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Título: Halting the isolation of jaguars: where to act locally to sustain connectivity in their southernmost population

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Animal Conservation. Print ISSN 1367-9430

Halting the isolation of jaguars: where to act locally to
sustain connectivity in their southernmost population
J. Martinez Pardo1,2, A. Paviolo1,2, S. Saura3 & C. De Angelo1,2
, Argentina
1 Instituto de Biolog ıa Subtropical, CONICET-Universidad Nacional de Misiones (UNaM), Puerto Iguazu
n Civil Centro de Investigaciones del Bosque Atl antico (CeIBA), Puerto Iguazu
, Argentina
2 Asociacio
3 European Commission, Joint Research Centre (JRC), Directorate D: Sustainable Resources, Ispra (VA), Italy

Atlantic Forest; graph-based models; habitat
connectivity; landscape-scale management;
jaguar; territorial planning; large carnivores.
Julia Martinez Pardo, Instituto de Biolog ıa
Subtropical, Universidad Nacional de
Misiones and CONICET (UNaM-CONICET),
n Civil Centro de Investigaciones
del Bosque Atl
antico (CeIBA), Bertoni 85,
, Misiones,
(N3370AIA) Puerto Iguazu
Argentina. Tel: +54 3757 423511.
Editor: Julie Young
Associate Editor: A. M arcia Barbosa
Received 30 December 2016; accepted 27
March 2017

Habitat loss and fragmentation are among the major threats to the conservation of
biodiversity. Improvement of landscape connectivity becomes one of the main
strategies for alleviating these threats and is an increasingly used target in management policies worldwide. However, implementation of connectivity principles
in local management actions often implies great difficulties derived from the different criteria used by connectivity analysts and policy makers. We generated a
management tool to incorporate connectivity criteria for large carnivores in landscape conservation planning at a local scale. Focusing on the southernmost population of jaguars Panthera onca, we use a graph-based connectivity approach to
(1) analyze habitat connectivity and availability in five areas previously identified
as main corridors; (2) detect priority forest patches for maintaining connectivity,
and (3) propose specific management strategies for each area matching the relative
importance and role of the forest patches in it. For this purpose, we defined the
patches as the local land management units (properties) and used information on
land cover and jaguar movement for determining the probabilities of connectivity
metric. We identified the key patches that represent 90% of the total contribution
to connectivity in the study areas; these patches were less than half of the total
number of patches in each corridor. Based on this forest patch prioritization, we
identified the most critical areas and specific patches where urgent conservation
measures need to be implemented. The percentage of patches and the total area
they covered varied among the five analyzed corridors showing contrasting situations for connectivity management and highlighting the importance of the proposed approach to understand the impact of patch-level actions in a broader
connectivity context. This approach might serve as a model to account for habitat
connectivity for large carnivores in the design of landscape management and landuse plans at a local scale.

The intensification of human activities has accelerated habitat
loss and fragmentation, becoming major threats for biodiversity conservation (Sanderson et al., 2002b; Pimm, 2008;
Haddad et al., 2015). Through these processes, natural areas
are distributed in small and isolated patches separated by an
altered landscape matrix (Saunders, Hobbs & Margules,
1991; Ruediger, 2004). Therefore, the species tend to be
restricted to patches with different degrees of isolation,
increasing their vulnerability to genetic drift, climate change,
and demographic and environmental stochasticity (Frankham,
Ballou & Briscoe, 2002; Gaggiotti & Hanski, 2004).
Increasing landscape connectivity, understood as the
degree to which the landscape facilitates species movements
(Taylor et al., 1993), may contribute to the mitigation of
Animal Conservation (2017) – ª 2017 The Zoological Society of London

these potentially adverse effects (Opdam, Vos & Foppen,
2002; Ara
ujo & Rahbek, 2006). Improvement of landscape
connectivity has consequently become one of the main challenges for biodiversity conservation (Rubio et al., 2014).
Connectivity maintenance and restoration is an increasingly
important goal in conservation management policies worldwide (Heller & Zavaleta, 2009). However, the implementation of these principles in specific management actions often
faces considerable difficulties, caused for example by the different scales, approaches or interests that connectivity planning and the actual implementation of policies have (Knight
et al., 2008).
The approaches and indices designed for measuring landscape connectivity are increasingly numerous and varied
(Urban et al., 2009; Saura & Rubio, 2010). Graph-based
indices, for example, have a good balance between the

