Evaluating Landscape Connectivity for

Puma concolor and Panthera onca Among

Atlantic Forest Protected Areas

Camila S. Castilho, Vivian

C. S. Hackbart, Vânia R. Pivello & Rozely
F. dos Santos

Environmental Management

ISSN 0364-152X

Environmental Management
DOI 10.1007/s00267-015-0463-7

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 Author's personal copy
Environmental Management
DOI 10.1007/s00267-015-0463-7
Evaluating Landscape Connectivity for Puma concolor
and Panthera onca Among Atlantic Forest Protected Areas
Camila S. Castilho1 • Vivian C. S. Hackbart1 • Vânia R. Pivello1 • Rozely F. dos Santos1
Received: 8 April 2014 / Accepted: 21 March 2015
Ó Springer Science+Business Media New York 2015
Abstract Strictly Protected Areas and riparian forests in
Brazil are rarely large enough or connected enough to
maintain viable populations of carnivores and animal
movement over time, but these characteristics are funda-
mental for species conservation as they prevent the ex-
tinction of isolated animal populations. Therefore, the need
to maintain connectivity for these species in human-
dominated Atlantic landscapes is critical. In this study, we
evaluated the landscape connectivity for large carnivores
(cougar and jaguar) among the Strictly Protected Areas in
the Atlantic Forest, evaluated the efficiency of the Mosaics
of Protected Areas linked to land uses in promoting land-
scape connectivity, identified the critical habitat connec-
tions, and predicted the landscape connectivity status under
the implementation of legislation for protecting riparian
forests. The method was based on expert opinion translated
into land use and land cover maps. The results show that
the Protected Areas are still connected by a narrow band of
landscape that is permeable to both species and that the
Mosaics of Protected Areas increase the amount of pro-
tected area but fail to increase the connectivity between the
forested mountain ranges (Serra do Mar and Serra da
Mantiqueira). Riparian forests greatly increase connec-
tivity, more than tripling the cougars’ priority areas. We
note that the selection of Brazilian protected areas still fails
Electronic supplementary material The online version of this
article (doi:10.1007/s00267-015-0463-7) contains supplementary
material, which is available to authorized users.
& Camila S. Castilho
cscastilho@ib.usp.br
1 cating the importance of maintaining ecological corridors
Laboratório de Ecologia da Paisagem e Conservação,
Departamento de Ecologia, Universidade de São Paulo, Rua
do Matão, 321 - Trav. 14, São Paulo, SP 05508-090, Brazil
to create connectivity among the legally protected forest
remnants. We recommend the immediate protection of the
priority areas identified that would increase the structural
landscape connectivity for these large carnivores, espe-
cially paths in the SE/NW direction between the two
mountain ranges.
Keywords Carnivore conservation
Cougars
Jaguars

