Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e61782, enero-diciembre 2025 (Publicado Feb. 24, 2025)

Two decades of jaguar and puma (Carnivora: Felidae)

activity in lowland forest of eastern Ecuador

John G. Blake1,2*; https://orcid.org/0000-0003-4668-2636

Diego Mosquera B.2,3; https://orcid.org/0000-0003-2169-4825

Gabriela S. Vinueza-Hidalgo2,4; https://orcid.org/0000-0002-2960-2836

Bette A. Loiselle2,5; https://orcid.org/0000-0003-1434-4173

1. Department of Wildlife Ecology and Conservation, University of Florida, Gainesville, Florida, 32611, United States of

America; john.blake@ufl.edu (*Correspondence)

2. Estación de Biodiversidad Tiputini, Colegio de Ciencias Biológicas y Ambientales, Universidad San Francisco de

Quito, Quito, Ecuador

3. Independent researcher; dimosb@rocketmail.com

4. Osa Conservation, Washington, D. C., United States of America; gabyvh@gmail.com

5. Department of Wildlife Ecology and Conservation and Center for Latin American Studies, University of Florida,

Gainesville, Florida 32611, United States of America; bloiselleb@latam.ufl.edu

Received 04-IX-2024. Corrected 28-XI-2024. Accepted 18-II-2025.

ABSTRACT

Introduction: Jaguars (Panthera onca) and pumas (Puma concolor) are the two largest terrestrial predators in

lowland Neotropical forests and as such, are important contributors to the ecosystem. Yet, long-term studies on

their temporal and spatial patterns of occurrence are not common.

Objectives: To update a previous eight year (2005-2012) camera-trap study on jaguars at Tiputini Biodiversity

Station, Yasuní Biosphere Reserve, with data from 2014 through 2023; and to add complementary information

on pumas.

Methods: We used camera traps set along trails or at mineral licks to document the occurrence of jaguars and

pumas. Individual jaguars were identified by their distinctive coat patterns.

Results: Capture rates from 2014 to 2023 varied from 0 to 2.94 images/100 trap days for jaguars and from 0.46 to

4.88 for pumas. These rates were similar or increased across all years for both species. We identified 28 individual

jaguars during the second sample period, including 18 males and seven females. Periods between captures ranged

from 1 to 84 months, with eight individuals recorded over at least 36 months. Including images from the first

period (2005-2012), when 21 individuals were identified, it is likely that ~50 individual jaguars have occurred in

or close to the research station over 19 years. Jaguars were primarily active during daylight hours, while pumas

were more active at night.

Conclusions: TBS is embedded within a large biosphere reserve but is too small (~670 ha) to cover the home

range of either species. Nonetheless, given the number of records and the fact that capture rates have not declined

in the past two decades, this region is important for the conservation of these two species and the many prey

they depend on.

Key words: camera trap; daytime activity; long-term studies; Neotropical; Tiputini Biodiversity Station; Yasuní.

https://doi.org/10.15517/rev.biol.trop..v73i1.61782

TERRESTRIAL ECOLOGY

2Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e61782, enero-diciembre 2025 (Publicado Feb. 24, 2025)

INTRODUCTION

Long-term (e.g., > 10 yr) studies on pat-

terns of occurrence of jaguars (Panthera onca,

Linnaeus, 1758) and other large predators

(e.g., pumas Puma concolor, Linnaeus, 1771)

in undisturbed lowland forest are not common

(Harmsen et al., 2017). Yet, such studies are

useful for evaluating the conservation poten-

tial of protected (and unprotected) regions.

Given that jaguars and pumas are the largest

terrestrial predators that coexist in lowland

forests of the Neotropics, their continued pres-

ence may serve as an indication of the health

of the forest (Espinosa et al., 2018; Terborgh

1988). If large predators are present in good

numbers, there are likely to be plenty of prey,

for example. Reduced numbers of jaguars may

reflect the impacts of human activities (e.g.,

hunting) on preferred prey items, such as pec-

caries, deer, agoutis, armadillos, and others,

rather than a consequence of direct hunting

by humans, although killings of jaguars do

occur (Espinosa et al., 2018; D. Mosquera,

personal observation).

Most studies of jaguars are conducted over

large areas given the large home ranges typi-

cal of large predators (Gonzalez-Borrajo et al.,

2016; Harmsen et al., 2020; Maffei et al., 2004;

Silver et al., 2004). Yet, most studies are relative-

ly short in duration (“snapshot surveys”; Harm-

sen et al., 2017), which may not provide a good

indication of population patterns. Although

home ranges of male and female jaguars are

known to overlap (e.g., Gonzalez-Borrajo et

al., 2016; Harmsen et al., 2017; Soisalo & Cav-

alcanti, 2006), we know relatively little about

the extent of temporal and spatial overlap of

individual jaguars at more local scales (Blake et

al., 2014; Emmons, 1987; Harmsen et al., 2009).

