Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e61727, enero-diciembre 2025 (Publicado Ago. 28, 2025)

Foraging patterns and spatial distribution of

synanthropic mammals and their interaction with dogs

Mariano Avendaño-Díaz1; https://orcid.org/0009-0001-5351-1109

Christian Delfín-Alfonso2*; https://orcid.org/0000-0003-3509-4601

Luís García-Feria3; https://orcid.org/0000-0002-0352-1851

Mircea Hidalgo-Mihart4; https://orcid.org/0000-0002-8779-6886

Omar Lagunes-Merino5; https://orcid.org/0000-0002-2933-4544

Jorge E. Morales-Mávil1*; https://orcid.org/0000-0001-9577-0777

1. Instituto de Neuroetología, Universidad Veracruzana, Dr. Luis Castelazo, Xalapa, CP. 91190, Veracruz, México; jormo-

rales@uv.mx (*Correspondence)

2. Instituto de Investigaciones Biológicas, Universidad Veracruzana, Dr. Luis Castelazo, Xalapa, CP. 91190, Veracruz,

México; cdelfin@uv.mx (*Correspondence)

3. Instituto de Ecología, Carretera antigua a Coatepec 351, El Haya, Xalapa, CP 91073, Veracruz, México.

4. Universidad Juárez Autónoma de Tabasco, Universidad s/n, Zona de la Cultura, Villahermosa, CP 86040, Tabasco,

México.

5. Facultad de Bioanálisis, Universidad Veracruzana, Médicos y Odontólogos, Xalapa, CP 91010, Veracruz, México.

Received 02-IX-2024. Corrected 27-II-2025. Accepted 12-VIII-2025.

ABSTRACT

Introduction: Synanthropic mammals benefit from food and shelter provided by green urban areas. However,

they frequently interact with predators such as dogs which may modify certain behaviours and their spatial dis-

tribution, compromising their survival.

Objective: To determine the effect of feral dogs’ presence on mesopredator synanthropic mammal feeding pat-

terns by analyzing the spatial distribution of three species, opossum (Didelphis marsupialis), ringtails (Bassariscus

astutus), and gray fox (Urocyon cinereoargenteus) in a green urban area in Mexico.

Methods: Camera traps and scent stations were used to record foraging patterns and spatial distribution. The

habitat was characterized and correlated with duration and frequencies of synanthropic mammal feeding patterns

and spatial distribution.

Results: Opossums were recorded more frequently inside vegetation compared to other areas of the park, while

dogs were recorded in grass-covered areas, on roads, and in recreation zones. Ringtails and gray foxes were

recorded inside vegetation and in open spaces. Feeding patterns were not affected by the presence or absence of

dogs. However, the presence of dogs affected opossum and gray fox vigilance frequency and was associated with

habitat characteristics. Ringtail vigilance was affected by dog presence and habitat characteristics.

Conclusion: The interaction between dogs and wildlife may generate species-specific behavioural respons-

es, allowing some species to be tolerant of risk while others may show spatial and temporal segregation.

Understanding the spatial distribution of dogs and their effect on wildlife inhabiting green urban areas will help

to improve control-impact programs of dogs, reducing predation events and improving the welfare of meso-

predator synanthropic mammals.

Key words: behaviour; camera traps; exotic predator; feeding habits; mesopredator; predation; scent stations;

vigilance.

https://doi.org/10.15517/rev.biol.trop..v73i1.61727

VERTEBRATE BIOLOGY

2Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e61727, enero-diciembre 2025 (Publicado Ago. 28, 2025)

INTRODUCTION

Animals inhabiting vegetation fragments

in urban and peri-urban areas often must

interact directly or indirectly with humans and

domestic animals and may be affected either

by the transmission of pathogens (Hernández

et al., 2021; Lange et al., 2016) or by predation

(Doherty et al., 2017; Guedes et al., 2021). In

green urban areas, the main potential predators

faced by synanthropic mammals are humans

(Cooper et al., 2008; Ritchie et al., 2013) and

their exotic pets, such as dogs (Canis lupus

familiaris Linnaeus, 1758) and cats (Campos

et al., 2007; Vanak & Gompper, 2009; Vanak

et al., 2013; Wierzbowska et al., 2016). Dogs

are often animals that have escaped or been

abandoned by their owners in vegetation frag-

ments in urban areas and can become feral

animals (Hughes & Macdonald, 2013; Young et

al., 2011). When several dogs are abandoned,

packs are formed that become more danger-

ous for native animals (Hughes & Macdonald,

2013; Mella-Méndez, Flores-Peredo, Pérez-Tor-

res et al., 2019). Similarly, feral dogs compete

for shelter and food with other species with

similar niches (Coronel-Arellano et al., 2021;

Young et al., 2011).

Dogs become a problem for wildlife

because they are important predators for ani-

mals of different taxa (Doherty et al., 2017;

Zamora-Nasca et al., 2021). Dogs may hunt or

harass synanthropic mammals (Villatoro et al.,

2019). Thus, some synanthropic mesopredator

mammals may change their behaviour due to

the presence of dogs (Guedes et al., 2021; Vanak

& Gompper, 2010). One way to explain this

interaction is the so-called the fear landscape

that represents the predation risk that an indi-

vidual perceives in their environment, which

modifies certain behaviours, such as foraging

patterns, feeding, and vigilance (Bedoya-Perez

RESUMEN

Patrones de alimentación y distribución espacial de los mamíferos sinantrópicos

y su interacción con los perros

Introducción: Los mamíferos sinantrópicos se benefician de la comida y el refugio que les proporcionan las áreas

verdes urbanas. Con frecuencia interactúan con depredadores como los perros, que pueden modificar ciertos

comportamientos y su distribución espacial, comprometiendo su supervivencia.

Objetivo: Determinar el efecto de la presencia de perros ferales en los patrones de alimentación de los mamí-

feros mesodepredadores sinantrópicos analizando la distribución espacial de tres especies, zarigüeya (Didelphis

marsupialis), cacomixtle (Bassariscus astutus) y zorra gris (Urocyon cinereoargenteus) en un área verde urbana de

México.

Métodos: Utilizamos cámaras trampa y estaciones de olor para registrar patrones de alimentación y distribución

espacial. Caracterizamos el hábitat y lo correlacionamos con la duración y la frecuencia de los patrones de alimen-

tación de los mamíferos sinantrópicos y la distribución espacial.

Resultados: Las zarigüeyas se registraron con mayor frecuencia dentro de la vegetación en comparación con otras

áreas del parque, mientras que los perros se registraron en áreas cubiertas de pasto, en caminos y en zonas de

recreación. Registramos zorras grises y cacomixtles dentro de la vegetación y en espacios abiertos. Los patrones

de alimentación no se vieron afectados por la presencia o ausencia de perros. Sin embargo, la presencia de perros

afectó la frecuencia de vigilancia de las zarigüeyas y zorras grises y se asoció con las características del hábitat. La

vigilancia de cacomixtles se vio afectada por la presencia de perros y las características del hábitat.

Conclusión: La interacción entre perros y vida silvestre puede generar respuestas conductuales especie-específi-

cas, lo que permite que algunas especies sean tolerantes al riesgo mientras que otras pueden mostrar segregación

espacial y temporal. Comprender la distribución espacial de los perros y su efecto sobre la vida silvestre que habita

en áreas urbanas verdes ayudará a mejorar los programas de control de impacto de los perros, reduciendo los

eventos de depredación y mejorando el bienestar de los mamíferos mesodepredadores sinantrópicos.