Where to act for connecting the southernmost jaguars

amount of input data they require and the detail in the information outputs they are able to provide (Calabrese & Fagan,
2004). A graph-based approach represents the landscape as a
set of nodes (habitat units) functionally connected by links
that join pairs of nodes (Urban & Keitt, 2001; Saura & Pascual-Hortal, 2007). This allows the characterization of the
landscape in a spatially explicit manner and the evaluation
of the relative importance of habitat patches for maintenance
of landscape connectivity. The graph-based prioritization of
patches can be incorporated into land management plans,
protected area planning or into conservation strategies for
threatened species (Minor & Urban, 2008; Saura & Rubio,
Large carnivores are among the first species to be threatened by habitat loss and fragmentation because they need
large territories with good prey availability for survival
(Cardillo et al., 2004; Ripple et al., 2014). Large carnivores
are often used as umbrella species in the design of conservation landscapes, under the assumption that if they are preserved, other species with less strict spatial and habitat
requirements will be also preserved (Ripple et al., 2014).
Consequently, these species are particularly suitable for guiding conservation efforts for the maintenance of landscape
connectivity (Crooks & Sanjayan, 2006).
The jaguar Panthera onca is the largest felid in America and
the top predator in the ecosystems where it occurs (Sunquist &
Sunquist, 2002). In the last two centuries, its range has undergone a large retraction (>54% of its area) due to habitat loss,
depletion of natural prey, and human persecution (Sanderson
et al., 2002a). This retraction has been especially severe in the
northern and southern extremes of its distribution, where surviving populations undergo increasing fragmentation and isolation levels (Rodr ıguez-Soto et al., 2011; De Angelo et al.,
2013; Quiroga et al., 2014). The southernmost population of
jaguars survives in the Green Corridor, a region situated in Misiones Province of Argentina and nearby areas of Brazil (De
Angelo et al., 2011). This population is estimated to be around
65 individuals and is the greatest of the entire Atlantic Forest of
South America (Paviolo et al., 2016). However, habitat loss
poses a high risk of subdividing it into several smaller subpopulations with increased isolation levels and extinction risk (Di
Bitetti et al., 2016).
In order to ensure jaguar conservation in the Green Corridor, a partnership of institutions developed a management
landscape for the species to focus conservation efforts in this
area (Schiaffino et al., 2011). This conservation strategy was
designed following a regional analysis of jaguar habitat for
the entire Upper Paran a Atlantic Forest (De Angelo et al.,
2013). The core areas in the management landscape for
jaguars contain the best habitat for this species. The main
corridors represent the areas that have the potential to connect the core areas in spite of containing lower-quality habitat. However, the main corridors are crossed by roads and
are subject to increasing human pressure, driving forest conversion into other land uses which likely reduce habitat connectivity for jaguars (De Angelo et al., 2013).
Although the management landscape for jaguars has identified these areas as important for preserving habitat

J. Martinez Pardo et al.

connectivity (Schiaffino et al., 2011), the conservation of all
the forest fragments remaining in those areas is very unlikely. The National Government of Argentina and the local
government in the area of the Green Corridor developed
laws for forest preservation and restoration, including in
their fundamentals the preservation of large carnivores
(National Law N 26.331 and Provincial Law XVI N 105).
However, territorial planning policies of forest conducted by
the government are applied at a property scale, which is a
very local scale in relation to the management landscape
developed for jaguars. For this reason, it is essential to
adjust the information from the management landscape for
jaguars to the scale and demands of the forest protection
laws, prioritizing specific areas to focus the government’s
conservation efforts.
Our main objective was to incorporate connectivity criteria
of large carnivores’ species in local landscape conservation
planning. We focus on the jaguar at the southern limit of the
species’ distribution in the Green Corridor. We use graph
theory as the analytical approach with the aim of: (1) analyzing habitat connectivity and availability in the areas identified as main corridors for the jaguar; (2) detecting priority
forest patches for maintaining connectivity of the species,
and (3) proposing specific management strategies for each
area matching the relative importance and role of the forest
patches in it. By doing so, we also demonstrate the applicability of this approach in territorial planning and conservation of large and endangered carnivores in other landscapes
and regions with similar pressures on connectivity.

