Whiskers in the city: domestic cat predation in Ecuadorian coastal cities and associated factors

Abstract

Domestic cats pose a latent threat to wildlife that lives within the remnants of natural vegetation in urban ecosystems. Both intrinsic (e.g., age, weight, sterilization status) and extrinsic factors (e.g., night confinement, interaction time with owners at home) can influence the number of prey items caught by cats. We assessed the fauna predation by domestic cats in three cities on the coast of Ecuador. We aimed to: (i) evaluate the composition of the prey brought home by cats, counting the taxa number and their capture frequency, as well as their conservation status, and (ii) identify the intrinsic and extrinsic factors that influence the quantity of prey brought home by cats (henceforth referred to as ‘prey captured’). A citizen science approach was employed to gather information about wildlife taxa caught and brought home by 100 cats in 50 households between March and October 2023. Cats captured 132 prey items, of which 53.8% were invertebrates, 27.3% reptiles, 8.3% birds, 6.8% small mammals, and 3.8% amphibians. These prey items belonged to 53 taxa, 56.6% native and 15.1% non-native. Non-native reptiles Hemidactylus sp. and Anolis sagrei were the most frequently captured taxa, and ten native taxa were among the most commonly captured, particularly odonates. This is the first study to register predation of cats on amphibians in northwestern South America. The capture by cats of Coniophanes dromiciformis, a vulnerable and probably endemic snake, is noteworthy. Three factors—age, nocturnal confinement, and the presence of toys in their homes—were the most important factors that contributed to predation events. We recommend controlling these factors to reduce the potential impacts caused by domestic cats on wildlife.

Full text

Urban Ecosystems (2025) 28:0
https://doi.org/10.1007/s11252-024-01628-9
has surely increased since then. Due to the lack of a specic
native distribution, domestic cats are considered non-native
fauna around the world (Loss and Marra 2017). Besides
providing companionship to humans, this species is valued
socially for its eectiveness in controlling pest populations
in urban environments (Hu et al. 2014; Crowley et al. 2020),
favoring a high density of cats in highly urbanized cities,
reaching up to 1,500 individuals/km2 (Liberg et al. 2000;
Baker et al. 2008), and raising concerns about interactions
and negative impacts on the wildlife inhabiting these places
(Trouwborst et al. 2020). As urban settlements continue to
expand, coexistence between humans, domestic cats, and
wildlife will be inevitable, making it essential to develop
comprehensive strategies that minimize the impact of cats
on wildlife (Schell et al. 2021).
Cats possess innate hunting abilities, demonstrat-
ing excellence in capturing a high diversity of species
(Turner and Meister 1988). The International Union for
Introduction
The domestication of cats (Felis catus Linnaeus, 1758)
occurred approximately 9,500 years ago in the Fertile
Crescent (Driscoll et al. 2007). Since then, humans have
facilitated the global expansion of cats, with the total world
population of cats being estimated at around 600 million in
2009 (Driscoll et al. 2009; Baker et al. 2010), a number that
Ileana Herrera
herrera.ita@gmail.com
1 Escuela de Ciencias Ambientales, Universidad Espíritu
Santo, Guayaquil 091650, Ecuador
2 Instituto Nacional de Biodiversidad (INABIO),
Quito 170501, Ecuador
3 Laboratorio de Ecología, Instituto de Investigaciones
Forestales, Universidad Veracruzana, Xalapa 91090, México
Abstract
Domestic cats pose a latent threat to wildlife that lives within the remnants of natural vegetation in urban ecosystems.
Both intrinsic (e.g., age, weight, sterilization status) and extrinsic factors (e.g., night connement, interaction time with
owners at home) can inuence the number of prey items caught by cats. We assessed the fauna predation by domestic
cats in three cities on the coast of Ecuador. We aimed to: (i) evaluate the composition of the prey brought home by cats,
counting the taxa number and their capture frequency, as well as their conservation status, and (ii) identify the intrinsic
and extrinsic factors that inuence the quantity of prey brought home by cats (henceforth referred to as ‘prey captured’).
A citizen science approach was employed to gather information about wildlife taxa caught and brought home by 100 cats
in 50 households between March and October 2023. Cats captured 132 prey items, of which 53.8% were invertebrates,
27.3% reptiles, 8.3% birds, 6.8% small mammals, and 3.8% amphibians. These prey items belonged to 53 taxa, 56.6%
native and 15.1% non-native. Non-native reptiles Hemidactylus sp. and Anolis sagrei were the most frequently captured
taxa, and ten native taxa were among the most commonly captured, particularly odonates. This is the rst study to register
predation of cats on amphibians in northwestern South America. The capture by cats of Coniophanes dromiciformis, a
vulnerable and probably endemic snake, is noteworthy. Three factors—age, nocturnal connement, and the presence of
toys in their homes—were the most important factors that contributed to predation events. We recommend controlling
these factors to reduce the potential impacts caused by domestic cats on wildlife.
Keywords Citizen science · Domestic animal impacts · Felis catus · Invasive species · Management policy · Pet
ownership
Accepted: 7 September 2024
© The Author(s), under exclusive licence to Springer Science+Business Media, LLC, part of Springer Nature 2024
Whiskers in the city: domestic cat predation in Ecuadorian coastal
cities and associated factors
KevinPanchana1· IleanaHerrera1,2 · AnahíVargas1· IsacMella-Méndez3· RafaelFlores-Peredo3
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Urban Ecosystems (2025) 28:0
Conservation of Nature (IUCN) considers them as one of
the most widespread invasive alien species in the world
(Lowe et al. 2000), posing a threat to wild species (Nogales
et al. 2013; Mahlaba et al. 2017; Reynolds et al. 2021). Pre-
dation behavior by these felines has been documented to
contribute to the extinction of 63 vertebrate species, and the
population decline of 367 species at global scale, several of
them with synanthropic habits (Doherty et al. 2016). Addi-
tionally, there is evidence that cats may have a signicant
impact on invertebrate populations (Gillies and Clout 2003;
Medina and García 2007; Krauze-Gryz et al. 2012; Woolley
et al. 2020). While most studies to date have focused on
the behavior and impact of feral cats (Graham et al. 2012;
Frank et al. 2014; Woolley et al. 2020; Doherty et al. 2021),
recently, there has been more exploration into the role of
owned domestic cats in wildlife decline in urban environ-
ments and the factors inuencing their predatory behavior
(Mori et al. 2019; Cecchetti et al. 2021a; Mella-Méndez et
al. 2022). Due to the loss of natural areas and the decrease
in biological diversity attributable to urban expansion, it
is crucial to determine the impacts of cats (feral cats and
owned cats) to establish wildlife conservation measures in
urban environments (Crowley et al. 2019, 2020).
