysis. We used the identification keys of De Angelo
et al. (2010) to differentiate first between felid and canid
tracks, and then between puma and jaguar tracks. When
the identification keys were unable to identify the tracks, we
used the complete identification discriminant models to
obtain a probability of a track being that of a puma or
jaguar. We only considered as presence records tracks with
. 80% probability of belonging to one of these species.
Because of the warm and wet weather in the study area we
selected only the best preserved faecal samples (fresh
collected and dried), and we extracted a piece from each of
these samples for specific DNA identification following the
methods developed by Haag et al. (2009). We extracted
DNA using specific kits following manufacturer protocols

(QIAamp DNA Stool Mini Kit and Puregene DNA Purification Kit from Qiagen Inc., Germantown, USA). The DNA
extracted from the samples was amplified by polymerase
chain reaction (PCR) using a primer pair (ATP6-DF2/ATP6DR1) designed for amplifying a short segment of 175 bp of the
mtDNA-ATP-synthase-subunit-6 (ATP6) gene. PCR products were purified and then sequenced in an automated
sequencer using the forward ATP6-DF2 primer. DNA sequences were aligned with the CLUSTALW algorithm implemented in MEGA v. 3.1 (The Biodesign Institute, Tempe,
USA). Finally, DNA sequences were compared with known
sequences of candidate species (jaguar Panthera onca, puma
Puma concolor, domestic dog and ocelot Leopardus pardalis)
and a complementary analysis was carried out with phylogenetic analysis (unweighted pair group method with arithmetic
mean assessing 1,000 bootstrap replications) using MEGA.
The faecal sample analysis was developed in Laboratorio de
Evolucio´n, Facultad de Ciencias Exactas y Naturales, University of Buenos Aires, Argentina, and Laborato´rio de Biologia
Genoˆmica e Molecular, Facultade de Biociencias, Pontificia
Universidad Cato´lica do Rı´o Grande do Sul, Brazil. The same
protocols were used in both laboratories.

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Monitoring of large carnivores
Appendix 4 Presence of pumas Puma concolor and jaguars Panthera onca in the protected areas (Fig. 1) monitored from June 2002 to
June 2008 in Argentina, Brazil and Paraguay.
Protected area (by country)
Argentina
´ rea Experimental Guaranı´
A
Monumento Natural Isla Palacios
Paisaje Protegido A. Giai
Parque Municipal L.H. Rolo´n
Parque Municipal Lote C
Parque Municipal P. los Indios
Parque Nacional Iguazu´
Parque Provincial Pto. Penı´nsula
Parque Provincial Caa Yarı´
Parque Provincial Cruce Caballero
Parque Provincial de la Araucaria
Parque Provincial de la Sierra
Parque Provincial del Teyu´ Cuare´
Parque Provincial Esmeralda
Parque Provincial Esperanza
Parque Provincial Fachinal
Parque Provincial H. Foerster
Parque Provincial Ing. Ag. Cametti
Parque Provincial Isla Caraguataı´
Parque Provincial Mocona´
Parque Provincial Pin˜alito
Parque Provincial Profundidad
Parque Provincial S. Welcz
Parque Provincial Salto Encantado
Parque Provincial Urugua-ı´
Parque Provincial Uruzu´
Parque Provincial V. del Cuna Piru´
Parque Provincial Yacui
Refugio Privado V. S. Lapacho Cue´
Reserva de Biosfera Yabotı´
Reserva Ecolo´gica Mbotabı´
Reserva ´Ictica de Caraguatay
Reserva ´Ictica de Corpus
Reserva N. y C. Papel Misionero
Reserva Nacional Iguazu´
Reserva Natural Estricta San Antonio
Reserva Natural Municipal Salto Kupper
Reserva Natural Municipal Yarara´
Reserva Privada "La Ponderosa"
Reserva Privada Aguara-i Mi
Reserva Privada Ing. Barney
Reserva Privada Itacuaraı´
Reserva Privada Julia´n Freaza
Reserva Privada Los Paraı´sos
Reserva Privada Puerto San Juan
Reserva Privada S. M. Aguaraı´ Minı´
Reserva Privada Santa Rosa
Reserva Privada Tomo
Reserva Privada UN La Plata
Reserva Privada V. S Timbo´ Gigante
Reserva Privada V. S. Caa´ Pora´
Reserva Privada V.S. Chachi
Reserva Privada V. S. Chancai
Reserva Privada V. S. El Yaguarete´