Materials and methods
Study area
The Atlantic Forest ecoregion extends across Brazil, Argentina and Paraguay, and is considered one of the world’s ‘biodiversity hotspots’ (Myers et al., 2000). The region has
undergone a severe transformation process, with only around
12% of the original native forest cover remaining (GalindoLeal & De Gusm~ao C^amara, 2003; Ribeiro et al., 2009).
Our study was conducted in the Green Corridor, located in
the southern portion of the Atlantic Forest (Fig. 1). The
dominant vegetation in the region is subtropical semi-deciduous forest with subtropical humid climate. The Green Corridor comprises the largest remaining area of continuous
Atlantic Forest in the world (approximately 1 000 000 ha)
and is one of the few areas with potential to preserve populations of large mammals in this ecoregion (Paviolo et al.,
This region is mainly covered by protected areas, native
forest in private lands, subsistence agriculture, plantations
and grasslands (Izquierdo, De Angelo & Aide, 2008). We
focused our work on the five most important areas classified
as main corridors in the management landscape for jaguars
(Fig. 1; Schiaffino et al., 2011). Forest cover and proportion
of different land-uses vary among these areas, providing different scenarios and challenges for the maintenance of habitat connectivity for the species.
Animal Conservation (2017) – ª 2017 The Zoological Society of London

J. Martinez Pardo et al.

Where to act for connecting the southernmost jaguars

Figure 1 The jaguar management landscape in the Green Corridor of the Atlantic Forest. The main map shows the five corridors analyzed:
(a) Foerster-Urugua- ı; (b) Yabot ı; (c) North; (d) Central and (e) South corridor. The upper left inset shows the location of the study area in
South America representing the Atlantic Forest in grey. [Colour figure can be viewed at]

Definition of forest habitat patches
First of all, we identified habitat patches and represented
them as nodes in the graph. We adapted the analysis to
the administrative scale used by the provincial government
to apply the Forest Law, selecting as nodes the properties
with native forest located in the five main corridors. Information about forest cover was obtained from aerial

Animal Conservation (2017) – ª 2017 The Zoological Society of London

photographs taken in 2009 and digitized by the Subsecretar ıa de Ordenamiento Territorial of the Ministry of Ecology of Misiones (MEyRNR). The property limits were
obtained from a vector layer containing cadastral information from the MEyRNR. We used the property layer
to divide forest fragments artificially and then measured
the area in hectares as the attribute selected for node


Where to act for connecting the southernmost jaguars

Links among habitat patches
Graph links represent the probability of direct movement
between each pair of nodes, which is obtained as a function
of the distance between them (Pascual-Hortal & Saura,
2006). We calculated this distance as an effective distance,
which is the accumulated cost of moving through the leastcost path between each pair of adjacent nodes (Adriaensen
et al., 2003). This approach considers variations in movement capacity and risk of mortality of a given species across
the different land uses occurring in the landscape matrix
(Adriaensen et al., 2003). For this calculation, we first constructed the friction surface. We developed land-cover layers
for each corridor based on Landsat 5 TM images obtained
from the National Institute for Space Research of Brazil
(INPE; see details in Supporting Information Appendix S1).
The resulting land-cover layers were transformed into friction
surfaces by assigning cost values to the different land uses
and covers present in the matrix separating forest patches in
each corridor. Connectivity analyses are sensitive to the
uncertainties and quality of the information used for determining the friction values (Zeller, Mcgarigal & Whitele,
2012). Although our analysis is not free from the potential
impacts of those uncertainties, we used the best available
information for the species. Values were assigned based on
records in the literature (Fu et al., 2010; Rabinowitz & Zeller, 2010; Gurrutxaga, Rubio & Saura, 2011; Saura et al.,
2011) and expert opinions from researchers that have been
working with jaguars in this region for over 13 years
(J. Martinez Pardo, A. Paviolo, S. Saura & C. De Angelo,
pers. comm.). The lowest friction values were assigned to
those land covers that had the best conditions for movement
and/or survival for the species and the highest values to the
least favorable land covers (Table AII in Supporting Information Appendix S1). Roads were incorporated into the friction surface since they pose an immediate mortality threat
(road kills), but also because they involve human presence,
provide access to hunting of jaguar or its prey, and the
expansion of the agricultural frontier (De Angelo et al.,
2013). The cell size used for all the raster layers was 100 m.
Data processing was developed using ArcGIS 10.1 (ESRI,
Redlands, CA, USA).
We used the friction surface to calculate effective distances among all the habitat units (nodes) of the five areas
analyzed. We obtained the effective distance values between
each pair of nodes using Linkage Mapper 0.9 (Mcrae &
Kavanagh, 2011).
The effective distances were transformed into probabilities
of movement for jaguars, which provided information on the
connectivity levels for the species as given by its homerange movements. In particular, we focused on evaluating
functional connectivity for (1) the species daily activities
rather than during occasional dispersal events, and for (2)
females, since their capacities for movement and dispersal
are more limited than those of males, and they are more relevant to species population viability than males (Eizirik,
Indrusiak & Johnson, 2002). We selected telemetry data of
the three adult females obtained from different studies

