Remote Sensing Applications: Society and Environment 34 (2024) 101155

Contents lists available at ScienceDirect

Remote Sensing Applications: Society and

Environment
journal homepage: www.elsevier.com/locate/rsase

Under pressure: suitable areas for neotropical cats within an under

protected biodiversity hotspot
Paula Ribeiro-Souza a, b, c, 1, *, Júlio Haji c, 1, Júlia Oshima c, d, Fernando Lima c,
Barbara Lima-Silva b, e, José Pires b, Milton Ribeiro c, f, Maurício Graipel b
a Programa de Pós Graduação em Ecologia e Recursos Naturais, Centro de Ciências Biológicas e da Saúde, Universidade Federal de São Carlos (UFSCar),
São Carlos, SP, 13565-905, Brazil
b Fauna Floripa Project, Departamento de Ecologia, Centro de Ciências Biológicas, Universidade Federal de Santa Catarina, Florianópolis, SC, 88040-9,

Brazil
c Spatial Ecology and Conservation Laboratory (LEEC), Programa de Pós Graduação em Ecologia, Evolução e Biodiversidade, Departamento de

Biodiversidade, Instituto de Biociências, Universidade Estadual Paulista (UNESP), Rio Claro, SP, 13506-900, Brazil
d Laboratório de Ecologia do Movimento, Instituto de Biociências, Departamento de Ecologia, Universidade de São Paulo (USP), São Paulo, SP, 05508-

090, Brazil
e Laboratório de Ecologia e Conservação (LAEC), Departamento de Biologia, Faculdade de Filosofia, Ciências e Letras de Ribeirão Preto, Universidade de

São Paulo (USP), Ribeirão Preto, SP, Brazil

f Centro de Estudos Ambientais (CEA), Instituto de Biociências, Universidade Estadual Paulista (UNESP), Rio Claro, SP, 13506-900, Brazil

ARTICLE INFO ABSTRACT

Keywords: Understand which factors influence the distribution of feline species can contribute to better
Atlantic forest planning of conservation strategies in a biodiversity hotspot. We modeled the potential distribu-
Biodiversity conservation tion of seven cat species occurring in the Atlantic Forest (AF). Here, we combined climatic and
Habitat loss landscape perspectives to determine the most suitable areas considering the taxonomic richness
Species distribution model
of these cats. We also assessed the ability of fully protected areas (FPAS) to protect these cat
Species richness
species. The results indicated that only 30% of the AF remnants are suitable for all species. Areas
with low species richness were located in Argentina and northeastern Brazil. For taxonomic rich-
ness, the Serra do Mar (45.89%) and the Araucaria (27.90%) sub-regions had the highest suitable
areas, followed by the Interior sub-region (21.89%). The Brejos Nordestinos and Pernambuco
sub-regions had less than 1% suitability. Considering taxonomic richness, only 9% of suitable ar-
eas are covered by FPAs. Leopardus emiliae (1.37%) and Panthera onca (1.97%) had the lowest val-
ues of suitable areas covered by FPAs. The other species of cats are also under low protection (L.
guttulus = 5.38%, L. wiedii = 5.71%, Herpailurus yagouaroundi = 6.70%, L. pardalis = 3.85%,
and Puma concolor = 4.94%). We reveal that a low percentage of suitable areas are currently
fully protected. This study also provides important conservation measures to be implemented in
different AF sub-regions. These findings may help in the planning, maintenance, and implemen-
tation of FPAs through restoration programs and the establishment of ecological corridors.

* Corresponding author. Postgraduate Program in Ecology and Natural Resources, Center of Biological and Health Sciences, Federal University of São Carlos
(UFSCar), São Carlos, SP, 13565-905, Brazil.
E-mail addresses: [email protected] (P. Ribeiro-Souza), [email protected] (J. Haji), [email protected] (J. Oshima), [email protected]
(F. Lima), [email protected] (B. Lima-Silva), [email protected] (J. Pires), [email protected] (M. Ribeiro),
[email protected] (M. Graipel).
1 PRS and JH should be considered joint first author.

https://doi.org/10.1016/j.rsase.2024.101155
Received 14 May 2023; Received in revised form 23 October 2023; Accepted 4 February 2024
Available online 15 February 2024
2352-9385/© 2024 Elsevier B.V. All rights reserved.
P. Ribeiro-Souza et al. Remote Sensing Applications: Society and Environment 34 (2024) 101155