With overlapping home ranges, many jaguars

and pumas may use the same areas of forest

(e.g., Gonzalez-Borrajo et al., 2016; Harmsen

et al., 2009; Harmsen et al., 2017; Scognamillo

RESUMEN

Dos décadas de actividad del jaguar y el puma (Carnivora: Felidae)

en los bosques de tierras bajas del oriente de Ecuador

Introducción: Los jaguares (Panthera onca) y los pumas (Puma concolor) son los dos mayores depredadores

terrestres de los bosques neotropicales de tierras bajas y, como tales, son importantes contribuyentes al eco-

sistema. Sin embargo, no son comunes los estudios a largo plazo sobre sus patrones temporales y espaciales de

presencia.

Objetivos: Actualizar un estudio previo de ocho años (2005-2012) con cámaras trampa sobre jaguares en la

Estación de Biodiversidad Tiputini, Reserva de la Biosfera Yasuní, con datos de 2014 a 2023; y agregar informa-

ción complementaria sobre pumas.

Métodos: Utilizamos cámaras trampa instaladas a lo largo de senderos o en saladeros para documentar la pre-

sencia de jaguares y pumas. Los jaguares individuales fueron identificados por sus patrones distintivos de pelaje.

Resultados: Las tasas de captura de 2014 a 2023 variaron de 0 a 2.94 imágenes/100 días de trampa para jaguares

y de 0.46 a 4.88 para pumas. Estas tasas se mantuvieron o incrementaron en todos los años para ambas especies.

Identificamos 28 individuos de jaguares durante el segundo período de muestreo, incluidos 18 machos y siete

hembras. Los períodos entre capturas variaron de 1 a 84 meses con ocho individuos registrados durante al menos

36 meses. Incluyendo imágenes del primer período (2005-2012) cuando se identificaron 21 individuos, es pro-

bable que ~50 jaguares individuales hayan aparecido en o cerca de la estación de biodiversidad durante 19 años.

Los jaguares estuvieron principalmente activos durante las horas del día. Los pumas fueron más activos durante

la noche.

Conclusiones: La TBS está dentro de una gran reserva de la biosfera, pero es demasiado pequeña (~670 ha) para

cubrir el área de distribución de ambas especies. No obstante, dada la cantidad de registros y el hecho de que las

tasas de captura no han disminuido en las últimas dos décadas, la región es importante para la conservación de

estas dos especies y las presas de las que dependen.

Palabras clave: cámara trampa; actividad diurna; estudios a largo plazo; Neotropical; Estación de Biodiversidad

Tiputini; Yasuní.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e61782, enero-diciembre 2025 (Publicado Feb. 24, 2025)

et al., 2003; Soisalo & Cavalcanti, 2006). How-

ever, individuals may use the same areas but at

different times (e.g., transients vs. permanent

residents; Harmsen et al., 2017) without neces-

sarily encountering one another, although that

is possible (Emmons, 1987).

Jaguars and pumas occur in a wide vari-

ety of habitats (Gonzalez-Borrajo et al., 2016)

although pumas have a much greater geograph-

ic range. In tropical regions, for example, jag-

uars occur in habitats as diverse as grasslands,

dry forests (Kelly 2003; Maffei et al., 2004;

Scognamillo et al., 2003; Soisalo & Cavalcanti,

2006), and lowland wet forests (Espinosa et

al., 2018; Harmsen et al., 2017; Harmsen et al.,

2020; Silver et al., 2004; Tobler et al., 2013; Wal-

lace et al., 2003). Although lowland Amazonian

forests are among the most important habitats

for jaguars (Tobler et al., 2013), there have been

relatively few studies on jaguars (or pumas) in

such habitats (Gonzalez-Borrajo et al., 2016).

Previously (Blake et al., 2014), we reported

on temporal and spatial patterns of occurrence

and activity of jaguars at Tiputini Biodiversity

Station (TBS), Ecuador. Located in lowland

Amazonian forest of Eastern Ecuador, TBS is

situated in the midst of extensive, relatively

undisturbed forest, in one of the most biodi-

verse regions in the world (Bass et al., 2010).

