Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e61917, enero-diciembre 2025 (Publicado Mar. 27, 2025)

Long-term monitoring of ocelots Leopardus pardalis

(Carnivora: Felidae) in Tortuguero National Park, Costa Rica

Stephanny Arroyo-Arce1*; https://orcid.org/0000-0003-0993-0071

Ian Thomson1; https://orcid.org/0009-0000-4714-005X

Kat Cutler1; https://orcid.org/0009-0005-0232-1822

Alexander Oakley1; https://orcid.org/0009-0007-3215-6017

1. Coastal Jaguar Conservation, 48 Milton Road, Kirkcaldy, KY1 1TL, Fife, Scotland; sturnina@gmail.com

(*Correspondence), ianitthomson@hotmail.com, kat_cutler@outlook.com, alex.aokley@gmail.com

Received 16-VII-2024. Corrected 11-XII-2024. Accepted 13-III-2025.

ABSTRACT

Introduction: The ocelot Leopardus pardalis, though currently listed as Least Concern on the IUCN Red List, has

a declining population, and faces threats such as habitat loss and fragmentation and hunting across the entirety of

its geographical range, from the Southwestern United States to Northern Argentina. Being a cryptic species and

with few studies monitoring ocelots over multiple years, there is a lack of information regarding ocelot ecology.

Objective: To further understand the behaviour and population dynamics of ocelots within the study area,

Tortuguero National Park (45 755 ha), Costa Rica, by analysing data from a long-term camera trapping study

(2011-2022).

Methods: Individual ocelots were identified using their unique spot markings. Sex ratio, daily activity patterns,

and behaviour were evaluated, whilst ocelot minimum age was estimated for each individual and designated as

either resident or transient based on the number of consecutive years observed in the study.

Results: A total of 30 individual ocelots were identified (12 females, 18 males). The daily activity pattern (noctur-

nal) and sex ratio (1 : 0.7) were consistent with that described for the species. The movement of one individual

from Tortuguero National Park to Pacuare Nature Reserve, 41 km South, highlights the importance of wildlife

corridors for the species. The study also revealed evidence of a female ocelot at least 14 years of age. There are

very few published occurrences of an ocelot this old in the wild. Scent marking behaviour was also recorded.

Conclusions: This is the longest monitoring effort for ocelots in the country, to our knowledge. Our results

highlight the importance of and need for further long-term studies of ocelots to enhance understanding of the

species and thus promote the conservation of the species and its habitat.

Keywords: camera trap; daily activity pattern; long-term monitoring; ocelot; protected area; Tropical Wet Forest.

RESUMEN

Monitoreo a largo plazo del ocelote Leopardus pardalis (Carnivora: Felidae)

en el Parque Nacional Tortuguero, Costa Rica.

Introducción: A pesar de ser considerado como una especie de Preocupación Menor según la Lista Roja de la

UICN, el ocelote Leopardus pardalis tiene una población en declive la cual enfrenta amenazas, como la pérdida

y fragmentación del hábitat natural, y la cacería, a lo largo de su distribución geográfica, desde el Suroeste de los

Estados Unidos hasta el Norte de Argentina. Al tratarse de una especie críptica y con pocos estudios disponibles

a largo plazo, se carece de información sobre su ecología.

https://doi.org/10.15517/rev.biol.trop..v73i1.61917

TERRESTRIAL ECOLOGY

2Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e61917, enero-diciembre 2025 (Publicado Mar. 27, 2025)

INTRODUCTION

The ocelot Leopardus pardalis is a Neo-

tropical spotted cat with a geographic distribu-

tion ranging from Southwestern United States

to Northern Argentina (Sunquist & Sunquist,

2002). The species is associated with areas of

dense vegetation cover, from scrublands to

tropical rainforest (Emmons, 1988; Emmons

et al., 1989; Sunquist & Sunquist, 2002) but

has been known to tolerate disturbed habitats

to some degree (Paviolo et al., 2024). It can

be found primarily at elevations below 3 000

m, although occasional sightings have been

documented above 3 000 m (Nowell & Jack-

son, 1996; Sunquist & Sunquist, 2002). While

the IUCN Red List of Threatened Species lists

the ocelot as Least Concern (Paviolo et al.,

2024), it is considered endangered in Costa

Rica. The primary threats to the ocelot across

its geographic range include habitat loss and

fragmentation, retaliatory killing due to depre-

dation of domestic animals and hunting for the

illegal wildlife trade across its geographic range

(Sunquist & Sunquist, 2002).