Several studies have evaluated the extrinsic (environ-
mental) and intrinsic (related to the cat itself) factors that
can favor or reduce cat predatory behavior in terms of the
number and types of prey they capture (Woods et al. 2003;
Lilith et al. 2006; Thomas et al. 2012; Cordonnier et al.
2022). Some extrinsic factors associated to cat predation
events include the time the cat spends outside its home
(Kays and DeWan 2004; McDonald et al. 2015; Mori et
al. 2019), nocturnal activity (Metsers et al. 2010; Thomas
et al. 2014; Linklater et al. 2019), distance to green areas
(Graham et al. 2013; McDonald et al. 2015), size of the sur-
rounding green areas (Dickman 1996), quality of shared
time between cat and owner (Cecchetti et al. 2021a), use of
play towers and other toys (Cecchetti et al. 2021a), use of
collars (Calver et al. 2007; Gordon et al. 2010), and the type,
origin, quality, and quantity of food (Silva-Rodríguez and
Sieving 2011; Piontek et al. 2021). Some intrinsic factors
that have been linked to the intensity of cat hunting activ-
ity are sex (Crowell-Davis et al. 2004), age (Barrat 1998;
McDonald et al. 2015), weight (Silva-Rodríguez and Siev-
ing; McDonald et al. 2015), sterilization status (Hall et al.
2016; Castañeda et al. 2019), and coloration (Mella-Méndez
et al. 2022). Understanding more about the consequences
of domestic cats on wildlife and its drivers (extrinsic and
intrinsic factors) is essential to prevent, manage and miti-
gate their impact on urban wildlife (Trouwborst et al. 2020).
Most studies on the impact of cats have focused on oce-
anic islands (Duy and Capece 2012; Flux 2017; Woinarski
et al. 2020; Woolley et al. 2020) or countries in temperate
climate regions such as those in Europe (Thomas et al.
2012; Krauze-Gryz et al. 2017; Mori et al. 2019; Piontek
et al. 2021) and North America (Loss et al. 2013; Loyd et
al. 2013; Kitts-Morgan et al. 2015; Flockhart et al. 2016).
However, research is scarce on this issue in urban areas of
the Neotropical region, which is of special interest due to its
high wild species richness and the high abundance of cats
(Reese 2005; Raven et al. 2020; Antonelli 2022). For exam-
ple, in Mexico, the domestic cat population is estimated to
be around seven million (Peña-Corona et al. 2022), with
documented evidence of their signicant impact on a wide
range of species, especially native species that play crucial
ecological roles (Mella-Méndez et al. 2022). In Brazil, pre-
dation by cats has been reported in insular regions (Ferreira
et al. 2014, 2019a; Ferreira and Genaro 2017), sites of con-
servation importance on the mainland (Assis et al.2023), and
urban areas (de Freitas 2023). In Colombia, a study carried
out in the tropical Andes, estimated that a population of 1.6
to 3.5 million cats could predate between eight to 29 million
of vertebrates annually (Sedano-Cruz 2022). These studies
highlight the urgent need for information about the potential
impacts that domestic cats can have on the biodiversity of
this valuable biogeographic region.
Despite Ecuador being considered one of the world’s
megadiverse countries (Lessmann et al. 2014), with nearly
20% of its territory protected through the National System
of Protected Areas, invasive species pose a prominent threat
to its biodiversity (Espinoza-Amén et al. 2021; Mestanza-
Ramón et al. 2023). Globally, impacts caused by introduced
fauna show greater severity in island ecosystems (Bonn-
aud et al. 2011; Medina et al. 2011). For this reason, recent
research in Ecuador on predation caused by cats has focused
on the Galápagos Islands, using fecal analysis (Carrión and
Valle 2018) and telemetry of feral cats (MacLeod et al.
2020). Although cats have been identied as a priority man-
agement species on mainland Ecuador (Espinoza-Amen et
al. 2021) as their population escalates to at least 2.4 million
(INEC 2022), there is limited information available about
the predation of local fauna by domestic and feral cats and
their impacts (Loss et al. 2022). Attacks by domestic cats
on wildlife have only been described and published for
the city of Quito, located in the northern Andes of Ecua-
dor (Díaz et al. 2023), while for cities located in the coastal
zone, no information has been documented or published.
Furthermore, this study provides, for the rst time, solid
information about predation by domestic cats in urban areas
surrounded by remnants of tropical dry forests, mangrove
forests, and semi-arid shrublands in the Neotropics (Rivas
et al. 2020; Morocho et al. 2022). Through a citizen science
study, the predation of urban wildlife by domestic cats was
assessed in three cities on the Ecuadorian coast (Guayaquil,
Daule, and Samborondón). For this purpose, two specic
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Urban Ecosystems (2025) 28:0
objectives were proposed to: (i) evaluate the composition
of the prey brought home by cats, counting the taxa number
and their capture frequency, as well as their conservation
status, and (ii) identify some intrinsic and extrinsic factors
that inuence the quantity of prey brought home by cats
(henceforth referred to as ‘prey captured’). Based on our
results, we also discuss some recommendations for the man-
agement of domestic cats on the Ecuadorian coast.
Methods
Study area
The study was conducted in the cities of Guayaquil (2° 11’
21” S, 79° 53’ 20” W), Daule (2° 02’ 40” S, 79° 53’ 50” W),
and Samborondón (2° 03’ 40” S, 79° 50’ 32” W) (Fig. 1),
located in the province of Guayas in the coastal region of
Ecuador. These three cities cover a total area of 3,000 km²
(2,430 km2 belonging to Guayaquil, 330 km2 of Daule, and
220 km2 of Samborondón) and are inhabited by approxi-
mately three million people in one million houses, resulting
in a population density of nearly 1,000 people per square
kilometer and a housing density of 330 houses per square
kilometer (INEC 2022). The elevation ranges from four to
Fig. 1 Spatial location of selected households (red diamonds) in the cities of Guayaquil, Daule, and Samborondón in Guayas, Ecuador. The orange
circles represent the area of inuence of each cat (~ 100 m radius)
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Urban Ecosystems (2025) 28:0
collected data in person and online through Google Forms
and social networks (email and WhatsApp). Also, we sent
monthly reminders to the owners to keep them informed
about the project’s progress.