1. Introduction
The Neotropics have a high degree of endemism and harbor the largest remnants of natural areas on the planet (Loyola et al.,
2009), including some global biodiversity hotspots (Oliveira et al., 2016). Among these hotspots, the Atlantic Forest (hereafter AF)
has been the focus of many studies due to its high species richness, endemism, and degree of threat (Mittermeier et al., 2011; Joly et
al., 2014). Species extinction in the AF is directly related to forest loss, urban and agriculture expansion, climate change, and invasive
species (Bellard et al., 2014; Estrada et al., 2017; Bello et al., 2021). Currently, the AF has about 26% of the original forest cover and
highly isolated fragments (Rezende et al., 2018). Nevertheless, the AF harbors about 321 mammal species, including 89 endemic
species (Graipel et al., 2017).
Human-caused environmental disturbances are the primary pressures responsible for the loss of species, ecological functions, and
ecosystem services in the AF (Bogoni et al., 2018, 2022; Cruz et al., 2018; de Lima et al., 2020). These disturbances affect species rich-
ness, distribution and abundance, as well as interspecific interactions (Fahrig, 2003; Gutiérrez et al., 2019; Lewis et al., 2015). More-
over, biodiversity information remains scarce, outdated, or inaccurate for many regions in the AF, including protected areas (Oliveira
et al., 2017). These issues make it difficult to efficiently assess the conservation status of species, particularly carnivorous mammals
(Oliveira et al., 2016). Historically, carnivorous mammals have undergone extinctions or declines in distribution and abundance in
many regions due to habitat loss, forest fragmentation, and illegal hunting (Valenzuela-Galván et al., 2008; Ceballos et al., 2015). Ac-
cordingly, there is great concern about the conservation status of neotropical felines, which are responsible for the maintenance of im-
portant ecological processes and ecosystem functions in tropical forests (Portela and Dirzo, 2020).
Neotropical wild cats (hereafter ‘cats’) are top predators, and by controlling prey populations, they influence vegetation structure,
composition, and abundance, as well as ecological succession (Terborgh et al., 2001; Estes et al., 2011; Morgan et al., 2017). Further-
more, larger body mass felines (e.g., the puma and jaguar) are considered umbrella species in defining priority areas for biodiversity
conservation (Jenkins et al., 2013; Ray et al., 2013). Seven species of Neotropical cats (Felidae) occur in the AF (Nagy-Reis et al.,
2020; IUCN, 2022), all of which are experiencing reductions in range and population size (IUCN, 2022). Most conservation studies
conducted in the AF have focused on the jaguar (Panthera onca) and puma (Puma concolor), which are the largest cats (see Lyra-Jorge
et al., 2010; Morato et al., 2016). Therefore, small, cryptic cats, such as the southern tiger cat (Leopardus guttulus), the margay (Leopar-
dus wiedii), and the jaguarundi (Herpailurus yagouaroundi) remain poorly studied and lack information regarding their biology, ecol-
ogy, and conservation (Macdonald et al., 2010; Nagy-Reis et al., 2017; IUCN, 2022).
According to the “insurance hypothesis”, species richness across their range is an important factor influencing the proper function-
ing of ecosystems (Yachi and Loreau, 1999). Therefore, conservation efforts should consider species richness to establish priority ar-
eas for conservation (Ceballos and Ehrlich, 2006). Most studies on the effects of habitat loss and forest fragmentation on species rich-
ness in the AF have been conducted at small (local or regional) scales. These results have been constantly used to infer large-scale im-
pacts on biodiversity (Sobral-Souza et al., 2021). Recent publications of species occurrence databases make large-scale studies possi-
ble, thus contributing to fill these knowledge gaps. Species Distribution Modeling (SDM) has been widely used to predict the occur-
rence and areas of higher richness (Fuller et al., 2007; Santos et al., 2020; Sobral-Souza et al., 2021) and to identify priority areas for
conservation (Brooks et al., 2006; Oshima et al., 2021; Mittermeier et al., 2005; Ribeiro-Souza et al., 2022). In general, SDM relates
occurrence records with environmental variables to identify suitable areas that allow populations of a given species to persist (Elith
and Leathwick, 2009).
Here, we model the potential distribution of seven cat species in the AF using climatic and landscape variables. We also analyze
the suitable areas most likely to harbor the highest taxonomic richness of these species. Moreover, we quantified the proportion of
these areas covered by fully protected areas (FPAs). Lastly, based on these data, we suggest strategies for the conservation of cats in
the AF areas identified as priorities.

2. Material and methods

2.1. Study area
Different territorial boundaries have been proposed for the AF, but in this work, we adopted the integrative boundary proposed by
Muylaert et al. (2018) (Fig. 1). According to this boundary, the AF covers 90% of its territory in Brazil, extending mainly along the
coast but also reaching the inland areas of the continent and parts of Paraguay and Argentina (Muylaert et al., 2018). The AF exhibits
a broad longitudinal, latitudinal, and altitudinal gradient, ranging from 0 to ∼3000 m, and extensive climatic diversity according to
the Köppen-Geiger classification. It records precipitation exceeding 1000 mm³/year, with higher indices in coastal regions and lower
ones in the inland areas, accompanied by widely variable temperatures (Peel et al., 2007). Additionally, the topography and soil types
of this ecoregion vary considerably (Eisenlohr and Oliveira-Filho, 2015). These variations have favored the formation of diverse
ecosystems characterized by distinct phytophysiognomies (Carnaval et al., 2009; Sobral-Souza et al., 2021; Veloso et al., 1991). It is
known that Protected Areas represent a cornerstone in regional biodiversity conservation strategies (Lewis et al., 2019). However, the
AF has a significant deficit of Protected Areas, which is insufficient for the conservation of many species (Lemes et al., 2014;
(Lourenço-de-Moraes et al., 2019; Ribeiro-Souza et al., 2022; Soares-Filho et al., 2014)).