Palabras clave: comportamiento; cámaras trampa; especies exóticas; hábitos alimentarios; mesodepredador;

depredación; estaciones de olor; vigilancia.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e61727, enero-diciembre 2025 (Publicado Ago. 28, 2025)

et al., 2013; Laundré et al., 2001; Sinclair & Arc-

ese, 1995; Tolon et al., 2009; van der Merwe &

Brown, 2008). The fear landscape considers the

predator identity (hunting habits) (Altendorf

et al., 2001) and prey (group living, vigilance,

and feeding behaviours) (Altendorf et al., 2001;

Stanford, 2002) both being integrated with

spatial and temporal habitat characteristics,

such as roost availability (Sweitzer, 1996; van

der Merwe & Brown, 2008), lunar light inten-

sity, and seasonal changes (Juliana et al., 2011;

Nowak et al., 2017; Prugh & Golden, 2014) to

obtain a representation of the most and least

risky sites according to prey perception. Thus,

it is possible to determine the decisions that

individuals make in their daily lives to satisfy

their needs and not die trying.

Individuals inhabiting or visiting risky

areas had increased vigilance and decreased

foraging times. Moose (Alces alces) reduce their

feeding time, inhabiting-sympatry with wolves,

resulting in a poor-quality diet, which might

compromise their survival populations in the

long term (Christianson & Creel, 2010; Creel et

al., 2009; Edwards, 1983; Hernández & Laun-

dré, 2005). Likewise, when prey avoid high-risk

sites and migrate to safer sites, they also restrict

their energy requirements; that is, although

they forage longer and keep watch less (Alten-

dorf et al., 2001), these are low quality food sites

(Creel, 2018).

Studies on the fear landscape in synan-

thropic mammals are scarcely relevant when

migration is not an option; that is, green urban

areas (urban parks, urban vegetation fragments,

municipal or state protected areas), unlike nat-

ural areas, are islands surrounded by the urban

matrix, a barrier that increases exposure to syn-

anthropic predators. Therefore, it is important

to understand how synanthropic mesopredator

mammals interact with predators, such as dogs,

and how this interaction might be influenced

by habitat, especially considering that whether

synanthropic mesopredator mammals exhibit

avoidance tactics against dogs has not been

extensively examined (Mitchell & Banks, 2005).

Our aim was to determine the effects of

dogs on synanthropic mesopredator mammal

feeding patterns by analyzing their spatial dis-

tribution, feeding and vigilance behaviours,

and habitat characteristics. We hypothesized

that the riskiest sites, those with the highest dog

presence, might be the ones most avoided by

synanthropic mammals and were thus expect-

ing the following observations at these sites:

1) fewer prey records, 2) longer vigilance than

feeding times, and 3) less vegetation and pro-

tective coverage availability against predators.

MATERIALS AND METHODS

Study area: Our research was carried out

in Natura Park in Xalapa City, Veracruz, Mex-

ico (Fig. 1), which has a 103 ha extension and

is surrounded by an urban matrix. The pre-

dominant vegetation is secondary vegetation

consisting of cloudy and lowland forests, cof-

fee plantations (Coffea arabica L.), and exotic

trees (p. ej. Spathodea campanulate, Jacaranda

mimosifolia, Eriobotrya japonica) (Secretaría de

Medio Ambiente [SEDEMA], 2001). This green

urban area is restricted to visitors with pet dogs;

however, the presence of feral dogs inside the

park has been recorded (Mella-Méndez, Flores-

Peredo, Bolívar-Cimé et al., 2019; Mella-Mén-

dez, Flores-Peredo, Pérez-Torres et al., 2019).

Medium-size wild mammal species previously

recorded in Natura Park are gray fox (Urocyon

cinereoargenteus Schreber, 1775), long-tailed

weasel (Mustela frenata Illiger, 1815), ringtail

(Bassariscus astutus Lichtenstein, 1830), rac-

coon (Procyon lotor Linnaeus, 1758), nine-

banded armadillo (Dasypus novemcinctus

Linnaeus, 1758), common opossum (Didelphis

marsupialis Linnaeus, 1758), Virginia opossum

(Didelphis virginiana Allen, 1900), and gray

four-eyed opossum (Philander opossum Lin-

naeus, 1758) (Mella-Méndez, Flores-Peredo,

Bolívar-Cimé et al., 2019).

Data collection: QGIS (QGIS Develop-

ment Team, 2020) was used to generate ran-

dom camera trap stations separated by 150

m to ensure spatial independence (Ramírez-

Cruz, 2020). We excluded camera trap sta-

tions located on main roads or sites that were

4Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e61727, enero-diciembre 2025 (Publicado Ago. 28, 2025)

highly exposed to avoid stolen camera traps.

We sampled 27 camera trap stations from 15

March to 8 May 2021. Due to the limited num-

ber of camera traps, each station was sampled

for 11 consecutive days, randomly rotating the

cameras until all 27 stations were covered. The

cameras were placed at a height of 50 cm above

the ground and were programmed to take pho-

tographs and video recordings for 30 s. Each

camera trap station had a feeding tray of 21.5 x

16.5 x 3 cm with a mix of 16 pieces of banana

and two boiled eggs, impregnated with vanilla

essence and covered with leaf litter to ensure

foraging behaviours (Bedoya-Perez et al., 2013).

The bait was placed only during the first two

days of each sampling period.

We reduced the sampling bias in open

sites, such as main roads and areas accessible

to visitors (sports fields, playgrounds, parking

areas, and guardhouses), by installing circular

1 m diameter olfactory stations baited with

sardines and vanilla. We reviewed the foot-

prints for the first two days of each sampling

period. The spatial design of the scent stations

Fig. 1. Samplings design of camera traps (yellow squares) and scent stations (red points) at Natura Park (green area), Xalapa,

Veracruz, México. Veracruz State is shown in shaded grey area on the small square. / Fig. 1. Diseño de muestreos de cámaras

trampa (cuadros amarillos) y estaciones de olor (puntos rojos) en el Parque Natura (área verde), Xalapa, Veracruz, México.

El estado de Veracruz se muestra en el área gris sombreada en el cuadrado pequeño.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e61727, enero-diciembre 2025 (Publicado Ago. 28, 2025)

was similar to that of the camera trap stations;

each camera trap station had its own olfactory

station. However, the olfactory stations were

not placed at the same location as the camera

traps, but rather on the roads or trails closest to

the camera trap stations (Fig. 1). To reduce the

spatial independence bias between the records

obtained by camera traps and olfactory stations,

the camera trap stations that were sampled dif-

fered from the olfactory stations during each

sampling period.

We characterized the vegetation surround-

ing each camera trap station using the point-

centered quadrant method (Mueller-Dombois

& Ellenberg, 1976) and selected the tree where

the camera trap was placed as the center. We

measured the nearest shrub and tree distance

(~5 cm in diameter at breast height), her-

baceous cover, and vegetation cover that we

considered as protection against predators. The

latter was estimated using a 1 m ruler marked

with black and white bands every 10 cm; the

ruler was positioned 10 m from the central tree

with the camera trap, and the percentage of

bands not visible or covered by vegetation was

estimated in each quadrant (Griffith & Youtie,

1988). The method used to characterize veg-

etation and cover against predators’ states that

four quadrants allow the variation in vegetation

around the central point to be captured, rather

than focusing on a single direction. We aver-

aged all measurements to avoid pseudo-replicas

and obtained a unique measurement at each

camera trap station.