Composition of the prey captured by cats
For eight consecutive months (from March 1 to October
31, 2023), encompassing both the rainy and dry seasons,
all animals captured by cats and reported by their owners
were meticulously recorded. For taxonomic identication of
the prey, guides specialized in invertebrates (Triplehorn and
Johnson 2005) and vertebrates (Ridgely et al. 2006; Tirira
2007; Valencia et al. 2008a, b) were used, along with digi-
tal resources such as the BIOWEB information repository
(https://bioweb.bio/) and the iNaturalist identication algo-
rithm (https://www.inaturalist.org/). For cases in which only
remains of the prey were found (e.g., feathers, fur, scales, or
exoskeletons), a stereoscope was used to assign the prey at a
more specic taxonomic level (Mella-Méndez et al. 2022).
In addition, specialists in the respective taxonomic group
were consulted for prey that were dicult to identify. The
prey identied at least at the genus level were categorized
as native or non-native, when their origin region could be
determined (Loss et al. 2013). If the prey could not be iden-
tied, the origin region for these taxa was categorized as
undetermined. Following this, the IUCN Red List (https://
www.iucnredlist.org/) and the National Red Lists (Carrillo
et al. 2005; Freile et al. 2019; Ortega-Andrade et al. 2021;
Tirira 2021) were consulted to determine the conserva-
tion status and natural distribution of the species captured
by cats. In situations where it was not possible to identify
a prey item, it was included in the total number of prey
but excluded from statistical analyses (Krauze-Gryz et al.
2017). The composition of the prey captured by cats does
not necessarily reect their real richness or abundance in
the environment, but it is useful to know the species most
susceptible to predation by cats.
Intrinsic and extrinsic factors of cats
Intrinsic factors of the cats, including age, weight, sex, color,
and sterilization status, were gathered through the surveys.
In addition, detailed information about extrinsic factors,
such as how much time cats spend outdoors, whether they
are indoors at night, their interaction time with owners at
home, the use of collars with bells, the presence of toys and
play towers, and specics about their diet, were collected
(Thomas et al. 2012; McDonald et al. 2015; Mella-Méndez
et al. 2022). Given that our study took place over eight
months, seasonality was initially considered an extrinsic
factor. However, we decided to remove this factor as 2023
600 m above sea level due to the presence of the Chongón
Colonche Range. The climate in the area is characterized
as an ecotone between the wet zones of Chocó and the arid
conditions of the Peruvian coast (Ministerio del Ambiente
del Ecuador 2013). The rainy season generally extends from
December to May, followed by the dry season from June
to November (Rossel and Cadier 2009). The average tem-
peratures range from 23 to 26 °C, with annual precipitation
between 500 and 1000 mm (Moran-Zuloaga et al. 2021).
The study area is part of the Tumbes-Chocó-Magdalena
Hotspot (Mittermeier et al. 2011), an area of great conser-
vation importance due to its high level of endemism and
biodiversity (Manchego et al. 2017). Despite its conserva-
tion signicance, the region has experienced signicant
changes in land use, marked by the expansion of buildings,
residential areas, roads, and suburban development, as well
as the allocation of land for agricultural, grazing, and aqua-
culture purposes (Delgado 2013). Predominant natural areas
include tropical dry forests, mangroves, and arid shrublands
(Arias de López et al. 2022), interspersed with croplands,
pastures and patches of gardens and ornamental tree and
shrub vegetation within urban areas. According to the last
census taken in Guayaquil, there are approximately 350,000
cats, of which 100,000 are strays (Municipalidad de Guaya-
quil 2022). This high presence of domestic cats throughout
the city and its surroundings can exert signicant pressure
on native urban fauna (e.g., Mella-Méndez et al. 2019).
Participants and citizen science survey
During January and February 2023, the project was pro-
moted through Facebook and Instagram to identify individ-
uals interested in participating, following the methodology
of Mori et al. (2019) and Mella-Méndez et al. (2022). Inter-
ested participants received a descriptive sheet containing
detailed information about the project (Fig. S1). All cats
that participated in the study had the full consent of their
respective owner. In total, 100 cats from 50 households par-
ticipated (2 cats per household) and were distributed across
the three cities (38 in Guayaquil, ve in Daule, and seven in
Samborondón). Cat owners were asked to collect any prey
brought home by their cats and place them in plastic bags
(Churcher and Lawton 1987; Barratt 1998; McDonald et
al. 2015). When we could not collect the prey or when a
participant chose not to retrieve it or allow home visits, we
provided a detailed guide for taking photographs of the prey
(Fig. S2), including information on their geographical coor-
dinates (Seymour et al. 2020).
To gather information about domestic cats, a modied
survey was conducted based on the proposals by Thomas
et al. (2012) and McDonald et al. (2015), as suggested by
Mella-Méndez et al. (2022) (Fig. S3). To achieve this, we
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Urban Ecosystems (2025) 28:0
of prey captured by cats. Households with multiple cats
were considered as a random factor (McDonald et al. 2015;
Doherty et al. 2021). Prey counts were modeled using the
Poisson distribution, and categorical variables were trans-
formed into numerical binary scales for testing (Zar 2010).
The models were built considering additive eects between
variables. Variables with missing values (i.e., cat weight,
quantity, and number of meals per day) were excluded from
the model. A multicollinearity analysis was performed to
ensure minimal correlation among the predictor variables,
using the variance ination factor (VIF) available in the car
package (Fox and Weisberg 2019). Variables with low cor-
relation to the response variable were removed, conrming
the absence of multicollinearity among the remaining vari-
ables (VIF < 2) (Zuur et al. 2010). The model incorporated
ten predictor variables, comprising three intrinsic and seven
extrinsic factors (Table 1).
