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Relative rarity of small wild cats in the Brazilian Pantanal

  • Institute for the Conservation of Neotropical Carnivores

Abstract and Figures

Small wild cats (SWC) are naturally cryptic species. The current study presents occurrence information and capture-rates on four SWC species found in the Brazilian Pantanal. The most commonly recorded SWC by camera trap was Leopardus pardalis . Leopardus colocolo and Puma yagouaroundi were relatively rare, while Leopardus guttulus was not recorded in any survey year. We interpret our findings based on the potential competitive influences of L. pardalis and practical implications of camera trap survey design. We recommend that future studies should design camera trap grids with spacing appropriate for SWC to more directly address questions on local population status and interspecific interactions.
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Short note
Fernando R. Tortato*, Allison L. Devlin, Ricardo L. P. Boulhosa and Rafael Hoogesteijn
Relative rarity of small wild cats in the Brazilian
Received December 13, 2019; accepted June 2, 2020; published online
July 29, 2020
Abstract: Small wild cats (SWC) are naturally cryptic
species. The current study presents occurrence information
and capture-rates on four SWC species found in the Bra-
zilian Pantanal. The most commonly recorded SWC by
camera trap was Leopardus pardalis.Leopardus colocolo
and Puma yagouaroundi were relatively rare, while Leop-
ardus guttulus was not recorded in any survey year. We
interpret our ndings based on the potential competitive
inuences of L. pardalis and practical implications of
camera trap survey design. We recommend that future
studies should design camera trap grids with spacing
appropriate for SWC to more directly address questions on
local population status and interspecic interactions.
Keywords: camera trap; Leopardus sp.; Ocelot effect;
Pantanal; Puma yagouaroundi; small wild cats (SWC).
Small wild cats (SWC) are cryptic and primarily nocturnal
species that naturally occur in low densities such that field
observations are rare (Hunter 2011; Sunquist and Sunquist
2002). Located in the center of South America, the Pantanal
oodplain is one of the largest inland wetlands in the world
(Harris et al. 2005; Junk et al. 2006) and is included in the
range of four SWC species: pampas cat Leopardus colocolo;
oncilla Leopardus guttulus; ocelot Leopardus pardalis; and
jaguarundi Puma yagouaroundi (Nascimento and Feijó
2017; Rodrigues et al. 2002). Although Geoffroys cat
Leopardus geoffroyi and margay Leopardus wiedii have
been recorded in this region, these species were not
included in the current study as they only occur along the
periphery of the Pantanal (Rodrigues et al. 2002; Tomás
et al. 2010).
One of the most effective methods to detect natu-
rally rare and elusive species includes the use of non-
invasive technology such as remotely triggered camera
traps (Karanth and Nichols 1998; Porfírio et al. 2018;
Srbek-Araujo and Chiarello 2005; Tobler et al. 2008;
Tomás et al. 2003). Over the past 16 years, camera trap
surveys conducted in the Pantanal have provided in-
sights into the distribution of L. colocolo (Godoi et al.
2010), temporal niche partitioning between L. pardalis
and P. yagouaroundi (Bianchi et al. 2016; Porfírio et al.
2018), and density estimates of L. pardalis (Trolle and
Kéry 2003; Trolle and Kéry 2005). Among these species,
L. guttulus is the most data decient (Tomás et al. 2010).
ThepresenceofL. guttulus in the Pantanal was rst
veried in 1988 with a skin (Figure 1) collected in the
Poconé region (Mato Grosso, Brazil) and currently
housed in the scientic collection of the Emili Goeldi
Museum (catalogue number MPEG22193; Belém, Brazil;
Nascimento and Feijó 2017). There has since been only
one veried photographic record of L. guttulus obtained
by camera trap (Trolle 2003).