Synanthropic mammal and dog occu-

pancy models: Single-species occupancy mod-

els were used to model the detection (p) and

occupancy (ψ) probabilities of each synan-

thropic mesopredator. We used the protection

cover against predators as a habitat covariate to

determine if it influenced detection probability,

whereas occupancy probability was modelled

as a function of all habitat covariates and the

presence or absence of feral dogs at each cam-

era trap station. To determine whether the

covariates represented our hypotheses, we gen-

erated null models in which the detection and

occupancy probabilities were not influenced

by any variable. We then compared the null

models and the models that included covariates

using the Akaike information criterion adjusted

for small samples (AICc) (MacKenzie et al.,

2006), and we selected the model that showed

ΔAICc < 2. When we obtained two or more

plausible models (ΔAICc < 2), we averaged

them using the MuMin package (Barton, 2020).

We constructed co-occurrence occupancy

models to determine whether the presence of

dogs affected occupancy probability and syn-

anthropic mammal detection using the wiqid

package (Meredith, 2022). These models esti-

mate occupation probability and species B

detection (subordinate, i.e., mesopredators)

based on the presence or absence of species A

(dominant, i.e., dogs) (Devarajan et al., 2020;

Richmond et al., 2010). We created two models,

the first was an unconditional model (ψBA =

ψBa), where mesopredator occupation is not

influenced by the presence/absence of dogs;

while the second model was conditional (ψBA

≠ ψBa), where mesopredator occupation is

influenced by the presence/absence of dogs.

Subsequently, AICc was used to evaluate the

models, and the model with the lowest value

was selected as the best fit. We used the best

model for estimating the species interaction

factor (SIF), which indicates occurrence proba-

bility of both species at a site compared to what

would be expected if both species occurred

independently (MacKenzie et al., 2006). SIF

values of 1 indicate that both species occur

independently; that is, there is no relationship

in the occupation of both species. SIF values >

1 indicate that the species co-occur more fre-

quently than expected under an independence

hypothesis, suggesting a positive association,

whereas SIF values < 1 indicate that the spe-

cies co-occur less frequently than expected

under an independence hypothesis, suggesting

a negative association, segregation, or evasion

(MacKenzie et al., 2006).

To estimate temporal overlap between

dogs and mesopredators, we used independent

records, considering those records that were

at least one hour apart or when two or more

6Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e61727, enero-diciembre 2025 (Publicado Ago. 28, 2025)

individuals were observed in the same photo-

graph. Delta 1 was used as the overlap estimator

because the species with the lowest number of

records had fewer than 50 independent records

(dogs). In addition, 95 % confidence intervals

were estimated. The overlap package in R soft-

ware (Meredith et al., 2024) was used.

Feeding patterns: We analyzed video

records obtained by the camera traps, and

the foraging duration and vigilance behaviour

were quantified in: 1) foraging, the individual

is observed manipulating, chewing, or con-

suming the food, using/or not using its front

limbs, and, 2) vigilance, the individual is in a

probable alert state, occasionally making lateral

head movements, and 3) rapid vigilance, we

only quantified the frequency, the individual

briefly interrupts its foraging activities and

quickly observes its surroundings. The videos

were analyzed using Boris software (Friard &

Gamba, 2016).

We reduced the bias in record frequency

obtained from camera traps and olfactory sta-

tions (11 and 2 days, respectively) using gener-

alized linear models with binomial distribution

(presence or absence of mesopredator or feral

dogs). We modelled the foraging duration and

vigilance recordings using generalized linear

models with a Gaussian distribution, whereas

the rapid vigilance event frequency was mod-

elled with a Poisson distribution. We incorpo-

rated two-factor interactions into all models

(binomial, poisson and gaussian generalized

linear model) with habitat variables and the

presence/absence of dogs. Statistical analyses

and occupancy models were performed using

R software (R Core Team, 2017), considering a

significance level of < 0.05. The significance of

the variables in the generalized linear models

was determined by the chi-square test.

RESULTS

Only three (opossums, ringtails, and gray

foxes) of the eight previously registered species

of medium-sized mammals were detected by

the trap cameras. Opossums and dogs differed

between camera traps and scent stations, opos-

sums were recorded more frequently by camera

traps (inside the vegetation, X2 = 26.16, df =

1, p < 0.001), while dogs were recorded more

frequently by the scent stations (outside the

vegetation, X2 = 7.28, df = 1, p = 0.006). No sig-

nificant differences were observed for ringtails

(X2 = 3.71, df = 1, p = 0.05), gray foxes (X2 =

0.06, df = 1, p = 0.79), or raccoons (X2 = 0.75, df

= 1, p = 0.38) between camera traps and scent

stations.

Single species and co-occurrence occu-

pancy models: Single species occupancy mod-

els showed that grass cover negatively affected

ringtail occupancy (ψ = -0.34, CI = -1.26-0.56);

however, no variable affected its detection

(Table 1). Considering the gray fox, protec-

tion from predators positively affected gray fox

detection (p = 0.27, CI = -0.54-1.30), while tree

distance positively affected its occupancy (ψ

0.38, CI = -0.54-1.30); whereas the presence of

dogs negatively affected gray fox occupation (ψ

-0.72, CI = -2.64-1.19) (Table 1). Only the pres-

ence of dogs positively affected the opossum

occupation probability (ψ 6.77, CI = -85.94-

98.60) (Table 1). The best-fitting co-occurrence

model was the unconditional model, showing

no interaction between synanthropic mammals

Table 1

Single species occupancy model of synanthropic mammals.

/ Tabla 1. Modelo de ocupación de especies de mamíferos

sinantrópicos.

Model AIC ΔAICc

Ringtail

p (.) ψ (.) 334 0

p (.) ψ (Grass cover) 336.8 1.97

Gray fox

p (.) ψ (.) 189.9 0

p (Protection from predators) ψ (.) 191.6 1.66

p (.) ψ (Tree distance) 191.7 1.78

p (.) ψ (Dogs) 191.9 1.96

Opossum

p (.) ψ (.) 408.4 0

p (.) ψ (Dogs) 410.2 1.84

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e61727, enero-diciembre 2025 (Publicado Ago. 28, 2025)

and dogs (Table 2). All the co-occurrence mod-

els had SIF values of 1.

Analysis of the temporal segregation of

mesopredator species with dogs showed spatial

segregation of the ringtail, grey fox and opos-

sum, with overlap below 0.5 in all three cases

(Fig. 2).

Feeding patterns: The foraging dura-

tion or interaction with habitation variables

were not affected by the presence or absence

of dogs; however, rapid vigilance frequency

showed significant differences with predator

protection and interaction with the presence/

absence of dogs (Table 3). Foraging duration

by gray foxes was not associated with habi-

tat variables or with the presence/absence of

dogs (Table 4). However, vigilance duration

increased at sites with dogs and was positively

associated with grass cover. In addition, there

Table 2

Co-ocurrence occupancy models between synanthropic

mammals and dogs. / Tabla 2. Modelos de ocupación de

co-ocurrencia entre mamíferos sinantrópicos y perros.

Modelo de co-ocurrencia AIC ΔAICc

Ringtail - Dog

Unconditional model: ψ BA = ψ Ba 439.44 0

Conditional model: ψ BA ≠ ψ Ba 442 2.54

Gray fox - Dog

Unconditional model: ψ BA = ψ Ba 294.58 0

Conditional model: ψ BA ≠ ψ Ba 297.40 2.81

Opossum – Dog

Unconditional model: ψ BA = ψ Ba 513.06 0

Conditional model: ψ BA ≠ ψ Ba 515.50 2.43

Table 3

Feeding patterns of opossums (Didelphis spp.) and their interaction with habitat variables and dogs. / Tabla 3. Patrones de

alimentación de zarigüeyas (Didelphis spp.) y su interacción con las variables hábitat y perros.