The station and surroundings have experi-

enced very little disturbance, except around

the main buildings, and human activity has

apparently had relatively little impact on ani-

mal activity (Blake & Mosquera, 2014; Blake,

Mosquera & Salvador, 2012). The station area

is characterized by an intact fauna, with all top

predators (e.g., jaguar, puma, ocelot Leopardus

pardalis Linnaeus, 1758, harpy eagle Harpia

harpyja¸Linnaeus, 1758) and prey (e.g., col-

lared peccaries Dicotyles tajacu, Linnaeus, 1758,

white-lipped peccaries Tayassu pecari, Link

1795; red brocket deer Mazama americana,

Erxleben, 1777; black agoutis Dasyprocta fuligi-

nosa, Wagler, 1832; and others) present; most

are frequently observed and/or captured in

camera-trap images (Blake, Mosquera, Loiselle

et al., 2012; Blake et al., 2014; Blake et al., 2016).

Nonetheless, the station and its surroundings

are likely subjected to external influences, such

as caused by climate change. For example,

substantial declines in bird populations over

the last decade (Blake & Loiselle, 2024), in the

absence of habitat change, suggest that external

forces have had an impact. Similarly, prey pop-

ulations may exhibit changes in abundance that

could influence predator populations (Espinosa

et al., 2018; Fragoso et al., 2022). White-lipped

peccaries, for example, have apparently mostly

disappeared from the station area based on

data from camera traps, with few if any images

since 2021 (J. G. Blake, unpublished data; see

Fragoso et al., 2022).

Our first study was based on camera-trap

images collected over an 8-yr sample period

(Blake et al., 2014). Here, we reexamine pres-

ence/activity of jaguars at the station over

10 additional years (i.e., ~20 years combined

across the two study periods, 2005-2012, 2014-

2023, longer than most studies on jaguars

(Harmsen et al., 2017), to determine (1) wheth-

er occurrence (e.g., capture rate), (2) daily

activity patterns (e.g., nocturnal vs diurnal),

and (3) presence and number of individuals

of jaguars have changed over time. Given that

jaguars and pumas are the two largest predators

in lowland tropical forest (Gonzalez-Borrajo et

al., 2016) and can overlap in diet and tempo-

ral/spatial occurrence (Harmsen et al., 2009;

Harmsen et al., 2011; Romero-Muñoz et al.,

2010), we compare capture rates and activity

between the two species. Unlike jaguars, how-

ever, we do not attempt to identify individual

pumas given their relative lack of distinguishing

marks, although some are identifiable based on

scars, bot-fly infestations, or other marks (Kelly

et al., 2008; Romero-Muñoz et al., 2010). There

have not been any significant changes in levels

of human activity at or near the station over

the past two decades so we predicted that there

would be little change in occurrence or activity.

As in our previous study (Blake et al., 2014), we

emphasize that we are not attempting to esti-

mate density of jaguars; our study area is much

too small to allow for such estimates (Harmsen

et al., 2020). Rather, we focus on the extent to

4Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e61782, enero-diciembre 2025 (Publicado Feb. 24, 2025)

which large cats use the same small area of for-

est across multiple years.

MATERIALS AND METHODS

Study area: Research was conducted at

Tiputini Biodiversity Station (TBS), Orella-

na Province, Ecuador (0°37’ S & 76°10’ W,

190-270 m.a.s.l.). TBS is located on the North

side of the Tiputini River, adjacent to Yasuní

National Park on a tract (~670 ha) of largely

undisturbed lowland rain forest within the

biologically diverse Yasuní Biosphere Reserve

(Bass et al., 2010). The station and nearby areas

are dominated by terra firme forest; várzea

forest, palm swamps, and various successional

habitats also are present. Additional descrip-

tions of the forest composition in the Yasuní

region can be found in Pitman et al. (2001) y

Pitman et al. (2002). Mean annual precipitation

at Yasuní Research Station, approximately 30

km WSW of TBS, is about 3 100 mm.

Camera traps: We used cameras triggered

by an infrared heat-and-motion detector to

capture images of jaguars (and other large

mammals and birds), starting in 2005. Here,

our main focus is on images collected at ten

locations during January to March from 2014

through 2023, to complement records from

2005 to 2012 (see Blake et al., 2014 for details

of the previous sampling activity). Data from

the previous study are included when appro-

priate for comparison. Pairs of cameras were

located approximately 1-1.2 km apart along

narrow (< 0.5 m) preexisting trails within terra

firme forest (see map in Blake et al., 2016). Two

cameras were placed at each location, on oppo-

site sides of the trail, approximately 0.5-0.75 m

off the ground, with the goal of ensuring that

both sides of jaguar individuals were photo-

graphed. Jaguar coat patterns differ by side so

having images of both sides improves chances

for individual identification. Vegetation was

cleared immediately in front of each camera,

but locations were not otherwise disturbed; no

attractants were used. Cameras remained con-

tinuously activated (except when malfunctions

occurred); date and time were automatically

stamped on each image. Cameras were set to

record five images when the sensors were acti-

vated with a minimum 5-min break between

successive triggers.