J. Martinez Pardo et al.

conducted in the Green Corridor (Crawshaw Jr, 1995; Morato et al., 2016). We used the value of the mean distance of
locations to the arithmetic center (MDLAC), which is the
average of distances between locations and the center of the
home range of each individual (Crawshaw Jr, 1995) and provides information about the distances moved by individuals
within their home range. To convert this distance (Euclidean)
to an effective distance (cost units), we calculated the mean
value of friction of the matrix within the home ranges of the
studied jaguar females and multiplied it by MDLAC. In that
way, we obtained an estimation of the mean effective distance moved by jaguar females within their home range,
which was equal to 58 225 cost units.
According to this, the probability of jaguar movement
between two patches was obtained from a negative exponential function of the effective distance separating those
patches. The decay parameter of this negative exponential
function gives the mean dispersal distance of the species
under consideration and was hence made equal to 58 225
cost units. The negative exponential function gives a probability of movement equal to 1 when the distance between
patches is equal to zero, with decreasing probability values
for larger distances separating the patches under consideration (Saura & Pascual-Hortal, 2007; Saura & Rubio, 2010).

Connectivity analysis: relative importance
of forest patches
We used the probability of connectivity index (PC) to analyze connectivity of jaguar’s habitat in the corridor areas.
This index is defined as the probability that two randomly
located points within the landscape are situated in connected
habitat units, for a given set of nodes (habitat patches) and
links (functional connections). This may occur if these two
points either fall within the same habitat patch or in two different patches that have a strong functional connection
(Saura & Pascual-Hortal, 2007). PC is based on the concept
of habitat availability, which considers a habitat patch itself
as an area where there is connectivity, and integrates the
intrapatch connectivity with the direct and indirect connections between different patches in a single measure (PascualHortal & Saura, 2006). PC is calculated via weighted graphs
and a probabilistic model of connectivity in which each connection between two patches is characterized by a given
probability of dispersal or movement. In our case, we used
the jaguar movement analysis explained in the previous section to estimate the interpatch connectivity for the PC calculation.
We prioritized forest patches by measuring the percentage
of variation in the PC index (dPC), that is the per cent
decrease in habitat connectivity and availability caused by
the loss of a given patch in the landscape, according to the
following formula:
dPC ¼ 100

PC PCelim

where dPC is the importance of a given patch in maintaining
habitat connectivity and availability according to this index,
Animal Conservation (2017) – ª 2017 The Zoological Society of London

J. Martinez Pardo et al.

Where to act for connecting the southernmost jaguars

PC is the index value in the original landscape (before
removing any patch), and PCelim is the index value after
removing the given patch (Saura & Pascual-Hortal, 2007).
We compared the values of the three fractions of dPC for
each corridor: dPCflux, dPCintra and dPCconnector. dPCintra
evaluates the contribution of a given patch in terms of area
connected within it, regardless of its position in the landscape network. dPCflux evaluates how well a given patch is
connected with the rest of the habitat patches in the landscape. dPCconnector evaluates how irreplaceable a given patch
is as a connecting element or stepping stone between the rest of
the habitat patches in the landscape (Saura & Rubio, 2010). We
decided to focus on the latter fraction (dPCconnector) for the
analyses in the main corridors given that the areas of the analyzed patches are too small for jaguars to maintain individual
territories and that the main aim of this work was to prioritize
the patches that served as stepping stones to other suitable
Additionally, we estimated the BC(PC) centrality index,
which was developed using the same probabilistic model as
that used for PC (Bodin & Saura, 2010). Both dPCconnector
and BC(PC) indices quantify the importance of patches as
connecting elements by measuring the degree of involvement
of a given patch in the movements between the remaining
patches of the landscape. However, BC(PC) index quantifies
this aspect in the intact landscape, without making patch
removal experiments, allowing to identify those patches that
play a role as stepping stones in the current landscape. In
contrast, dPCconnector quantifies the impact that the loss of a
patch would have on the maintenance of connectivity
between the remaining patches (Bodin & Saura, 2010). We
performed all connectivity analyses using the software Conefor 2.6 (; Saura & Torn e, 2009), which
computes the values of the PC and BC(PC), and the dPC
fractions: dPCflux, dPCintra and dPCconnector (see Saura &
Torn e, 2009 for more details about the procedures to obtain
these indices and fractions).
Finally, with the aim of facilitating the interpretation of
the results obtained from those indices and of summarizing
this information for management recommendations, we identified the patches with the highest contribution to habitat
connectivity (hereafter, key patches). We ranked all the
patches according to their contribution in maintaining the