The MuMIn package (Barton 2023) in R was used to
identify the most optimal model among the set of selection
models generated by the dredge function with the selected
factors. Models were ranked based on the Akaike informa-
tion criterion with second-order correction (AICc) due to
the limited sample size (Burnham et al. 2011). The impor-
tance of the models was evaluated with respect to a null
model where the dependent variable is the number of prey
items and the random factor is households with multiple cats
(number of prey items ~ (1|households). The importance of
each factor is interpreted through cumulative AICc weights
and the model-averaged eect sizes with their condence
intervals (α < 0.05). Cumulative AICc weights for each vari-
able represent the proportion of weight attributable to mod-
els containing that specic variable and are calculated by
summing the AICc model weights of all models containing
that variable. Continuous covariates were standardized to
have a mean of zero and a standard deviation of 1 to show
the relative strength of regression coecients (McDonald
et al. 2015; Mella-Méndez et al. 2022). This analysis only
included cases where the cat responsible for prey capture
was identied.
Results
Composition of the prey captured by cats
A total of 132 prey items were recorded as captured by the
cats in eight months (X
 = 1.3 prey per cat; SD = 2.6) (Table
S1), equivalent to an average of two prey items per cat per
year. Out of the 100 cats, 64 did not record any captures,
while 36 captured between one and 14 prey items. Con-
sidering only the cats that hunt, the average capture rate is
5.5 prey items per cat per year. The prey items belonged
was a climatologically atypical year for Ecuador, resulting
in an extended dry season that lasted nearly the entire year.
We conducted a spatial analysis to determine the dis-
tance from each cat’s residence to the nearest green area
and to estimate the size of this green area (Mella-Méndez
et al. 2022). For these measurements, the WGS84 system
and UTM zone 17 S projection were utilized. We adopted
the land cover classication proposed by the MapBiomas
Ecuador Project (v. 1.0) (https://ecuador.mapbiomas.org/
mapas-de-cobertura-y-uso/). For this analysis, six types of
land cover (equatorial dry forest, open forest, mangrove,
non-forest wetland, dry shrublands, and cropland/pasture)
were considered to be green areas. An inuence area of
36,000 m2 around each cat’s home was considered (Kays
et al. 2020) to determine the presence of green areas within
these zones. These analyses were carried out using the QGIS
3.24.2-Tisler free-access software.
Statistical analysis
Prey composition was described by assessing the relative
frequency of captured prey items per class and the num-
ber of taxa per class, presented using bar graphs. Addition-
ally, a histogram was used to describe the abundance of the
most frequently prey captured by cats. A generalized lin-
ear mixed model (GLMM) was employed using the lme4
package (Bates et al. 2014) in R version 4.3.1 to determine
the impact of intrinsic and extrinsic factors on the quantity
Table 1 Selected intrinsic and extrinsic factors for analyzing the quan-
tity of captured prey using a GLMM. For qualitative variables, the
number of cats in each category is given. For quantitative variables,
indicated with an asterisk (*), the average value and the minimum and
maximum values are provided in parentheses. VIF values, which mea-
sure the amount of multicollinearity in a regression model, are pre-
sented for each predictor factor
Type of
factor
Factor name Description VIF
Intrinsic Age * 55.8 (6–182) months 1.23
Color 45 Tabby, 55
Non-tabby
1.21
Sterilization 88 Yes, 12 No 1.26
Extrinsic Collar with bell 22 Yes, 78 No 1.18
Cat toys 74 Yes, 26 No 1.09
Time spent with owner
*
10 (0–24) hours 1.27
Food origin 81 Commercial, 19
Both commercial and
homemade
1.07
Nocturnal connement 82 Yes, 18 No 1.14
Size of the nearest
green area *
26.9 (0.00086–535.2)
km2
1.10
Green area within inu-
ence zone
31 Yesa, 69 No 1.22
a Cover percentage of green areas within cats’ inuence zone ranged
between > 0– 68.0%
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Urban Ecosystems (2025) 28:0
identied to the lowest possible taxonomic level. Of these,
56.6% were categorized as native and 15.1% as non-native.
The majority of taxa were invertebrates (33 taxa), followed
by reptiles (11 taxa) and birds (seven taxa), while amphib-
ians (three taxa) and small mammals (two taxa) exhibited
to 19 orders, 28 families, 35 genera, and 27 species. Over-
all, 38.6% of the captured individuals were native taxa and
36.4% were non-native. Of the total prey, 53.8% were inver-
tebrates, 27.3% reptiles, 8.3% birds, 6.8% small mammals,
and 3.8% amphibians (Fig. 2a). A total of 53 taxa could be
Fig. 2 Percentage of total prey captured (a) and total number of taxa per taxonomic group captured (b) by domestic cats in the study area. The preys
are categorized as native, non-native, or undetermined origin
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Urban Ecosystems (2025) 28:0
We identied 10 native invertebrates and 11 native ver-
tebrates up to species level. Most of the species (8/11) of
vertebrates were categorized at both national and global
scales as “Least Concern” (Table 2). Although these spe-
cies are not in a threatened category, some of them have a
restricted distribution, such as Campylorhynchus fasciatus,
which is an endemic bird in the Tumbesian region. Other
species with distributions restricted to specic geographic
areas are C. buckleyi, Gonatodes caudiscutatus, and Scinax
quinquefasciatus. Notably, we recorded the snakes Conio-
phanes dromiciformis, globally categorized as “Vulnerable”
and locally as “Near Threatened,” and M. pulchriceps, cat-
egorized as “Near Threatened” at local scale (Table 2).
Eect of intrinsic and extrinsic factors
Based on the cumulative AICc weights, it was found that
age (c = 0.98), nocturnal connement (c = 0.79), and the
lower representativeness (Fig. 2b). All amphibians and the
majority of invertebrates (45.4% of prey items; 54.5% of
taxa) and birds (81.8% of prey items; 71.4% of taxa) cap-
tured were native, whereas all small mammals and the
majority of reptiles (86.1% of prey items; 50.0% of taxa)
were either non-native or their origin could not be deter-
mined (Fig. 2a, b).