The previous study (Blake et al., 2014)

was based on results from a combination of

filmbased (Highlander Photoscout, PTC Tech-

nologies, Huntsville, Alabama; 2005-2008) and

digital camera traps (Cuddeback Capture, Cud-

deback, Green Bay, Wisconsin 2010-2012). The

current study is based on Reconyx Hyper-

fire cameras (Reconyx, Holmen, Wisconsin).

According to manufacturers’ information, all

cameras had similar reaction times of ~ 0.2-

0.5 sec. Based on video records of jaguars and

pumas walking along the trails, it is clear that

they walk relatively slowly, and all camera

models should, therefore, be able to capture

their images. This is further supported by

the fact that multiple images are obtained for

each event.

In addition to images captured by cameras

located along trails, other images were captured

by cameras located on two 100 ha study plots

(Blake & Loiselle, 2018) and at four mineral

licks (saladeros). Two licks were within the sta-

tion boundaries but not along trails and two

were located near (2 to ~5 km) but outside

the station boundaries south of the Tiputini

River. Further, separate cameras captured video

records of jaguars along trails on the station.

We use images from these additional sites in

analyses that focus on daily activity patterns

and to document presence in the study area

(e.g., number of months over which an individ-

ual occurred within the study area) but do not

use them for analyses based on capture rates.

All capture-rate analyses for the current study

are based on cameras located at the ten sites

sampled in each year; capture rate data from

the earlier study (Blake et al., 2014) were based

on nine to twelve sites from each year, including

the locations used in the present study.

Analyses: We summarized images by spe-

cies (and individuals of jaguars, identified,

when possible, by the distinctive patterns on

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e61782, enero-diciembre 2025 (Publicado Feb. 24, 2025)

coats), location, date, and time. We classified

images as belonging to independent records if

more than 30 min had elapsed between con-

secutive photographs of the same species at

the same location (Datta et al., 2008; O’Brien

et al., 2003). Activity was evaluated in terms of

the number of independent images / 100 trap-

days (hereafter referred to as capture rates; i.e.,

captures of images). Previous studies have dem-

onstrated that capture rates can reflect changes

in actual abundance (Carbone et al., 2001;

Kuprewicz, 2012; Rovero & Marshall, 2009) but

we do not claim that capture rates are necessar-

ily an accurate indication of changes in actual

abundance. We calculated number of trap days

from the time the camera was placed in opera-

tion until it was removed or, if malfunctions

occurred (e.g., batteries failed), until the last

image was recorded (based on date and time

stamps on the image). We combined records

within a year. We classified records by two-hour

blocks, starting at midnight, to examine hourly

patterns of activity (e.g., 0200 h would include

records from 0001 h to 0200 h).

We used correlation analyses to compare

capture rates, hourly activity patterns, and spa-

tial distribution patterns between jaguars and

pumas. Similarly, we used correlation analyses

to examine capture rates over time (years).

We further used linear regression to examine

the change in capture rates from 2005 to 2023.

Two-sample t-tests were used to compare cap-

ture rates between the first (2005-2012) and

second (2014-2023) sample periods for both

jaguars and pumas. We used paired t-tests to

compare capture rates of jaguars and pumas

over time, from 2005 to 2023, and to compare

numbers of images at different locations. We

used chi-square tests to compare the distribu-

tion of numbers of months individual jaguars

were present between the first and second

sample periods and to compare diurnal vs noc-

turnal records for both species. All statistical

analyses were run with Statistix 10.0 (Analytical

Software, 2013) except for the analyses of over-

lap described below.

To further evaluate hourly activity pat-

terns between the two species, we estimated the

coefficient of overlapping Δ (Ridout & Linkie,

2009) using package overlap in R (Meredith &

Ridout, 2014; R Core Team, 2024). This coef-

ficient provides a descriptive measure of the

similarity in two Kernel density curves and

ranges from Δ = 0 (no overlap; different activity

patterns) to Δ = 1 (complete overlap; identical

activity patterns). We used the estimator Δ^

when sample sizes for jaguar and puma were

less than 50 and Δ^

4 when sample sizes were

greater. Confidence intervals (95 %) for the

overlap estimator were calculated using boot-

strap resampling of the data set of detections of

each species with 999 iterations, recalculating Δ

each time (Ridout & Linkie, 2009). To evaluate

whether the pairs of activity patterns were sig-

nificantly different, we used the compareCkern

function in the package activity (Rowcliffe,

2023) in R. This function is a randomization

test to determine if two sets of circular observa-

tions differ from each other (Lee et al., 2024).