Permeability
Riparian Forest
Environmental planning
Introduction
The cougar (Puma concolor) and jaguar (Panthera onca)
are the largest Brazilian terrestrial carnivores and have
been included at endangered species lists, as least concern
(cougar) and near threatened (jaguar) by the International
Union for Conservation of Nature—IUCN (2013) and as
vulnerable (both species) on the national and regional red
lists of threatened fauna (MMA 2008; Bressan et al. 2009).
They still can be found in the Brazilian Atlantic Forest,
which is a highly fragmented biome. Only 11.2 % of this
biome’s original area remains and 1.6 % is under legal
protection (Ribeiro et al. 2009). The majority of Strictly
Protected Areas (areas determined by Brazilian law as re-
stricted to human use) are not large enough to sustain long-
term viable populations for either species (Beier 1993;
Sollmann et al. 2008). Therefore, maintaining landscape
connectivity in this scenario is a high priority.
Protecting large carnivores requires protecting land-
scape connectivity between protected areas (Newmark
1995; Soulé and Noss 1998, Di Minin et al. 2013), indi-
and connectivity paths (Zeller et al. 2011). This assertion
especially applies to jaguar, which are now extremely rare
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(Galetti et al. 2013), and is at a high risk of extinction
within the next 88 years in parts of the Atlantic Forest
(Ferraz et al. 2012). For the entire Atlantic Forest biome,
the conservation status of these populations may be even
worse (Ferraz et al. 2012).
Until a few years ago, Brazilian legal decisions had not
taken into consideration the concept of connectivity when
establishing protected areas. For example, the protected
areas are concentrated in areas of no interest in terms of
human use such as mountain ranges with nutrient-poor
soils (Terra and Santos 2012). Only after year 2000
(Brazilian law 9 985 July 2000) the concept of landscape
connectivity has been incorporated into protected areas
strategies, primarily by establishing mosaics of protected
areas and ecological corridors. However, these measures
are still extremely modest and the decisions are based on
‘‘structural connectivity,’’ referring to the number of
habitat patches and their spatial configuration in the land-
scape (Baudry and Merriam 1988), rather than ‘‘functional
connectivity’’ that refers to the ability of individuals to
move through the landscape.
Riparian forests are extremely important for the land-
scape connectivity of different Brazilian biomes for several
species (Metzger 2010), considered as potential ecological
corridors for jaguar (Silveira et al. 2014) and have been
shown to be preferred by cougar in their movements
through the fragmented landscapes (Dickson and Beier
2002). Riparian forests acquired legal protection and
regulation in 1965 (Federal Law 4 771/1965), but in spite
of their importance for conservation a recent change in the
federal law has reduced the width requirement for protec-
tive corridors from 30 m to 15 m on each side of the river
(Brazil’s Federal Law 12 651/2012). Moreover, in practical
terms, the implementation of riparian forest protection has
rarely been followed regardless of the penalties mandated
by the law (Federal Law 4 771/1965, Federal Law 6
905/1998 and more recently Federal Law 12 651/2012).
Has the allocation of protected areas ensured adequate
landscape connectivity for protecting jaguar and cougar
over time? How much landscape connectivity remains?
How much riparian forests contribute for landscape con-
nectivity? We use real land use/land cover information and
simulated riparian forest defined according to Brazilian
laws in combination with expert opinion on habitat per-
meability to identify connectivity among two sets of pro-
tected areas: (i) only Strictly Protected Areas (National and
State Parks) and (ii) Mosaics of Protected Areas (which
combine fully protected and sustainable-use areas). We
believe that the obtained results will be very useful to de-
cision makers because they contribute to landscape man-
agement strategies for maintaining and improving the
connectivity for large carnivores in the extremely frag-
mented Atlantic Forest.
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Methods
Study Area
The study area includes regions of the two major remnants
of the Atlantic Forest in São Paulo State (southeastern
Brazil) located between two major rugged mountain ranges
(Serra do Mar and Serra da Mantiqueira), 1,428,194.5 ha in
area (Fig. 1). The area is important for water resources and
biodiversity conservation. Since the creation of the first
Brazilian National Park in 1937, other important protected
areas have been established in the region (Table 1). The
Atlantic Forest is an endangered biome with less than 12 %
of its original area remaining (Ribeiro et al. 2009), and it
still receives only limited legal protection (Ribeiro et al.
2009).
The National System of Protected Areas, SNUC
(Brazilian Law 9 985 18 July 2000), has defined several
categories of protected areas in Brazil. In this study, we
refer directly to the Strictly Protected Areas and the Mo-
saics of Protected Areas. The first category is included in
the group of indirect use areas, in which the primary goal is
nature conservation. The Mosaics of Protected Areas in-
tegrate a set of neighboring protected areas that may (or
may not) belong to different categories, including those
areas for sustainable use. Mosaics aim to combine both
conservation and social objectives in a regional context.
Land Cover Mapping
The regional land use/land cover was mapped in ArcGIS
10 (Environmental Systems Research Institute, Inc.) using
SPOT satellite images (2007 and 2009, supplied by the
Environmental Secretariat of São Paulo State-SMA) with
RGB color band composition and 2.5 m of resolution,
orthorectified and georeferenced. We adopted the Univer-
sal Transverse Mercator (UTM) coordinate system, zone
23S, and Geodesic System SIRGAS 2000. Shapefiles with
hydrography, roads, and protected areas were obtained
from free databases maintained by the Brazilian Environ-
ment Ministry-MMA (http://mapas.mma.gov.br, accessed
11 June 2013) and the National Water Agency-ANA
(http://www.ana.gov.br, accessed 11 June 2013). We
mapped at a 1:50,000 scale in the preview display and
generated a final map at 1:100 000.
To evaluate the riparian forest, we created buffers of
15 m and 30 m along both sides of all rivers, which is
consistent with the current minimum width of the protected
riparian forest belt defined by the federal environmental law
(Federal Law 12 651—May 25, 2012). We substitute the
real land use/land cover for secondary forest at intermediate
and advanced stages, simulating a continuous corridor.
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Fig. 1 The study area: 1–6 state and national Strictly Protected range, while Protected Areas 2, 3, 4, and the Bocaina Mosaic (B) are
Areas; A–B Mosaics of Protected Areas. Protected Areas 1, 5, 6, and in the Serra do Mar mountain range
the Mantiqueira Mosaic (A) are in the Serra da Mantiqueira mountain