Among the 22 most frequently captured taxa (capture
frequency > 1 prey item), the taxa most captured by cats
were the non-native reptiles Hemidactylus sp. (9.9%) and
Anolis sagrei (9.1%), followed by two invertebrates, Gryl-
lidae (8.3%) and Periplaneta sp. (7.6%), and one mammal,
Mus musculus (5.3%). The most frequently captured native
taxa were six invertebrates (Erythemis vesiculosa, Pantala
avescens, Erythrodiplax umbrata, Schistocerca sp., Eryth-
rodiplax sp., and Blaberus sp.) and four vertebrates (Colum-
bina buckleyi, Scinax quinquefasciatus, Mastigodryas
pulchriceps, and Leptodactylus sp.) (Fig. 3).
Fig. 3 Capture frequency of the 22 taxa with the highest capture frequency by domestic cats in the study area
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Urban Ecosystems (2025) 28:0
eect of these three factors on the number of captured prey
items is signicant (Fig. 4; Table S2). The eect of age and
nocturnal connement factors is negative, whereas the pres-
ence of toys has a positive eect on the number of captured
prey (Fig. 4). Younger cats that spend the night outdoors
and have toys in their homes are associated with a higher
number of captured prey items.
presence of toys (c = 0.76) are the variables contributing
most signicantly to the models. On the other hand, the size
of the nearest green area (c = 0.41), color (c = 0.39), time
spent with the owner (c = 0.38), the use of a collar with a
bell (c = 0.32), presence of green areas in the cat’s zone of
inuence (c = 0.30), sterilization (c = 0.26), and commercial
food (c = 0.26) carry lesser weight. For three of the ten fac-
tors examined (age, nocturnal connement, and presence
of toys), the condence intervals of average β coecient
values from the models did not include zero. Therefore, the
Fig. 4 Average β coecient values from the models with their 95%
condence intervals indicate the direction and magnitude of the eects
of intrinsic and extrinsic variables inuencing the quantity of prey.
Asterisks (*) highlight the signicant factors with the highest AICc
weight within the model combinations
Class Species Red list Distribution
IUCN National
Aves Columbina buckleyi LC LC Colombia, Ecuador, Peru
Campylorhynchus fasciatus LC LC Ecuador, Peru
Polioptila bilineata NE NE From Mexico to northwestern Perua
Sporophila corvina LC LC From southern Mexico to north-
western Perua
Troglodytes aedon LC LC From Canada to South America
Reptilia Mastigodryas pulchriceps LC NT Colombia, Ecuador, Peru
Coniophanes dromiciformis VU NT Ecuador, Peru (doubtful)
Gonatodes caudiscutatus LC LC Ecuador, Peru
Iguana iguana LC LC From Mexico to Paraguay and
southeastern Brazil
Amphibia Scinax quinquefasciatus LC LC Colombia, Ecuador
Leptodactylus melanonotus LC LC From southern Mexico to Ecuador
Table 2 Conservation status
and distribution of the 11 native
vertebrate species captured by
domestic cats
NE = “Not Evaluated”, LC =
“Least Concern”, NT = “Near
Threatened”, VU = “Vulner-
able”
a Distribution data retrieved
from eBird (https://ebird.org/
species/)
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Urban Ecosystems (2025) 28:0
or left on-site outside the home, aspects not accounted for in
household owner counts; therefore, the capture rate obtained
in our study is probably underestimated.
A recent study carried out in a neotropical city of Mexico
(Xalapa), employing a similar owner survey methodology
to ours, reports a capture rate of 4 prey items per cat per
year (Mella-Méndez et al. 2022), which is high compared to
that estimated in our study, namely two prey items per cat
per year considering all cats in the study. The discrepancy
observed can be primarily attributed to the type of ecosys-
tem (mountain cloud forest) surrounding the urban areas.
Rainforests are known for harboring higher biodiversity per
square meter than other ecosystems (Abrams and Abrams
2020). Specically, mountain cloud forest ecosystems con-
tain unique ora and fauna characterized by high levels of
richness and abundance (Almazán-Núñez et al. 2018). Since
our study focuses on remnants of tropical dry forests and
xerophytic shrublands, our ecosystems are less diverse than
montane cloud forests, and a lower capture of prey and taxa
is expected. Despite the lower capture rate in the study area,
it is worth noting that the Ecuadorian coast holds the last
threatened remnants of tropical dry forests and xerophytic
shrublands characterized by their high degree of endemism
(Cuesta et al. 2017). Estimating the wildlife captured by cats
is crucial, as it provides insights into the potential impacts
of cat predation on local ecosystems. Discrepancies in cap-
ture rates between regions underscore the importance of
considering ecosystem types and biodiversity levels when
evaluating these impacts, highlighting the need for targeted
conservation eorts in areas like the Ecuadorian coast, to
preserve unique and threatened ecosystems. No previous
study has examined the impacts of cat depredation on wild-
life in tropical dry ecosystems; hence, our study represents
the rst contribution to lling this information gap.
Cats generally hunt animals smaller than themselves, as
shown by the high number of small preys found in the lit-
erature (Lepczyk et al. 2004; Kutt 2012; Paul and Friend
2020). Most of the studies report that birds and small mam-
mals predominate among the prey items and species cap-
tured by domestic cats (Morgan et al. 2009; van Heezik et
al. 2010; Díaz et al. 2023; Lepczyk et al. 2023). Unexpect-
edly, most of the prey items and taxa captured by cats in
our study consist of invertebrates. These variations could
be related to dierences in prey population dynamics, as the
presence and abundance of dierent species can vary sig-
nicantly depending on local environments and the avail-
ability of resources (Lepczyk et al. 2023). In other words, if
there is a higher presence of accessible invertebrates in the
environment, cats may adjust their hunting patterns towards
this taxonomic group. This observation aligns with studies
conducted in some tropical islands, where cats prominently
capture invertebrates, attributed to seasonal variations in
Discussion
In the present study, we employed a citizen science approach
to provide a baseline list of species aected by cat preda-
tion, their capture rate, and critical insights for policymak-
ers in an urban area in the Neotropical region. While similar
studies have been conducted in other cities surrounded by
humid forests in the Neotropics (Assis et al. 2023; Mella-
Méndez et al. 2022), this is the rst report of the impact of
cats on wildlife in dry ecosystems in this region. We esti-
mated that domestic cats have an average capture rate of
two prey items per cat per year. Considering only the cats
that hunt, the capture rate estimated is 5.5 prey items per cat
per year, capturing a wide variety of prey (53 taxa) in the
study area, with invertebrates (53.8%; n = 71) as the most
common, followed by reptiles (27.3%; n = 36). Our results
suggest that three factors favor the rate of prey capture by
domestic cats: age, nocturnal connement, and the presence
of toys in their homes. Specically, nocturnal connement
stands out as an essential factor to be considered in develop-
ing public policy that regulates the responsible ownership
of domestic cats, reducing the impacts on wildlife caused
by these pets.
