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 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.
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
ﬂoodplain 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 Geoffroy’s 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 deﬁcient (Tomás et al. 2010).
ThepresenceofL. guttulus in the Pantanal was ﬁrst
veriﬁed in 1988 with a skin (Figure 1) collected in the
Poconé region (Mato Grosso, Brazil) and currently
housed in the scientiﬁc collection of the Emili Goeldi
Museum (catalogue number MPEG22193; Belém, Brazil;
Nascimento and Feijó 2017). There has since been only
one veriﬁed 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 2–3 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,
E-mail: email@example.com. https://orcid.org/0000-0003-1901-
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 ﬁrst collected in 1988 (Figure 1; Nascimento
and Feijó 2017); all subsequent visual records of L. guttulus
occurred during ﬁeld 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
conﬁrmation 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 conﬁrmed.
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 ﬁrst conﬁrmed only in 2001 (Pacheco
et al. 2001) and 2012 (Luque et al. 2012), respectively. In
Colombia, between 1970 and 2011 there were only 16
conﬁrmed 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-speciﬁc 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 deﬁciency, the distri-
bution of L. guttulus in the Pantanal is difﬁcult 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 intraspeciﬁc con-
ﬂict 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
difﬁcult. 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). Intraspeciﬁc variations in detection rates
have been found between males and females of P. onca,
whereby males were signiﬁcantly 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-speciﬁc
to range wide distribution and density, population rates and
traits, and interspeciﬁc 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 scientiﬁcally 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.
Conﬂict of interest statement: The authors declare no
conﬂicts of interest regarding this article.
Arita, H.T., Robinson, J.G., and Redford, K.H. (1990). Rarity in
Neotropical forest mammals and its ecological correlates.
Conserv. Biol. 4:181–192.
Bianchi, R.C., Oliﬁers, N., Gompper, M.E., and Mourão, G. (2016).
Niche partitioning among mesocarnivores in a Brazilian wetland.
PLoS ONE 11:e0162893.
Bianchi, R.C. (2009). Ecologia de Mesocarnívoros em uma Área no
Pantanal Central, Mato Grosso do Sul, Ph.D. dissertation.
Bolzan, A. (2011). Relação Entre a Estimativa da Abundância de
Mamíferos Terrestres de Médio e Grande Porte e Variáveis
Ambientais em uma Área do Pantanal de Mato Grosso do Sul.
Graduate thesis. Universidade Federal do Rio Grande do Sul, Rio
Grande do Sul, Brazil.
Carbone, C., Christie, S., Conforti, K., Coulson, T., Franklin, N., Ginsberg,
(2001). The use of photographic rates to estimate densities of tigers
and other cryptic mammals. Anim. Conserv. 4:75–79.
Godoi, M.N., Teribele, T., Bianchi, R., Olﬁers, N., Concone, H.V.B., and
Xavier-Filho, N.L. (2010). New records of pampas cat for Mato
Grosso do Sul State, Brazil. Cat News 52:28–29.
Harmsen, B.J., Foster, R.J., Silver, S., Ostro, L., and Doncaster, C.P.
(2010). Differential use of trails by forest mammals and the
implications for camera-trap studies: a case study from Belize.
Harris, M.B., Tomás, W., Mourão, G., Silva, C.J., Guimarães, E., Sonoda,
F., and Fachim, E. (2005). Safeguarding the Pantanal wetlands:
threats and conservation initiatives. Conserv. Biol. 19:714–720.
Hunter, L. (2011). Carnivores of the world. Princeton University Press,
Princeton, NJ, USA.
Junk, W.J., Nunes da Cunha, C., Wantzen, K.M., Petermann, P.,
Strüssmann, C., Marques, M.I., and Adis, J. (2006). Biodiversity
and its conservation in the Pantanal of Mato Grosso, Brazil.
Aqua. Sci. 69:278–309.
Karanth, K.U. and Nichols, J.D. (1998). Estimation of tiger densities in
India using photographic captures and recaptures. Ecology 79:
Luque, J.A.D., Beraud, V., Torres, P.J., Kacoliris, F.P., Daniele, G.,
Wallace, R.B., and Berkunsky, I. (2012). First record of Pantanal
cat, Leopardus colocolo braccatus, in Bolivia. Mast. Neotrop. 19:
MacKenzie, D.I. and Royle, J.A. (2005). Designing occupancy studies:
general advice and allocating survey effort. J. Appl. Ecol. 42:
Nascimento, F.O. and Feijó, A. (2017). Taxonomic revision of the tigrina
Leopardus tigrinus (Schreber, 1775) species group (Carnivora,
Felidae). Pap. Avul. de Zool. (São Paulo) 57:231–264.
Oliveira, T.G. and Pereira, J.A. (2013). Intraguild predation and
interspeciﬁc killing as structuring forces of carnivoran
communities in South America. J. Mamm. Evol. 21:427–436.
Oliveira, T.G., Tortato, M.A., Silveira, L., Kasper, C.B., Mazim, F.D.,
Lucherini, M., Jácomo, A.T., Soares, J.B.C., Marques, R.V., and
Sunquist, M.E. (2010). Ocelot ecology and its effect on the small-
felid guild in the lowland Neotropics. In: Macdonald, D.W. and
Loveridge, A.J. (Eds.), Biology and conservation of wild felids.