Herein, we analyzed camera trap data from long-
term annual surveys to estimate the capture rate
(([photographic records/survey effort] ×100); Carbone
et al. 2001) of SWC at two cattle ranches (Fazenda São
Bento, Mato Grosso do Sul, Brazil; Fazenda Jofre Velho,
Mato Grosso, Brazil) in the Porto Jofre region of the
northern Brazilian Pantanal. Camera trap surveys were
designed for the study of jaguars Panthera onca;
(i.e., minimum distance of 2,500 m between each sta-
tion or 23 cameras per female jaguar home range;
camera trap height 40 cm above ground level; Soisalo
and Cavalcanti 2006) and covered a total area of up to
300 km2. We only counted independent records of SWC
(i.e., photographs recorded at 1 h intervals) and
compared our records with those from other studies
conducted in the Pantanal (Table 1). As it is considered
*Corresponding author: Fernando R. Tortato, Panthera, 8 West 40th
St. 18th Floor, New York, NY 10018, USA,
Allison L. Devlin: Panthera, 8 West 40th St. 18th Floor, New York, NY
10018, USA; Wildlife Biology Program, W.A. Franke College of Forestry
and Conservation, University of Montana, 32 Campus Drive, Missoula,
MT 59812, USA
Ricardo L. P. Boulhosa: Instituto Pró-Carnívoros, Av. Horácio Neto,
1030, Atibaia, SP, 12945-010, Brazil
Rafael Hoogesteijn: Panthera, 8 West 40th St. 18th Floor, New York,
NY 10018, USA
Mammalia 2020; aop
the rarest SWC species in the Pantanal, we also
included visual observations of L. guttulus made by felid
researchers between 1994 and 2018 in the same region
sampled by camera trap surveys.
We conducted 17 independent surveys from 2011
through 2018 with a total effort of 7559 trap nights. We
recorded 1236 independent photos of L. pardalis,20of
P. yagouaroundi, and one of L. colocolo. Between 1994 and
2018, 13 visual records of L. guttulus occurred in the same
region sampled by camera traps (Table 2). All observations
were less than 60 km from the site where the individual
L. guttulus was rst collected in 1988 (Figure 1; Nascimento
and Feijó 2017); all subsequent visual records of L. guttulus
occurred during eld surveys (i.e., transects; night
spotlighting) along the edge of forested areas. Ecotourism
guides in the region (Tortato and Izzo 2017) reported ob-
servations of L. guttulus in lodges near the study area.
However, the authors did not receive photographic
conrmation of the sightings and thus did not include the
observations in the current study. Due to the phenotypic
similarity between spotted SWC species L. guttulus and
juvenile L. pardalis, our 13 direct visual records of L. gut-
tulus are considered likely but not conrmed.
Capture rates for L. colocolo,L. guttulus, and
P. yagouaroundi estimated in this study were similar to
those from other areas of the Pantanal (Table 1). Among all
studies, L. pardalis was one of the most frequently recorded
carnivore species (Porfírio et al. 2018; Trolle, 2003). The
capture rate of L. pardalis in this study, however, was 2.45
times higher than reported in the literature (Table 1).
According to Arita et al. (1990), the rarity of Neotrop-
ical forest mammals can be divided into four categories:
restricted distribution and high density; wide distribution
and high density; restricted distribution and low density;
and wide distribution and low density. The majority of SWC
species in the Pantanal most closely align with the category
of wide distribution and low density (Arita et al. 1990). The
rarity of L. guttulus and other SWC observed in the Pantanal
were similarly found in Bolivia, where Leopardus tigrinus
and L. colocolo were rst conrmed only in 2001 (Pacheco
et al. 2001) and 2012 (Luque et al. 2012), respectively. In
Colombia, between 1970 and 2011 there were only 16
conrmed records of L. tigrinus (Payán-Garrido and
Figure 1: Leopardus guttulus skin collected in 1988 from the
Pantanal of Poconé, Mato Grosso state, and housed in the Emilio
Goeldi Museum (catalogue number MPEG22193; Belém, Brazil).
Photo credit: Fábio Nascimento.
Table :Summary of study design and species-specic capture rates ((photographic records/survey effort) ×) from studies conducted on
each small wild cat (SWC) species in the Pantanal.
Study details Study site
Distance between stations (m) NI  , , ,
Sampled area (km)   NI 
Grid-based sampling No Yes Yes No Yes
Leopardus pardalis  (.) (.) (.) (.), (.)
Puma yagouaroundi ()(.)(.) (.) (.)
Leopardus colocolo ()(.)(.)()(.)