Habitat variables Estimate Standard error P value

Duration of feeding

Tree distance 0.21 0.88 χ2 = 0.92, gl = 1, p = 0.70

Grass cover -0.04 0.03 χ2 = 3.16, gl = 1, p = 0.49

Protection from predators 0.06 0.02 χ2 = 22.90, gl = 1, p = 0.06

Tree distance x dogs -0.84 3.08 χ2 = 0.03, gl = 1, p = 0.94

Grass cover x dogs 0.06 0.20 χ2 = 5.28, gl = 1, p = 0.37

Protections from predators x dogs -0.11 0.21 χ2 = 1.92, gl = 1, p = 0.59

Duration of vigilance

Tree distance -0.84 1.30 χ2 = 0.92, gl = 1, p = 0.20

Grass cover 0.02 0.05 χ2 = 3.16, gl = 1, p = 0.38

Protection from predators 0.00 0.04 χ2 = 22.90, gl = 1, p = 0.96

Tree distance x dogs 0.12 4.56 χ2 = 0.03, gl = 1, p = 0.74

Grass cover x dogs 0.23 0.30 χ2 = 5.28, gl = 1, p = 0.97

Protections from predators x dogs -0.25 0.31 χ2 = 1.92, gl = 1, p = 0.42

Frequency of fast vigilance

Tree distance -0.19 0.27 χ2 = 0.78, gl = 1, p = 0.37

Grass cover -0.00 0.01 χ2 = 2.44, gl = 1, p = 0.11

Protection from predators 0.01 0.00 χ2 = 2.80, gl = 1, p = 0.09

Tree distance x dogs 0.36 1.10 χ2 = 0.1.29, gl = 1, p = 0.25

Grass cover x dogs 0.18 0.06 χ2 = 0.48, gl = 1, p = 0.48

Protections from predators x dogs -0.17 0.06 χ2 = 26.16, gl = 1, p = 0.00*

*Significance less than 0.05. / *Significancia menor a 0.05.

8Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e61727, enero-diciembre 2025 (Publicado Ago. 28, 2025)

was a positive association with arboreal dis-

tance, which increased substantially at the sites

where dogs were recorded. Rapid vigilance

frequency increased at sites with dogs, whereas

the interaction with tree distance and the pres-

ence/absence of dogs decreased when dogs

were present and increased when dogs were

absent. Likewise, this behaviour showed a posi-

tive association with grass cover, and although

in sites with and without dogs, the positive

association was maintained, it increased sub-

stantially at sites where dogs were recorded.

Fig. 2. Temporal segregation plots are shown for the ringtail (a), grey fox (b) and opossum (c). In all cases the overlap was

less than 0.5. / Fig. 2. Se muestran los gráficos de segregación temporal del cacomixtle (a), zorra gris (b) y zarigüeya (c). En

todos los casos el traslape fue por debajo del 0.5.

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Foraging duration and vigilance as well as the

rapid vigilance frequency of ringtails did not

show significant associations with habitat vari-

ables or their interaction with the presence or

absence of dogs (Table 5).

DISCUSSION

Predation risk in a fear landscape includes

the individual environmental perception by

the prey, as well as predator identification,

which interferes with prey vigilance, feeding

behaviours, and environmental use. Differ-

ences observed in the probability of opossums

occurring between camera traps and olfac-

tory stations may indicate spatial segregation

associated with dogs. In urban reserves, opos-

sums are a frequent prey for dogs (Mella-Mén-

dez, Flores-Peredo, Bolívar-Cimé et al., 2019)

because of their reduced mobility (Arcangeli,

2014; Campos et al., 2007; Carrasco-Román

et al., 2021). During fieldwork, dogs and their

traces (tracks and droppings) were recorded

on trails and main roads, as well as a dead D.

virginiana individual that had multiple bites

caused by dogs, an event that appears to be

frequent at Natura Park (Mella-Méndez, Flores-

Peredo, Bolívar-Cimé et al., 2019), in this case,

this interaction may be more of a competition,

as there was no real predation.. The low occur-

rence of opossums in open areas suggests that

they avoid dogs because this marsupial does not

have efficient escape routes such as nearby trees

or holes, thus representing easy prey for dogs.

However, during night tours, we observed some

opossums using roads and open spaces, which

suggests that these sites necessarily represent

connection zones between vegetation patches

within Natura Park.

Table 4

Feeding patterns of gray foxes (Urocyon cinereoargenteus) and their interaction with habitat variables and dogs. / Tabla 4.

Patrones de alimentación de zorras grises (Urocyon cinereoargenteus) y su interacción con variables de hábitat y perros.

Habitat variables Estimate Standard error P value

Duration of feeding

Tree distance -3.85 4.05 χ2 = 57.13, gl = 1, p = 0.34

Grass cover 0.38 0.55 χ2 = 3.45, gl = 1, p = 0.81

Protection from predators 0.18 0.21 χ2 = 14.78, gl = 1, p = 0.62

Tree distance x dogs -15.27 11.73 χ2 = 115.94, gl = 1, p = 0.17

Grass cover x dogs -0.70 0.70 χ2 = 63.30, gl = 1, p = 0.31

Protections from predators x dogs NA NA NA

Duration of vigilance

Tree distance 0.57 1.17 χ2 = 1.61, gl = 1, p = 0.58

Grass cover 0.04 0.15 χ2 = 26.28, gl = 1, p = 0.02*

Protection from predators 0.09 0.06 χ2 = 9.20, gl = 1, p = 0.19

Tree distance x dogs 20.33 3.40 χ2 = 191.73, gl = 1, p < 0.01

Grass cover x dogs 0.00 20 χ2 = 0.01, gl = 1, p = 0.96

Protections from predators x dogs NA NA NA

Frequency of fast vigilance

Tree distance -0.34 0.28 χ2 = 15.37, gl = 1, p = 0.006*

Grass cover 0.15 0.06 χ2 = 19.42, gl = 1, p < 0.001*

Protection from predators 0.03 0.01 χ2 = 1, gl = 1, p = 0.48

Tree distance x dogs -20.44 5768.21 χ2 = 45.18, gl = 1, p < 0.001*

Grass cover x dogs -0.22 7.96 χ2 = 2.57, gl = 1, p = 0.003*

Protections from predators x dogs NA NA NA

*Significance less than 0.05. / *Significancia menor a 0.05.

10 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e61727, enero-diciembre 2025 (Publicado Ago. 28, 2025)

The absence of differences observed in the

ringtail and gray fox occurrences between trap

cameras and scent stations might reflect similar

ecological preferences because they usually use

open spaces. Carnivores in urban reserves use

vegetation and urban structures for resting and

feeding (Castellanos & List, 2005; Castellanos-

Morales et al., 2008; Castellanos-Morales et

al., 2009; Ramírez-Cruz, 2020). Furthermore,

ringtails and gray foxes use road edges to make

their latrines (Barja & List, 2006; Ramírez-

Cruz, 2020) and feed on anthropogenic waste

in garbage dumps (Castellanos & List, 2005;

Castellanos-Morales et al., 2009). Even dur-

ing fieldwork, we observed both carnivores

walking on the park main roads and trails on

several occasions. We also recorded their feces

on the roadsides, which has been documented

in other studies (Ramírez-Cruz, 2020). Thus,

the presence of both carnivores in the scent

stations might reflect the frequent use of these

open spaces, despite them being spaces with a

predation risk.

The greater occurrence probability of dogs

observed at scent stations associated with roads

and ways has been reported in other studies on

the habitat preferences of this canid (Manjar-

rés-Rodríguez, 2010; Reatiga-Parish, 2015; Tor-

res & Prado, 2010). For example, the presence

of roads was positively associated with canid

detectability in the Alerce Costero National

Park and Valdivian Coastal Reserve in South

America (Silva-Rodríguez & Sieving, 2012).