Table 1 Categories of protected areas established by law in the Serra do Mar and Serra da Mantiqueira (São Paulo State, Brazil)
Protected Area Site (mountain range) Decree Area Legal instrument
(year) (hectares)

Strictly protected areas

National areas
Itatiaia National Park Serra da Mantiqueira 1937 28,084.100 Decree n8 1.713—June 14th 1937
Decree n8 87.586—September 20th 1982
Serra da Bocaina National Park Serra do Mar 1971 104,044.89 Decree n8 68.172—February 04th 1971
Decree n8 70.694—June 08th 1972
State Areas
Bananal Ecological Station Serra do Mar 1964 884 Decree 26.890—April 03rd 1964
Decree. 43.193—March 12th 1987
Campos do Jordão State Park Serra da Mantiqueira 1941 8341.00 Decree-Law 11.908—March 27th 1941
Serra do Mar State Park Serra do Mar 1977 315,390.69 Decree 10.251—August 30th 1977
Decree 13.313—March 06th 1979Decree 19.448—
August 30th 1982
Mananciais de Campos do Jordão Serra da Mantiqueira 1993 502.96 Decree 37.539—September 27th 1993
State Park
Mosaics
Bocaina Mosaic Serra do Mar 2006 221,754 Ordinance MMA n°349—December 11th 2006
Mantiqueira Mosaic Serra da Mantiqueira 2006 445,615 Ordinance MMA n°351—December 11th 2006