Capture rate, composition, and conservation status
of prey
The capture rate in our study (5.5 prey items per hunting
cat per year) is signicantly lower than the global average
(39 prey items per hunting cat per year; Legge et al. 2020).
However, these rates dier because the global average was
obtained from a meta-analysis of 47 studies, some of which
employed more precise methodologies (e.g., crittercams
and molecular analysis of stomach contents and scats). For
instance, a study conducted in the temperate rainforests of
Chile found that 168 vertebrates were captured per hunt-
ing cat per year using the scat analysis methodology (Silva-
Rodríguez and Sieving 2011). Another study conducted in
two rural areas of Poland utilizing scat and stomach con-
tents reported a capture rate of 443 vertebrates per hunting
cat per year (Krauze-Gryz et al. 2019). Additionally, a study
carried out in Georgia (USA) highlights the inuence of the
methodology used on the values of estimated capture rate by
cats (Lloyd et al. 2013). These authors using animal-borne
video cameras documented a capture rate of 125 prey items
per hunting cat per year; in contrast, counting only the prey
brought home by cats, they estimated a capture rate four
times less (a rate of 28.7 prey items per hunting cat per year
Lloyd et al., 2013). According to some studies, domestic
cats typically bring home only 10 to 23% of the prey they
capture (Loyd et al. 2013; Krauze-Gryz et al. 2019; Sey-
mour et al. 2020). The remaining prey are either consumed
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Urban Ecosystems (2025) 28:0
and the threats posed by urbanization, including habitat
loss, fragmentation, and degradation (Ortega-Andrade et al.
2021; Hamer and McDonnell 2008). Nevertheless, captured
amphibian species (S. quinquefasciatus and L. melano-
notus) exhibit varied responses to urbanization, inuenc-
ing their presence and abundance in the study area. For
instance, the locally abundant treefrog S. quinquefasciatus,
known for its wide distribution and tolerance to anthropo-
genic activities, is found in urban areas of the Ecuadorian
coast (Ron et al. 2018), making it a potential prey for cats.
Nevertheless, some of the amphibian species registered (i.e.,
Leptodactylus spp.) may be abundant but also exhibiting
anti-predatory behavior such as changing their body shape
or inating themselves, mouth gaping, and jumping away
from predators (see Ferreira et al. 2019b), which can help
them in avoiding and escaping cat attacks. These ndings
emphasize the interplay between cat predation and amphib-
ian responses, requiring further research.
Non-threatened local species are generally more abun-
dant and may be more accessible than threatened species
to opportunistic predators such as domestic cats (Loss and
Marra 2017; Mori et al. 2019). The record of cats prey-
ing on various common native dragonies (E. vesiculosa,
E. umbrata, and P. avescens) in large quantities could be
linked to their frequent occurrence in anthropogenic water
bodies within urban areas, such as pools and garden foun-
tains, particularly during dry seasons when natural water
sources are scarce (Perron and Pick 2020; Husband and
McIntyre 2021). Another common species that cats capture
is the Ecuadorian ground dove (C. buckleyi), which is locally
adapted and abundant, given its adaptability to urban areas.
This species can utilize urban plants to feed on grains and
seeds, primarily foraging in lower strata (Allieri and Man-
Ging 2015; Carrión-Zambrano et al. 2022), making them
available and exposed to urban predators such as cats. How-
ever, the record of threatened poorly studied snakes like C.
dromiciformis, which is globally “Vulnerable” and “Near
Threatened” locally, along with M. pulchriceps, which can
be found solely in forest remnants (Amador-Oyala, 2015),
contrasts with the common idea that domestic cats focus
only on abundant prey (Dickman and Newsome 2015; Yip
et al. 2015). This nding suggests that predation of threat-
ened species by cats, in synergy with other pressures (e.g.,
habitat loss, road deaths), may even contribute to the decline
of their populations (Medrano-Vizcaíno et al. 2023). The
interactions between cats and local species, both common
and threatened, highlight potential impacts on general local
wildlife conservation.
Five out of the 11 native vertebrate species reported in
our study, including the white-browed gnatcatcher (P. bilin-
eata), the variable seedeater (S. corvina), and the house
wren (T. aedon), have wide distributional ranges inhabiting
prey abundance (Ferreira et al. 2014; Ferreira and Genaro
2017; Bruce et al. 2019). In urban settings, a higher abun-
dance of captured invertebrates by cats is expected not only
due to their high global diversity (Stork et al. 2015) but also
to the habitat heterogeneity present in cities, such as parks
and gardens, which oer suitable foraging and nesting sites
for these animals (Hennig and Ghazoul 2012; Jones and
Leather 2012). Considering that the predatory behaviors of
cats towards wildlife in urban environments are inuenced
by various factors, such as prey abundance and ease of cap-
ture, making their understanding and management complex,
it is crucial to address the variables comprehensively.
Cat owners often allow their cats to roam freely, employ-
ing them for pest control, since they frequently capture
non-native animal species (Crowley et al. 2020; Trouwborst
et al. 2020). This perception is partially supported by our
results, which show that cats capture a signicant number
of species considered invasive species or pests in the study
area, including brown lizards, house geckos, crickets, cock-
roaches, mice, and rats to a greater extent. The prey most
captured were two non-native reptiles, the brown lizards (A.
sagrei) and the house geckos (Hemidactylus sp.), given their
higher abundance resulting from adaptability to urban envi-
ronments. These species are frequently found in residential
areas and parks, making them more susceptible to preda-
tors such as cats, as they require thermoregulation to remain
active (Borroto-Páez and Pérez 2020; Horváth et al. 2020;
Reynolds et al. 2021). Also, two non-native mammals, rats
(R. rattus) and mice (M. musculus), were some of the most
captured preys because they are abundant in urban areas,
where they nd food sources and refuge (Gillies and Clout
2003; Maeda et al. 2019). Predation by cats has also been
demonstrated to impact native fauna. Our results showed
that 56.6% of all prey taxa targeted by cats are native, align-
ing with ndings in other Neotropical regions. In a study
in a southern Brazilian municipality, for example, 64.0%
of species reported as preyed upon by domestic cats were
native (de Freitas et al. 2023), while in a southeastern Mexi-
can city, 93.5% of the registered prey taxa were also native
(Mella-Méndez et al. 2022). These results underscore the
importance of considering the dual role of cats, both as
contributors to controlling non-native pests and potential
threats to native wildlife.