The ocelot has been the subject of numer-

ous studies in Costa Rica. The predominant

approach has involved employing non-invasive

techniques, such as camera traps, to gather

information about this elusive species. Topics

of study have included behaviour (scent mark-

ing: Cove et al., 2014; King et al., 2017; activ-

ity patterns: Botts et al., 2020; Herrera et al.,

2018), population density (González-Maya &

Cardenal-Porras, 2011; Vargas et al., 2023), and

habitat use (Cambronero et al., 2023; De Olivei-

ra et al., 2010; Montalvo et al., 2015; Montalvo

et al., 2023; Yaap et al., 2015). Other research

has focused on the ocelot’s diet (Chinchilla,

1997; González-Maya et al., 2010; Montalvo

et al., 2020), illegal trade (Kelly, 2018), genetic

diversity and population structure (Ruiz-García

et al., 2012; Salom-Pérez et al., 2022), the pres-

ence of infectious diseases in wild and cap-

tive felids (Avendaño et al., 2016; Blanco et

al., 2011; Romero-Vega et al., 2024), and the

rehabilitation and reintroduction of orphaned

ocelots into the wild (Montalvo et al., 2022). It

is important to highlight that although these

studies have been conducted in different areas

of Costa Rica, none of them have focused in

Tortuguero National Park.

In Tortuguero National Park, our long-

term wildlife monitoring programme has main-

ly focused on studying the local jaguar Panthera

onca population. Through this monitoring

effort, three additional species of wild cat have

been documented inhabiting the area. These

comprise sporadic records of the puma Puma

concolor and the margay Leopardus wiedii, as

Objetivo: Aumentar el conocimiento sobre el comportamiento y la dinámica poblacional de los ocelotes en el

Parque Nacional Tortuguero (45 755 ha), Costa Rica, mediante el análisis de datos de un estudio a largo plazo con

cámaras trampa (2011-2022).

Métodos: Los ocelotes se identificaron con base en el patrón de manchas. Para cada individuo se calculó la edad

mínima, y se designó como residente o transeúnte en función del número de años consecutivos observados en el

estudio. También se estimó la proporción de sexos y el patrón de actividad diaria, y se brindaron observaciones

sobre el comportamiento de la especie.

Resultados: Se identificaron 30 ocelotes (12 hembras, 18 machos). El patrón de actividad diaria (nocturno) y

la proporción de sexos (1 : 0.7) concuerda con lo descrito para la especie. El desplazamiento de un individuo

desde el Parque Nacional Tortuguero hasta la Reserva Natural Pacuare, 41 km al sur, resalta la importancia de los

corredores biológicos. Se destaca la presencia de un ocelote hembra de al menos 14 años. Hay pocas ocurrencias

publicadas de ocelotes salvajes de esa edad. También se registraron comportamientos de marcaje olfativo.

Conclusiones: Es el estudio de mayor esfuerzo de monitoreo del país, bajo nuestro conocimiento. Nuestros resul-

tados destacan la importancia de los estudios a largo plazo para incrementar el conocimiento sobre el ocelote, y

así promover la conservación de la especie y de su hábitat natural.