2.2. Selection of occurrence locations

We compiled 20,000 feline occurrence records for the entire Neotropical region. For this, we used the open databases Atlantic
Mammals (Souza et al., 2019), Atlantic Camtraps (Lima et al., 2017) and Neotropical Carnivores (Nagy-Reis et al., 2020); data avail-
able in the literature (see Sartor et al., 2021; Nascimento and Feijó, 2017); and the virtual databases GBIF (www.gbif.org), SiBBr

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P. Ribeiro-Souza et al. Remote Sensing Applications: Society and Environment 34 (2024) 101155

Fig. 1. Atlantic Forest domain (Muylaert et al., 2018) and its biogeographic sub-regions (adapted from Silva and Casteleti, 2003).

(www.sibbr.gov.br), PortalBio (portaldabiodiversidade.icmbio.gov.br) and speciesLink (specieslink.net). We only considered records

obtained through camera traps, roadkill, radiotelemetry, active search, transects, and museum records. We only considered data col-
lected from the year 2000 onwards and with precise geographic coordinates (better than 1 km precision). Next, we removed duplicate
or overlapping occurrence records for each species. To minimize spatial autocorrelation, which could inflate model predictions
(Araújo and Guisan, 2006; Veloz, 2009), we selected only one occurrence record within each 1 km2 pixel for the small cats (H.
yagouaroundi, Leopardus emiliae, L. guttulus, L. pardalis, and L. wiedii) and 5 km2 pixels for the P. onca and P. concolor. The occurrence
records that did not meet our criteria were disregarded from this study. This selection was necessary because large felid species such
as P. concolor and P. onca have larger home ranges than small felids (Crawshaw et al., 2004; Lindzey et al., 1987) and can have a scale
of effect (Alegre et al., 2023; Zeller et al., 2014) that is greater than other small felids when we consider resource selection analysis.
Therefore, large felids can cover long distances per day, and the same individual may be recorded in different camera traps. At the end
of the data selection, we obtained 5284 cat occurrence records for the Neotropical region (see https://github.com/LEEClab/ms_
neotropical_cats_suitability). Within these occurrence records for the Neotropical region, the following were used in the climatic mod-
els: 23 for L. emiliae, 381 for L. guttulus, 693 for L. wiedii, 870 for H. yagouaroundi, 1,821 for L. pardalis, 1,779 for P. concolor, and 677
for P. onca. Within the expanded limit of the AF (Muylaert et al., 2018), we obtained 2263 records (Figure appendix B.1). To generate
landscape models within the AF, we used 19 occurrence data for L. emiliae, 381 for L. guttulus, 244 for L. wiedii, 422 for H.
yagouaroundi, 597 for L. pardalis, 464 for P. concolor, and 73 for P. onca. (Figure appendix B.1).

2.3. Climate, environmental and landscapes predictors

We obtained 19 bioclimatic variables from WorldClim v2.1 (www.worldclim.org). These variables were generated from interpo-
lated data recorded between 1950 and 2000 (Hijmans et al., 2005). In addition to climatic variables, we also used topographic, land-
scape, and human influence variables.
The topographic variables were the terrain ruggedness index (median and standard deviation of the terrain ruggedness index,
Amatulli et al., 2018) and terrain slope (median and standard deviation of the slope, Amatulli et al., 2018). The landscape variables
included the percentage of tree canopy cover (Hansen et al., 2013), habitat heterogeneity (coefficient of variation of Enhanced Vege-
tation Index data and Shannon index of Enhanced Vegetation Index data, Tuanmu and Jetz, 2015), and water frequency (Feng et al.,
2016). The human influence variable was human density (population density per 1 km2; https://sedac.ciesin.columbia.edu/data/
collection/gpw-v4/documentation) and human footprint (Venter et al., 2016). We used the topographic, landscape, and human influ-
ence variables to compose our landscape model and the bioclimatic variables to compose the climatic model. All layers were at 30 arc
sec resolution (∼1 km at the equator). To reduce the degree of uncertainty associated with the model results (Dormann et al., 2008),
we tested all variables for multicollinearity using a variance inflation factor (VIF <2.0; Dormann et al., 2013) (Table appendix A.1).
Five variables were selected for the landscape models: terrain ruggedness, percentage of tree canopy cover, habitat heterogeneity, wa-
ter frequency and human density. Four variables were selected for the climatic models: mean temperature of the wettest quarter
(BIO8), mean temperature of the driest quarter (BIO9), precipitation of the wettest month (BIO13) and precipitation of the coldest
quarter (BIO19) (Table 1).

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Table 1
Species studied, climatic and landscape variables names and description.