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Expert Consultation
To assign permeability values to each land use/land cover,
we selected experts (according LaRue and Nielsen 2008,
2011) who have developed studies on the cougar and
jaguar in the Atlantic Forest that focused on the species’
habitat and landscape use. To obtain the researchers’ names
and addresses, we used search tools such as Web of
Knowledge, Science Direct, Google Scholar, Elsevier,
Scielo, and Lattes-CNPq. We excluded individuals who
only worked occasionally with the species, as well as un-
dergraduate students.
Eleven specialists were selected and consulted by email
about the landscape permeability for cougar and jaguar.
They received explanations about the project and a table to
be completed with values varying from (-10) to (10),
where (-10) corresponds to the least permeable land
use/land cover (higher resistance to animal movement) and
(10) corresponds to the most permeable (lower resistance
to animal movement). To obtain more consistent results,
we eliminated the highest and the lowest values for each
experts for each land use/land cover type and worked with
the mean of the remaining intermediate values (Goh et al.
1996) standardized on a 0–100 scale (Legendre and Le-
gendre 1998).
Landscape Connectivity Analysis
We used isolation-by-resistance and least-cost-corridor
analyses to determine the level of connectivity of the
considered protected areas for the focal species. We chose
to perform both analyses because circuit theory (the basis
of the isolation-by-resistance method) complements the
least-cost analysis and helps prioritize the important areas
for conservation according to connectivity (McRae et al.
2008).
An isolation-by-resistance analysis was conducted using
the program CIRCUITSCAPE 3.5.8 (McRae 2006) and the
land use/land cover maps and permeability values as inputs
to generate current maps that identified all the paths that
were potentially used by cougar and jaguar. The isolation-
by-resistance analysis uses circuit theory to predict con-
nectivity in heterogeneous landscapes for individual
movement, gene flow, and conservation planning. Circuit
theory includes a theoretical basis in random walk theory
and an ability to evaluate contributions of multiple dis-
persal pathways. We chose the one-to-all (McRae and Shah
2009) mode and the option to connect a cell with eight of
its neighbors. In the one-to-all mode, connectivity is cal-
culated between all focal regions. In each interaction, one
focal region is connected to a source, which represents in
this study the Strictly Protected Areas or Mosaics while the
other places are considered inactive. We used land use/land
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cover raster cell sizes of 50 m due to software limitations.
However, we felt confident with this cell size based on
McRae et al. (2008), which evaluated different cell sizes
and concluded that as long as the cell size remains fine
enough to capture relevant landscape elements, similar
results will be obtained even with an increased cell size.
A least-cost-corridor analysis was performed using the
Linkage Mapper 1.0.2 ArcGIS tool (McRae and Kavanagh
2011). This analysis shows the relative value of each grid
cell in providing connectivity among focal areas (connec-
tivity among Strictly Protected Areas or among Mosaics of
Protected Areas) and identifies the routes that facilitate or
impede species movement among the focal areas according
to the software guidelines (McRae and Kavanagh 2013). In
this analysis, we used land use/land cover map raster cell
sizes of 10 m.
Both analyses used a habitat raster map that specified
the conductivity or resistance of each cell (raster pixel) in
the landscape (values for the cougar and jaguar were de-
termined by the specialists for each land use/land cover
type) and a map of focal regions (Strictly Protected Areas
or Mosaics of Protected Areas) to identify the connections
between the focal regions (Mcrae and Shah 2011; McRae
and Kavanagh 2013).
To determine the critical habitat connections, we used
Pinchpoint Mapper, part of the ArcGIS Linkage Mapper
Toolkit (McRae and Kavanagh 2011). Pinchpoint Mapper
links CIRCUITSCAPE to the maps produced by Linkage
Mapper (McRae 2012), providing maps that identify pinch
points in the least-cost corridors and the effective resis-
tance values for the Linkage Mapper maps to complement
the least-cost-corridor results (McRae 2012). The Pinch-
point map identifies regions where the loss of a small area