connectivity in each corridor, doing that independently for
the different indices: the fraction dPCconnector, BC(PC) and
dPC. Then, for each index we used the corresponding ranking to select the minimum number of patches (MNPC)
needed to maintain the 90% of the connectivity: the MNPC
according to dPCconnector, the MNPC according to BC(PC)
and the MNPC according to dPC. All the patches included
at least in one MNPC was considered a ‘key patch’, but we
classified them into three conservation categories: ‘maximum
priority’ were those key patches included in the group of
MNPC for the dPCconnector (i.e. irreplaceable connecting
patches); ‘high priority’, were those key patches not classified as maximum priority but that were included in the
MNPC for BC(PC) index (i.e. other important stepping-stone
patches), and ‘medium priority’ were those key patches not
classified as maximum or high priority (i.e. other important
patches according to their impact in the probability of connectivity – dPC). ‘Low priority’ patches were those that
were not included in the MNPC for none of the mentioned
indices, and consequently were not considered as key

For the five areas analyzed, the resulting number of nodes
ranged between 478 (Foerster-Urugua- ı corridor) and 1281
(South corridor). On average, the Central corridor showed
the largest patches followed by the North corridor, with the
most permeable matrix found in the Yabot ı corridor but with
a very high variation (i.e. the lowest mean CWD/ED
ratio SD, Table 1). The Foerster-Urugua- ı corridor presented the smallest patches which were separated by the less
permeable matrix (Table 1).
Overall, we found a wide variation in the relative importance to maintain the connectivity of the patches in each corridor according to the PC index, with patches showing
different relevance in relation to each component of the
index (Fig. 2 and see more details in Supporting Information
Appendix S2). The relevance of the three components also
varied among the analyzed corridors (Table 1). The dPCflux
fraction of the PC index had a higher share in the habitat
availability than the other two fractions (Table 1), with this
fraction being more important in the corridors with larger

Table 1 Characterization of the analyzed corridors regarding their patches (number and mean area), the matrix (mean ratio among the cost
weighted distance -CWD- and the Euclidean distance -ED- separating each pair of nodes), the connectivity and habitat availability analyses
(percentage contributions of the three components of the dPC index; Saura & Rubio, 2010), and the minimum number of patches
(expressed as percentages of the total number of patches in each corridor) which together comprise 90% of the connectivity evaluated by
dPC, BC(PC) and dPCconnector

Number of

Mean patch
area SD (ha)

ratio ( SD)







Foerster-Urugua- ı
Yabot ı










( 13)
( 503)
( 390)
( 601)
( 88)

( 278.4)
( 2091.4)
( 250.2)
( 128.4)
( 186.7)

Animal Conservation (2017) – ª 2017 The Zoological Society of London


Where to act for connecting the southernmost jaguars

J. Martinez Pardo et al.

Figure 2 Importance of each habitat patch in the five corridors analyzed in terms of its individual contribution to the maintenance of overall
landscape connectivity as measured by dPCconnector. The colors in the legend represent the differences in relative importance from the highest (dark red = 8) to the lowest (light yellow = 1).


