Frontiers

https://www.frontiersin.org/guidelines/author-guidelines

Title: The impact of COVID-19

pandemic restrictions on feral dogs in
Arequipa, Peru
Authors:
Micaela De la Puente-León 1,2, Elvis W. Díaz 1, Brinkley Raynor Bellotti 3,4, Guillermo Porras
1
, Katty Borrini 1, Olimpia Chuquista-Alcarraz 1, Valerie A. Paz-Soldán 1,5, Michael Z. Levy 1,6,
Ricardo Castillo-Neyra 1,4,6 *

1 Zoonotic Disease Research Lab, One Health Unit, School of Public Health and
Administration, Universidad Peruana Cayetano Heredia, Lima, Perú

2 School of Health Sciences, Universidad San Ignacio de Loyola, Lima, Perú

3 Department of Internal Medicine - Infectious Diseases, Wake Forest University School of

Medicine, Winston-Salem, North Carolina, United States of America.

4 Department of Pathobiology, School of Veterinary Medicine, University of Pennsylvania,

Philadelphia, Pennsylvania, United States of America

5 Department of Tropical Medicine and Infectious Disease, Celia Scott Weatherhead Tulane
University School of Public Health and Tropical Medicine, New Orleans, Louisiana, United
States of America.

6 Department of Biostatistics, Epidemiology & Informatics, Perelman School of

Medicine at University of Pennsylvania, Philadelphia, Pennsylvania, United States of
America

* Corresponding author: RCN [email protected]

Abstract (max. 350 words)

Background. Feral dogs in Arequipa, Peru, inhabit caves in periurban areas and primarily
rely on organic waste from landfills and livestock from backyard farms. These feral dogs
frequently attack small and medium farm animals and occasionally people, posing significant
public health risks. Beyond the physical injuries resulting from the attacks, these dogs pose
a threat for dog-mediated human rabies as there is active virus transmission in the dog
population of Arequipa city. The COVID-19 pandemic restrictions in Arequipa, including
restaurant closures, led to a decline in backyard farming and organic waste, thereby
reducing food availability for feral dogs.
Methods. We longitudinally (2019–2022) examined the impact of pandemic restrictions on
feral dog presence in the periurban areas of Arequipa. Monthly surveys recorded direct and
indirect evidence of feral dog presence in caves. An interrupted time series (ITS) analysis
evaluated changes in the number of caves with evidence of feral dogs before and after
pandemic restrictions. In addition, we conducted in-depth interviews with local farmers to
understand the interactions and conflicts between feral dogs and human populations in
those periods.
Results. Over 29 months (7 pre-pandemic, 22 during/post-pandemic), an average of 16.42
caves per month showed evidence of feral dog presence, mostly in the form of indirect
evidence. Following the pandemic restrictions, the total number of occupied caves
decreased by 42% (p<0.010), with a 41% reduction in indirect evidence (p=0.012) and a
striking 76% decrease in direct evidence of feral dogs (p<0.001). Farmers reported a
decrease in dog attacks after the pandemic restrictions.
Conclusions. The observed population decline suggests that reduced food availability
impacted the local feral dog population. The greater reduction in direct evidence compared
to indirect signs indicates a decline in active feral dog presence. These findings highlight the
strong dependency of feral dogs on human-generated food sources and emphasize the
influence of environmental changes on their population dynamics. Despite the ongoing
canine rabies epidemic in Arequipa, feral dog ecology remains poorly understood. Our study
underscores the potential of appropriate solid waste management as an effective strategy to
control feral dog populations through decreased food availability.

Keywords: Dog population management, Feral dogs, One Health, Pandemic, Population
dynamics, Rabies, Zoonosis.

Introduction
A persistent dog rabies epidemic has affected Arequipa city, Peru, at least since 2015 when
the rabies virus reintroduction was detected (Castillo-Neyra, Zegarra, et al., 2017) . Periurban
communities on the outskirts of the city face persistent geographic, economic, and social
vulnerabilities that increase their risk of canine rabies, particularly due to limited access to
post-exposure prophylaxis following dog bites
(Castillo-Neyra, Zegarra, et al., 2017; De la Puente-León et al., 202
. In these same periurban areas, fieldwork
conducted in 2019 revealed a previously undocumented issue: feral dogs inhabiting caves in
the surrounding landscape (Castillo-Neyra et al., 2025) .