could seriously jeopardize the landscape connectivity
(McRae 2012). We used 5 km width cutoff for pinch
points, as used by McRae (McRae 2012), and 10 m cells.
The final maps were visualized in ArcGIS 10.
We repeated the three analyses (isolation-by-resistance,
least-cost-corridor, and pinch point) for a legally protected
riparian forest belt of 15 and 30 m along both sides of the
river. The guidelines are detailed on a flowchart (Fig. 2)
showing cost surface and focal regions inputs and analysis.
In order to compare remaining scenarios, we add up all
priority connectivity areas identified by pinchpoint analysis
in one unique region. These regions are also used for
measure connectivity paths width and landscape metrics
analysis. The connectivity areas used for metric analysis
were 11,517.7, 48,835.2, and 59,616.04 ha, for scenarios
real, simulated 15 m and simulated 30 m, respectively. We
only used cougar areas on this analysis since jaguar areas
are part of cougar’s. We have interest in knowing the land
use/land cover present at landscape connectivity paths at
each scenario (real, simulated 15 and 30 m) in order to
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Fig. 2 Flowchart of the method
facilitate landscape management decisions. Therefore, to
understand the landscape structural heterogeneity (Chapin
et al. 2012) on the connectivity priority areas (pinch points)
for cougar in all scenarios, we applied the following
landscape metrics, calculated in Patch Analyst 4.0 (ArcGIS
10), that could express landscape fragmentation and con-
nectivity (Noss 1990; Gustafson and Parker 1992; Shu-
maker 1996): patch density (PD), number of patches
(NumP), mean patch size (MPS), total edge (TE), edge
density (ED), mean patch edge (MPE), patch shape (PS),
and mean shape index (MSI).
Results
The expert consultation inquiries had a return rate of 64 %
(seven experts responded). As expected, human-altered
environments were less permeable to jaguar than to cougar,
and wet environments were more resistant to cougar
(Fig. 3). The values of land use/land cover permeability
attributed by the experts were transferred to the respective
polygons of the land use/land cover map and revealed the
identity of classes favorable to the movement of cougar and
jaguar (Figs. 4, 5, 6, 7; Table 2). The isolation-by-resis-
tance and least-cost-corridor results indicated that forest
remnants protected by Strictly Protected Areas (Table 1)
are still connected by a permeable landscape for both
cougar and jaguar (Figs. 4A, 5A) but only in a small por-
tion of the study area (Figs. 4A, 5A). Although the
delimited landscape has more than 450,000 ha of Strictly
Protected Areas, allowing species movement in a SW/NE
direction in Serra do Mar, no significant links have been
legally established to enable movement from Serra do Mar
to Serra da Mantiqueira in a SE/NW direction (Fig. 1).
The establishment of the Mosaics of Protected Areas
increased the protected area by 46 % (although a part of
this area is under a lower degree of protection); however, it
only increased the connectivity and protection of animal
movement in a SW/NE direction in the Mantiqueira Mo-
saic. A lack of connectivity between the mountain ranges
still remains (Figs. 6, 7). All the remaining connectivity
links between both Mosaics prioritized for cougar
(Fig. 6A) consist of a narrow area that is 11 517.7 ha, or
0.8 % of the study area, for jaguar are even narrower, with
5489.00 ha, or 0.4 % of the study area.
Other results using mosaics as Input Focal Regions
consist on simulated connectivity paths using simulated
riparian forests of 15 m for cougar (Fig. 6B) and jaguar
(Fig. 7B) and 30 m for cougar (Fig. 6C) and jaguar
(Fig. 7C). The inclusion of the scenario that utilizes a 15 m
riparian forest buffer (according to Law 12 651/2012) in-
creased the connectivity for the two species in terms of
both the least-cost-corridor strategy and the pinch points
adding several new paths that create a wider connectivity
area (Figs. 4B, 5B, 6B, 7B). This option provides an ex-
tended movement area for cougar, 48 835.2 ha, a 324 %
increase in the priority connectivity area (Fig. 6B) com-
pared with real state of land use/land cover (Fig. 6A).
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Fig. 3 Mean values of permeability (±standard deviation, SD) of the different land use/land cover classes assigned by experts (n = 7, 64 % of
all Brazilian experts consulted) for Puma concolor (cougar) and Panthera onca (Jaguar)