Our study documents, for the rst time, predation by
cats on native amphibians in northwestern South America.
The absence of previous records of cats preying on amphib-
ians aligns with the general trend of declining diversity and
abundance of native herpetofauna caused by the expansion
of urbanized areas (Hamer and McDonnell 2010; Faeth et
al. 2011; Ackley et al. 2015). The limited capture of amphib-
ian species was anticipated, given their lower diversity in
Guayas, the least diverse vertebrate class in the province,
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Urban Ecosystems (2025) 28:0
stimuli but also olfactory, auditory, and tactile senses (Cinel
et al. 2020). Moreover, some lizard species rely on their
olfactory sense to identify predators’ scents through chemi-
cal cues rather than visual recognition (Ortega et al. 2018;
Fernández-Rodríguez and Braña 2022). Consequently, fac-
tors associated with visual stimuli, such as cat color, appear
less inuential than, for instance, exposure to cat scent as
a triggering element for antipredator behavior (Cli et al.
2022). This distinction likely aects the hunting success of
cats with these taxa.
Among the extrinsic factors, the presence of toys in cats’
homes had a signicant weight in the model. Toys are an
essential part of the environmental enrichment that every
cat should have (Ellis 2009). The literature suggests that
cats with access to an enriching environment with toys that
simulate common prey, scratchers, or towers can channel
their energy into play and simulated hunting activities,
which may reduce the need to hunt wild animals (Cecchetti
et al. 2021a). Surprisingly, our results suggest that pro-
viding toys to domestic cats could increase the number of
captured prey, as these toys can serve as tools for training
activities to make their hunting events more ecient. This
result may be supported by the fact that for cats, playing
with objects and predation can share a motivational basis
(Biben 1979; Hall and Bradshaw 1998). In this context, it
could be inferred that providing toys (especially those that
simulate birds, mice, and small reptiles) that produce rapid
movements could motivate cats to engage in hunting behav-
iors and actively predate more (Chávez-Contreras 2016).
However, other factors to assess the quality of play, such as
the quantity and type of toys in their homes, positive hunt-
ing reinforcement, and playtime, should also be considered
for a more comprehensive explanation of this phenomenon
(Strickler and Shull 2014; Delgado and Hecht 2019; Cec-
chetti et al. 2021a).
Nocturnal connement has been shown to decrease the
amount of prey hunted by cats, consistent with ndings in
other studies (Robertson 1998; Mella-Méndez et al. 2022).
Since it represents a reduction in encounter opportunities
between cats and prey, cats that spend 12 h outdoors will
interact with less prey than those that spend 24 h outdoors.
Considering that in highly urbanized areas, cats are more
active at night (Bennett et al. 2021), it is during noctur-
nal hours when cats would have a greater opportunity to
catch prey (e.g., Denny and Dickman 2010). Hence, noc-
turnal connement is crucial to prevent cats from roaming
and encountering nocturnal native wildlife (e.g., snakes),
which are not typically found inside homes (Lilith et al.
2006). Even diurnal or crepuscular animals that change their
behaviors because of light sources present in cities (Russart
and Nelson 2018) or natural moonlight (Brisbane and Van
Den Burg 2020) can be vulnerable to nocturnal predators.
a variety of habitats and climates throughout the Americas.
However, there are unique species in South America, such as
the fasciated wren (C. fasciatus) and the shield head gecko
(G. caudiscutatus), which have geographical distributions
restricted to the northwestern part of the continent, which are
also aected by cat predation. These ndings illustrate the
potential interaction between cats and locally adapted spe-
cies, which can carry consequences for the function of eco-
systems (Kitts-Morgan 2015; Escribano-Avila et al. 2017).
A recent meta-analysis reported 89 native species as prey of
cats for the South American continent (Lepczyk et al. 2023),
while we recorded 30 native taxa (21 native species) solely
in the study area, which has an area of approximately 3,000
km2. These results suggest that many more studies in South
America are needed to reveal the true extent of the impact of
cats on all native and endemic predated wildlife.
Intrinsic and extrinsic factors
Of the intrinsic factors considered for the model, only age
was a signicant factor in hunting activities, consistent with
previous studies (Barrat 1998; McDonald et al. 2015). The
positive association between the youth of cats and prey cap-
ture is consistent with the existing literature, suggesting that
younger cats often exhibit more active and exploratory hunt-
ing behaviors than older cats (Fitzgerald and Turner 2000;
Philippe-Lesare et al. 2024). Indeed, the majority (77.8%)
of actively hunting cats in our study are kittens or young
adult cats (< 6 years), according to the Quimby et al. (2021)
life stage classication. The youngest cat, in particular, cap-
tured the largest quantity of prey items. As a cat becomes
older, its predatory behavior may be reduced because of
changes in their sensory systems and cognitive and physi-
ological functions (Bellows et al. 2016; Delgado and Hecht
2019; Cecchetti et al. 2021b). Our data does not support the
idea that sterilization inuences the number of prey items
captured by a cat, which is consistent with certain studies
(Guttilla and Stapp 2010; Hall et al. 2016). However, this
contrasts with other research ndings that suggest that non-
sterilized cats may roam more extensively, providing them
with increased opportunities for predatory activity (Kitts-
Morgan et al. 2015; Ferreira et al. 2020). Further investiga-
tion into the correlation between home range and predation
is necessary to comprehend this discordance better.
Contrary to what was documented by Mella-Méndez
et al. (2022), the tabby color of cats did not prove to be
a determining intrinsic factor in prey capture rates. This
result could be attributed to the primary mechanisms of
predator recognition observed in the most captured taxa
reported in the present study (insects and reptiles), which
are not primarily based on the visual sense. In the case of
insects, predator-induced responses involve not only visual
1 3
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Urban Ecosystems (2025) 28:0
p.m. to 6 a.m.). Several studies suggest nocturnal conne-
ment as a practical measure (Lilith et al. 2006; Linklater et
al. 2019; Cecchetti et al. 2021b; Mella-Méndez et al. 2022).