Similarly, the distance from trails to scent sta-

tions was one of the variables that explained

the potential distribution of dogs in the upper

basin of the Otún River, Colombia (Manjarrés-

Rodríguez, 2010), whereas in reserves in Brazil,

Table 5

Feeding patterns of gray ringtails (Bassariscus astutus) and their interaction with habitat variables and dogs. / Tabla 5.

Patrones de alimentación del cacomixtle (Bassariscus astutus) y su interacción con variables de hábitat y perros.

Habitat variables Estimate Standard error P value

Duration of feeding

Tree distance -1.33 1.82 χ2 = 10.00, gl = 1, p = 0.54

Grass cover -0.03 0.07 χ2 = 12.88, gl = 1, p = 0.49

Protection from predators 0.04 0.06 χ2 = 6.50, gl = 1, p = 0.62

Tree distance x dogs -1.56 7.21 χ2 = 0.23, gl = 1, p = 0.92

Grass cover x dogs -0.11 0.44 χ2 = 17.20, gl = 1, p = 0.42

Protections from predators x dogs -0.01 0.44 χ2 = 0.03, gl = 1, p = 0.97

Duration of vigilance

Tree distance 0.68 0.63 χ2 = 0.19, gl = 1, p = 0.80

Grass cover 0.01 0.02 χ2 = 2.10, gl = 1, p = 0.42

Protection from predators 0.02 0.02 χ2 = 1.78, gl = 1, p = 0.46

Tree distance x dogs 0.03 2.51 χ2 = 1.73, gl = 1, p = 0.46

Grass cover x dogs 0.09 0.15 χ2 = 5.05, gl = 1, p = 0.21

Protections from predators x dogs -0.15 0.15 χ2 = 3.42, gl = 1, p = 0.30

Frequency of fast vigilance

Tree distance 0.32 0.27 χ2 = 0.45, gl = 1, p = 0.50

Grass cover 0.00 0.01 χ2 = 0.17, gl = 1, p = 0.67

Protection from predators 0.01 0.01 χ2 = 0.16, gl = 1, p = 0.68

Tree distance x dogs -1.12 1.15 χ2 = 0.19, gl = 1, p = 0.66

Grass cover x dogs 0.06 0.06 χ2 = 2.57, gl = 1, p = 0.10

Protections from predators x dogs -0.10 0.07 χ2 = 2.19, gl = 1, p = 0.13

*Significance less than 0.05. / *Significancia menor a 0.05.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e61727, enero-diciembre 2025 (Publicado Ago. 28, 2025)

a higher frequency of dogs was documented

outside secondary forest patches (Torres &

Prado, 2010).

Interaction between synanthropic mam-

mals and dogs: We hypothesized that syn-

anthropic mammals would show a positive

association with vegetation, providing protec-

tion from predators (Bonnot et al., 2013; Creel

et al., 2005; Donadio & Buskirk, 2016). Simi-

larly, we expected spatial segregation in dogs,

which has been documented (Silva-Rodríguez

& Sieving, 2012; Silva-Rodríguez et al., 2010;

Vanak & Gompper, 2010); however, this only

occurred in opossums, suggesting species-spe-

cific responses.

The low spatial segregation between ring-

tails and dogs in our study may be because

we recorded dogs during the day, while we

recorded ringtails at night, at twilight, and only

one recording during the day. This difference

in activity period has also been documented in

other urban reserves. In the Pedregal San Ángel

Ecological Reserve, Mexico City, Ramírez-Cruz

(2020) suggested that dogs accompanied by

their owners did not affect ringtail spatial dis-

tribution, although they frequently occurred

in the same places because of the daytime

and nocturnal activities of dogs and ringtails,

respectively. Likewise, the negative association

of ringtails with herbaceous cover has already

been registered previously, which responds to

their preference for open spaces or conspicuous

sites, which are used for depositing their feces

and for communication between conspecifics

(Ramírez-Cruz, 2020).

The spatial segregation observed between

gray foxes and dogs is consistent with other

studies that found that dogs displace smaller

canids (Vanak et al., 2013; Vanak & Gompper,

2009; Vanak & Gompper, 2010). Gray foxes are

prey that occasionally appear in the dog’s diet

(Carrasco-Román et al., 2021), thus safer area

use, in addition to vegetation structure, given

the positive association with protection against

predators, are factors that reduce predation

risk. Protection against predators represents

the interaction between herbaceous, shrubby,

and tree cover, which together comprise the

sites with the greatest amount of vegetation

that are frequently used by gray foxes in natural

and urban ecosystems (Castellanos-Morales et

al., 2009; Servín et al., 2014), both for shelter

and food. The positive association with tree

distance may reflect the use of open sites, such

as roads, grasslands, and some buildings within

the park. In urban reserves, foxes sometimes

tend to use open spaces more than expected

(Castellanos-Morales et al., 2008; Castellanos-

Morales et al., 2009; Rountree, 2004), which

may be because of the benefits in bringing

them closer to human activity, even when

observed in the periphery of some buildings

(Rountree, 2004).

Opossums are synanthropic mammals

with the highest mortality associated with dogs

within urban reserves; therefore, we expected

a greater occupation of sites with vegetation

cover, which was not recorded in our study.

Similar results were observed in D. virginiana

inhabiting urbanized landscapes in the Chi-

cago, Illinois, U.S.A. metropolitan area, because

this marsupial did not show a specific associa-

tion with vegetation cover (Fidino et al., 2016),

which may be due to its ability to occupy a

wide variety of habitats (Cruz-Salazar et al.,

2016). We believe that the positive association

between opossum occupation and the presence

of dogs does not correspond to direct interac-

tion; instead, it might reflect the use of open

spaces by opossums and dogs, mainly because

of high food supply availability at those sites.

Feeding habits: We expected that vigilance

times would be greater at sites where dogs

were present, and feeding would be reduced,

which was not observed with the opossum. Our

results coincide with those of some studies that

have quantified the foraging and vigilance times

of marsupials living in sympatry with dogs, in

which no effects were observed on either behav-

iour (Cortés-Alfonso et al., 2021; López-Barra-

gan & Sánchez, 2017; Rodríguez-Matla, 2016).

It has been suggested that dogs do not represent

a potential threat to these marsupials (Cortés-

Alfonso et al., 2021); however, based on the

12 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e61727, enero-diciembre 2025 (Publicado Ago. 28, 2025)

spatial segregation observed between camera

traps and scent stations, it would be convenient

to explore foraging and vigilance behaviours in

open spaces such as roads and trails, which are

sites with a higher frequency of dogs, because

we only observed these behaviours within veg-

etation patches using camera traps.

The positive association between the fast

vigilance frequency and arboreal distance

observed in opossums represents a behaviour

that does not compromise daily energy require-

ments while reducing predation risk in open

spaces. We believe that rapid vigilance events,

in addition to reducing predation risk within

vegetation patches, also help reduce encounters

with other mesopredators at foraging stations.

Foraging was the main behaviour in which

opossums spent most of their time at the sta-

tions, a behaviour mostly recorded in our study

area in previous research (Rodríguez-Matla,

2016). One aspect to highlight is that opos-

sums were also vigilant during feeding states;

that is, both behaviours were not temporarily

excluded, which allowed them to obtain the

maximum nutritional benefit inside the vegeta-

tion patch while reducing predation risk.

Regarding ringtail and gray fox foraging

patterns, Rodríguez-Matla (2016) documented

in sites impregnated with dog urine, a decrease

in feeding times and increase in ringtail deter-

rence, whereas our study did not show any

association with dogs and vegetation structure.