RESULTS

Capture rates: Cameras along trails (n =

10 sites) captured 65 independent images of

jaguars and 91 of pumas. Capture rates from

2014 to 2023 (Table 1, Fig. 1) varied from 0.0 to

2.94 images/100 trap days for jaguars and from

0.46 to 4.88 for pumas. The high value (4.88)

for pumas was from 2016 when more than

twice as many images were recorded compared

to other years. Across all years, from 2005 to

2023 (Fig. 1), mean capture rates were 1.20 (SE

= 0.20, CV = 65.1) for jaguars and 1.25 (SE =

0.29, CV = 93.9) for pumas and did not differ

between species (paired t-test, t = 0.17, d.f. =

15, p = 0.87). Mean capture rates of jaguars

did not differ between the first set of samples

(2005-2012, 0.97, 0.23 SE) and the second

(2014-2023, 1.38, 0.30 SE; t = 1.03, d.f. = 14, p

= 0.32, variances not different, F8,6 = 2.13, p =

0.19). In contrast, capture rates of pumas were

significantly higher during the second period

(0.55, 0.07 SE and 1.80, 0.45 SE; t = 2.76, d.f.

= 8.4, p < 0.05, unequal variances F8,6 = 49.3,

p < 0.001).

6Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e61782, enero-diciembre 2025 (Publicado Feb. 24, 2025)

Number of images and number of indi-

viduals of identified jaguars are given first for

cameras located along trails (i.e., those used

to calculate capture rates) and for all camera

locations, in parentheses, including those not

on trails.

Regression analyses (Fig. 1) indicated

that capture rates increased for both species,

although the rate of increase was not great

(slope = 0.074, 0.03 SE, p < 0.05 for jaguars;

slope = 0.091, 0.049 SE, p = 0.086 for pumas).

When data from 2016 was omitted because of

the unusual number of records for pumas, the

rates of increase were similar and significant

for both species (slope = 0.076, 0.031 SE for

jaguars; 0.075, 0.025 SE for pumas; p < 0.05

for both species. Capture rates of jaguars and

pumas were not correlated across years (r =

0.23, p = 0.40).

Individual jaguars: Including images from

cameras not located along the main trails,

there were 121 independent records of jaguars

(Table 1) that represented 28 individuals (plus

four photographs that could not be assigned

to a specific individual), including 18 males

and seven females; 12 images of at least three

individuals could not be assigned to a sex

(Table 2). One male was melanistic. Number

of photographs per individual jaguar ranged

from one or two (12 individuals) to 12 or 13 (3

individuals, Table 2). Capture periods (number

of months from first to last photograph of an

individual) ranged from one to three months

(i.e., all within a one-year sample, 15 individu-

als) to 36 or more (i.e., over a ≥ 3-year period,

eight individuals) from 2014 to 2023 (Fig. 2).

Five males and two females were recorded

over a period of at least 36 months; these two

females were present at TBS for at least 72 and

84 months (Table 2). The distribution of indi-

viduals across different numbers of months did

not differ between 2005-2012 and 2014-2023

(Fig. 2, χ2 = 0.83, d.f. = 4, p = 0.94).

Diurnal activity patterns: Jaguars

were primarily active during daylight hours

Table 1

Summary of sampling effort by year; cameras operating Jan-March. Number of sites, total number of trap nights, total

number of independent photos, number of identified individual jaguars, and capture rates.

Ye a r Sites Trap nights Jaguars Pumas

Images Individuals Rate Images Rate

2014 10 652 0 (4) 0 (4) 0.00 3 0.46

2015 10 529 4 (13) 2 (5) 0.95 3 0.57

2016 10 594 6 (18) 3 (7) 1.01 29 4.88

2017 10 557 4 (8) 1 (3) 0.72 11 1.97

2018 10 623 12 (28) 5 (9) 1.93 8 1.61

2019 10 562 6 (9) 3 (5) 1.07 13 2.31

2020 10 578 16 (24) 7 (8) 2.94 13 2.25

2022 10 447 8 3 1.79 6 1.34

2023 10 558 9 4 1.61 5 0.90

Fig. 1. Capture rates (independent images) for jaguars and

pumas at Tiputini Biodiversity Station, Ecuador, from 2005

to 2023. Regression lines were based on data with 2016

excluded (see text).