Variable Abbreviation

Coefficient of Variation of EVI data - heterogeneity Habitat heterogeneity*

Water Frequency - Global Inland Water Water Frequency*
2
Population density per 1 km Human density*
Standard deviation terrain ruggedness index Terrain ruggedness*
Percent of treecover from 2000 Tree cover*
Mean Temperature of Driest Quarter BIO09*
Precipitation of Wettest Month BIO13*
Precipitation of Coldest Quarter BIO19*
Mean Diurnal Range BIO02
Mean Temperature of Wettest Quarter BIO08*
Isothermality BIO03

3. Modeling procedures
We separately developed landscape and climate-based SDMs for each species. To include the full climatic niche of the species, the
climatic models were generated for the entire Neotropical region following the delimitation proposed by Morrone (2014). After-
wards, they were clipped to fit the expanded limit of the AF in Brazil, Argentina, and Paraguay, as well as the landscape models
(Muylaert et al., 2018). We used the ensemble forecasting approach to test different methodologies on a set of predictions (Araújo and
New, 2007). To accomplish this, we employed three algorithms utilizing varying techniques: (i) machine learning method that esti-
mates species distributions by finding the maximum entropy distribution, MaxEnt (Phillips et al., 2006); (ii) random forest, an artifi-
cial intelligence method that combines predictive regression tree predictors (Breiman, 2001); and (iii) multiple discriminant analysis
(MDA) method, a classification based on multiple models (Hastie et al., 1994). We selected 1000 background points, 70% of occur-
rence data for training the algorithm and 30% for external testing, randomized 10 times to generate less biased models (k-folding
technique, k = 2). In the end, we obtained 30 climatic and 30 landscape models for each species (10 × 3 algorithms). The efficiency
of the models was evaluated using the True Skill Statistic (TSS) test (Allouche et al., 2006). The TSS test generates values between −1
and 1, with values closer to 1 indicating perfect agreement, and values of zero or less indicating a performance no better than random
(Allouche et al., 2006). The area under the ROC curve (AUC; Hanley and McNeil, 1982) values were also presented as an additional
assessment (Table 0.2). Models with AUC values > 0.90 are excellent; 0.80–0.90 are good; 0.70–0.80 are reasonable; 0.60–0.70 are
poor, and 0.50–0.60 are failing (Araujo et al., 2005). All models were run using the ‘sdm’ package (Naimi and Araújo, 2016) in the R
environment. From the generated models, we estimated the suitable areas for each species from a maximum decision threshold that
maximizes the sum of sensitivity and specificity (Max SSS) (Manel et al., 2001). This approach is one of the most recommended for
measuring the performance of SDMs (Liu et al., 2005, 2013; Shabani et al., 2018). Following the maximization of the sum of sensitiv-
ity-specificity, the higher the sensitivity, the lower the false absence rate, and the higher the specificity, the lower the false presence
rate. Thus, we selected areas with higher climate and landscape suitability as priorities for conservation. We then constructed binary
maps, in which 0 = areas with inadequate climate and landscape and 1 = areas with adequate climate and landscape.

3.1. EcoLand and cat species richness in the AF

To measure the area with climatic and landscape suitability for each species, we ran an EcoLand analysis (Sobral-Souza et al.,
2021), in which we constructed a composite map and a comparative map of the results of the two models generated separately. For
this, we created maps with four different habitat suitability classes per species using the following criteria: (i) high climatic and land-
scape suitability — areas (pixel) with values > 0.5 for both models, (ii) high climatic and low landscape suitability — areas with val-
ues > 0.5 for the climatic model and ≤0.5 for the landscape model, (iii) high landscape and low climatic suitability — areas with val-
ues > 0.5 for the landscape model and ≤0.5 for the climate model, and (iv) low climatic and landscape suitability — areas with val-
ues ≤ 0.5 for both models.
To elaborate the taxonomic richness maps along the AF, we summed the binary maps of the seven species separately for each suit-
ability type (climate and landscape). Next, we summed and reclassified the two resulting maps (one of richness based on climate and
another based on landscape), generating a map with the estimated number of species (richness) per pixel (Figure appendix B.2). In the
richness scatterplot, the X-axis represents the richness predicted by the climate, and the Y-axis the richness predicted by the land-
scape. In the EcoLand scatterplot, we created four categories based on climate and landscape (Table 2).

Table 2
Species richness criteria for the EcoLand scatterplot categories.

Category Climate richness Landscape richness

Low climate and low landscape <4 species <4 species

Low climate and high landscape <4 species ≥4 species
High climate and low landscape ≥4 species <4 species
High climate and high landscape ≥4 species ≥4 species

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We calculated the areas predicted by the final taxonomic richness maps in km2, selecting only suitable areas for more than four
species. We also calculated the areas using the final EcoLand maps, considering only areas with high climatic and landscape suitabil-
ity for each species using QGIS. The calculations were then tabulated considering the total suitable area for taxonomic richness, val-
ues by AF sub-regions (Silva and Casteleti, 2003, Fig. 1), and FPAs. The following categories were included: Ia (Strict Nature Reserve),
Ib (Wilderness Area), II (National Park), and III (Monument or natural feature) (IUCN, 2001). Data on protected areas were obtained
from protectedPlanet (Juffe-Bignoli et al., 2014; https://www.protectedplanet.net/en).

4. Results
The potential species distribution models demonstrated performance with TSS ranging from 0.36 to 0.73 (climate) and 0.34 to
0.92 (landscape). Regarding our AUC results, we obtained models with reasonable performance for climate (AUC = 0.73) and land-
scape (AUC = 0.71) and good performance for climate (AUC = 0.82) and landscape (AUC = 0.89) (Table appendix A.2).