Fig. 4 Landscape connectivity for cougar (Puma concolor) in the B scenario simulating 15 m riparian forest and C scenario simulating
study area according to the pinch point analysis showing connectivity 30 m riparian forest
paths among the Strictly Protected Areas. A real remaining paths;

Moreover, for jaguar, simulated riparian forest of 15 m compared with real state of land use/land cover (Fig. 7A).
provide an extended movement area, 26,344.1 ha, a 480 % We also included an analysis of the scenario that included a
increase in the priority connectivity area (Fig. 7B) minimum 30 m riparian forest (according to a previous

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Fig. 5 Landscape connectivity for jaguar (Panthera onca) in the
study area according to the pinch point analysis, showing connectivity
paths among the Strictly Protected Areas. A real remaining paths;
Brazilian law from 1965) (Figs. 4C, 5C, 6C, 7C). For this
scenario, the connectivity area linking Mosaics compared
with real land use/land cover area increased by more than
400 % for cougar, and more than 500 % for jaguar, re-
sulting in a connectivity area of 59,616.04 ha (6C com-
pared with 6A) and 28,217.9 ha (7C compared with 7A) for
cougar and jaguar, respectively.
We observe that there is a great improvement of link-
ages between Strictly Protected Areas with simulated ri-
parian forest of 15 m and 30 m (Figs. 4B, C, 5B, C) when
comparing with the real scenario (Figs. 4A, 5A) or Mosaics
(Figs. 6, 7). We believe that this result is due to the high
amount of river courses at the study area and no-existence
of riparian forest at real scenario. Although we identified a
substantial improvement in the availability of movement
paths, especially those connecting, several of these paths
would cross large agricultural and urban areas on the
border of Highway BR-116. Therefore, this narrow band of
B scenario simulating 15 m riparian forest and C scenario simulating
30 m riparian forest
riparian forest would most likely be ineffective in sup-
porting the connectivity of the habitat. An increase in
movement paths generated by linking the Mosaics
(Figs. 6B, C, 7B, C) is most likely the best connectivity
option.
The landscape metrics of the different land use/land cover
based on the cougar’s connectivity priority areas (Table 2)
showed that secondary forest at intermediate and advanced
successional stages, secondary forest at the initial succes-
sional stage, forestry, and grassland prevailed in the land-
scape with a 15 m, 30 m, or no legal band of protected
riparian forest (representing 96.3, 97 and 94.7 % of the total
connectivity priority areas, respectively) (Table 2). Forestry
was the dominant land use/land cover class when consider-
ing the priority areas without the riparian forest bands,
comprising large patches and intermediate edge values,
whereas secondary forest at intermediate and advanced
stages showed a larger number of patches and a greater edge
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Fig. 6 Landscape connectivity for cougar (Puma concolor) in the
study area according to the pinch point analysis showing connectivity
paths among the Protected Areas Mosaics. A real remaining paths;
density, indicating the high fragmentation level of these
classes (Table S1). The grassland class coverage of the ri-
parian forest band increased to 44.6 % in the landscape sce-
nario (Table 2) and to 40.3 % in the simulated 30 m riparian
band scenario, whereas secondary forest at intermediate and
advanced successional stages maintained a large number of
patches, and the Forestry land use/land cover class lost its
previous importance (Table S1). Secondary forest at inter-
mediate and advanced successional stages, added to riparian
forest, maintained a cover percentage of 27.4 % (3155.9 ha)
in the real status scenario, 26.1 % (12,745.9 ha) in the 15 m
riparian band scenario, and 33.9 % (20,209.8 ha) in the 30 m
simulated scenario, where we observed a greater amount of
mature forest available (Table S1). The main difference be-
tween the scenarios with 15 m and 30 m of simulated riparian
forest was the amount of secondary forest at the intermediate
and advanced stages available, which increased by 20 % in
the connectivity path area.
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B scenario simulating 15 m riparian forest and C scenario simulating
30 m riparian forest. The widths of the connectivity paths are
indicated
The most dramatic contrasts in our results were ob-
served when comparing the land use/land cover of the
actual scenario with both of the simulated scenarios that
included riparian forests. In these comparisons, we ob-
served and concluded that no riparian forest, neither the
previous 30 m minimum (law from 1967 to 2012) nor the
current 15 m minimum (law 2012 until today), is present in
the study area. The lack of any amount of legally protected
riparian forest is the most striking result from our scenario
comparison. The existing connectivity path (union of
pinchpoint analysis results) is 6 km wide at its largest
width and only 1 km wide at its narrowest width for cougar
and 2.5 km and 0.7 km for jaguar, and the 15 m and 30 m
simulated scenarios with riparian forest have widths that
are 16 km and 20 km for cougar (Fig. 6), 8 km and 11 km
for jaguar (Fig. 7).
The other main results are the very narrow connectivity path
remaining between the protected areas (Figs. 4A, 5A, 6A, 7A).
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Fig. 7 Landscape connectivity for jaguar (Panthera onca) in the B scenario simulating 15 m riparian forest and C scenario simulating
study area according to the pinch point analysis, showing connectivity 30 m riparian forest. The widths of the connectivity paths are
paths among the Protected Areas Mosaics. A real remaining paths; indicated

Table 2 Principal land use/land cover classes for two scenarios

Scenario Total area (ha) Secondary forest: intermediate Secondary forest: Forestry Grassland
and advanced stages initial stage
A P A P A P A P

Without Riparian Forest buffer 11,515.70 3,157.5 27.4 2,327.3 20.2 2,194.1 19.0 3,234.8 28.1
15 m of Riparian Forest buffer 48,835.23 12,742.50 26.1 7,243.5 14.8 5,307.2 10.9 21,781.7 44.6
30 m of Riparian Forest buffer 59,616.04 20,200.70 33.9 7,992.55 13.4 5,625.2 9.4 23,998.5 40.3
A area in hectares, P class percentage in the landscape

Discussion connectivity in the region is narrow and lacks legal pro-

The forest remnants of Serra do Mar and Serra da Man-
tiqueira are located in a human-dominated area in the most
highly developed region of Brazil. Nevertheless, this study
shows that both areas maintain structural landscape con-
nectivity for cougar and jaguar even though the SE/NW
tection (Figs. 4A, 5A, 6A, 7A). The strategy adopted to
select and establish Strictly Protected Areas in Brazil,
where mountain ranges are generally preferred for this
designation because of their low economic value, created
isolated and restricted protected areas. More recently, there
has been an increasing concern about preventing the