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(06:00-18:00, 77 % of photographs with all years

combined) whereas pumas were most active at

night (18:00-06:00, 65 %) (Fig. 3). Number of

nocturnal vs diurnal records differed signifi-

cantly between species (χ2 = 42.1, d.f. = 1, p <

0.001). Number of records per 2-hour blocks

were negatively correlated between jaguars and

pumas (r = -0.61, p < 0.05). Results were simi-

lar when compared across 1-hour blocks (r =

-0.49, p < 0.05). The coefficient of overlapping

(Δ) indicated some overlap in activity between

the two species (Δ^

= 0.59, 0.49-0.70, 95 % CI)

with data from all years combined but the activ-

ity patterns were, based on the compareCkern

function, significantly different (p < 0.001).

Both species increased diurnal activity some-

what from the first period (2005-2012, jaguars

68 % diurnal, pumas 27 %) to the second period

Table 2

Occurrences of 28 individually identified jaguars at Tiputini Biodiversity Station, Ecuador, from 2014 through 2023.

ID Sex1Number of Photos Number of Trap Sites2Number of Months3First Month Last Month

14M 2 2 36 2/2012 2/2015

2 M 5 3 13 1/2022 2/2023

35F 6 6 72 2/2014 2/2020

46M 5 4 2 1/2022 2/2022

55M 13 10 36 3/2014 3/2016

6 M 2 2 1 1/2022 1/2022

74F 2 3 84 2/2010 2/2016

8 M 1 1 1 1/2023

95M 5 4 1 2/2017 3/2017

10 F 1 1 1 1/2023

11 M 1 1 1 1/2018

125,7 M 7 5 48 3/2016 3/2020

135,7 M 13 6 12 2/2018 2/2019

145,7 F 12 7 24 2/2018 2/2020

155,7 M 3 2 2 1/2018 2/2018

165F 2 2 1 1/2016

175M 5 4 12 1/2019 1/2020

185,7 M 3 3 12 1/2017 1/2018

195F 1 1 1 2/2017

205M 4 3 55 2/2014 9/2018

215M 5 5 24 2/2014 2/2016

225M 1 1 1 2/2016

235,7 ? 3 2 2 1/2018 3/2018

24 ? 1 1 1 1/2019

25 ? 2 2 1 1/2020

265F 2 2 3 1/2020 4/2020

27 M 5 2 2 1/2020 3/2020

285M 5 4 54 9/2018 2/2023

Does not include 4 images that could not be assigned to an individual, typically because only a small part of the animal

was visible. Number of photos refers to images separated by at least 30 min and/or at a different trap site. Trap sites include

those regularly used along trails as well as extras (see text). / 1M = male; F = Female; ? = Unknown. / 2Number of distinct

locations, including cameras located along trails, on plots, and at other locations in the Tiputini area, including sites across

the Tiputini River. / 3Number of months from first to last image. / 4Also photographed in period covered by Blake et al.

(2014). / 5Includes photos from extra locations, not long-term sites along trails. / 6Melanistic individual. / 7Includes photos

from extra locations on the south side of the Tiputini River.

8Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e61782, enero-diciembre 2025 (Publicado Feb. 24, 2025)

(2014-2023, jaguars 82 %, pumas 39 %) but the

change in the distribution of diurnal vs noctur-

nal records was not significant for either species

(jaguar: χ2 = 2.88, d.f. = 1, p = 0.09; puma: χ2 =

1.28, d.f. = 1, p = 0.26). Similarly, the number

of nocturnal and diurnal records still differed

significantly between species (2005-2012, χ2 =

10.91, d.f. = 1, p < 0.001; 2014-2023, χ2 = 31.6,

d.f. = 1, p < 0.001). Coefficients of overlapping

were similar during the first and second periods

(Δ^

= 0.58, 0.40-0.74 CI and Δ^

= 0.55, 0.43-0.67

CI, respectively). In both periods, the activity

patterns were significantly different between

jaguars and pumas (p < 0.002 and p < 0.001, for

the two periods respectively).

Spatial patterns: Number of images per

camera-trap site (n = 10) ranged from one to 15

for jaguars and from one to 19 for pumas (Fig.

4) but number of images per site did not differ

between species (paired t-test, t = 1.41, d.f. = 9,

p = 0.19). Number of images per trap site was

not correlated between jaguars and pumas (r =

0.33, p = 0.35) indicating that the two species

differed in their spatial pattern of occurrence

among camera locations. For example, jag-

uars were most frequently captured at M4200

whereas pumas were more frequent at M2200

(Fig. 4). These two sites are approximately 1.5

km apart (see map in Blake et al., 2016). Three

sites, P150, P1000, P2450, are in a peninsula

of forest and typically had fewer captures of

jaguars when compared to the other seven sites

(t = 3.33, d.f. = 8, p < 0.05), similar to results

from the previous study (i.e., from 2005-2012).