4.1. Species richness and protected areas

In the climatic models, areas with the highest richness indicated suitability for seven species, which were concentrated mainly in
small patches within the Interior sub-region of southern Brazil. Areas with the second highest richness (six species) were observed in
the Araucaria, Serra do Mar, and Interior sub-regions. The results predicted by the landscape also estimated suitable areas for up to
seven species, especially in the Misiones region in Argentina and the Serra do Mar and Araucaria sub-regions in Brazil (Fig. 2).
The results considering species richness, classified by the EcoLand method, indicated that only 30% of the AF remnants are suit-
able for all analyzed species. For seven cat species, suitable areas were concentrated mainly in remnants within the Serra do Mar sub-
region (from southeastern to southern Brazil) and in some patches on the border of Brazil and Argentina. Areas with low species rich-
ness (1–3 species) were located in Argentina and southern and northeastern Brazil (Fig. 3). For the areas with higher taxonomic rich-
ness (>4 species), the Serra do Mar (45.89%) and Araucaria (27.90%) sub-regions had the highest suitable areas. The Brejos Nordes-
tinos and Pernambuco sub-regions had less than 1% suitability. Considering this higher taxonomic richness, only ∼9% of suitable ar-
eas are covered by FPAs (Table 3). L. emiliae (1.37%) and P. onca (1.97%) had the lowest values of suitable areas covered by FPAs. The
other species are also under low protection (L. guttulus = 5.38%, L. wiedii = 5.71%, H. yagouaroundi = 6.70%, L. pardalis = 3.85%,
and P. concolor = 4.94%; Table appendix A.3).

5. Discussion
In this study, we present SDMs of suitable areas predicted by climate and landscape for seven cat species in the AF. Our results
show that only ∼30% of the AF remnants are suitable for cat species. Furthermore, over 90% of suitable areas are located outside
FPAs and in small, isolated forest patches. These findings reinforce the impacts of habitat loss and fragmentation in the AF and high-

Fig. 2. Distribution of Neotropical cat richness in the Atlantic Forest according to climatic (left) and landscape (right) models. Observe a gradient from 0 to 7 represent-
ing the number of species.

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Fig. 3. Distribution of Neotropical cat richness in the Atlantic Forest according to the EcoLand model.

Table 3
Suitable areas and percentage predicted by the climate and landscape (EcoLand) for higher taxonomic richness (≥4 species) areas in the Atlantic Forest (AF). We
present the total suitable areas for each AF sub-region and the total suitable areas covered by fully protected areas (FPAs).

Atlantic Forest sub-region Total FPAs

2 2
km % km %

Climate 415,219 100.0 16,330 3.93

Landscape 351,159 100.0 11,383 3.24
EcoLand
Araucaria 83,849 27.90 1395 1.66
Bahia 10,647 3.54 204 1.92
Brejos Nordestinos 972 0.32 20 2.06
Serra do Mar 137,917 45.90 19,774 14.34
Pernambuco 1547 0.51 37 2.39
Interior 65,632 21.80 6819 10.39
Total 300,564 100.0 28,249 9.40

light their negative effects on the conservation of wild cats in this ecoregion. Based on our findings, we classify regions with different
suitability levels and suggest punctual and essential measures. Our results address several goals of the National Action Plan for the
Conservation of Small Cats (NAP Small Cats) and the National Action Plan for the Conservation of Big Cats (NAP Big Cats). Above all,
our results contribute to filling existing knowledge gaps in the AF by identifying potential habitats for forest-dependent cats based on
data compiled at the biome level and related to climatic and landscape variables. Thus, we provide new information that is essential
for decision-making regarding cat conservation.
Although higher resolution landscape data could provide a more detailed perspective of cat habitat suitability in our study, which
can be considered a limitation of it, the ocurrence data currently available for modeling the distribution of cats in the AF is not yet sys-
tematized along the entire biome. Therefore, higher resolution occurrence data and frequent monitoring will also be necessary for
more detailed predictions. To our knowledge, our predictions provide the best current overview of the status of suitable areas for
neotropical cats in the AF considering the datasets based on presence-only data.

5.1. Climate and landscape based models

Camera traps are potentially more effective at night (Srbek-Araujo and Chiarello, 2005), and most felids are predominantly noc-
turnal (Graipel et al., 2019); however, the species H. yagouaroundi, is predominantly diurnal (Giordano 2016). Our study includes 870