Palabras clave: cámaras trampa; patrón de actividad diaria; monitoreo a largo plazo; ocelote; área protegida;

bosque tropical húmedo.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e61917, enero-diciembre 2025 (Publicado Mar. 27, 2025)

well as more frequent records of the ocelot

(Arroyo-Arce & Salom-Pérez, 2014; Arroyo-

Arce et al., 2019). Despite being the second

most recorded felid in the national park after

the jaguar (I. Thomson pers. obs.), little is

known about the population and behavior of

the ocelot in the area. Our study aims to further

our understanding of this species by primarily

examining population dynamics, with obser-

vations on the behaviour of the species, using

camera trap data from the 12-year monitoring

program (2011-2022) in Tortuguero National

Park, Costa Rica. To our knowledge, this is

the longest monitoring effort for ocelots in the

country to date and one of the largest moni-

toring efforts across the species’ geographic

range (Haines et al., 2006; Satter et al., 2018;

Veals et al., 2022).

MATERIALS AND METHODS

Study site: Tortuguero National Park is

located on the Northeast Caribbean coast of

Costa Rica (10°32’28” N & 83°30’08” W, Fig. 1)

and encompasses an approximate terrestrial

area of 45 755 ha. The predominant ecosystem

is Tropical Wet Forest (Holdridge, 1969), and

elevation ranges from 0 m to 311 m above

sea level. The average temperature is between

25 and 30 oC, with a mean annual precipita-

tion of 6 000 mm (Bermudez & Hernandez,

2004). The national park is bordered to the

Northwest by the Barra del Colorado Wildlife

Refuge and Tortuguero Protected Zone, all

of which are part of Tortuguero Conserva-

tion Area (ACTo). The Western and Southern

edges of the national park are bordered by

Fig. 1. Camera trap locations across the study site of Tortuguero National Park, Costa Rica, between 2011 and 2022.

4Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e61917, enero-diciembre 2025 (Publicado Mar. 27, 2025)

communities economically dependent on crop

farming (mainly banana and pineapple), exten-

sive livestock farming (meat and milk) and to a

lesser extent, tourism (Bermudez & Hernandez,

2004; Ling, 2002).

Data collection: Ocelot presence was

recorded at camera trap stations from January

2011 to November 2022. Each station consisted

of a digital camera trap (Bushnell and Moultrie

models) set to take photos (three successive

photos per trigger event with 3 s delay between

triggers) or videos (20 s video per trigger event

with 10 s delay between triggers) and placed

in areas where ocelot detection probability is

considered to be high (e.g. trails). A total of 43

camera trap stations were continuously active

for three months to 10 years, with seven located

at the Tortuguero village boundary, nine in

the forest interior, 10 near canals, and 17 in

the coastal habitat (Fig. 1). The date, time and

geographic coordinates were recorded for each

photo/video.

Data analysis: Individual ocelots were

identified from their unique spot pattern by

cross-referencing the images and videos with

an existing photographic database created by

Coastal Jaguar Conservation from the research

conducted in Barra del Colorado Wild-

life Refuge (2014), Pacuare Nature Reserve

(2015-2016), and Tortuguero National Park

(2011-2022).

Each identifiable individual was classi-

fied as male or female based on the presence

or absence of testicles. To estimate the ocelot

minimum age, it was assumed that all adult

individuals recorded were at least a year and a

half old when first detected, as kittens usually

spend the first 12-18 months of life with their

mother (Hunter, 2015; Laack et al., 2005).

The sex ratio was expressed as the number

of males per female (male : female; Pérez-Irineo

& Santos-Moreno, 2014). A Chi-square test was

performed to determine if the ratios differed

significantly from a 1:1 ratio. Additionally, all

individuals were categorised as either ‘resi-

dent’ or ‘transient’ according to Harmsen et al.

(2017), where a resident is an individual who

has been detected in the study area for at least

three consecutive years, whereas a transient has

been detected for less than two years.

Daily activity pattern was estimated using

the kernel density estimation method, with the

von Mises distribution for circular data (Ridout

& Linkie, 2009) and following the recommen-

dations of Peral et al. (2022). All statistics were

performed using the package “Activity” (Row-

cliffe, 2023) in R version 4.4.1 (R Core Team,

2024), following the R-Code developed by

Andrade-Ponce et al. (2022). Additional infor-

mation was obtained through the camera trap

records (photos or videos) when possible (e.g.

health condition, scent-marking behaviour).