Leopardus guttulus (.)()()()()
Total number of records  ,  , ,
Species-specific values are reported as total number of records with capture rates in parentheses. Study sites: A (Trolle ); B (Bianchi );
C (Bolzan ); D (Porfírio et al. ); and E (this study). NI: Not Informed.
2F.R. Tortato et al.: Small wild cats in the Pantanal
González-Maya 2011). Due to its morphological similarity
with other species of spotted cats (e.g., juvenile L. parda-
lis), cryptic habits, and overall data deciency, the distri-
bution of L. guttulus in the Pantanal is difcult to accurately
map (Rodrigues et al. 2002). However, the visual observa-
tions of L. guttulus occurred in similar geographic region
and habitat composition as the specimen collected in 1988.
Such observations can assist in directing future survey ef-
forts to better understand the occurrence and ecology of
this rare species.
The relatively high density of L. pardalis in the Pan-
tanal (Trolle and Kéry 2005) may be a contributing factor to
the low detectability and density of other SWC species,
likely due to potential intraguild competition termed the
Ocelot effect(Oliveira et al. 2010). Prior studies indicate
that L. pardalis may be a competitor or even be a potential
predator of L. guttulus,L. colocolo, and P. yagouaroundi
(Oliveira and Pereira 2013). For example, L. guttulus shifts
activity patterns to avoid overlap with L. pardalis (Oliveira-
Santos et al. 2012), indicating potential intraspecic con-
ict or predator effect.
Survey design may have also contributed to the
observed relative rarity of the SWC species. Minor changes
in the spacing of sampled sites can influence species-
specific detection rates (Srbek-Araujo and Chiarello 2013;
Trolle and Kéry 2005). In our review (Table 1), each study
used a different survey design, thus making comparisons
difcult. For logistic reasons, most studies deploy camera
traps along trails and roads, thus neglecting a question- or
design-driven distribution of cameras across the landscape
(e.g., random or grid-based sampling for density or occu-
pancy analyses; MacKenzie and Royle 2005; Sollmann
et al. 2012; Tobler and Powell 2013). This bias in sampling
design may affect estimates for elusive species like wild
cats (Tobler and Powell 2013). The majority of studies
conducted in the Pantanal did not use a random sampling
design. In Asia, Wearn et al. (2013) compared data from
random and non-random camera trap placement, and
found that the non-random survey design resulted in
underestimated abundances of Catopuma badia.
Placement of camera traps on- versus off-road can
also influence the probability of detection. In the Neo-
tropics, roads and trails are differentially used by species
including P. onca and Puma concolor,wherebyP. onca
used roads more frequently than P. concolor (Harmsen
et al. 2010). Intraspecic variations in detection rates
have been found between males and females of P. onca,
whereby males were signicantly more detectable on
roads versus females (Sollmann et al. 2011). In SWC, the
detection rate of L. pardalis was higher on roads versus
forested trails (Srbek-Araujo and Chiarello 2013).
The present study provides support for potential
contributing factors to the relative rarity of L. colocolo,
L. guttulus,andP. yagouaroundi in the Pantanal. Due to low
capture rates, future research should design surveys with
camera trap spacing and efforts appropriate for SWC. In-
sights gained through camera trap surveys will provide
better understanding of SWC ecology, including site-specic
to range wide distribution and density, population rates and
traits, and interspecic interactions. Ultimately,
Table :Visual records of Leopardus guttulus in the northern region of the Pantanal from  to .
Date Site Time (: h) Coordinates (WGS decimal degrees)
 Rio Cassange* - . .
May  Estrada da Base Night . .
May  Fazenda São João : . .
July  Fazenda São João : . .
July  Fazenda São João : . .
August  Transpantaneira : . .
February  Transpantaneira : . .
July  Fazenda São João : . .
July  Fazenda São João : . .
November  Fazenda Santa Inês : . .
July  Fazenda São Bento : . .
September  Fazenda São Bento : . .
August  Fazenda São Bento : . .
October  Estrada da Base : . .
The first confirmed record of L. guttulus in the Pantanal was an adult male collected in  and deposited at the Emilio Goeldi Museum
(catalogue number MPEG; Belém, Brazil; Nascimento and Feijó, ).