ª 2011 Fauna & Flora International, Oryx, 1–4

Puma*

Jaguar*

Confirmed
Confirmed
No evidence
No evidence
No evidence
No evidence
Confirmed
Confirmed
Surroundings
No evidence
No evidence
Confirmed
No evidence
Confirmed
No evidence
Confirmed
Surroundings
No evidence
No evidence
Confirmed
Confirmed
Surroundings
Surroundings
No evidence
Confirmed
Surroundings
Confirmed
No evidence
No evidence
Confirmed
No evidence
No evidence
No evidence
Confirmed
Confirmed
No evidence
No evidence
No evidence
No evidence
Confirmed
Surroundings
Surroundings
No evidence
No evidence
No evidence
No evidence
No evidence
No evidence
Confirmed
No evidence
Surroundings
No evidence
No evidence
No evidence

Confirmed
Confirmed
No evidence
No evidence
No evidence
No evidence
Confirmed
Confirmed
Confirmed
Surroundings
No evidence
No evidence
No evidence
Confirmed
No evidence
No evidence
No evidence
Confirmed
No evidence
Confirmed
No evidence
No evidence
No evidence
Surroundings
Confirmed
Surroundings
Confirmed
No evidence
No evidence
Confirmed
No evidence
No evidence
No evidence
Surroundings
Confirmed
No evidence
No evidence
No evidence
No evidence
Surroundings
No evidence
No evidence
No evidence
No evidence
No evidence
No evidence
No evidence
No evidence
No evidence
No evidence
Surroundings
No evidence
No evidence
Surroundings

3

4

C. De Angelo et al.

Appendix 4 (Continued)
Protected area (by country)
Reserva Privada V. S. Urugua-i
Reserva Privada V. S. Yacutinga
Reserva Privada Yaguarundı´
Reserva Uso Mu´ltiple A. Orlof Salt
Reserva Uso Mu´ltiple EEA C Azul
Reserva Uso Mu´ltiple EEA Victoria
Reserva Uso Mu´ltiple F. Basaldua´
Brazil
Estaça˜o Ecologica Mico-Leao-Preto
Estaça˜o Ecologica do Caiua´
Parque Estadual Ivinhema
Parque Estadual de Ampora
Parque Estadual do Turvo
Parque Estadual Morro do Diabo
Parque Nacional de Ilha Grande
Parque Nacional do Iguacu
Reserva Biolo´gico das Perobas
Reserva Particular de Patrimonio Natural Santa
Marı´a
Paraguay
Monumento Cientı´fico Moise´s Bertoni
Parque Nacional Cerro Cora
˜ acunday
Parque Nacional N
Parque Nacional Caazapa´
Refugio Biolo´gico Pikyry
Refugio Biolo´gico Tatı´ Yupı´
Reserva Biolo´gico Itabo´
Reserva Biolo´gico Limoy
Reserva Biolo´gico Mbaracayu´ (Paraguay/Brazil)
Area de Reserva para Pque. Nac. San Rafael
Reserva Nacional Kuriy
Reserva Natural Bosque Mbaracayu´
Reserva Natural Privada Morombı´
Reserva Natural Privada Tapyta´
Reserva Natural Privada Ypetı´
Reserva Privada Arroyo Blanco
Reserva Privada Kai Ragu¨e

Puma*
Confirmed
Confirmed
No evidence
No evidence
No evidence
No evidence
No evidence

Jaguar*
Confirmed
No evidence
No evidence
No evidence
No evidence
No evidence
No evidence

Confirmed
Confirmed
Confirmed
Surroundings
Surroundings
Confirmed
Confirmed
Confirmed
Surroundings
Confirmed

Confirmed
Confirmed
Confirmed
No evidence
Confirmed
Confirmed
Confirmed
Confirmed
Confirmed
Confirmed

No evidence
Confirmed
No evidence
No evidence
No evidence
No evidence
Confirmed
Confirmed
Surroundings
Confirmed
No evidence
Confirmed
Confirmed
Confirmed
No evidence
No evidence
No evidence

Surroundings
No evidence
No evidence
No evidence
No evidence
No evidence
Confirmed
Confirmed
No evidence
Confirmed
No evidence
Confirmed
Confirmed
No evidence
No evidence
No evidence
No evidence

*Confirmed, records obtained inside a protected area; Surroundings, records obtained in nearby areas at a distance less than half the mean maximum
distance moved calculated in camera-trap surveys for pumas and jaguars in the Upper Parana´ Atlantic Forest; No evidence, protected area included in the
surveyed area but where no evidence was found for species’ presence

ª 2011 Fauna & Flora International, Oryx, 1–4


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