When prey detect clues left by their predators,

such as feces or urine, they may modify their

foraging patterns (Kats & Dill, 1998; Rosell

& Czech, 2000) or even remain unchanged

(Rosell & Czech, 2000). A plausible explana-

tion for our results could be the low territorial

marking frequency of dogs during their walk-

ing through the feeding stations, which could

be true because in the video recordings, we

scarcely observed individuals marking the site.

Furthermore, most of the individuals spent

very little time at the feeding stations, suggest-

ing that vegetation usage might correspond to

transit areas.

The interaction between dogs and wild-

life may generate species-specific behavioural

responses, allowing some species to be tolerant

to risk while others may show spatial and tem-

poral segregation. Our study analyses aspects

of feral dogs and synantropic mammals and

we suggest that in Natura Park it is necessary

to implement protocols with implementing a

specific action to control feral fauna consider-

ing that this is a protected area.

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.

ACKNOWLEDGMENTS

We thank the CONACyT for the financial

support of the “Proyecto Ciencia de Frontera”

clave 64358 “Funcionalidad Socio-Ecológica de

Áreas Verdes Urbanas Neotropicales” and the

SEDEMA for the permits granted to work in

the Natura Park. We thank Fernando Martínez

Barradas, Pedro Pareja Badillo, Saul Hernán-

dez Carmona and volunteers during the field

work. We thank Gabriel Andrade Ponce for

assisting with occupancy models. This research

was funded by CONAHCyT, scholarship num-

ber 785744 (MAD). Permission for this study

was provided by the Federal Government

of Mexico’s Secretariat of Environment and

Natural Resources (SEMARNAT No. SGPA/

DGVS/02771/21).

REFERENCES

Altendorf, K. B., Laundré, J. W., López-González, C. A.,

& Brown, J. S. (2001). Assessing effects of preda-

tion risk on foraging behavior of mule deer. Jour-

nal of Mammalogy, 82(2), 430–439. https://doi.

org/10.1644/1545-1542(2001)082%3C0430:AEOPRO

%3E2.0.CO;2

Arcangeli, J. (2014). Manejo de crías de zarigüeya (Didel-

phis virginiana) en cautiverio. Revista Electrónica de

Veterinaria, 15(9), 1–13.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e61727, enero-diciembre 2025 (Publicado Ago. 28, 2025)

Barja, I., & List, R. (2006). Faecal marking behaviour

in ringtails (Bassariscus astutus) during the non-

breeding period: spatial characteristics of latrines and

single faeces. Chemoecology, 16, 219–222. https://doi.

org/10.1007/s00049-006-0352-x

Barton, K. (2020). MuMIn: Multi-Model Inference (Ver-

sion 1.47.5) [Computer software]. CRAN. https://

CRAN.R-project.org/package=MuMIn

Bedoya-Pérez, M. A., Carthey, A. J. R., Mella, V. S. A.,

McArthur, C., & Banks, P. B. (2013). A practical guide

to avoid giving up on giving-up densities. Behavioral

Ecology and Sociobiology, 67, 1541–1553. https://doi.

org/10.1007/s00265-013-1609-3

Bonnot, N., Morellet, N., Verheyden, H., Cargnelutti, B.,

Lourtet, B., Klein, F., & Hewison, A. J. M. (2013).

Habitat use under predation risk: Hunting, roads and

human dwellings influence the spatial behaviour of

roe deer. European Journal of Wildlife Research, 59,

185–193. https://doi.org/10.1007/s10344-012-0665-8

Campos, C. B., Esteves, C. F., Ferraz, K. M. P. M. B.,

Crawshaw Jr., P. G. C., & Verdade, L. M. (2007).

Diet of free-ranging cats and dogs in a subur-

ban and rural environment, south-eastern Brazil.

Journal of Zoology, 273(14), 14–20. https://doi.

org/10.1111/j.1469-7998.2007.00291.x

Carrasco-Román, E., Medina, J. P., Salgado-Miranda, C.,

Soriano-Vargas, E., & Sánchez-Jasso, J. M. (2021).

Contributions on the diet of free-ranging dogs (Canis

lupus familiaris) in the Nevado de Toluca Flora and

Fauna Protection Area, Estado de México, Mexico.

Revista Mexicana de Biodiversidad, 92, 1–11. https://

doi.org/10.22201/ib.20078706e.2021.92.3495

Castellanos-Morales, G., García-Peña, N., & List, R. (2008).

Uso de recursos del cacomixtle (Bassariscus astutus) y

la zorra gris (Urocyon cinereoargenteus) en una reser-

va urbana de la Ciudad de México. En L. Consuelo, E.

Espinoza, & J. Ortega (Eds.), Avances en el estudio de

los mamíferos (II, pp. 377–390). Asociación Mexicana

de Mastozoología, A. C.

Castellanos-Morales, G., García-Peña, N., & List, R. (2009).

Ecología del cacomixtle (Bassariscus astutus) y la

zorra gris (Urocyon cinereoargenteus). En A. Lot & Z.

Canos-Santana (Eds.), Biodiversidad del ecosistema

del Pedregal de San Ángel (pp. 371–381). Universidad

Nacional Autónoma de México (UNAM).

Castellanos, G., & List, R. (2005). Área de actividad y uso

de hábitat del cacomixtle (Bassariscus astutus) en “El

Pedregal de San Ángel”. Revista Mexicana de Masto-

zoología (Nueva época), 9(1), 113–122. https://doi.

org/10.22201/ie.20074484e.2005.9.1.365

Christianson, D., & Creel, S. (2010). A nutritionally medi-

ated risk effect of wolves on elk. Ecology, 91(4),

1184–1191. https://doi.org/10.1890/09-0221.1

Cooper, C. A., Neff, A. J., Poon, D. P., & Smith, G. R. (2008).

Behavioral responses of Eastern Gray Squirrels in

suburban habitats differing in human activity levels.

Northeastern Naturalist, 15(4), 619–625. https://www.

jstor.org/stable/25177145

Coronel-Arellano, H., Rocha-Ortega, M., Gual-Sill, F.,

Martínez-Meyer, E., Ramos-Rendón, A. K., González-

Negrete, M., Gil-Alarcón, G., & Zambrano, L. (2021).

Raining feral cats and dogs? Implications for the

conservation of medium-sized wild mammals in

an urban protected area. Urban Ecosystems, 24(1),

83–94. https://doi.org/10.1007/s11252-020-00991-7

Cortés-Alfonso, D., Valenzuela, A. E. J., & Sánchez-Barrera,

F. A. (2021). Percepción del riesgo de depredación y

uso de hábitat del fara Didelphis pernigra (Didelphi-

morphia: Didelphidae) en un área exurbana andina.

Caldasia, 43(2), 331–342. https://doi.org/10.15446/

caldasia.v43n2.84851

Creel, S. (2018). The control of risk hypothesis: reactive vs.

proactive antipredator responses and stress-mediated

vs. food-mediated costs of response. Ecology Letters,

21(7), 947–956. https://doi.org/10.1111/ele.12975

Creel, S., Winnie Jr., J., Maxwell, B., Hamlin, K., & Creel,

M. (2005). Elk alter habitat selection as an antipreda-

tor response to wolves. Ecology, 86(12), 3387–3397.

https://doi.org/10.1890/05-0032

Creel, S., Winnie, J. A., & Christianson, D. (2009). Gluco-

corticoid stress hormones and the effect of predation

risk on elk reproduction. Proceedings of the National

Academy of Sciences of the United States of Ameri-

ca, 106(30), 12388–12393. https://doi.org/10.1073/

pnas.0902235106

Cruz-Salazar, B., Ruiz-Montoya, L., Navarrete-Gutiérrez,

D., & Vázquez, L. B. (2016). Influence of the com-

position and structure of modified landscapes on

abundance of two marsupials during the dry sea-

son. Therya, 7(3), 393–406. https://doi.org/10.12933/

therya-16-401

Devarajan, K., Morelli, T. L., & Tenan, S. (2020). Multi-

species occupancy models: review, roadmap, and

recommendations. Ecography, 43, 1612–1624. https://

doi.org/10.1111/ecog.04957

Doherty, T. S., Dickman, C. R., Glen, A. S., Newsome, T.