Animal Conservation (2017) – ª 2017 The Zoological Society of London

J. Martinez Pardo et al.

Where to act for connecting the southernmost jaguars

main potential connection pathways linking the main core
areas of this region (Fig. 3a). Interestingly, in Yabot ı corridor the small or medium-sized properties (Table 3) were
mostly those that established the main connection of high
priority patches between Yabot ı Biosphere Reserve and the
central zone of the main forest block of the Green Corridor
(Fig. 3b). The situation of the three remaining corridors was
very variable, with areas where connectivity was highly
threatened, such as the South corridor and Foerster-Urugua- ı
and others where remnant forest cover was greater, such as
the Central corridor (see more details in Supporting Information Appendix S3).

patches (e.g. Central corridor). The dPCconnector fraction
showed, however, relatively high contributions to global connectivity (Table 1). It was most important in areas with the
highest number of patches of relatively small size (South
corridor, Fig. 2e), and it contributed the least in the Central
corridor (Table 1; Fig. 2d). Regarding dPCintra, it showed a
low weight in all the corridors (Table 1).
We found a number of key patches that represent 90% of
connectivity according to the different indices, but that constitute a relatively low percentage (<50%) out of the total
number of patches in each corridor (Table 1). According to
the BC(PC) index or the dPCconnector fraction, the number of
key patches was even lower, and in general proportion of
key patches was larger in the corridors with smaller patches
(Table 1).
Based on forest patch prioritization and classification we
found that from the total area comprised with the patches of
all corridors, 47% was categorized as high or maximum priority (Table 2). However, patch percentage and area for the
three conservation categories varied among corridors showing contrasting situations (Table 3; Fig. 3 and see more
details in Supporting Information Appendix S3). In the North
corridor, for example, the highest priority patches were large
patches belonging to large properties (Table 3), with two

Using the jaguar as focal species and a graph-based connectivity analysis, we characterized the situation in five corridors in
terms of habitat availability and connectivity and identified the
priority areas where management and conservation actions
should be implemented at a scale relevant to the local application authority. Our results show that the five analyzed areas present different conditions and need distinct management actions
in order to maintain habitat connectivity for the jaguar. These
dissimilarities are due to the variability in the configuration and

Table 2 Description of the prioritization categories of forest patches to preserve jaguar habitat connectivity, including the main management
recommendations, the percentage of patches and total area covered by each category summing the patches from the five corridors
Total area
Percentage (ha)


172 262



213 224



220 551



219 921

Main management recommendation

Forest patches that play a key role
as irreplaceable connecting
patches whose loss cannot be
compensated by others
Forest patches that play a key role
as stepping stones in the current

Preserve all forest coverage. Reduce all sources of jaguar mortality
and poaching of prey. Livestock activities should not be allowed.
Implement road effects mitigation. Urgently promote habitat
restoration in adjacent patches.
Preserve all forest coverage. Reduce all sources of jaguar mortality
and poaching of prey. Reduce the conflict with livestock owners.
It is desirable to implement road effects mitigation. Promote
habitat restoration in adjacent patches.
Forest patches that play a key role
Logging activities should be carried out following low-impact
as reservoirs of habitat
techniques. Reduce all sources of jaguar mortality and poaching
prey. Reduce the conflict with livestock owners.
Forest patches that are not within
It would be advisable to maintain as much forest cover as possible.
the group of patches that comprise
Reduce sources of jaguar mortality and poaching of prey.
90% of corridor connectivity
Reduce the conflict with livestock owners.

Table 3 Mean patch area and number of patches (expressed as a percentage of the total number of patches) for the four categories of
forest patches based on their priority for conservation in each of the corridors
Maximum priority

High priority

Medium priority

Low priority


Mean patch
area (ha)


Mean patch
area (ha)


Mean patch
area (ha)


Mean patch
area (ha)


Foerster-Urugua- ı
Yabot ı

8. 7








( 14.3)
( 182.3)
( 907.7)
( 308.4)
( 129.6)

( 14.2)
( 212.0)
( 630.1)
( 406.3)
( 94.3)

Animal Conservation (2017) – ª 2017 The Zoological Society of London

( 13.3)
( 204.5)
( 745.9)
( 392.3)
( 56.2)

( 13.3)
( 199.7)
( 602.1)
( 390.8)
( 41.7)


Where to act for connecting the southernmost jaguars

J. Martinez Pardo et al.

Figure 3 Importance of each habitat patch in North corridor (a) and Yabot ı corridor (b) according to their prioritization category. The turquoise
arrows indicate the potential jaguar movements.

number of patches, the patch area and the matrix heterogeneity
of each corridor, which is typical of highly human-modified
landscapes, such as the Atlantic Forest (De Gusm~ao C^amara,
2003). These findings highlight the importance of conducting
detailed analyses, as those performed in this work, in order to
target management actions efficiently according to the specific
situation in each area.