*Nascimento and Feijó .
F.R. Tortato et al.: Small wild cats in the Pantanal 3
understanding SWC distribution and the factors that
contribute to their persistence across a given landscape will
help provide more scientically accurate conservation as-
sessments and guide future management plans.
Acknowledgments: The authors would like to thank the
ranch employees of Fazenda São Bento (MS, Brazil) and
Fazenda Jofre Velho (MT, Brazil) who assisted in the field
activities. We thank Fabio Nascimento for kindly providing
the photo of Leopardus guttulus from the collection of the
Emilio Goeldi Museum (Belém, Brazil).
Author contribution: All the authors have accepted
responsibility for the entire content of this submitted
manuscript and approved submission.
Research funding: None declared.
Conict of interest statement: The authors declare no
conicts of interest regarding this article.
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Although known globally for its biodiversity, only around 5% of the Brazilian Pantanal is protected. The Network for Protection and Conservation of Amolar Mountain Ridge is an informal initiative that legally protects over 2000 km2 of the Pantanal biome. Several camera-trapping surveys were carried out at Amolar Mountain Ridge from August 2011 to September 2013 in order to increase our knowledge of the species occurrence and its ecological requirements. The aims of this study were : 1) to inventory the carnivore species occurring within this network of protected areas; 2) to describe their activity patterns and 3) to discuss threats for those species’ conservation in the region. We used the Kernel density method to describe the species’ activity patterns. We obtained 764 records (from 12703 camera-days) of eight carnivores, including endangered species in Brazil, such as the jaguar (Panthera onca), puma (Puma concolor), and ocelot (Leopardus pardalis), that were among the most frequently recorded by camera traps. The other species detected were the South America coati (Nasua nasua), the tayra (Eira barbara), the crab-eating raccoon (Procyon cancrivorus) and the jaguarundi (Puma yagouaroundi). We provided information on activity patterns of the jaguar and puma, which exhibited cathemeral activity patterns, on the ocelot and crab-eating fox, which were mostly nocturnal, and on the Southern coati and jaguarundi, which were diurnal. Scansorial and species that occur naturally in low densities as the tayra and the crab-eating raccoon were difficult to be detected with the used camera trapping setting. However, due to the natural characteristics of the study area, camera trapping is among the most appropriate tools for providing data about carnivores and their prey. This information is essential to delineate conservation plans for Amolar Mountain Ridge.
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The tigrina Leopardus tigrinus (Schreber, 1775) is a small-sized Neotropical spotted cat found from northern Argentina and southern Brazil to Costa Rica. Four subspecies are traditionally recognized: L. t. tigrinus (Schreber, 1775) from northern Brazil, the Guianas and eastern Venezuela; L. t. pardinoides (Gray, 1867) from western Venezuela, Colombia, Ecuador and Peru; L. t. guttulus (Hensel, 1872) from southern Brazil, Paraguay and northern Argentina; and L. t. oncillus (Thomas, 1903) from Costa Rica. We studied external and craniodental morphology in quantitative and qualitative terms from 250 specimens in order to clarify the taxonomic status of tigrina. Based on the characters analyzed in this study, we recognize three diagnosable morphogroups, each with a distinct geographic distribution: northern/northwest-ern/west (samples from northern Brazil, the Guianas, Venezuela, Colombia, Ecuador, Peru, northwestern Argentina and Costa Rica), eastern (samples from northeastern and central Bra-zil), and southern (samples from southern Brazil, Paraguay and northeastern Argentina). Taking into account the morphologic evidence presented here, supported by biogeographic data and molecular studies available, we recognize three full species for tigrinas: L. tigrinus (including the putative subspecies L. t. pardinoides and L. t. oncillus as junior synonyms) for northern/ northwestern/west group; L. emiliae (Thomas, 1914) for eastern group; and L. guttulus for southern group.
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Wildlife tourism has grown in recent years and in many countries represents a major economic activity. The observation of wildlife in Brazil, despite its great biodiversity potential, is underexploited. Here we discuss the importance of jaguar-tourism as an economic and ecological activity, contributing to the jaguar conservation in the Pantanal, a region with abundant fauna and recognized as an important ecotourism destination in Brazil. We argue that the absence of management plans for protected areas in the Pantanal are impeding the development of this activity.