Concurrently, this could be combined with other measures
used for cat identication (microchips) and population con-
trol through sterilization (Grayson and Calver 2004), along
with the implementation of cat-exclusion zones (Metser et
al. 2010; Herrera et al. 2022). Promoting these measures is
essential for owners of younger cats. Environmental enrich-
ment is crucial for cats’ well-being (Grigg and Kogan 2019).
Hence, before any restrictions are implemented, there
should be further investigation into whether cats that play
with toys that simulate their prey (birds, mice, and little rep-
tiles) increase their prey capture.
Cat population control through sterilization is one of the
primary practices regulated and accepted in urban areas
(Cecchetti et al. 2021b), including the study area. However,
more research about public perceptions and social attitudes
toward responsible cat ownership and anti-predation strate-
gies is needed, mainly focusing on nocturnal connement.
Additionally, outreach and environmental education proj-
ects regarding the impact of domestic cats on wildlife are
necessary to foster greater acceptance by people of move-
ment restriction policies for cats in urban areas. In this con-
text, environmental awareness is necessary to convey the
information obtained (Thomas et al. 2012; Woolley and
Hartley 2019) and to counter misinformation (Loss et al.
2018; Gow et al. 2022).
Conclusions
Our study provides the rst documented evidence of the
impact of domestic cat predation on wildlife in a Neotropical
dry ecosystem. Most prey and taxa captured were inverte-
brates, followed by reptiles, birds, mammals, and amphibi-
ans. While two non-native reptile species (Hemidactylus sp.
and Anolis sagrei) were commonly preyed on, a signicant
proportion of the prey were native species, highlighting the
potential threat posed by cats to local biodiversity. We docu-
mented predation by cats on amphibians for the rst time
in northwestern South America and the capture of the vul-
nerable and possibly endemic snake species Coniophanes
dromiciformis. Both intrinsic and extrinsic factors inuence
the hunting behavior of domestic cats in the study area.
Cat age was identied as the most signicant factor, with
younger cats exhibiting higher hunting activity compared
to older cats. Among extrinsic factors, nocturnal conne-
ment and the presence of toys in cats’ homes were identied
as factors to consider in the management of owned cats. In
particular, nocturnal connement of domestic cats stands
out as a measure that could be eectively included in public
For instance, brown anoles (A. sagrei) are diurnal lizards
that engage in foraging and reproductive activities at night
inuenced by urban lights (Rojas-González 2021), exposing
them to cats.
Unexpectedly we found no eect of cat’s collar with bell,
despite some studies suggesting that bells may decrease pre-
dation on wildlife (Calver et al. 2007; Gordon et al. 2010).
Furthermore, time spent with the owner was hypothesized to
decrease the capture rate (Cordonnier et al. 2023), as owners
actively engaged with their cats could potentially prevent
hunting events. However, our analysis did not detect any
signicant eect. Additionally, the origin of cat food, asso-
ciated with nutritional contributions, did not signicantly
aect cats’ hunting tendencies (Cecchetti et al. 2021a). Con-
trary to previous ndings (Dickman 1996; Barrat 1998), our
analysis of green areas did not reveal a correlation between
the presence of green areas in the cat’s inuence zone or
the larger size of the nearest green area and an increase in
captured prey. The selected “zone of inuence” for cats is
based on data from non-tropical countries (UK, USA, Aus-
tralia, and New Zealand) (Kays et al. 2020), which means
that actual cat ranges may dier substantially across land-
scape types. Other types of articial green spaces present
in urban environments, such as gardens and parks, have
been documented to oer suitable habitats for domestic cats
and should be considered in further studies. These specic
green areas may provide opportunities to nd potential prey
(Pavisse et al. 2019), but they were not considered in the
analysis.
Implications for the management
The management and control of cats undoubtedly represent
a signicant challenge in any neotropical city. The manage-
ment of feral or ownerless cats falls under the responsibil-
ity of local authorities, while the care of owned cats is not
directly included in their duties. However, eorts to prevent
or mitigate any impact of cats on wildlife require knowledge
of the situation and awareness strategies regarding the man-
agement and control of such impact (Crowley et al. 2019).
Some studies express skepticism about the negative impact
of domestic cats on biodiversity (Lynn et al. 2019; Turner
2022). However, searching for comprehensive strategies
that reduce the impact of cats on wildlife and disseminating
ecient mechanisms for controlling these animals in areas
of high biodiversity is necessary (Trouwborst 2020).
Among the three variables that signicantly aect the
hunting behavior of the sampled cats, nocturnal conne-
ment is the one that can be most eectively controlled
through public policies. The measures that should be imple-
mented involve nocturnal connement for cats, which
should remain indoors during a set schedule (e.g., from 6
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policies to diminish the impact of domestic cats on wildlife.
Further research is needed on owner acceptance of using
and implementing anti-predation strategies.
Supplementary Information The online version contains
supplementary material available at https://doi.org/10.1007/s11252-
024-01628-9.
Acknowledgements We thank all the participants who generously
volunteered their time and valuable records for this study. We would
like to express our gratitude to independent researchers Julian Perez-
Correa, Mario H. Yánez-Muñoz, Marissa Barreno, Stefania Cuadrado,
and Andrea Narvaez, who assisted us in identifying the captured prey.
A big thank you to Jemedier Benites for the design and illustrations
featured in the supplementary material. We would like to express our
gratitude to the anonymous reviewers for providing us with valuable
suggestions.
Author contributions All authors contributed to the study conception
and design. Material preparation and data collection were carried out
by KP. All authors contributed to the data analysis and interpretation.
The rst draft of the manuscript was written by KP and IH and all au-
thors commented on previous versions of the manuscript. All authors
read and approved the nal manuscript.
Funding This research was funded by the Centro de Investigaciones
(CIN) of Universidad Espíritu Santo (UEES)—Ecuador (project num-
ber: 2023-ING-001 to IH).
Data availability Data is provided within the manuscript or supple-
mentary information les.
Declarations
Ethical approval No ethical approval was required as this was an ob-
servational study.
Competing interests The authors declare no competing interests.
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