In a graph-based view of our corridors, we found that the
PC was dominated by the dPCflux fraction. This is an
expected result since the analysis was conducted in relatively
small areas (distance between patches <30 km) and we
focused our analysis on a species of large movement capacities (Crawshaw Jr, 1995). For large movement capacities
species, the loss of a single patch is unlikely to completely

Animal Conservation (2017) – ª 2017 The Zoological Society of London

J. Martinez Pardo et al.

hinder the possibility of moving using other patches or alternative paths (Hodgson et al., 2009; Saura & Rubio, 2010),
which results in high dPCflux values. Stepping-stone patches
were, however, significantly important for jaguar habitat connectivity in the area; jaguar movements between the core
habitat areas depended to some extent, even if not completely, on individual forest patches that could be used as
intermediate connectors along the way. Indeed, the comparison of our dPCconnector values with those reported elsewhere
(Saura & Rubio, 2010; Gurrutxaga et al., 2011) shows that
our values are relatively high in all the corridors, except for
the Central corridor, which has a greater proportion of forest
and larger patches. These results suggest that connectivity in
the Central corridor should be less limited and less dependent on the possible loss of small habitat patches, thereby
explaining the lower dPCconnector values. Conversely, values
of dPCintra were lower than those of the two mentioned fractions for all corridors, suggesting that patch connectivity
plays a fundamental role in maintaining total habitat availability for jaguars in the study region, and that partial or
total loss of connections would have a great impact on the
populations living in this region (Saura & Rubio, 2010).
We also detected that a great part of the connectivity
(90%) is concentrated in a low percentage of key forest
patches. This result is especially useful for territorial and
conservation planning since it allows decision makers to concentrate conservation efforts in the areas of the greatest
importance in maintaining habitat connectivity for this species. Likewise, our results highlight that some patches are of
higher priority than others in a given area, and the loss of
those key patches would have a greater impact on the functioning of the network of all the remaining patches.
The observed variability in the percentage and mean size
of key patches may serve as a first approximation to establish the most suitable management strategies. On one hand,
there are corridors with a low percentage of maximum or
high priority patches, but with those patches having large
areas, such as in the North corridor, where most of the land
belongs to forestry companies. These patches should be designated as protected areas, preferably with high protection
levels. If that would not be possible, an alternative would be
that they are declared as ‘areas of high conservation value’
by the companies that run them. Such designation has
already been used by some forestry companies working in
the area and it is promoted by the principles of the Forest
Stewardship Council (FSC, 1996). On the other hand, in
areas such as Yabot ı corridor, where the percentage of key
patches is considerably higher and their mean patch area is
smaller, corridors should be implemented preserving the set
of stepping-stone patches by a regional program promoting
jaguar habitat conservation through economic incentives provided by the government (Trainor et al., 2013). This initiative could easily be included in the current National and
Provincial legislation, which already considers this type of
incentives in native forest areas of great importance for conservation (Laws N 26.331 and XVI N 105, respectively).
Two of the analyzed areas deserve special attention since
they maintain key connections for jaguar habitat conservation
Animal Conservation (2017) – ª 2017 The Zoological Society of London