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isolation of Strictly Protected Areas (Brazilian Law 9
985/2000). After the 2000s, this concern motivated the
establishment of Mosaics of Protected Areas as an attempt
to increase landscape connectivity and promote species
movements among the protected areas with various degrees
of protection. Additionally, these mosaics are intended to
integrate the protection of highly restrictive and less re-
strictive protected areas due to their various uses.
We observed several SW/NE paths of connectivity in
the area protected by the Mantiqueira Mosaic, although the
resulting almost 50 % increase in protection area (after the
implementation of mosaics) did not ensure an increase in
paths of connectivity and failed to effectively guarantee
protection of the SE/NW connection between the Serra da
Mantiqueira and Serra do Mar remnants. As a result, le-
gally protected landscape connectivity was not present
between these important biodiversity sites.
Comparing scenarios that include and do not include the
riparian forest in the landscape showed that although these
forests have been protected—in theory—by the Brazilian
law since 1965, they have not been effectively protected in
the study area. A remarkable increase in landscape con-
nectivity could have been achieved if the reduced riparian
forest band that was mandated by the recent revision of the
federal law (in 2012) had been established; the connectivity
priority areas for cougar would have increased by more than
300 % with the inclusion of a 15 m riparian forest and by
over 400 % with the inclusion of a 30 m riparian forest.
However, this would likely be an increase in structural
connectivity. Brazilian researchers argue that, regardless of
the biome, the taxonomic group, soil or type of topography
for faunal, a minimum of 50 m on each side of the river, is
necessary for providing functional connectivity on Brazilian
biomes (Metzger 2010), and regarding water quality and
sediment retention, at least 52 m on each side of the river is
needed (Sparovek et al. 2002). Therefore, based on the
current riparian forest condition, an increase of 300–400 %
in the connectivity paths linking mosaics would be a good
start in improving large carnivore protection and establish-
ing the credibility of the environmental policies, although
based on existing research for full functional connectivity
riparian forests should be even larger that 30 m.
The 300–400 % increase could mean connectivity areas
that are 16–20 km wide, whereas the real existing con-
nectivity areas are 1–6 km wide for cougar (Fig. 6), for
jaguar an increase of 400–500 % could mean connectivity
areas 8–11 km width, whereas the real connectivity area is
0.7–2.5 km width (Fig. 7). Rodrı́guez-Soto et al. (2013)
suggested a minimum width of 14 km for viable corridors
and a 2.5 km width for potential corridors for jaguar in
Mexico. Both of the simulated scenarios with riparian
forest for cougar were larger than 14 km, although it is
possible that even they could not provide functional
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Environmental Management
connectivity. For jaguar, the connectivity paths are nar-
rower than 14 km and possibly a riparian forest larger than
30 m is needed to provide connectivity areas wide as Ro-
drı́guez-Soto et al. (2013) suggested for jaguar in Mexico.
Metrics analysis was used to support management de-
cisions and actions. The four land use/land cover classes
that dominated the critical habitat connections in the
landscape (secondary forest in intermediate and advanced
successional stages, secondary forest in the initial stage of
succession, forestry, and grassland) are still viable for
management strategies. We consider it an advantage that
these habitat areas do not contain consolidated urban areas
or mines. This advantageous situation may facilitate the
implementation of restoration of riparian forest and other
conservation strategies. One of these strategies is to man-
age the initial successional stages of secondary forest to
encourage its development to the intermediate and ad-
vanced stages and prevent fragmentation and habitat loss.
These procedures would increase the persistence of
populations of larger animals—including the species that
need large territories—as well as improve the ecosystem
functions they support (Shumaker 1996).
In the study area, even in the case of small patches,
mature forests are able to maintain landscape connectivity,
most likely due to their spatial arrangement as areas located
within the forestry (monoculture plantations) (Garmendia
et al. 2013) land use/land cover class, producing a landscape
pattern that is suitable for large felines (Mazzolli 2010).
Forestry is a land use/land cover class subject to regular
management, and the landscape changes greatly when trees
are removed or planted. However, despite the temporary
exclusion of the protective vegetation and the occurrence of
a reasonable level of disturbance due to the continuous
presence of employees, the presence of forestry employees
discourages harvesting and poaching, and thus indirectly
provides wildlife protection (Mazzolli 2010). A decrease in
native species is usually observed when the planted trees are