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records of this species. Previous studies indicate that H. yagouaroundi is a habitat generalist and is associated with open areas (Caso et
al., 2015; Regolin et al., 2017), which may explain the lower performance of our landscape models. There are few records of the oc-
currence of H. yagouaroundi obtained using camera traps in open areas, especially due to the risk of loss of such equipment (Magioli et
al., 2022).
The wide geographic distribution may have influenced the lower TSS values for the climatic models of H. yagouaroundi, L. pardalis,
P. onca, and P. concolor. Since climatic variables influence the distribution of these species more strongly at first-order selection
(Johnson, 1980), landscape variables are essential to predict suitable remnant areas at finer, second-order levels. Our results uncover
climatically suitable areas for L. emiliae in the northern AF and L. guttulus in the southern AF (Figure appendix B.17 and Figure B.18,
respectively). These findings further support the definition of the two valid species, L. guttulus and L. tigrinus (Nascimento, 2010; Trigo
et al., 2013). The existence of large suitable fragments (in terms of climate and landscape) makes Serra do Mar a priority area for the
protection of Brazilian endangered species, such as L. guttulus and H. yagouaroundi.
For L. wiedii, we detected (as predicted by the landscape models) large continuous fragments in the Serra do Mar potentially suit-
able for the species (Figure appendix B.19). In contrast, these same fragments have low climatic suitability for the species, making it
necessary to monitor these areas under future climate change scenarios (Figure appendix B.5). For L. guttulus, our models suggest suit-
able areas more concentrated in the Serra do Mar and Araucaria (Figure appendix B.18). However, this species has already been
recorded in altered areas, as long as there was adjacent native vegetation that allowed its movement across the landscape (Oliveira-
Santos et al., 2012). In this case, an important conservation action could be to focus on forest restoration to increase the functional
connectivity between the forest fragments where the species still occur.
Although P. concolor occurs throughout most of South America and shows more generalist habitat selection, our models indicate
that suitable areas are limited to the large continuous fragments in the Serra do Mar and small portions in the Araucaria and Interior
sub-regions. The greater use of landscapes with forest cover was shown in another study carried out in the AF (Regolin et al., 2017). In
the Interior sub-region in southeastern Brazil, there is high climatic suitability for the species, but fragments with high landscape and
climatic suitability are smaller and far apart. In northeastern Brazil, the models for many regions indicate low climatic suitability, and
the remaining fragments (Bahia, Brejos Nordestinos, and Pernambuco sub-regions) with landscape suitability occurring close to the
coast (Figure appendix B.22).
Our model exhibits few suitable areas for L. emiliae predicted by climate and landscape (Figure appendix B.17), thus requiring
greater attention for its conservation status. Leopardus pardalis is widely distributed throughout South America (except Chile), but our
models indicate a considerable reduction of areas that are originally suitable. The areas with high climatic and landscape suitability
for this species are located in the Serra do Mar, Araucaria, Bahia, and Interior sub-regions (Figure appendix B.21). Despite the reduc-
tion of originally suitable areas, our results show that L. pardalis is still the Neotropical cat with the largest suitable area (high climatic
and landscape suitability; area = 305,000 km2) in the AF (Table appendix A.3), which reinforces its important role as a mesopreda-
tor in this ecoregion (Moreno et al., 2006; Silva-Pereira et al., 2011).
Finally, our models for P. onca indicate a suitable area (high climatic and landscape suitability) of only 6546 km2 (Figure appendix
B.23). According to Paviolo et al. (2016), P. onca has lost about 85% of its habitat and has practically disappeared from northeastern,
southeastern, and southern Brazil (Sanderson et al., 2002; Tôrres et al., 2008). In addition to the great loss of suitable forest habitats,
the remaining areas are highly fragmented, reaching 93% loss of its connectivity, mainly in Brazil and Paraguay (Pardo et al., 2023).

5.2. Species richness and protected areas

Areas with high richness (>4 species) and high climatic and landscape suitability are located in the AF's most continuous or most
connected remnants. This is the case of the Serra do Mar, which is considered one of the most preserved sub-regions in the AF with a
high fauna species richness (Carnaval et al., 2009; Silva and Casteleti, 2003) and a center of endemism for species of marsupials, pri-
mates, and rodents (Costa et al., 2000). Our findings reinforce the importance of the Serra do Mar sub-region, especially considering
the ecological importance of cats and their degree of threat, mainly due to habitat loss and hunting (Bogoni et al., 2022; Valenzuela-
Galván et al., 2008). Although this sub-region is the most covered by FPAs in the AF, the current protected area (∼14% of the total
area) may be insufficient to conserve its biodiversity. The expansion of FPAs in the Serra do Mar could enable the maintenance of suit-
able areas, thus achieving one of the goals for the conservation of P. onca and P. concolor (Brasil, 2018).
The suitable areas in the Araucaria, the second sub-region with the highest potential richness of cats, are very fragmented and
poorly protected (∼2% of FPAs). Some of the FPAs are located on the border of Brazil and Argentina, which may make the conserva-
tion of cat species beyond borders unfeasible. Therefore, both countries must cooperate to conserve this important region. The main-
tenance of P. onca populations in the southern Brazilian AF is probably accomplished by migrating individuals from the Misiones re-
gion, where a larger amount of suitable habitat is still available. Associated with measures to mitigate poaching and conflicts arising
from the predation of domestic livestock (Mazzolli et al., 2002), the areas bordering Argentina can be recolonized by young dispers-
ing individuals. In fact, this scenario has been identified in the Iguaçu National Park in recent decades. In the Bahia sub-region, we ob-
served a continuous fragment with climatic and landscape suitability. This area is more isolated from the other suitable areas in the
southern AF and is not covered by FPAs. Therefore, the persistence and dispersal of cats among these regions may be impaired, espe-
cially for small-sized forest species such as L. guttulus and L. wiedii (Cruz et al., 2019). The Interior sub-region, with only 7% of the
original forest cover (Ribeiro et al., 2009), is dominated by areas of low potential climatic and landscape richness. The exceptions are
areas near the Serra do Mar, where there is high climatic suitability between the states of São Paulo and Minas Gerais and some
patches of remnant fragments near Diamantina and Bahia.
The Diamantina, Pernambuco, and São Francisco sub-regions have small, fragmented areas with high landscape suitability, low
climatic suitability, and few FPAs. Moreover, these sub-regions are transition zones between biomes such as Caatinga and Cerrado,