Feral dogs—free-roaming dogs that have reverted to a wild state—form packs, avoid human
interaction, and subsist independently, often hunting or scavenging
(Boitani et al., 2016; Miternique & Gaunet, 2020; Smuts, 2010)
. These dogs pose a significant threat to public
health, livestock, and wildlife in Arequipa (Castillo-Neyra et al., 2025) . Their exclusion from
mass vaccination and sterilization campaigns, combined with potential for long-range
mobility and high intra-group contact, presents a challenge to reaching the 70% canine
vaccination threshold recommended by WHO (or 80% by PAHO) necessary for rabies
elimination
(Norma Técnica de Salud Para La Prevención y Control de Rabia Humana En El Perú, 2017; Schneider et
.
While data on the ecology of feral dogs in Arequipa remain limited
(Castillo-Neyra et al., 2025)
, studies in similar urban-adapted wildlife cities (e.g., raccoons, foxes), suggest that
feral dogs populations often rely on human-generated food waste, which enables their
persistence in periurban environments
(Duke et al., 2013; Dürr & Ward, 2014; Raynor et al., 2020)
. In periurban Arequipa, solid waste is often managed informally (UNV, 2014) .
Residents may pay waste pickers to collect trash (Espinosa-Aquino et al., 2023) , but
disposal commonly occurs in nearby water channels
(Andina.pe, 2025; Defensoría del Pueblo, 2022)
, geographic features that have also been associated with rabies virus
transmission (Castillo-Neyra, Zegarra, et al., 2017) . Formal municipal waste collection
remains inconsistent due to budgetary and infrastructural limitations (Alegre, 2021) .

Backyard livestock farming is common in periurban areas of Arequipa. Animals are often fed
with organic scraps, obtained through informal supply chains, particularly from restaurants
and markets (Saikia & And Bhar, 2010; Tripathi & Lingaraju Balaraju, 2016) . Animal remains
and waste from these systems contribute to the diet of feral dogs
(Castillo-Neyra et al., 2025)
. Disruptions to these food sources—whether due to economic shifts or environmental
change—could significantly impact the viability of feral dog populations.

In this study, we leverage the restrictions imposed during the COVID-19 pandemic
(El Peruano, 2020; MINSA, 2025)
as a natural experiment to assess the impact of
anthropogenic environmental change on feral dog populations. These restrictions
substantially reduced the availability of organic waste in landfills and informal animal farms in
the periurban areas of Arequipa (Malone et al., 2021; Perfecto et al., 2022) , where cave-
dwelling feral dogs are commonly found. Our objective was to evaluate whether there was
an association between the timing of the pandemic restrictions and the number of caves
exhibiting evidence of feral dog presence in periurban communities. We hypothesize that
reduced food availability during the pandemic led to a decline in feral dog activity, as
indicated by decreased signs of cave use, highlighting the potential role of food waste
management in the control of free-roaming dog populations.

Material and methods

Ethics statement
Ethical approval was obtained from Universidad Peruana Cayetano Heredia (approval
number: 012 - 04 - 22).

Study settings
This longitudinal study was conducted from 2019 to 2022 in the periurban areas of the Alto
Selva Alegre (ASA) district in Arequipa. The periurban areas of ASA were established on the
outskirts of the city of Arequipa over the past few decades
(Bayer et al., 2009; Levy et al., 2014)
. Periurban areas are characterized by unplanned and rapid growth, often comprised
by rural-urban migrants, low SES, very limited infrastructure, and high environmental
vulnerability (e.g., poor solid waste management). New settlers usually come together from
the same rural town, occupy a discrete and continuous piece of land, and request formal
land tenure from the district and city authorities. This discrete geographical area is called a
locality, and each locality is engaged in a specific economic activity, such as rock extraction
or the raising of domestic farm animals (MDASA, 2016, 2017) . Moreover, each locality is
characterized by a different level of backyard livestock keeping. The study population
consisted of feral dog and their caves located in four periurban localities: the San Isidro
Labrador Pig Farmers Association (APSIL), San Luis Gonzaga Zone A, San Luis Gonzaga
Zone D, and El Roble (Fig 1). These caves, which vary in depth, are found in the ground or
walls along hillside paths. Feral dogs use them for resting and reproduction
(Castillo-Neyra et al., 2025)
. Some caves are dug by the dogs themselves, while others are natural
formations or landscape features that dogs utilize (Castillo-Neyra et al., 2025) .