Pumas also were less likely to occur at these

three sites than at other sites, but the difference

was not significant (t = 1.88, d.f. = 8, p = 0.10).

Fig. 2. Numbers of months individual jaguars were

recorded at or near Tiputini Biodiversity Station, Ecuador,

during two different study periods.

Fig. 3. Hourly activity (2-hour periods) of jaguars and

pumas at Tiputini Biodiversity Station, Ecuador, from 2014

to 2023.

Fig. 4. Number (percentage) of images of jaguars and pumas

from 10 Camera-trap Sites within Tiputini Biodiversity

Station, from 2014 to 2023.

DISCUSSION

Long-term studies (i.e., > 10 years) of jag-

uars and other predators of Neotropical forests

are not common (Harmsen et al., 2017). The

current study encompasses 19 years which

allows us to evaluate variation in occurrence

of jaguars and pumas over time. Capture rates

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e61782, enero-diciembre 2025 (Publicado Feb. 24, 2025)

of both species varied across years, with puma

capture rates somewhat more variable (higher

CV), but neither species showed any evidence

of declines; in fact, capture rates increased

slightly over the years for both species. Con-

tinued presence of jaguars and pumas in the

TBS area suggests that their abundance, or at

least activity as indexed by capture rate, may

be relatively stable. This in turn suggests that

human activities in the region have not had a

negative impact on populations of these preda-

tors and that prey abundance has remained at

a high enough level to support them (Espinosa

et al., 2018). In fact, capture rates of many typi-

cal prey items, such as collared peccaries, red

brocket deer, pacas (Cuniculus paca Linnaeus

1766, black agoutis, and nine-banded armadil-

los (Dasypus novemcinctus Linnaeus 1758) but

not white-lipped peccaries or tapirs (Tapirus

terrestris Linaeus 1758), were positively corre-

lated with year (J. G. Blake, unpublished data).

Jaguar capture rates were not significantly cor-

related with any of the prey species whereas

pumas were positively associated with collared

peccaries, red brocket deer, and nine-banded

armadillos (J. G. Blake, unpublished data).

Both jaguars and pumas were recorded at

all camera locations within the station bound-

aries, with no significant differences in overall

capture rates. The two species did, however,

differ somewhat in their activity (number of

records) at specific camera locations. This

might indicate some spatial segregation in

activity. Both species were less likely to be cap-

tured in cameras that were located within an

area of forest bounded closely by the Tiputini

River; a similar result was seen for jaguars in

our earlier study (Blake et al., 2014).

Jaguars are known to be good swimmers

(Emmons, 1987; Da Silveira et al., 2010; Duarte

et al., 2022) and several individuals were record-

ed at sites located on both sides of the Tiputini

River, up to 5 km from camera locations within

the station boundaries. The two sites on the

south side of the river were saladeros (mineral

licks) that attract a variety of species known to

be prey to jaguars (e.g., peccaries, deer; Blake

et al., 2011). Similarly, images of jaguars were

captured at two mineral licks found within

the station boundaries, perhaps because of the

presence of potential prey.

Jaguars and pumas showed a clear sepa-

ration in hourly activity in the current study,

with jaguars primarily active during daylight

hours while pumas were mostly active at night.

This was true both during the first years of the

study (2005-2012) and the latter years (2014-

2023) suggesting that these activity patterns are

relatively consistent over time. This pattern of

activity differs from some previous studies that

found both species to be primarily nocturnal

(Scognamillo et al., 2003) without clear dif-

ferences in hourly activity. Foster et al., (2013)

found that the two species were primarily noc-

turnal and crepuscular in closed, grasslands,

and scrubby forest habitats and were more

diurnal in pantanal. The two species showed

little temporal segregation but did have a sig-

nificant overlap with activity of their main prey.

Similarly, Harmsen et al., (2009) found that

both were predominantly nocturnal with a high

correlation in capture rates at different loca-

tions and significant overlap with their main

prey items (Harmsen et al., 2011). In contrast,

Romero-Muñoz et al. (2010) found that the

two species varied in hourly activity depend-

ing on the study site, showing significant tem-

poral segregation in some dry forest sites but

considerable overlap in others. The extent of

nocturnal vs diurnal activity also varied among

sites (Romero-Muñoz et al., 2010), suggesting

that their behavior was flexible. The temporal

differences in activity seen at TBS might sug-

gest that jaguars and pumas may differ in pri-

mary prey items. Such differences also might

reflect competitive avoidance of jaguars by the

relatively smaller puma at TBS, although more

study is needed to confirm this hypothesis

regarding diet and biotic interactions. Clearly,

hourly activity patterns of the two species can

vary substantially depending on habitat and

study region and likely with the distribution

and abundance of different prey species.