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We investigated the home range size, habitat selection, as well as the spatial and activity overlap, of four mid-sized carnivore species in the Central Pantanal, Mato Grosso do Sul, Brazil. From December 2005 to September 2008, seven crab-eating foxes Cerdocyon thous, seven brown-nosed coatis Nasua nasua, and six ocelots Leopardus pardalis were radio-collared and monitored. Camera trap data on these species were also collected for the crab-eating raccoon Procyon cancrivorus. We hypothesized that there would be large niche differentiation in preferred habitat-type or active period between generalist species with similar diet, and higher similarity in habitat-type or activity time between the generalist species (crab-eating foxes and coatis) and the more specialized ocelot. Individual home ranges were estimated using the utilization distribution index (UD– 95% fixed Kernel). With data obtained from radio-collared individuals, we evaluated habitat selection using compositional analysis. Median home range size of ocelots was 8 km². The proportion of habitats within the home ranges of ocelots did not differ from the overall habitat proportion in the study area, but ocelots preferentially used forest within their home range. The median home range size of crab-eating foxes was 1.4 km². Foxes showed second-order habitat selection and selected savanna over shrub-savanna vegetation. The median home range size for coati was 1.5 km². Coati home ranges were located randomly in the study area. However, within their home range, coatis occurred more frequently in savanna than in other vegetation types. Among the four species, the overlap in activity period was the highest (87%) between ocelots and raccoons, with the least overlap occurring between the ocelot and coati (25%). We suggest that temporal segregation of carnivores was more important than spatial segregation, notably between the generalist coati, crab-eating fox and crab-eating raccoon.
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The distribution of species and population attributes are critical data for biodiversity conservation. As a tool for obtaining such data, camera traps have become increasingly common throughout the world. However, there are disagreements on how camera-trap records should be used due to imperfect species detectability and limitations regarding the use of capture rates as surrogates for abundance. We evaluated variations in the capture rates and community structures of mammals in camera-trap surveys using four different sampling designs. The camera traps were installed on internal roads (in the first and fourth years of the study), at 100-200 m from roads (internal edges; second year) and at 500 m from the nearest internal road (forest interior; third year). The mammal communities sampled in the internal edges and forest interior were similar to each other but differed significantly from those sampled on the roads. Furthermore, for most species, the number of records and the capture success varied widely among the four sampling designs. A further experiment showed that camera traps placed on the same tree trunk but facing in opposing directions also recorded few species in common. Our results demonstrated that presence or non-detection and capture rates vary among the different sampling designs. These differences resulted mostly from the habitat use and behavioral attributes of species in association with differences in sampling surveys, which resulted in differential detectability. We also recorded variations in the distribution of records per sampling point and at the same spot, evidencing the stochasticity associated with the camera-trap location and orientation. These findings reinforce that for species whose specimens cannot be individually identified, the capture rates should be best used as inputs for presence and detection analyses and for behavior inferences (regarding the preferential use of habitats and activity patterns, for example). Comparisons between capture rates or among relative abundance indices, even for the same species, should be made cautiously.
The Pantanal floodplain of southwestern Brazil, with its mosaic of marshes,seasonally flooded savannas, woodlands and forests, is known as one of thewildlife hotspots of South America. The region harbors a rich mammal fauna andis a stronghold for a number of mammal species; however, still very few thoroughmammal surveys have been conducted in the Pantanal. This is the first mammalsurvey from the southeastern part of the region. An intensive,three-month study of the medium to large, non-volant species wasconducted at a location in the upper Rio Negro Basin, Aquidauana region, MatoGrosso do Sul. Thirty species of mammals were recorded, including 5 xenarthrans,1 primate, 10 carnivores, 7 ungulates and 5 caviomorph rodents. It was possibleto analyze the field biology of many of the species to a level not previouslypublished for the Pantanal. Recommendations are given for camera trapping in thePantanal and the Amazon. Finally, the effects of intensive cattle ranching onconservation in the Pantanal are discussed.