M., Nimmo, D. G., Ritchie, E. G., Vanak, A. T., &

Wirsing, A. J. (2017). The global impacts of domestic

dogs on threatened vertebrates. Biological Conserva-

tion, 210(Part A), 56–59. https://doi.org/10.1016/j.

biocon.2017.04.007

Donadio, E., & Buskirk, S. W. (2016). Linking predation

risk, ungulate antipredator responses, and patterns of

vegetation in the high Andes. Journal of Mammalogy,

97(3), 966–977. https://doi.org/10.1093/jmammal/

gyw020

14 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e61727, enero-diciembre 2025 (Publicado Ago. 28, 2025)

Edwards, J. (1983). Diet shifts in moose due to preda-

tor avoidance. Oecologia, 60, 185–189. https://doi.

org/10.1007/BF00379520

Fidino, M. A., Lehrer, E. W., & Magle, S. B. (2016).

Habitat dynamics of the Virginia Opossum in a

highly urban landscape. The American Mid-

land Naturalist, 175(2), 155–167. https://doi.

org/10.1674/0003-0031-175.2.155

Friard, O., & Gamba, M. (2016). BORIS: a free, versa-

tile open-source event-logging software for video/

audio coding and live observations. Methods in Ecol-

ogy and Evolution, 7(11), 1325–1330. https://doi.

org/10.1111/2041-210X.12584

Griffith, B., & Youtie, B. A. (1988). Two devices for estimat-

ing forage density and deer hiding cover. Wildlife

Society Bulletin, 16(4), 206–210.

Guedes, J. J. M., Assis, C. L., Feio, R. N., & Quintela, F. M.

(2021). The impacts of domestic dogs (Canis familia-

ris) on wildlife in two Brazilian hotspots and impli-

cations for conservation. Animal Biodiversity and

Conservation, 44(1), 45–58. https://doi.org/10.32800/

abc.2021.44.0045

Hernández, F. A., Manqui, J., Mejías, C., & Acosta-Jamett,

G. (2021). Domestic dogs and wild foxes interactions

in a wildlife-domestic interface of North-Central

Chile: implications for multi-host pathogen trans-

mission. Frontiers in Veterinary Science, 8, 631788.

https://doi.org/10.3389/fvets.2021.631788

Hernández, L., & Laundré, J. W. (2005). Foraging in the

landscape of fear and its implications for habitat

use and diet quality of elk Cervus elaphus and bison

Bison bison. Wildlife Biology, 11(3), 215–220. https://

doi.org/10.2981/0909-6396(2005)11[215:FITLOF]2.0

.CO;2

Hughes, J., & Macdonald, D. W. (2013). A review of the

interactions between free-roaming domestic dogs and

wildlife. Biology Conservation, 157, 341–351. https://

doi.org/10.1016/j.biocon.2012.07.005

Juliana, J. R. S., Kotler, B. P., Brown, J. S., Mukherjee, S.,

& Bouskila, A. (2011). The foraging response of

gerbils to a gradient of owl numbers. Evolutionary

Ecology Research, 13, 869–878. https://hdl.handle.

net/10027/9673

Kats, L. B., & Dill, L. M. (1998). The scent of death: chemo-

sensory assessment of predation risk by prey animals.

Écoscience, 5(3), 361–394. https://doi.org/10.1080/11

956860.1998.11682468

Lange, M., Kramer-Schadt, S., & Thulke, H. H. (2016).

Relevance of indirect transmission for wildlife disease

surveillance. Frontiers in Veterinary Science, 3, 110.

https://doi.org/10.3389/fvets.2016.00110

Laundré, J. W., Hernández, L., & Altendorf, K. B. (2001).

Wolves, elk, and bison: reestablishing the “landscape

of fear” in Yellowstone National Park, U.S.A. Cana-

dian Journal of Zoology, 79(8), 1401–1409. https://doi.

org/10.1139/z01-094

López-Barragán, C. N., & Sánchez, F. (2017). Food selection

and predation risk in the Andean white-eared opos-

sum (Didelphis pernigra Allen, 1900) in a suburban

area of Bogotá, Colombia. Mammalian Biology, 86,

79–83. https://doi.org/10.1016/j.mambio.2017.07.001

MacKenzie, D. I., Nichols, J. D., Royle, J. A., Pollock, K. H.,

Bailey, L., & Hines, J. E. (2006). Occupancy estimation

and modeling: Inferring patterns and dynamics of spe-

cies occurrence (2nd ed.). Academic Press.

Manjarrés-Rodríguez, T. S. (2015). Distribución y uso de

hábitat del perro (Canis familiaris) en la cuenca alta

del río Otún (Risaralda-Colombia) [Tesis de mae-

stría, Pontificia Universidad Javeriana]. Repositorio

Institucional PUJ. https://repository.javeriana.edu.co/

items/8aec4062-8ba5-4f9c-b390-aa5a37f98a20

Mella-Méndez, I., Flores-Peredo, R., Bolívar-Cimé, B., &

Vázquez-Domínguez, G. (2019). Effect of free-rang-

ing dogs and cats on medium-sized wild mammal

assemblages in urban protected areas of a Mexican

city. Wildlife Research, 46(8), 669–678. https://doi.

org/10.1071/WR19074

Mella-Méndez, I., Flores-Peredo, R., Pérez-Torres, J.,

Hernández-González, S., González-Uribe, D. U., &

Bolívar-Cimé, B. S. (2019). Activity patterns and

temporal niche partitioning of dogs and medium-

sized wild mammals in urban parks of Xalapa, Mex-

ico. Urban Ecosystems, 22, 1061–1070. https://doi.

org/10.1007/s11252-019-00878-2

Meredith, M. (2022). Wiqid package: Quick and dirty

estimates for wildlife populations. R package (Ver-

sion 0.3.3) [Computer software]. CRAN. https://doi.

org/10.32614/CRAN.package.wiqid

Meredith, M., Ridout, M., & Campbell, L. A. D. (2024).

Overlap: Estimates of coefficient of overlapping for ani-

mal activity patterns. R package (Version 0.3.9) [Com-

puter software]. CRAN. https://doi.org/10.32614/

CRAN.package.overlap

Mitchell, B. D., & Banks, P. B. (2005). Do wild dogs exclude

foxes? Evidence for competition from dietary and

spatial overlaps. Austral Ecology, 30(5), 581–591.

https://doi.org/10.1111/j.1442-9993.2005.01473.x

Mueller-Dombois, D., & Ellenberg, H. (1976). Aims and

method of vegetation ecology. Geographical Review,

66(1), 114–116. https://doi.org/10.2307/213332

Nowak, K., Wimberger, K., Richards, S. A., Hill, R. A., &

le Roux, A. (2017). Samango monkeys (Cercopithecus

albogularis labiatus) manage risk in a highly seasonal,

human-modified landscape in Amathole Mountains,

South Africa. International Journal of Primatology, 38,

194–206. https://doi.org/10.1007/s10764-016-9913-1

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e61727, enero-diciembre 2025 (Publicado Ago. 28, 2025)

Prugh, L. R., & Golden, C. D. (2014). Does moonlight

increase predation risk? Meta-analysis reveals diver-

gent responses of nocturnal mammals to lunar cycles.

