Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e2025-702, enero-diciembre 2025 (Publicado Set. 30, 2025)

Top predator feces: behavioral consequences

for coexisting species

Danelly Solalinde1; https://orcid.org/0009-0004-0578-6759

Cuauhtémoc Chávez2*; https://orcid.org/0000-0003-2201-4748

Gerardo Ceballos1; https://orcid.org/0000-0001-8374-2656

Francisco Palomares3; https://orcid.org/0000-0002-4655-7205

1. Wildlife Ecology and Conservation Department, Ecology Institute, National Autonomous University of Mexico,

Mexico City, Mexico; ddsolalinde@gmail.com, gceballo@ecologia.unam.mx

2. Environmental Sciences Department, Metropolitan Autonomous University, Mexico State, Mexico; j.chavez@correo.

ler.uam.mx (*Correspondence)

3. Biology Conservation Department, Doñana Biological Station, CSIC, Seville, Spain; ffpaloma@ebd.csic.es

Received 18-XI-2024. Corrected 11-III-2025. Accepted 18-IX-2025.

ABSTRACT

Introduction: Deposition of feces in the environment is a key behavior in the ecology of predator carnivores,

which promotes behavioral changes in animals, influencing the ecology, abundance, and distribution of the spe-

cies with which they coexist.

Objective: To analyze whether jaguar feces are detected by other jaguars and mammal species and if they have

behavioral effects.

Methods: We recorded the behavior of jaguar (Panthera onca), puma (Puma concolor), mesocarnivores

(Leopardus pardalis, Leopardus wiedii, Herpailurus yagouaroundi), and prey species using camera traps, at

marked sites (with jaguar feces; n = 28) and control sites (without jaguar feces; n = 10). Using the records, we

analyzed, for rainy and dry seasons: (1) the detection of jaguar feces by animals and (2) whether animals modify

their exploration, remarking, vigilance and foraging behavior after detecting jaguar feces.

Results: The detection of jaguar feces by jaguars and pumas was similar and the detection was higher in dry

season. Regarding the behaviors analyzed, jaguars explored 15 times more and remarked 13 more times, marked

sites than control sites. Pumas explored eight times more marked sites than control sites. Mesocarnivores

explored marked sites 17 times more than control sites. Prey explored 41 times more and increased their vigilance

24 times more at marked sites compared to control sites times.

Conclusions: Jaguar feces are detected by conspecifics and other mammal species, and they have behavioral

effects on them. This can trigger changes in the abundance and distribution of populations and may be one of

the ways that large predators shape ecosystems.

Keywords: scent marks; feces detection; animal behavior; top carnivores.

RESUMEN

Heces de depredadores tope: consecuencias conductuales para la coexistencia de las especies

Introducción: La deposición de excremento en el ambiente es un comportamiento clave en la ecología de los

carnívoros depredadores, que promueve cambios conductuales en los animales, influyendo en la abundancia y

distribución de las especies con las que coexisten.

https://doi.org/10.15517/m5j8t121

VERTEBRATE BIOLOGY

2Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e2025-702, enero-diciembre 2025 (Publicado Set. 30, 2025)

INTRODUCTION

Top carnivores influence the behavior of

animals, and it affects the ecology, abundance,

and distribution of the species with which they

coexist (Epperly et al., 2021; Hoeks et al., 2020;

Srivathsa et al., 2023; Wooldridge et al., 2019).

One of the most frequent ways carnivores

modify animal behavior is through their scent

marks, be such as urine, feces, and/or glandular

secretions (Allen, Gunther & Wilmers, 2017;

Rafiq et al., 2020; Sunde et al., 2022). Carni-

vores deposit their scent marks in the environ-

ment, and the presence of these marks has the

potential to modify animal behavior, includ-

ing conspecifics and other species (Cornhill

&Kerley, 2020; Edwards et al., 2022). The scent

marks of carnivores primarily have intraspecific

functions, such as searching for and selecting a

reproductive partner, and defending territories

(Miaretsoa et al., 2022; Morehouse et al., 2021).

But when the scent marks of top predators are

detected by other mammal species, changes or

adjustments in the behavior of these animals

are activated; which can affect from popula-

tions to the functioning of ecosystems (Rafiq et

al., 2020; Sunde et al., 2022).