Figure 1. (A) Peru, the country where the study was conducted. (B) Arequipa Region, the
canine rabies endemic area in Peru. (C) Alto Selva Alegre District, located within the city of
Arequipa.

Cave surveillance
At each study locality, caves were identified by field teams following visible paths created by
regular movement of feral dogs. These paths were readily apparent both in the field and on
satellite maps. Monthly surveillance visits were conducted to monitor the formation of new
caves as well as to collect longitudinal data on the use of caves by feral dogs. Detailed
information on the monthly surveys is provides elsewhere (Castillo-Neyra et al., 2025) .
Surveillance was conducted from September 2019 to March 2022. This time frame includes
7 months before the pandemic restrictions were instituted (September 2019 - March 2020)
and 22 months during and after the pandemic restrictions (June 2020 - March 2022).

Cave survey data collection

Field teams collected data through a standardized mobile data collection form developed in
the World Veterinary Service (WVS) application (Mission Rabies, n.d.) . The form in the WVS
app was designed and piloted by the research team and included fields for cave registration
and the presence or absence of various indicators of feral dog activity (Supplement 1).

For each monthly visit, data on feral dog evidence was recorded. Observed evidence was
categorized as direct or indirect. Direct evidence of feral dog presence included: sighting of
solitary dogs, dog packs, puppy litters, or dead dogs inside or around the cave opening (Fig
2). Indirect evidence of feral dog presence included canine feces, fresh or desiccated,
canine tracks, canine scratch marks, and prey carcases (Fig 3). Additional data were
recorded along trails—regardless of cave presence—on dog packs, solitary dogs, and
carcasses. No animal handling (e.g., capture, restraint, or sample collection) was conducted
in this study. Each cave visit was conducted by two observers with equivalent experience
levels, working together to increase detection sensitivity for both caves and animals
(Castillo-Neyra et al., 2025).

Figure 2. Direct evidence recorded in and around caves. (A) dog packs, (B) solitary dogs,
(C) dead dogs, and (D) litters.
Figure 3. Indirect evidence found in and around cave dogs. (A) Paw prints, (B) dog feces,
(C) claw marks in caves, and (D) prey carcass of a pig.

Statistical analysis
The primary outcome analyzed was the count of caves with general evidence of feral dogs
per month. Secondary outcomes included the count of caves with direct and indirect
evidence, and the number of litters, solitary live dogs, packs, and dead dogs per locality (i.e.,
all those associated with a cave and those observed not associated with a cave). We
compared these primary and secondary outcomes before and after/during the COVID-19
pandemic restrictions. We used Student’s t-tests for mean comparisons and Wilcoxon rank-
sum tests for median comparisons. Simple generalized linear models (GLMs) were
constructed to explore potential associations of time period (pre- or post- pandemic
restrictions) with the primary and secondary outcomes. We used GLMs with a negative
binomial family due to overdispersion in the data. In addition, given that a higher number of
caves visited increases the likelihood of detecting caves with evidence of feral dogs, we
included the number of caves visited each month in each locality as an offset term in our
models and evaluated its inclusion using Akaike Information Criterion (AIC) (Supplement 2).