Oxford University Press, Oxford, UK, pp. 563–574.
Oliveira-Santos, L.G.R, Graipel, M.E., Tortato, M.A., Zucco, C.A.,
Cáceres, N.C., and Goulart, F.V.B. (2012). Abundance changes
and activity ﬂexibility of the oncilla, Leopardus tigrinus
(carnivora: Felidae), appear to reﬂect avoidance of conﬂict.
Pacheco, L.F, Guerra, J.F., Deem, S.L., and Frías, P.C. (2001). Primer
registro de Leopardus tigrinus (Shreber, 1775) en Bolivia. Ecol. en
Payán-Garrido, E. and González-Maya, J.F. (2011). Distribución
geográﬁca de la oncilla (Leopardus tigrinus) en Colombia e
implicaciones para su conservación. Rev. Latin. de Conserv.
Porfírio, G, Foster, V.C., Sarmento, P., and Fonseca, C. (2018). Camera
traps as a tool for carnivore conservation in a mosaic of protected
areas in the Pantanal wetlands, Brazil. Nat. Conserv. Res.
Rodrigues, F.H.G, Medri, I.M., Tomás, W.M., and Mourão, G.M. (2002).
Revisão do conhecimento sobre ocorrência e distribuição de
Mamíferos do Pantanal. Embrapa Pantanal Documentos 38,
Brazil, Available at: https://ainfo.cnptia.embrapa.br/digital/
bitstream/item/81202/1/DOC38.pdf (Accessed 05 November
Soisalo, M.K. and Cavalcanti, S.M.C. (2006). Estimating the density of
a jaguar population in the Brazilian Pantanal using camera-traps
and capture-recapture sampling in combination with GPS radio-
telemetry. Biol. Conserv. 129:487–496.
Sollmann, R., Gardner, B., and Belant, J.L. (2012). How does spatial
study design inﬂuence density estimates from spatial capture-
recapture models?. PLoS ONE 7:e34575.
Sollmann, R., Furtado, M.M., Gardner, B., Hofer, H., Jácomo, A.T.A.,
Torres, N.M., and Silveira, L. (2011). Improving density estimates
for elusive carnivores: accounting for sex-speciﬁc detection and
movements using spatial capture-recapture models for jaguars
in central Brazil. Biol. Conserv 144:1017–1024.
4F.R. Tortato et al.: Small wild cats in the Pantanal
Srbek-Araujo, A.C. and Chiarello, A.G. (2005). Is camera-trapping an
efﬁcient method for surveying mammals in Neotropical forests? A
case study in south-eastern Brazil. J. Trop. Ecol. 21:1–5.
Srbek-Araujo, A.C. and Chiarello, A.G. (2013). Inﬂuence of camera-trap
sampling design on mammal speciescapture ratesand community
structures in southeastern Brazil. Biota Neotrop. 13:51–62.
Sunquist, M. and Sunquist, F. (2002). Wild cats of the world. The
University of Chicago Press, Chicago, IL, USA.
Tobler, M.W. and Powell, G.V.N. (2013). Estimating jaguar densities with
camera traps: problems with current designs and
recommendations for future studies. Biol. Conserv. 159:109–118.
Tobler, M.W., Carrillo-Percastegui, S.E., Pitman, R.L., Mares, R., and
Powell, G. (2008). An evaluation of camera traps for inventorying
large and medium sized terrestrial rainforest mammals. Anim.
Tomás, W.M., Miranda, G.H.B., and Rudran, R. (2003). Uso de
armadilhas fotográﬁcas em levantamentos populacionais. In:
Cullen, Jr., L. and Valadares-Padua, C. (Eds.), Métodos de
Estudos em Biologia da Conservação e Manejo da Vida Silvestre.
Editora UFPR, Curitiba, Brazil, pp. 243–265.
Tomás, W.M., Cáceres, N.C., Nunes, A.P., Fischer, E., Mourão, G.M.,
and Campos, Z.Z. (2010). Mammals in the Pantanal wetland,
Brazil. In: Junk, W.J., Silva, C.J., Cunha, C.N., and Wantzen, K.M.
(Eds.), The Pantanal: Ecology, biodiversity and sustainable
management of a large Neotropical seasonal wetland. Pensoft
Publishers, Soﬁa, Bulgaria, pp. 127–141.
Tortato, F.R. and Izzo, T.I. (2017). Advances and barriers to the
development of jaguar-tourism in the Brazilian Pantanal. Persp.
Ecol. Cons 15:61–63.
Trolle, M. and Kéry, M. (2003). Estimation of ocelot density in the
Pantanal using capture–recapture analysis of camera-trapping
data. J. Mammal. 84:607–614.
Trolle, M. and Kéry, M. (2005). Camera-trap study of ocelot and other
secretive mammals in the northern Pantanal. Mammalia 69:3–4.
Trolle, M (2003). Mammal survey in the southeastern Pantanal, Brazil.
Biodivers. Conserv. 12:823–836.
Wearn, O.R., Rowcliffe, J.M., Carbone, C., Bernard, H., and Ewers, R.M.
(2013). Assessing the status of wild felids in a highly-disturbed
commercial forest reserve in Borneo and the implications for
camera trap survey design. PLoS ONE 8:e77598.
F.R. Tortato et al.: Small wild cats in the Pantanal 5