Scent marks are involved and play a deter-

mining role in the carnivore ecology. Top

carnivores are characterized by being solitary

species, having large territories and tending

to avoid each other most of the time. In this

context, indirect communication between con-

specifics mediated by scent marks is convenient

(Elbroch & Quigley, 2017; Wooldridge et al.,

2019). Scent marks contain information about

identity, sex, health, age, and dominance sta-

tus, and their deposition in the environment

is strategic. Carnivore species deposit their

scent marks in spatio-temporal patterns that

maximize the transmission of the informa-

tion contained in the marks and increase the

probability of being found by other individuals

(Allen, Gunther & Wilmers, 2017).

Deposition of scent marks is a behavior

involved in the entire carnivore ecology and

often has effects on other species (Allen et

al., 2023; Hansen et al., 2024). Scent marks

of top predators have profound consequences

for co-predators and mesocarnivores (Prugh

& Sivy, 2020). Subordinate carnivore species

can detect the predator presence through their

scent marks; when it occurs, avoidance behav-

ior is activated to reduce intraguild predation

(Samuel et al., 2020; Sheriff et al., 2020). Avoid-

ance behavior is associated with changes that

promote spatial, temporal, and dietary niche

Objetivo: Analizar si el excremento de jaguar es detectado por sus conespecíficos y otros mamíferos, y si tiene

efectos en el comportamiento de estos.

Métodos: Registramos la conducta del jaguar (Panthera onca), puma (Puma concolor), mesocarnívoros

(Leopardus pardalis, Leopardus wiedii, Herpailurus yagouaroundi) y presas, mediante cámaras trampa, en sitios

marcados (con excremento de jaguar; n = 28) y sitios control (sin excremento de jaguar; n = 10). Con los regis-

tros analizamos, para lluvias y secas: (1) la detección de los excrementos de jaguar por animales y, (2) si después

de detectar los excrementos de jaguar, los animales modifican su comportamiento de exploración, observación,

vigilancia y forrajeo.

Resultados: Pumas y jaguares detectan el excremento con la misma frecuencia y la detección es mayor en tem-

porada seca. En cuanto a las conductas analizadas, los jaguares exploraron 15 veces más y remarcaron 13 veces

más, los sitios marcados que los sitios control. Los pumas exploraron ocho veces más los sitios marcados que los

sitios control. Los mesocarnívoros exploraron los sitios marcados 17 veces más que los sitios control. Las presas

exploraron 41 veces más y aumentaron su vigilancia 24 veces en los sitios marcados que en los sitios de control.

Conclusiones: El excremento del jaguar es detectado por conespecíficos y otras especies de mamíferos y tiene

efectos en su conducta. Esto puede desencadenar cambios en la abundancia y distribución de las poblaciones y

puede ser una forma en que los carnívoros depredadores moldean los ecosistemas.

Palabras clave: marcas olfativas; detección de heces; conducta animal; carnívoros grandes.

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segregation (Kemna et al., 2020; Russell et al.,

2009; Wooldridge et al., 2019).

Co-predators, mesocarnivores and prey

show specific adaptations that allow them to

detect predators through scent marks and acti-

vate behavioral changes that enable animals to

evade predators. Avoidance behavior involves

restricting activities at safer areas where top

predator scent marks are absent or scarce; gen-

erally, these areas are suboptimal sites, and it is

more difficult for animals to satisfy their needs

(Apfelbach et al., 2005; Sheriff et al., 2020).

This compromises the animal fitness and con-

sequently, their abundance, distribution, and

ecological functions can be affected (Allen et

al., 2024; Wang et al., 2020).

Detecting predator scent marks is essen-

tial for the survival, reproduction and fitness

of animals so they invest time and energy

in detection (Allen et al., 2024; Tallian et al.,

2021). Invest in detecting scent marks of preda-

tors is positively associated with the risk of

predation; when the risk of being predated

is higher, the scent marks detection and the

time of scent marks exploration increases. In

environments with contrasting seasonality, the

risk of predation is higher in the dry season

than rainy season, so in dry season the detec-

tion of scent marks predator by potential prey

increases (Herman & Valone, 2000).