RESULTS

After a total of 38 151 camera trap nights,

we were able to identify 30 adult ocelots (12

females and 18 males) across 25 camera trap

stations including two at Tortuguero village

boundary, four in the forest interior, five near

canals, and 14 in the coastal habitat (Table 1,

Fig. 1). Although we did not record the pres-

ence of cubs, we observed four females (two

resident and two transient individuals) show-

ing signs of pregnancy or having recently given

birth (e.g. saggy belly; Table 2). The minimum

age of the ocelots recorded by the end of the

study varied from 3 years old up to the oldest

individual recorded, F02, estimated to be at

least 14 years of age (Table 2)

The sex ratio was 1:0.7, which is not sig-

nificantly different from 1:1 (c2 = 1.2; d.f. =

1; p = 0.2733). Seven of the individuals were

residents (three females and four males) and

23 transients (nine females and 14 males; Table

2). Our data indicated a stable trend of resi-

dent ocelots throughout the study period, with

varying numbers of transient individuals. The

year with the highest record of individuals was

2020 with 11 individuals (six residents and five

transients) followed by 2013 with 10 individu-

als (two residents and eight transients; Fig. 2).

It is important to highlight that one transient

male, M08, was first detected in Tortuguero

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National Park in November 2012, and on a

later date, July 2015, was identified at Pacuare

Nature Reserve, approximately 41 km South of

Tortuguero National Park (Fig. 1). After this

date, this individual was not recorded again in

our study.

The ocelots within the study showed a pre-

dominantly nocturnal activity pattern (Fig. 3).

All individuals appeared in good physical con-

dition with one exception, M19, a male resident

who was recorded from 2019 to 2022. On June

6, 2022, he was observed with eyeshine only

from his right eye (Table 2). Scent-marking

behaviour was exhibited exclusively by resident

individuals and was recorded seven times dur-

ing the study; defecation by F02, urine spraying

by M15 and M21, and the spraying of glandular

secretions by M19 on four different occasions in

the same location (Table 2). All scent-marking

behaviour occurred within the coastal habitat

along a human-made trail parallel to the beach.

DISCUSSION

The local ocelot population includes both

resident and transient individuals. It is impor-

tant to note that these terms, as described by

Harmsen et al. (2017), apply within our specific

Table 1

Ocelot Leopardus pardalis monitoring sampling effort conducted in Tortuguero National Park, Costa Rica, between 2011

and 2022.

Sector Camera trap

stations

Camera trap

days

Camera trap

images

Individuals

recorded*

Females

recorded

Males

recorded

Tortuguero village boundary 72 046 7 (3) 2 0 2

Forest interior 91 764 58 (25) 5 2 3

Canals 10 2 086 16 (9) 4 2 2

Coastal habitat 17 32 255 657 (367) 19 8 11

Tot al 43 38 151 738 (404) 30 12 18

*All individuals were classified as adults. In brackets: number of images used for individual identification.

Fig. 2. The number of ocelot Leopardus pardalis individuals identified from camera trap stations in Tortuguero National Park,

Costa Rica, between 2011 and 2022.

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sampling area. Therefore, some individuals we

identify as transient might be residents out-

side our sampling area. It is also possible that

residents have a larger territory than the area

covered by our study (Harmsen et al., 2017). To

gain a better understanding of the local ocelot

population, future studies should focus on the

species’ spatial ecology, particularly its territo-

rial behaviour and habitat use in relation to its

home range.

Table 2

Ocelot Leopardus pardalis individuals identified from camera trap stations in Tortuguero National Park, Costa Rica, between

2011 and 2022.