Where to act for connecting the southernmost jaguars

not only in this region of the Atlantic Forest but also
throughout its distribution range. These areas are the South
and Yabot ı corridors, which maintain connections with the
protected areas that form the southernmost distribution of the
jaguar (De Angelo et al., 2011). In both cases, connectivity
is seriously threatened. In Yabot ı area, the estimated jaguar
densities are the lowest values detected for the region, with
only 10 individuals persisting (Paviolo et al., 2008). This
estimation indicates that, despite its large area (more than
200 000 ha), Yabot ı can hardly maintain by itself a viable
jaguar population and, therefore, maintaining connectivity
between this area and the rest of the Green Corridor is crucial for jaguar survival. Implementing measures to mitigate
the barrier effect of the road extending across this area is
urgent. In addition, it would be necessary to promote restoration projects of native forest in areas adjacent to maximum
or high priority patches.
The South corridor connects a protected area of
13 000 ha in the south-west extreme of the Green Corridor.
Considering the large territories of jaguars, the size of this
area is relatively small and therefore, only a few individuals
can survive in this region and possibly their home ranges
include the analyzed area. The main land use in this corridor
is pine plantations and, therefore, as proposed for the North
corridor, actions should be conveyed with the companies that
exploit these areas. Big companies or pools of companies
that exploit large areas can use these prioritization strategies
for regional planning. The Sustainable Forest Mosaics Initiative is an example of the implementation of such strategies
in other regions of the Atlantic Forest (Mesquita et al.,
A common challenge in animal conservation research is to
develop results that can be easily adapted and incorporated
into decision-making and management strategies. For our
analysis, we divided remnant forest patches ‘artificially’
using property limits; this procedure is of great advantage
since by linking the nodes directly with the land ownership
we gave priority to the units that can (or cannot) be truly
influenced by management actions. However, this procedure
also implies that most patches were adjacent to one another,
evidenced complex shapes resulting from cadastral errors,
and had much smaller areas than those of the patches
defined exclusively based on the continuity of forest cover.
The adopted graph-based approach is able to reduce the
potential impacts on the spatial prioritization resulting from
using land properties, rather than forest continuity when
defining the patches (nodes) for the analyses. On one hand,
Saura & Pascual-Hortal (2007) showed that the landscapelevel value of the PC index is not affected by the presence
of adjacent habitat patches imposed by ownership, administrative or management limits (property 13 of the PC index
as described in that paper). On the other hand, BlazquezCabrera, Bodin & Saura (2014) showed that the spatial prioritization of connectivity areas provided by the PC index is
robust against different scales or hierarchical levels used in
defining the habitat patches. While it is important to take
into account the aspects related to the patch definition and
scale of analysis when interpreting the results, we could here

Where to act for connecting the southernmost jaguars

address a specific management need at the real working scale
of the government or local organizations by defining patches
based on land properties. This is a fundamental issue that
facilitates the implementation of recommendations that
emerge from our analysis as viable management and conservation measures, reducing the gap that often exists between
the generated knowledge and planning and implementation
on the ground (Opdam et al., 2002).
Maintaining habitat connectivity is one of the main strategies proposed for large carnivores conservation at the global
level (Ripple et al., 2014) and, indeed, some conservation
plans have already included this action (e.g. Global Tiger
Initiative, Paseo Pantera, etc.). There are initiatives promoting connectivity improvement for the jaguar throughout its
distribution range that have identified potential corridors
among the main populations of the species (Rabinowitz &
Zeller, 2010). Following the same objective, some works
have been conducted at the scale of the entire Atlantic Forest
(Paviolo et al., 2016) and in the southern part of this region
(De Angelo et al., 2013). Given the resolution and the aims
of those works, connectivity analyses were of structural type
and did not consider the functional aspect related to the
movement capacity of jaguars. Here, we incorporated this
aspect and provided detailed information to improve connectivity for the jaguar in specific and strategic portions of its
In broad terms, the actions we suggest for preserving
habitat connectivity for the jaguar in this region (Table 2)
agree with recommendations proposed for other large carnivorous species (Chapron et al., 2014; Ripple et al., 2014) but
they are prioritized spatially and indicated in land management units. We hope that the specific approach we adopted
for applying the graph-based methodology might serve as an
example for other cases in which large carnivores and landuse planning are involved and where it is necessary to bring
the analytical results closer to the scale and needs of the
actual implementation of conservation management measures
on the ground.

Fundaci on Vida Silvestre Argentina, U.S. Fish & Wildlife
Service and CONICET provided the funding to this project,
and the Society for Conservation GIS, the ESRI Conservation Program and Planet Action provided the software and
hardware for developing the analysis. We also want to thank
the help and data received from the Ministry of Ecology and
Natural Resources of Misiones province, especially from
J.M. D ıaz, J. Solari and V. Centuri on. Additionally, we are
grateful to the associate editor and two anonymous reviewers that contributed greatly to the improvement of the

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Supporting information
Additional Supporting Information may be found in the
online version of this article at the publisher’s web-site:
Appendix S1. Land-use analysis.
Appendix S2. Importance of each habitat patch in the five
corridors analyzed in terms of its individual contribution to
the maintenance of overall landscape connectivity as measured by BC(PC), dPC, dPCflux and dPCintra.
Appendix S3. Importance of each habitat patch in the Foerster-Urugua- ı corridor (a), Central corridor (b) and South corridor (c) according to its prioritization category.

Animal Conservation (2017) – ª 2017 The Zoological Society of London

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