Top predator interactions influence the

abundance and distribution of species, as well

as the structure and functioning of ecosystems

(Estes et al., 2011; Lu et al., 2023; Sommers

& Chesson, 2019). Frequently, their interac-

tions are based on scent marks that have the

potential to modify the behavior of other spe-

cies: co-predators, mesocarnivores and prey.

Consequently, the effects of scent marks on

animal behavior are considered a way through

which top predators affect the abundance and

distribution of the species with which they

coexist (Ripple et al., 2014; van Beeck Calkoen

et al., 2021). However, the mechanism by which

scent marks modify animal behavior has been

studied little. This may be due to the difficulty

of studying large carnivore predators, which

tend to have extensive spatial requirements and

be evasive animals, as well as the complexity

of recording and analyzing animal behavior in

wildlife (Allen, Allen et al., 2017; Allen, Gun-

ther & Wilmers, 2017; Ford, 2017).

The jaguar (Panthera onca) is the largest

feline and the top predator of the Neotropics; the

most frequent route of intraspecific and inter-

specific interactions is the use of scent marks

(Ceballos et al., 2016; Medellín et al., 2016;

Palomares et al., 2018). Despite the relevance of

scent marks in the ecology of this carnivore, the

behavioral effects of scent marks on the behav-

ior of species and their potential implications

have been little explored. We analyzed whether

jaguar feces are detected by conspecifics, co-

predators, mesocarnivores, and prey and the

potential behavioral effect on them.

Our hypothesis is that mammals can detect

the presence of predators though scents marks

and respond by modifying their behavior. Sea-

sonality influences this interaction, with detec-

tion frequency varying between dry and rainy

seasons. We predict that: 1) jaguar conspecifics,

co-predators, mesocarnivores, and prey species

will detect jaguar feces; 2) detection frequency

will be higher during the dry season compared

to the rainy season; 3) animals will modify their

behavior in response to detecting jaguar feces.

The behavioral changes depend on their inter-

action with this carnivore. If jaguar conspecifics

detect jaguar scent marks, we predict that they

increase exploration of feces and show remark-

ing behavior as part of their communication

behavior. If the jaguar scent marks are detected

by individuals of puma (Puma concolor), a co-

predator, competitor, and subordinated species,

or by individuals of mesopredators, that are

also subordinated species with risk of intraguild

predation, we predict only an increase in explo-

ration behavior and no remarking behavior.

Finally, if any prey species detects jaguar scent

marks, we predict an increase in exploration

and vigilance behaviors, and a reduction in

foraging behavior.

4Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e2025-702, enero-diciembre 2025 (Publicado Set. 30, 2025)

MATERIALS AND METHODS

Study area: The study area covers 90 Km2

and includes the Ecological Reserve “El Zapo-

tal” and adjacent areas located in the Northeast-

ern part of the Yucatan Peninsula, Mexico (Fig.

1). El Zapotal is a private reserve acquired for

conservation purposes and managed by a non-

governmental organization (PRONATURA).

The extreme coordinates of the study area are:

upper right point 21°22’55’’ N & 87°31’59’’

W and lower left point 21°16’52.73’’ N &

87°36’48.823’’ W. The bioclimatic system is dry

forest (Holddridge, 1967). The climate is tropi-

cal sub-humid with an annual average tempera-

ture of 25.8 to 26.3 °C and a marked seasonality,

rainy season from June to November and dry

season for the year, from December to May

(Köppen, 1900).

Feces collection: The search for feces was

conducted by slowly walking (2-3 km/hr) along

all the trails in the study area (a trail system

that total 142 km), twice in each season, with

a time interval of 60 days. Feces were searched

through trails because that is where jaguars

and other carnivores frequently deposit them

(Palomares et al., 2018). Each feces found was

georeferenced and for genetic analysis, a small

sample (less than 20 % of the scat) was collected

using latex gloves.

Sample genetic identification: Feces sam-

ples were preserved in silica gel until their

Fig. 1. Study area, in Northern Yucatan Peninsula, Mexico and location of marked (n = 28) and control (n = 10) monitoring

sites through trail system.

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analysis in the laboratory. DNA was extracted

using protocols based on the GuSCN/silica

method (Frantz et al., 2003). Each sample was

genetically identified at the species level using

previously developed species-specific primers,

following the method described by Roques et

al. (2011). The feces were analyzed to determine

sex using the technique described by Pilgrim et

al. (2005). Microsatellite markers were used for

individual genotyping, as described in Roques

et al. (2011). Molecular analysis was under-

taken in the Molecular Ecology Laboratory of

de Doñana Biological Station (LEM-EBD).