small. Nevertheless, forestry lands usually have a very
positive effect on wildlife movement (Lyra-Jorge et al.
2008; Mazzolli 2010). In combination with secondary forest
at intermediate or advanced stages of succession, forestry
usually creates good cover for large felines.
Jaguar prefer landscapes with less human impact (Cul-
len et al. 2005; Cullen et al. 2013) and are more sensitive to
disturbed areas and habitat loss. Therefore, they are more
prone to extinction (Mazzolli and Hammer 2008). In our
study, we observed that critical landscape connections for
jaguar are generally included in the priority connectivity
areas for cougar. In this case, implementing management
strategies that maintain or increase the landscape connec-
tivity for cougar in the study area would also be effective
for jaguar. Critical habitat connections—or pinch points—
between two habitat patches are essential sites for species
 Author's personal copy
Environmental Management
movement (McRae et al. 2008). Accordingly, the loss of
these paths could isolate Strictly Protected Areas and the
Mosaics of Protected Areas that we analyzed (McRae
2012). These pinch points should be a conservation priority
that includes strategies to maintain these areas and recover
riparian forest, which should be immediately implemented.
It is important to note the limitation resulting from using
expert consultation, a subjective measure. Conversely, ex-
pert consultation as a methodology strategy has been suc-
cessfully used in carnivore conservation (Clevenger et al.
2002; Doswald et al. 2007) and more specifically in cougar
(Thatcher et al. 2006, 2009; LaRue and Nielsen 2008, 2011)
and jaguar research (Rabinowitz and Zeller 2010). Addi-
tionally, Beier et al. (2009) analyzing the uncertainty in
designing wildlife corridors found that corridor designs us-
ing a least-cost-corridor approach are not highly sensitive to
uncertainty as long as the resistance values and factor
weights are arrayed in the correct rank order. Following this
logic, we were careful to examine and verify that the class
hierarchy obtained from the expert consultation was sup-
ported by the existing literature. Although most of the
studies did not analyze all classes in the same study, we
could identify the used hierarchy when cross-referencing
works using telemetry (Beier et al. 1995; Dickson and Beier
2002; Dickson et al. 2005; Dickson and Beier 2007; Maz-
zolli 2010; Kertson et al. 2011; Cullen et al. 2013), camera
traps and tracks (Lyra-Jorge et al. 2008; Mazzolli and
Hammer 2008; Mazzolli 2010; Cherem et al. 2011), tracks,
road kill, predation events and interview (Mazzolli 1993;
Zúñiga et al. 2009; De Angelo et al. 2011a, 2011b; Ro-
drı́guez-Soto et al. 2011; Miotto et al. 2012), and expert
consultation (LaRue and Nielsen 2008, 2011; Rabinowitz
and Zeller 2010; Zarco-González et al. 2013). Finally, the
standard deviations calculated for the four land use/land
cover classes that dominated the critical habitat connections
in the landscape were low, indicating a low dispersion from
the average or consistency in the expert opinion on the
importance of these cover land types for both species.
It is important to point out that our study deals with
structural connectivity. Therefore, we recommend the im-
plementation of GPS telemetry studies in jaguar and cougar
research programs in the study area to establish the effec-
tive movement patterns and the existence of functional
connectivity between the protected areas at real remaining
connectivity paths of 1–6 km wide.
Conclusions
The remaining, very narrow priority areas of connectivity
for large felines were identified in this study and are fun-
damental for maintaining their movement throughout the
habitat remnants in the Serra do Mar and Serra da
Mantiqueira mountain ranges as these areas are most likely
the last links among these forest remnants. The areas
identified in this study should be of high priority for legal
protection, which is in urgent need of implementation.
Moreover, as demonstrated in this study, the legal
minimum of riparian forest bands are lacking, and we urge
the immediate implementation of the environmental law
concerning the protection of riparian forests, especially
between the Serra do Mar and Serra da Mantiqueira in the
SE/NW direction.
Acknowledgments The authors acknowledge the support received
from FAPESP (Fundação de Amparo à Pesquisa do Estado de São) in
the form of scholarship and financial support (process 2011/10.791-0)
and to the Secretaria do Meio Ambiente do Estado de São Paulo-SMA
for allowing the use of satellite images. We also thank Agustı́n J.
Paviolo, Ph.D.; Beatriz M. Beisiegel, Ph.D.; Carlos D. De Angelo,
Ph.D.; Peter G. Crawshaw Jr., Ph.D.; Renata A. Miotto, Ph.D.;
Rogério C. de Paula, M.Sc. and Ronaldo G. Morato, Ph.D. for par-
ticipating as consultant specialists.
Ethical Statement Funding: This study was funded by Fundação de
Amparo à Pesquisa do Estado de São Paulo - FAPESP (Grant Number
2011/10.791-0).
Conflict of Interest The authors declare that they have no conflict
of interest.
Studies with Human Participants or Animals This article does
not contain any studies with human participants or animals performed
by any of the authors.
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