After establishing a potential association between the outcomes and the pandemic
restrictions, we conducted an interrupted time series (ITS) analysis as our primary analytical
framework. ITS is a method used in longitudinal studies to assess whether a response
variable changes, and if that change is immediate or progressive, following an intervention.
In this study, the onset of COVID-19 restrictions was treated as the intervention, and
changes in the number of caves with feral dog evidence were evaluated across this temporal
breakpoint. The ITS model tested whether the onset of pandemic restrictions was associated
with changes in the number of caves showing evidence of feral dogs before and during the
restrictions. ITS used three variables besides the response variable (number of caves with
evidence): a time variable since the beginning of the study, a time variable since the onset of
the intervention (0 previous the intervention) and a dichotomous variable before/after the
onset of the intervention. ITS results indicate whether there was an immediate and/or
progressive change in the number of caves with evidence at the start of the restrictions, and
whether such a change was statistically significant. The ITS analysis provided information
about pre-intervention trends and whether those trends changed following the intervention.
For the regression model we used a negative binomial family based on our simple GLMs
results. The final parsimonious model was selected by comparing versions of the model with
and without the locality variable using log-likelihood estimation. All statistical analyses,
tables, and figures were prepared using R version 4.2.0 (R, 2024) .

Interviews to dwellers and farmers

In order to understand the interactions and conflicts between feral dogs and human
populations, we conducted in-depth interviews with dwellers and farmers from periurban
communities. These interviews were aimed at capturing lived experiences, farming
practices, and community-level responses to the presence of feral dogs. In particular, we
sought to explore how changes in food systems—such as the availability of organic waste,
fluctuations in livestock production, and informal feeding practices—could contribute to
shaping the ecological dynamics of feral dog populations. In the interviews, we explored the
effect of the pandemic restrictions on farming activities and the indirect impact on periurban
food systems that sustain feral dog populations. By examining how human activities related
to food production and waste disposal influence dog behavior and survival, we aimed to
generate insights into the socio-ecological drivers of human–dog interactions in rapidly
changing periurban environments

Interview Sampling & Recruitment

Purposive sampling was used to select participants across all four study localities. All
participants were residents and farmers from periurban neighbourhoods. Members of the
research team recruited participants through door-to-door visits, during which they obtained
contact information (e.g., cell phone numbers). To minimize face-to-face interaction and
reduce the risk of COVID-19 transmission, interviews were conducted a few days later by
phone.

Qualitative Data Collection

Interview guides were developed to explore four main topics: farm animal husbandry,
changes in feed and animal prices, interactions with feral dogs, and the impact of the
COVID-19 pandemic on farming practices. Two members of the research team conducted
the interviews: a Peruvian disease ecology researcher (MDLP, DVM, PhD in Epidemiology)
and a Peruvian veterinary epidemiologist (OCA, DVM, MSc).
Qualitative Data Management and Analysis
All interviews were digitally audio-recorded and transcribed, with detailed notes taken
throughout. Centering on the four main topics mentioned above, an inductive coding
approach was employed. Initially, the research team immersed themselves in the data to
allow key topics to emerge organically, without preconceptions (Bernard, 2006) . Codes were
then developed based on the emerging themes. Data was imported into Dedoose
(Dedoose, 2022)
and double-coded by two members of the research team. Any discrepancies in coding
were discussed thoroughly until consensus was reached, with new codes added as
necessary. All transcripts were re-coded using the finalized coding scheme. To address the
primary research question, all data coded under themes related to feral dogs were
summarized in tables, and representative quotes were selected for presentation.

Results
Study population and cave areas
The study covered 3.91 km² across the four periurban localities. Between September 2019
and March 2022, field teams recorded 9,114 cave observations. In nearly every monthly
visit, at least one cave with evidence of feral dog activity was detected, except for two
months in one locality (San Luis Gonzaga D), which had the fewest number of farms even
though it was the largest locality. On average, 16.42 caves per month showed evidence of
feral dogs’ presence, primarily indirect (e.g. paw prints, claw marks, feces) with direct signs
(e.g. live dogs, litters) observed less frequently (mean=2.06 caves/month). Paw prints were
the most prevalent indirect indicator, found in 99% of observations containing evidence.
Claw marks were present in 91%. Direct sightings of dogs near caves were reported in 10%
of cave observations. A total of 46 litters of puppies, 166 dead dogs, 42 solitary adult dogs,
and 97 packs were recorded throughout the study. Within or in proximity to the caves, 8
litters of puppies, 25 observations of adult dogs inside caves, and 176 observations of dogs
near caves were documented. Estimated average litter size was 7 puppies; average pack
size was 11 individuals.