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P. Ribeiro-Souza et al. Remote Sensing Applications: Society and Environment 34 (2024) 101155

which also harbor Neotropical cat populations (Oliveira et al., 2016). The Brejos Nordestinos region has a relatively small area com-
pared to other sub-regions and a naturally dispersed distribution (Ribeiro et al., 2009), presenting mostly areas of low climatic and
landscape suitability, small remaining forest fragments, and few FPAs.

5.3. Implications for conservation and protected areas

Increased human pressure on biodiversity conservation has caused impacts such as changes in the activity pattern of mammals
(Gaynor et al., 2018), roadkill (Abra et al., 2021), poisoning and slaughter (Lima et al., 2017), and poaching (Peres et al., 2016). Con-
sequently, biodiversity loss has increased in areas where species still occur (Galetti et al., 2017; Peres et al., 2016). Thus, strategies
must be created to halt the loss of biodiversity by planning priority areas for conservation and creating new FPAs. Recent studies have
addressed the prioritization of large-scale conservation areas, greatly contributing to the goals of biodiversity conservation and the
understanding of their limitations. These studies suggest that, by expanding conservation efforts, part of the AF will hold great poten-
tial to increase biodiversity protection (Pouzols et al., 2014; Luby et al., 2022). In view of our results, we suggest some specific mea-
sures for the conservation of cats in the AF.
Areas with high climatic and low landscape suitability (Araucaria, Serra do Mar, and Interior) need restoration of natural forest
cover to prevent loss of biodiversity in the AF. Moreover, creating functionally connected ecological corridors is necessary to increase
the connectivity and species dispersal between fragments, especially in the Bahia and Interior sub-regions, thus increasing the persis-
tence of forest species populations (Frankham, 1996). In the Araucaria sub-region, we observed large areas with climatic suitability
for cat richness among fragments suitable for landscape. The restoration of the landscape and the creation of ecological corridors will
enable the connection between patches in the Araucaria, according to priority conservation actions previously suggested (Brasil,
2018). Thus, there will be a large extension suitable for the climate and landscape near the Iguaçu National Park and the Araucárias
National Park. The same is true for the Interior sub-region, especially in southeastern Brazil, where there is a large area (orange area,
Fig. 3) near FPAs such as Campos do Jordão State Park, Itatiaia National Park, and Cantareira State Park. A recent study has shown
the importance of small habitat patches for biodiversity (Riva and Fahrig, 2023). We consider that expanding the connection between
small fragmented areas and protected areas is essential to enable the rescue effect and the recomposition of communities through the
top-down effect promoted by cats in the AF (Elton, 1927; Gonzalez et al., 1998).
Many species may migrate in search of climatically suitable habitats under climate change scenarios (Singh, 2020), which will re-
quire the existence of other suitable landscapes to enable dispersals. Areas with low climatic and high landscape suitability, such as
Misiones (Argentina), Araucaria, Bahia, and part of the Serra do Mar (Brazil), may become climatically suitable in the future. There-
fore, additional studies on the effects of climate change on cat species in the AF are needed. Butti et al. (2022) showed that carnivores
are undergoing the greatest habitat loss compared to other mammalian orders. Areas with low climatic and high landscape suitability
(green areas - Fig. 3) are located mainly in southern Brazil, close to FPAs. These areas are extremely important because they provide
forest cover, primarily for widely distributed Neotropical cats, covering a wide climatic range (Macdonald et al., 2010).
Human-induced landscape changes compromise ecosystem stability and resilience due to reduced species richness and functional
groups (Tscharntke et al., 2012). To avoid such problems and enable the persistence of wild cats, preserving areas that have both cli-
mate and landscape suitability in the AF is necessary. We consider the regions with high habitat suitability for more species as priori-
ties for conservation. These conservation measures can inhibit the loss of suitable habitat, the main factor influencing the local extinc-
tion of about 75% of mammal species (Bogoni et al., 2022). New FPAs should be strategically created, considering the expansion of
biodiversity protection and connectivity between existing FPAs.
Our results endorse the classification of the Serra do Mar sub-region as an extremely high-priority area for conservation (Brasil,
2018). Suitable habitats that are connected and protected by FPAs will reduce hunting pressure, one of the main drivers of the local
extinction of cat species (Bogoni et al., 2022; Gallego-Zamorano et al., 2020), and will enable increased gene flow and effective popu-
lation sizes (Fahrig, 2003). We also draw attention to the priority areas for conservation on the border between Brazil and Argentina,
which have high suitability predicted by climate and landscape and harbor prey populations and potential areas for their distribution
(Paviolo et al., 2018). The protection of habitats of threatened species (e.g., Ribeiro-Souza et al., 2022) contributes to the conserva-
tion of other species dependent on the same habitat, such as cats, which are a key group in an ecosystem. Studies conducted in the
Americas have shown the importance of maintaining the connectivity among protected areas beyond national borders (Dudley et al.,
2014; Thornton et al., 2020; Ribeiro-Souza et al., 2022). The Iguaçu National Park (between Brazil and Argentina), the Mesoamerican
Biological Corridor (Hilty et al., 2012), and the transboundary conservation measures in of the Gran Chaco (https://
www.wwf.org.py/que_hacemos/proyectos/pacha/) are prime examples of this type of conservation.