Experimental design: Animal behavior

was recorded using camera traps placed at

two types of monitoring stations: (1) marked

sites by jaguar or sites with jaguar feces (n =

28). We identified these sites during the search

for feces along the trails, based on our experi-

ence in identifying tracks, and subsequently we

genetically confirmed that the feces belonged

to jaguar (Roques et al., 2011). The minimum

distance between marked sites was one kilo-

meter, and (2) Control sites or unmarked sites

(without jaguar feces or n = 10). The criteria

for determining these sites were the absence

of feces within a one-kilometer radius (Fig. 1).

We installed one camera trap (Cuddeback

1 279 model, 20 MP with Infrared Flash for

night detection) per monitoring station. At

the marked sites, we focus the cameras on the

feces to observe in detail the behavior of the

animals when they detect it. Cameras traps

were programmed to obtain a 30-second video

at detection; for 25 days following the initial

feces search. Based on the videos obtained and

considering independent records (with a mini-

mum interval of one hour or when different

individuals were recorded), animal behavior

was analyzed.

Data analysis: We analyzed whether jag-

uar feces are detected by: congeners (jaguar;

Panthera onca); competitor species: puma

(Puma concolor); mesocarnivores: ocelot (Leop-

ardus pardalis), margay (Leopardus wiedii) and

jaguarundi (Herpailurus yagouaroundi); and

prey: eight mammal species were considered

as prey, these species represents 83 to 95 % of

jaguar diet in similar sites and close to the study

area: central american red brocket (Mazama

temama), white-tailed deer (Odocoileus virgin-

ianus), white-nosed coati (Nasua narica), low-

land paca (Cuniculus paca), collared peccary

(Tayassu tajacu), nine-banded armadillo (Dasy-

pus novemcinctus), gray fox (Urocyon cinereoar-

genteus), central american agouti (Dasyprocta

punctata; Ávila-Najera et al., 2018).

Feces detection: To determine whether

animals detect jaguar feces, we reviewed vid-

eos from camera traps placed at marked sites,

for both seasons. We calculated the average

number of jaguar feces detection events per day

for each animal group. That is, we divided the

total number of feces detection events in each

camera trap during the sampling period by the

number of sampling days (n = 25). We consid-

ered as detection event the direct sniffing of

feces or approximately one meter away, in both

cases we considered that the sniffing should last

≤ 5 seconds.

We analyzed for each animal group wheth-

er the average number of jaguar feces detection

events per day changes between seasons using

t-student test with independent samples.

Behavioral responses: To determine

whether animals modify their behavior after

detecting jaguar feces, we reviewed the vid-

eos from each camera trap placed at marked

sites by jaguars and unmarked sites. For each

group of animals, in both seasons, we deter-

mined the number of events of four behaviors.

(1) Exploration: any site investigation activ-

ity > 5 seconds, without surveillance status,

with or without movement (Murphy, 1978),

(2) remarking: the activity of depositing feces,

urine or glandular secretions, forming scratches

or scratching surfaces, rubbing some part of

the body or rolling (Allen et al., 2016), (3)

vigilance: when animals carefully observed the

environment in an alert state (Houtman, 2003;

Kimbrell et al., 2007) and (4) foraging: when

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feed consumption was observed (Creel et al.,

2014; Spalinger & Hobbs, 1992).

We calculated, for each group of animals,

the average number of occurrences of each

behavior per day; for this, we divided the total

number of occurrences of each behavior in

each camera trap during the sampling period

by the number of sampling days (n = 25). We

analyzed each animal group if the average num-

ber of occurrences of each behavior per day is

different between marked and unmarked sites

using t-student test with independent samples.

All statistical analysis were fitted with RStudio

4.4.2 program.

RESULTS

We found a total of 93 feces, 80 % belong

to jaguars, 5 % were margay or ocelot, 2 % were

puma and 13 % could not be identified. We

found 1.3 times more jaguar feces in dry season

than in rainy season (42 and 32 feces respec-

tively). The jaguar feces belong to five male

individuals, which deposited their feces during

both seasons.