Temporal patterns in feral dog evidence

Following the onset of COVID-19 restrictions, a statistically significant and sharp decline was
observed in the number of caves with general, indirect and direct evidence of feral dog
presence, evaluated with simple GLMs. Median monthly caves count with general evidence
fell from 31.5 to 10 (p<0.001), while direct evidence declined from 2 to 0 (p<0.001) (Table 1).
Additional indicators such as litters (p<0.001), solitary dogs (p<0.001), dead dogs (p<0.001),
and packs (p<0.001) also showed statistically significant reductions. However, no significant
differences were found in the number of puppies per litter (p=0.542) or the number of dogs
per pack (p=0.142) (Table 2). In addition to measures of central tendency, bar graphs show
these trends clearly (Fig 4). A gradual decrease was observed in the number of litters (Fig.
4A), dead dogs (Fig. 4B), and packs (Fig. 4D) at the onset of the pandemic restrictions,
along with a substantial drop in the number of live dogs (Fig. 4C).
 Table 1. Number of caves with direct and indirect evidence of feral dogs before and during
the COVID-19 pandemic restrictions.
COVID-19 pandemic restrictions
Variables p-value *
Without restrictions With restrictions
Caves with any evidence - monthly median (IQR) 31.50 (22.75 - 35.75) 10.00 (4.75- 19.00) <0.001
Caves with indirect evidence - monthly median (IQR) 31.50 (22.75 - 35.75) 10.00 (4.75 - 19.00) <0.001
Caves with direct evidence - monthly median (IQR) 2.00 (0.25-6.75) 0.00 (0.00-2.00) <0.001
* Wilcoxon rank sum test; IQR: Interquartile range.

Table 2. Population indicators of feral dogs in caves, trails, and open fields before and
during the COVID-19 pandemic restrictions.
COVID-19 pandemic restrictions
Variables p-value *
Without restrictions With restrictions
Litters - monthly median (IQR) 0.50 (0.00-1.00) 0.00 (0.00-0.00) <0.001
Puppies - monthly median (IQR) 6.00 (3.00-13.00) 5.00 (3.00-6.75) 0.542
Dead dogs - monthly median (IQR) 2.50 (0.00-6.00) 0.00 (0.00-1.00) <0.001
Solitary live dogs - monthly median (IQR) 0.50 (0.00-2.00) 0.00 (0.00-0.00) <0.001
Dog packs - monthly median (IQR) 2.00 (0.00-3.00) 0.00 (0.00-1.00) <0.001
Number of dogs in packs - monthly median (IQR) 11.50 (6.75-18.75) 7.00 (4.00-16.00) 0.142
* Wilcoxon rank sum test; IQR: Interquartile range.
Figure 4. Monthly counts of A) litters, B) dead dogs, C) live dogs and D) packs.
The red vertical line indicates the start of the lockdown due to the COVID-19
pandemic.

The ITS models for presence of feral dogs showed a statistically significant decrease in the
number of caves with general evidence, indirect evidence, and direct evidence at the onset
of COVID-19 restrictions (p<0.010, p=0.012, and p=0.002, respectively; Table 3). Compared
to the pre-restriction period, at the onset of restrictions, caves with general evidence
decreased by 42%, caves with indirect evidence decreased by 41%, and caves with direct
evidence decreased by 76% (Fig. 5). Temporal trend variables— “visit number since the
start of the study” and “visit number since the onset of restrictions”—were significant for
direct evidence. Prior to restrictions, caves with direct evidence increased by 25% per
month. After restrictions were implemented, this trend reversed, with the number of caves
decreasing by 25% per month. Temporal variables were not statistically significant for
general and indirect evidence. In these full multivariable models, locality was also a
significant predictor. For all types of evidence, each locality experienced a more pronounced
decline following the onset of pandemic restrictions compared to APSIL, the most densely
populated human settlement in the study area.