6. Conclusion
We show for the first time how integrated landscape and climatic models can be used to estimate habitat suitability and important
conservation areas for Neotropical cats in the AF. We reveal that only a low percentage of suitable areas are currently fully protected.
This study also provides important conservation guidelines based on suitability models for specific sub-regions in the AF that could be
used by stakeholders. These findings may help in planning the maintenance and implementation of FPAs through restoration pro-
grams and the establishment of ecological corridors in this decade. We also suggest that future conservation efforts could be focused
not only on the conservation of cat richness suitable areas but also on new modeling approaches considering the presence and poten-
tial distribution of prey, improving functional connectivity, and considering possible future climate and landscape change scenarios
to strengthen the knowledge about suitable areas in the AF.

8
P. Ribeiro-Souza et al. Remote Sensing Applications: Society and Environment 34 (2024) 101155

Funding
This work was supported by Fundação Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - CAPES (process
#88887.519051/2020-00), Conselho Nacional de Desenvolvimento Científico e Tecnológico - CNPq (process #130769/2021-5;
#442147/2020-1; #440145/2022-8; #402765/2021-4; #313016/2021-6 and #440145/2022-8; 420094/2023-7) and Fundação de
Amparo à Pesquisa do Estado de São Paulo - FAPESP (processes #2013/50421-2; #2020/01779-5; #2021/06668-0; #2021/08322-3;
#2021/08534-0; #2021/10195-0; #2021/10639-5; #2022/10760-1; #2014/23132-2 and #2021/02132-8). This study is also part
of the Centro de Pesquisa em Dinâmica da Biodiversidade e Mudanças Climáticas, which is financed by the FAPESP.

Ethical statement
All authors declare that all ethical practices have been followed in relation to the development, writing, and publication of the ar-
ticle.

Author contribution
Conceptualization: Paula Ribeiro-Souza, Júlio Haji, Julia Oshima, Fernando Lima, Milton Ribeiro, Maurício Graipel, Barbara
Lima-Silva and José Pires. Data curation: Paula Ribeiro-Souza and Júlio Haji. Formal analysis: Paula Ribeiro-Souza and Júlio Haji.
Funding acquisition: Paula Ribeiro-Souza, Júlio Haji and Milton Ribeiro. Investigation: Paula Ribeiro-Souza. Methodology: Paula
Ribeiro-Souza, Júlio Haji, Julia Oshima, Fernando Lima, Milton Ribeiro, Maurício Graipel, Barbara Lima-Silva and José Pires. Pro-
ject administration: Paula Ribeiro-Souza and Júlio Haji. Resources: Paula Ribeiro-Souza, Júlio Haji, Maurício Graipel and Milton
Ribeiro. Software: Paula Ribeiro-Souza. Supervision: Paula Ribeiro-Souza and Júlio Haji. Validation: Paula Ribeiro-Souza, Júlio
Haji, Julia Oshima, Fernando Lima, Milton Ribeiro, Maurício Graipel, Barbara Lima-Silva and José Pires. Visualization: Paula
Ribeiro-Souza, Júlio Haji, Julia Oshima, Fernando Lima, Milton Ribeiro, Maurício Graipel, Barbara Lima-Silva and José Pires. Writ-
ing – original draft: Paula Ribeiro-Souza and Júlio Haji. Writing – review & editing: Paula Ribeiro-Souza, Júlio Haji, Julia Os-
hima, Fernando Lima, Milton Ribeiro, Maurício Graipel, Barbara Lima-Silva and José Pires.

Data accessibility statement

The feline occurrence data used in the analyses and the richness and ecoland maps are available at the link: https://
datadryad.org/stash/share/7MpwfQWHDmImnFBhV_Czejjq-_QW4_5CRWv-fu51f6g; All variables used are third party data freely
available online. Climate data are available on the Worldclim website (https://www.worldclim.org/data/worldclim21.html). The
landscape variables are available on the following sites: Habitat heterogeneity (http://www.earthenv.org/texture), Water frequency
(DOI: 10.1080/17538947.2015.1026420), Human density (http://sedac.ciesin.columbia.edu/data/set/gpw-v4-population-density-
rev10), Terrain ruggedness (http://www.earthenv.org/topography) and Tree cover (https://doi.org/10.1126/science.1244693).

Declaration of competing interest

The authors declare that they have no known competing financial interests or personal relationships that could have appeared to
influence the work reported in this paper.

Data availability
We shared the link with the data in the revised version of the manuscript.

Acknowledgements
We thank all the researchers who contributed to the cat database we used for the analyses.

Appendix A. Supplementary data

Supplementary data to this article can be found online at https://doi.org/10.1016/j.rsase.2024.101155.

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