Journal of Animal Ecology, 83(2), 504–514. https://doi.

org/10.1111/1365-2656.12148

QGIS Development Team. (2020). QGIS Geographic Infor-

mation System (Version 3.4) [Computer software].

Open Source Geospatial Foundation Project. http://

qgis.org

R Core Team. (2017). R: A language and environment

for statistical computing (Version 3.3-4) [Computer

software]. R Foundation for Statistical Computing.

https://www.r-project.org/

Ramírez-Cruz, G. A. (2020). Analysis of the effect of

recreational dog walking on the occupancy prob-

ability of the ringtail Bassariscus astutus (Carnivora:

Procyonidae) within an urban ecosystem. Urban

Ecosystems, 23, 107–115. https://doi.org/10.1007/

s11252-019-00922-1

Reatiga-Parrish, J. F. (2015). Determinación del efecto de per-

ros ferales (Canis lupus familiaris) sobre los mamíferos

del Parque Nacional Natural Chingaza, mediante foto-

trampeo [Tesis de Licenciatura, Pontificia Universidad

Javeriana]. Repositorio de la Pontificia Universidad

Javeriana. http://hdl.handle.net/10554/17913

Richmond, O. M., Hines, J. E., & Beissinger, S. R. (2010).

Two-species occupancy models: A new parameter-

ization applied to co-occurrence of secretive rails.

Ecological Applications, 20(7), 2036–2046. https://doi.

org/10.1890/09-0470.1

Ritchie, E. G., Dickman, C. R., Letnic, M., & Vanak, A. T.

(2013). Dogs as predators and trophic regulators. In

M. E. Gompper (Ed.), Free‐ranging dogs and wildlife

conservation (pp. 55–68). Oxford University Press.

Rodríguez-Matla, M. (2016). Efecto de la orina de perros en

la conducta de alimentación de mamíferos medianos

silvestres en parques ecológicos urbanos de Xalapa,

Veracruz, México [Tesis de licenciatura no publicada].

Universidad Veracruzana.

Rosell, F., & Czech, A. (2000). Responses of foraging

eurasian beavers Castor fiber to predator odours.

Wildlife Biology, 6(1), 13–21. https://doi.org/10.2981/

wlb.2000.033

Rountree, G. H. (2004). Comparative study of the home

range and habitat usage of red foxes and gray foxes

in an urban setting: A preliminary report [Confer-

ence presentation]. In W. W. Shaw, L. K. Harris, & L.

VanDruff (Eds.), Proceedings of the 4th International

Urban Wildlife Symposium (pp. 238–244). 4th Inter-

national Urban Wildlife Symposium, Tucson, AZ,

United States.

Secretaría de Medio Ambiente. (2001). Área natural prote-

gida El Tejar-Garnica: Programa de manejo. SEDE-

MA, Gobierno del Estado de Veracruz, México.

Servín, J., Bejarano, A., Alonso-Pérez, N., & Chacón, E.

(2014). El tamaño del ámbito hogareño y el uso

de hábitat de la zorra gris (Urocyon cinereoargen-

teus) en un bosque templado de Durango, México.

Therya, 5(1), 257–269. https://doi.org/10.12933/

therya-14-174

Silva-Rodríguez, E. A., & Sieving, K. E. (2012). Domes-

tic dogs shape the landscape-scale distribution of

a threatened forest ungulate. Biological Conserva-

tion, 150(1), 103–110. https://doi.org/10.1016/j.

biocon.2012.03.008

Silva-Rodríguez, E. A., Ortega-Solís, G. R., & Jiménez, J.

E. (2010). Conservation and ecological implications

of the use of space by chilla foxes and free-ranging

dogs in a human-dominated landscape in southern

Chile. Austral Ecology, 35(7), 765–777. https://doi.

org/10.1111/j.1442-9993.2009.02083.x

Sinclair, A. R., & Arcese, P. (1995). Population conse-

quences of predation-sensitive foraging: the Seren-

geti wildebeest. Ecology, 76(3), 882–891. https://doi.

org/10.2307/1939353

Stanford, C. B. (2002). Avoiding predators: expectations

and evidence in primate antipredator behaviour.

International Journal of Primatology, 23, 741–757.

https://doi.org/10.1023/A:1015572814388

Sweitzer, R. A. (1996). Predation or starvation: consequenc-

es of foraging decisions by porcupines (Erethizon

dorsatum). Journal of Mammalogy, 77(4), 1068–1077.

https://doi.org/10.2307/1382787

Tolon, V., Dray, S., Loison, A., Zeileis, A., Fischer, C., &

Baubet, E. (2009). Responding to spatial and tempo-

ral variations in predation risk: space use of a game

species in a changing landscape of fear. Canadian

Journal of Zoology, 87(12), 1129–1137. https://doi.

org/10.1139/Z09-101

Torres, P. C., & Prado, P. I. (2010). Domestic dogs in a

fragmented landscape in the Brazilian Atlantic Forest:

abundance, habitat use and caring by owners. Brazil-

ian Journal of Biology, 70(4), 987–994. https://doi.

org/10.1590/S1519-69842010000500010

van der Merwe, M., & Brown, J. S. (2008). Mapping the

landscape of fear of the cape ground squirrel (Xerus

inauris). Journal of Mammalogy, 89(5), 1162–1169.

https://doi.org/10.1644/08-MAMM-A-035.1

Vanak, A. T., & Gompper, M. E. (2009). Dogs Canis famil-

iaris as carnivores: Their role and function in intra-

guild competition. Mammal Review, 39(4), 265–283.

https://doi.org/10.1111/j.1365-2907.2009.00148.x

Vanak, A. T., & Gompper, M. E. (2010). Interference

competition at the landscape level: the effect of free-

ranging dogs on a native mesocarnivore. Journal

of Applied Ecology, 47(6), 1225–1232. https://doi.

org/10.1111/j.1365-2664.2010.01870.x

16 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e61727, enero-diciembre 2025 (Publicado Ago. 28, 2025)

Vanak, A. T., Dickman, C. R., Silva-Rodriguez, E. A., But-

ler, J. R. A., & Ritchie, E. G. (2013). Top-dogs and

under-dogs: competition between dogs and sympatric

carnivores. In M. E. Gompper (Ed.), Free-Ranging

Dogs and Wildlife Conservation (pp. 69–93). Oxford

University Press.

Villatoro, F. J., Naughton-Treves, L., Sepúlveda, M. A.,

Stowhas, P., Mardones, F. O., & Silva-Rodríguez, E.

A. (2019). When free-ranging dogs threaten wild-

life: public attitudes toward management strategies

in southern Chile. Journal of Environmental Man-

agement, 229(1), 67–75. https://doi.org/10.1016/j.

jenvman.2018.06.035

Wierzbowska, I. A., Hędrzak, M., Popczyk, B., Okarma,

H., & Crooks, K. R. (2016). Predation of wildlife by

free-ranging domestic dogs in Polish hunting grounds

and potential competition with the grey wolf. Biologi-

cal Conservation, 201, 1–9. https://doi.org/10.1016/j.

biocon.2016.06.016

Young, J. K., Olson, K. A., Reading, R. P., Amgalanbaatar,

S., & Berger, J. (2011). Is wildlife going to the dogs?

Impacts of feral and free–roaming dogs on wildlife

populations. BioScience, 61(2), 125–132. https://doi.

org/10.1525/bio.2011.61.2.7

Zamora-Nasca, L., di Virgilio, A., & Lambertucci, S. A.

(2021). Online survey suggests that dog attacks on

wildlife affect many species and every ecoregion

of Argentina. Biological Conservation, 256, 109041.

https://doi.org/10.1016/j.biocon.2021.109041