Over the course of ~20 years, ~50 indi-

vidual jaguars have been present at some point

within the boundaries of Tiputini Biodiversity

10 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e61782, enero-diciembre 2025 (Publicado Feb. 24, 2025)

Station. There were 21 individuals identified

from images taken from 2005 to 2012 (Blake et

al., 2014) and 28 from 2014-2023; two of the 28

(one male, one female) from the current period

also were recorded during the first study. In

addition, there were several images during

both sample periods that could not be assigned

to a specific individual, so the actual num-

ber that have occurred at the station is likely

greater than the number identified. That only

two individuals were recorded in both sample

periods indicates substantial turnover in the

identities of jaguars found within the boundar-

ies of the station.

Many individuals were recorded relatively

few times, with 11 during the first period

and 15 during the second period recorded in

only one year, often in only one month. Most

of these individuals likely were simply tran-

sients, passing through Tiputini, or individuals

whose home range lies primarily outside the

station boundary. On the other hand, some

individuals were recorded over many months,

more than 72 (6 years) in some cases, suggest-

ing that they are resident in the area. During

the current study, five males and two females

were recorded over more than three years,

with the two females present in more months

than any of the males. One female, that could

not be identified, was captured on video with

two cubs during April 2015 (D. Mosquera,

unpublished data), the only instance when cubs

were documented. Although not identified,

this likely was one of the resident females and

provides further evidence of the viability of

the population. During the first study, 4 males

and 1 female were recorded over more than 3

years, with two males and one female recorded

over at least 6 years (72-81 months; Blake et

al., 2014). No cubs were photographed during

that period. TBS is only ~670 ha so clearly is

not large enough to encompass the entire home

range of an individual jaguar. Instead, most of

these individuals likely have home ranges that

include some or all of TBS property. Harmsen

et al. (2017), in a 14-year study in Belize, iden-

tified 105 individual jaguars in a study area

of ~100 km2, including 57 males, 31 females,

and 17 whose sex was not determined. They

suggested that individuals recorded over < 3

years should be considered transients with

those recorded over longer periods considered

residents. Their data indicated a maximum age

of 14 years for males and 13 for females. If the

same age structure applies to individuals in

the Tiputini area, this could explain why only

two individuals were recorded during both

sample periods.

Tiputini Biodiversity Station is in the midst

of extensive, relatively undisturbed lowland

forest. Despite its relatively small size, jaguars

and pumas, as well as other predators (e.g.,

ocelots) are regularly present within and near

the station. Results demonstrate the large over-

lapping ranges of jaguars, based on individual

identifications, and the spatial overlap of the

Neotropics largest terrestrial predators, jaguars

and pumas. Nearly 25 % of jaguar individuals

occurred within the station across more than 3

years, some for at least 6 years. Temporal diver-

gence in daily activity patterns, together with

some spatial separation in sites most frequently

encountered, suggest possible behavioral avoid-

ance or differences in diet between the two

species. The continued presence within the

station boundaries of the two largest predators

in Neotropical lowland forest provides a posi-

tive indication of the conservation status and

importance of the region. It suggests that prey

populations (e.g., many ungulates) are suffi-

cient and that human activities have not had, to

date, a large negative impact on either predators

or prey. Whether this will remain true in the

future remains to be seen as climate and other

anthropogenic changes (e.g., deforestation, oil/

gold mining) continue to affect many parts of

the Amazon basin.

Ethical statement: the authors declare that

they all agree with this publication and made

significant contributions; that there is no con-

flict 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

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e61782, enero-diciembre 2025 (Publicado Feb. 24, 2025)

section. A signed document has been filed in

the journal archives.

ACKNOWLEDGMENTS

We thank the staff of the Tiputini Biodiver-

sity Station, especially J. Guerra, C. de Romo, D.

Romo, K. Swing, S. Arroyo, C. Valle and all oth-

ers who have made visits to the site so reward-

ing. We thank two anonymous reviewers whose

comments helped improve this manuscript.

Approval for this research was obtained from

Institutional Animal Care and Use Committee,

University of Florida (IACUC202300000605

and earlier ones). Work at Tiputini Biodi-

versity Station was conducted in accordance

with research permit number MAATE-ARS-

FC-2023-0285 (and earlier ones), Ministerio

del Ambiente, Agua, y Transición Ecológica,

República del Ecuador.

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