Figure 5. Estimates and trends in TRI analyses before and after COVID-19 pandemic
restrictions for A) Indirect evidence and B) Direct evidence of dogs in the cave dogs area.
Table 3. Parsimonious model for the number of caves with direct evidence of feral dogs in periurban areas of Arequipa.
Variables Direct evidence* Indirect evidence* General evidence*
p-
Est 95% CI p-value Est 95% CI p-value Est 95% CI value
Monthly visit since study (0.93 - (0.93 -
1.02 0.669 1.02 0.677
beginning 1.25 (1.06 - 1.49) 0.015 1.11) 1.11)
(0.39 - (0.39 -
0.59 0.012 0.58 0.010
COVID-19 restrictions 0.24 (0.10 - 0.56) 0.002 0.87) 0.86)
Monthly visit since restrictions (0.87 - (0.87 -
0.95 0.238 0.95 0.247
beginning 0.75 (0.63 - 0.90) 0.003 1.03) 1.04)
Locality
APSIL REF - - REF - - REF - -
San Luis Gonzaga A <0.001 (0.79- (0.89-
1.03 0.845 1.02 0.876
0.24 (0.14- 0.41) 1.33) 1.32)
El Roble <0.001 (0.51 - (0.51 -
0.66 0.002 0.66 0.002
0.13 (0.07 - 0.23) 0.87) 0.86)
San Luis Gonzaga D <0.001 (0.52- (0.53 -
0.72 0.035 0.72 0.039
0.02 (0.001- 0.09) 0.98) 0.98)

* considering the number of caves visited in each location and each month as the offset.
Human and Feral Dog Interactions

We recruited 64 farmers during the door-to-door visits; when called, only 41 accepted to
participate and were interviewed. All the interviewees raised or had raised farm animals and
20 of them were women. The interviewed farmers mentioned that the pandemic affected the
prices of their animals, forcing them to reduce the number of animals they were raising—or,
in some cases, to stop raising them altogether. This decline in small-scale animal husbandry
not only impacted household income and food security, but also had potential ecological
consequences, such as reduced availability of organic waste that previously served as a
food source for feral dogs.

"I used to raise a larger number of animals, mainly pigs. Now, with the pandemic,
business has declined and there's not much demand, so the number of animals has
decreased." - Man, San Luis Gonzaga A

"I used to raise my chickens and ducks, but after the pandemic, that started to
disappear." - Woman, APSIL

Farmers noted that animal farming can serve as a food source for feral dogs, due to
inadequate disposal of animal carcasses, which are easily accessible to roaming dogs.

"Wild dogs are large and hang around the pig farms; sometimes pigs are thrown into
the [dry] water channels, and the dogs feed on them." - Man, ASPIL

The reduction in solid waste during the pandemic, a primary food source for dogs, was
associated with increased starvation among feral dog populations. Unexpectedly, despite
mostly unfavourable discussions regarding feral dogs, one woman reported that some
people actually felt sorry for them:

“There were other neighbours who would go all the way to the market to bring
scraps, intestines, to help feed the dogs because they were completely skinny—they
looked pitiful. During the lockdowns there was no garbage to feed the dogs.” -
Woman, APSIL

Multiple dimensions of human–animal conflict emerged during the in-depth interviews with
local farmers, mostly describing concerns about safety of themselves, their children, or their
farm animals encountering hungry feral dogs. The identified dimensions and their
descriptions are presented in Table 4. Interestingly, fecal contamination was not mentioned,
despite the widespread presence of feces observed in the study area.

Table 4. Dimensions of human-dog conflict in the periurban areas of Arequipa, Peru, 2015.

Dimensions Quotes

Attacks on livestock "I mostly stay here on my farm because that's where my
animals are—I have to watch over them, because if I'm
not here, there are also wild dogs around that come to eat
the animals."
 “One day, the neighbor forgot to close the pen and left it
open. Around 1 a.m., I heard the pigs screaming. The next
morning, I saw that all eight pigs had been eaten.”

Attacks on people "Our children go to the store to buy things, and sometimes
the dogs are hungry… they go with a stick for
protection…"

"I walk through where the dogs are, even with my child. I
have a six-year-old, and we have to pass through there
holding a stone in hand for protection."