Individual 2011 2012 2013 2014 2015 2016 2017 2018 2019 2020 2021 2022

Number of

Years Recorded

Estimated

Minimum Age

M01 5 7

F02 PRGN PRGN PRGN, SMB PRGN PRGN 12 14

M03 1 3

M04 2 4

F05 1 3

M06 2 4

M07 2 4

M08 CONNT 1 3

M09 1 3

F10 1 3

F11 PRGN 1 3

F12 1 3

F13 1 3

M14 1 3

M15 SMB SMB 5 7

F16 PRGN PRGN 5 7

M17 1 3

M18 2 4

M19 SMB SMB, BLIND 4 6

M20 1 3

M21 SMB 4 6

F22 1 3

F23 3 5

F24 PRGN 1 3

F25 1 3

M26 1 3

M27 1 3

M28 1 3

F29 1 3

M30 1 3

M: male. F: female. SMB: scent-marking behaviour. PRGN: female with signs of pregnancy or recently given birth (e.g. saggy

belly). CONNT: individual recorded in Pacuare Nature Reserve in July 2015 (Arroyo-Arce et al. 2017). BLIND: eyeshine only

from the right eye as from June 6, 2022. Dark grey bars are resident individuals (recorded for ≥ 3 consecutive years). Light

grey bars are transient individuals (recorded for ≤ 3 consecutive years).

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e61917, enero-diciembre 2025 (Publicado Mar. 27, 2025)

We were able to estimate the minimum age

of the individuals, one of whom was at least 14

years old, representing one of the oldest record-

ed ocelots in the wild worldwide. Ocelots can

live up to 20 years in captivity (Carolina Tiger

Rescue, 2024; Wilson & Mittermeier, 2009)

with some extraordinary cases of up to 28 years

(Weigl, 2005). Although long-term studies on

ocelot populations are rare, it has been estimat-

ed that longevity in the wild is likely between

seven to 10 years (Laack, 1991; Wilson & Mit-

termeier, 2009), with some authors recording

ocelots living 10 years (Lombardi et al., 2022),

and 14 years (Muskat, 2024) in North America.

The sex ratio of ocelots observed in our

study is consistent with expectations for the

species and aligns with findings from other

studies across the species’ geographic range

(Laack et al., 2005; Pérez-Irineo & Santos-

Moreno, 2014). However, some studies have

reported skewed sex ratios (Di Bitetti et al.,

2006; Maffei et al., 2005; Magalhães & Srbek-

Araujo, 2022; Satter et al., 2018, Sternberg et

al., 2023). These variations are possible as sex

ratios can be influenced by factors such as

social organisation (e.g. solitary species, home

range sizes varying with sex), habitat character-

istics (e.g. dry versus wet season), and popula-

tion change (births, deaths, in-migration, and

emigration) which may vary across space and

time (Di Bitetti et al., 2006; Kappeler, 2017;

Maffei et al., 2005; Magalhães & Srbek-Araujo,

2022; Pérez-Irineo & Santos-Moreno, 2014).

It is important to also consider the sampling

design used in each study, as this can impact the

observed sex ratio (Magalhães & Srbek-Araujo,

2022). For example, males are more likely to use

trails compared to females (Satter et al., 2018),

influencing detection rates when camera traps

are positioned mainly on trails.

The ocelot M08 covered approximately

41 km, comparable to distances recorded in

literature (Booth-Binczik, 2007; Crawshaw,

1995; Jacob, 2002). Although it is unknown

which route the ocelot traversed, it is possible

the individual used the Colorado-Tortuguero

Biological Corridor and the Moín-Tortugue-

ro Biological Corridor, which connects Tor-

tuguero National Park and Pacuare Nature

Reserve through fragmented landscape using

Fig. 3. Daily activity pattern of the ocelot Leopardus pardalis in Tortuguero National Park, Costa Rica. Yellow: daytime; dark

yellow: twilight; grey: nighttime; blue bars: number of camera trap records.

8Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e61917, enero-diciembre 2025 (Publicado Mar. 27, 2025)

the remaining forest patches along the Carib-

bean coast (Acevedo, 2013). This is the first

evidence of wildlife movement between Tor-

tuguero National Park and Pacuare Nature

Reserve, highlighting the significance of wild-

life corridors for the long-term conservation of

the species (Arroyo-Arce et al., 2017).