Feces detection: Jaguar and puma detected

jaguar feces with similar frequency (tjaguar-puma-

detection = -2.0, d.f. = 17, p = 0.05). Moreover,

both felines significantly increased their detec-

tion frequency of feces in dry season compared

to rainy season (tjaguar-season detection = - 2.2, d.f. =

19.1 p < 0.05; tpuma-season detection = - 2.1, d.f. = 17,

p < 0.05; Fig. 2).

Behavioral responses: For jaguars, the

recorded behaviors were exploration and

remarking, both behaviors were observed only

at marked sites. The frequency of exploration

and remarking behaviors of jaguars is different

significantly between marked and unmarked

sites (Student’s t test: tjaguar-exploration = 2.7, d.f.

= 26, p < 0.01; tjaguar-remarking = 3.1, d.f. = 26, p

< 0.01; respectively; Fig. 3). For puma, explo-

ration behavior was significantly higher in

marked sites than in unmarked sites (Student’s

t test: tpuma-exploration = 2.9, d.f. = 8, p < 0.01;

Fig. 2. Number of detections of jaguar feces by group of mammals per day. Light gray = rainy season and dark gray = dry

season. Median; Quartiles; Maximum–Minimum.

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Fig. 3); while remarking behavior does not dif-

fer between sites (Student’s t test: tpuma-remarking

= 1, d.f. = 8, p = 0.3; Fig. 3).

For mesocarnivores, the only behavior

detected was exploration and its frequency was

significantly higher in marked sites by jaguars

than in unmarked sites (Student’s t-test: tmesocar-

nivores-exploration = 2.6, df = 20.9, p < 0.01; Fig. 3).

Prey species significantly increased their explo-

ration and vigilance behaviors in marked sites

by jaguars compared to unmarked sites (Stu-

dent’s t-test; tprey-exploration = 3.8, df = 26, p < 0.01;

tprey-vigilance = 4.2, df = 26, p < 0.01, respectively;

Fig. 3). In contrast, prey foraging behavior had

similar frequency in both sites (Student’s t test;

tprey-foraging = 1.3, df = 4.3, p = 0.2; Fig. 3).

DISCUSSION

We found that all jaguar feces belonged

to male individuals, and that one to four indi-

viduals deposit their feces in the same site. This

could be related to the defense of territories

through scent marks, which is a typical behav-

ior of male carnivores; although this, as in

others researches, opens the question of where

carnivore females deposit their feces (Allen et

al., 2015; Palomares et al., 2018). The deposi-

tion of scent marks in common sites, in some

carnivore species to facilitate communication

between individuals and these sites are often

located on the boundaries of territories to

reduce intraspecific competition (Buesching &

Jordan, 2022; Smith et al., 1989).

Indirectly, the deposition of scents marks

in common places promotes spatial and tem-

poral segregation of individuals, which has

effects on abundance and distribution of lower

trophic levels species (Allen et al., 2015; Allen,

Gunther & Wilmers, 2017; Estes et al., 2011).

It has been observed in carnivore species such

as pumas (Puma concolor), cheetahs (Acinonyx

jubatus) and European badgers (Meles meles)

(Allen et al., 2015; Buesching & Jordan, 2022;

Cornhill & Kerley, 2020). Our results suggest

that jaguar deposit feces strategically to reduce

competition with neighbors. As a predator, this

behavior involves profound ecological effects

and to understand them properly it is necessary

to study these topics carefully at different scales

(Allen et al., 2016; Cornhill & Kerley, 2020).

We observed that (1) jaguar feces are

detected by jaguars, pumas, mesocarnivores

Fig. 3. Behaviors recorded per day in marked (dark grey) and unmarked (light grey) sites by jaguars. Median;

Quartile; Maximum–Minimum.

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and prey; (2) animals modify their behavior

after detecting jaguar feces. Pumas, a co-pred-

ator species, detected jaguar feces and explored

them with similar frequency to jaguars, but we

observed that remarking behavior was different

between both carnivores. Remarking behavior

was exclusive to jaguars; we did not record

remarking behavior in pumas. The behavior

observed in pumas, when they detected jaguar

feces is consistent with what has been reported

in others researches about the dominance of

jaguar over puma (Elbroch & Kusler, 2018;

Harmsen et al., 2009).