The ocelot’s nocturnal habits found in

the study mirror those observed in similar

areas along the Caribbean coast of Costa Rica,

including the Barra del Colorado Wildlife Ref-

uge (Arroyo-Arce et al., 2016) and Pacuare

Nature Reserve (Arroyo-Arce et al., 2017). This

is consistent with previous descriptions of the

species (Wilson & Mittermeier, 2009), likely

co-ordinating with the activity patterns of its

main prey, small and medium-sized mammals

(Ludlow & Sunquist, 1987; Moreno et al., 2006).

It also serves as a strategy to avoid interspecific

competition and predation by the predomi-

nantly crepuscular, larger cat species, such as

jaguars and pumas (Gómez et al., 2005; Maffei

et al., 2005), both of which are found in Tortu-

guero National Park (Arroyo-Arce & Salom-

Pérez, 2014; Arroyo-Arce & Thomson, 2023).

Scent marking is a prevalent behaviour

among wild cats (Mellen, 1993), yet it poses

challenges in terms of documentation, par-

ticularly for cryptic species such as the ocelot

(King et al., 2017; Moreno & Giacalone, 2006;

Rodgers et al., 2014). Despite these challenges,

our camera traps captured seven instances of

scent marking, including the deposition of

faeces and urine marking. As noted by previous

researchers, scent marking serves as an olfacto-

ry mechanism for both intra- and interspecific

communication, conveying information about

an individual’s sex, reproductive status, terri-

tory, and movement patterns (King et al., 2017;

Mellen, 1993; Moreno & Giacalone, 2006).

The ocelot population exhibited signs of

reproduction, with females showing signs of

pregnancy, despite the absence of observed off-

spring or young individuals. The overall health

of the individuals appeared to be robust, as no

external signs of sickness or injury were evi-

dent, except for one ocelot, identified as M19,

which displayed eyeshine from only one eye,

possibly indicative of damage to the tapetum

lucidum. This could have been caused by vari-

ous factors, such as an injury from a prey ani-

mal defending itself (Kays, 2016). Although no

mortalities were documented during the study,

it is imperative for future research to investigate

the threats faced by ocelots, not only in Tortu-

guero National Park, but also in the Tortuguero

Conservation Area. Potential threats include

conflict with local communities (e.g. ocelot

predation on domestic animals), illegal wild-

life trade, road mortality (ocelots are the most

frequently road-killed wild cat in Costa Rica),

and habitat fragmentation (Corrales-Gutiérrez,

2016; Kelly, 2018; Salom-Pérez et al., 2022).

Understanding these threats is crucial for mak-

ing informed management decisions, necessary

for the long-term survival of the species at both

local and regional levels.

Our research is, to our knowledge, the

only study of its kind conducted in Tortuguero

National Park and represents the longest moni-

toring effort for ocelots in Costa Rica. It is also

among the longest studies of ocelots in the

Americas (Haines et al., 2006; Satter et al., 2018;

Veals et al., 2022). Consequently, our findings

provide critical insights into the ecology of this

species. Additionally, our study raises impor-

tant questions regarding various population

demographic parameters of ocelots, such as

abundance, density, and survival rate. Address-

ing these questions will enhance our under-

standing of ocelots at both local and regional

levels. Finally, our study emphasises the sig-

nificance of long-term monitoring projects in

achieving a deeper, more robust understanding

of the ocelot and the conditions necessary for

the species to thrive.

Ethical statement: The authors declare

that they all agree with this publication and

made significant contributions; that there is no

conflict of interest of any kind; and that we fol-

lowed all pertinent ethical and legal procedures

and requirements. All financial sources are fully

and clearly stated in the acknowledgments sec-

tion. A signed document has been filed in the

journal archives.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e61917, enero-diciembre 2025 (Publicado Mar. 27, 2025)

ACKNOWLEDGMENTS

Our research was conducted under the

permits required by the Costa Rican govern-

ment (ACTo-SINAC). We are thankful for the

support provided by Global Vision Internation-

al, Sea Turtle Conservancy, The Rufford Small

Grants Foundation, and Idea Wild, that allowed

us to conduct our research. Special thanks to

the reviewers for their insightful comments,

which significantly contributed to the improve-

ment of our manuscript.

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