Scent marks are useful for co-predator spe-

cies to avoid encounters with dominant species

and reduce the risk of physical confrontations.

Avoiding dominant species promotes niche seg-

regation (spatial, temporal and dietary), which

is a fundamental mechanism for coexistence

of species with similar ecology (Karanth et al.,

2017 Kemna et al., 2020; Müller et al., 2022;

Palomares et al., 2016). We observed that puma

detects and explores jaguar feces; according to

the theory, jaguar feces could allow pumas to

recognize the spatiotemporal presence of the

jaguar, triggering avoidance behaviors. In this

context feces could be considered as an element

that promotes the segregation of these large

felines and facilitates their coexistence.

Intraguild competition has a strong

impact on population dynamics and ecosys-

tem structure (Bai et al., 2021; Davis et al.,

2021). Top predators are responsible for one

third of mesocarnivores mortality, and to avoid

top predators, mesocarnivores have developed

avoidance behaviors that are activated when

they detect predation risk (Allen et al., 2024;

Prugh & Sivy, 2020). Predator scent marks

are interpreted by mesocarnivores as signals

of predation risk, allowing them to detect and

recognize the presence of top predators (Prugh

& Sivy, 2020). Upon finding scent marks of

predators, mesocarnivores modify their behav-

ior to avoid predators, including changes in

activity patterns, spatial movements, home

ranges, foraging behavior and even diet compo-

sition. (Haswell et al., 2018). These behavioral

changes reduce the risk of predation but imply

negative effects on their survival and reproduc-

tion chances, reducing fitness (Ruprecht et al.,

2021; Sheriff et al., 2020).

We observed that mesocarnivores (L.

pardalis, L. wiedii, H. yagouaroundi) detected

and explored jaguar feces, which suggests that

they could detect and recognize the jaguar

presence through their feces, as predicted by

theory. However, after feces detection, we did

not observe changes in their vigilance and

foraging behavior. Nevertheless, it is expected

that changes will occur in behaviors that we did

not evaluate or even some components of vigi-

lance and foraging behavior that were not the

focus of this study. Further research is needed

to understand the effect of jaguar feces on the

behavior and fitness of mesocarnivores. Study-

ing behavioral responses of mesocarnivores to

avoid predation and its consequences is crucial

and the first step to understand the importance

of intraguild predation in ecosystem function-

ing (Haswell et al., 2018).

Prey detected jaguar feces, and it is very

likely that prey perceives it as an indicator of

predation risk, as prey increased their explora-

tion and vigilance in marked sites by jaguars.

Increase exploration and vigilance behavior

helps prey avoid predation but also leads to

changes in animal ecology, such as spatiotem-

poral movements, habitat use, animal nutrition

or reproduction rates, which frequently have

negative effects on animal fitness (Altendorf

et al., 2001; Laundré et al., 2001). Although

we found that foraging frequency was similar

in sites marked by jaguars and control sites,

other components of foraging behavior, such as

search or consumption time, may be affected

by predation pressure. Examining the role of

predator scent marks in predator-prey interac-

tions would be useful to understand how preda-

tors influence patterns of prey abundance and

distribution (Suraci et al., 2016; Taylor, 2013).

Our results are consistent with other

research that show that scent marks of top pred-

ators can be detected by other species, inducing

behavioral changes. We also found that jaguar

feces are detected by 1) puma, the subordinate

sympatric predator, 2) mesocarnivores, and

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e2025-702, enero-diciembre 2025 (Publicado Set. 30, 2025)

3) prey. We found that the detection of jaguar

feces generates behavioral changes, according

to the ecological interaction that each species

has with the jaguar. This study is the first to

analyze the effects of jaguar feces on the behav-

ior of other species. However, more research

is needed to understand the behavioral effects

that triggers by jaguar feces and their implica-

tions at different ecological levels.

The presence of top predators such as the

jaguar causes cascading effects down food webs

potentially critical for conserving ecosystem

function (Berger, 2010; Suraci et al., 2016). Cas-

cading effects are triggered by direct killing of

prey and by modifying the behavior of prey that

perceive their presence (Mpemba et al., 2019;

Schmitz et al., 1997; Schmitz et al., 2008). In

this context, the feces deposition by top preda-

tors such as jaguar and its potential to modify

the behavior of other species have great scope

in terms of conservation.

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.

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