Neglect and abandonment "The dogs were abandoned, they had their puppies who
grew up, and so on. And these dogs survive by stealing
animals—chickens, sometimes they even come in and eat
the cats."

Scavenging and waste "The dogs usually come around here because, as this is a
dispersion farm, garbage and food waste are sometimes discarded,
which attracts wild dogs that then gather in groups."

Fear and reduced mobility “You walk down the street and see dogs everywhere."

"I avoid walking through those places because it scares

me."

Animal abuse and culling "We scare them off with arrows and sticks, and we chase
them with rocks so they stop coming around."

"They [other farmers] have tried to poison them, but the

dogs don’t eat it. A neighbor gave them poison and said
the wild dogs didn’t eat it, but his own puppies ended up
dying."

Discussion
Since 2015, a canine rabies epidemic has continued in Arequipa, Peru, with 394 confirmed
cases reported by April 2025 (DGE, 2015, 2016, 2017, 2018, 2019, 2021, 2022, 2023). To
effectively address this crisis, it is essential to understand the dynamics of the city’s dog
population, particularly feral dogs, which operate outside traditional ownership and exhibit
distinct ecological behaviors
(Boitani et al., 2016; Dürr & Ward, 2014; Fanaro et al., n.d.; Miternique & Gaunet, 202
Although previously unreported, a study
published in this special issue confirms the presence of feral dogs in periurban areas. These
areas are highly vulnerable to rabies due to high dog bite rates, low vaccination coverage,
limited health service access, and a dense population of free-roaming dogs
(Castillo-Neyra, Brown, et al., 2017; Castillo-Neyra et al., 2019; Ca
. The presence of feral dogs exacerbates
these risks. Although feral dog populations are often assumed to be negligible relative to
owned free-roaming dogs due to presumed low carrying capacity—defined as the maximum
population size a habitat can support—our findings challenge this assumption
(Boitani & Ciucci, 1995; Nicholson, 2014; Santos B
. We demonstrate that peri-
urban areas of Arequipa can support substantial feral dog populations, although these
populations remain sensitive to fluctuations in local carrying capacity.

Evidence gathered in and around caves confirms that periurban regions on the outskirts of
Arequipa city offer high habitat suitability for feral dogs. During the study, over 90% of
surveyed caves contained paw prints and scratch marks, while more than half had feces,
indicating frequent use. Direct sightings were limited—likely due to the dogs’ avoidance of
humans and daytime survey hours—but key indicators such as pups, solitary individuals,
packs, and carcasses were documented. The presence of pups points to ongoing
reproduction, while dog packs suggest structured social behavior. Carcasses, averaging 3.5
per month, further indicate regular access to farm animals. Globally, feral dogs have been
documented in diverse countries like Chile, Brazil, Italy, and India
(Hennelly et al., 2015; Young et al., 2011)
. Feral dogs in Arequipa city show similar behavioral traits reported in
other countries—such as avoidance of humans and group living
(Boitani & Ciucci, 1995; Spotte, 2012)
. Pack sizes ranged from 2 to 22 dogs, with an average of 5, aligning with
international data
(Boitani et al., 2016; Boitani & Ciucci, 1995; Miternique & Gaunet, 2020; Smuts, 2010)
. Dogs feed primarily on household waste dumped in open fields and
supplement their diets by preying on backyard livestock or scavenging from carcasses
(Castillo-Neyra et al., 2025) , behaviors that mirror those reported in other countries
(Young et al., 2011)
and have prompted community responses such as fencing, traps, poisoning,
and guard dogs—highlighting the economic and social impact of these animals.

The COVID-19 pandemic introduced drastic ecological changes that affected these dogs'
food system, behavior, and interactions with humans. Lockdowns reduced organic waste
production and disrupted commercial supply chains for animal feed. As a result, backyard
farmers became more dependent on household waste to feed livestock, intensifying
competition with feral dogs. Reports of increased livestock predation during this time reflect
how reduced environmental carrying capacity may have forced feral dogs to shift their
behavior. In our study, we observed a decline in both live sightings and cave-related
evidence during the pandemic, consistent with a reduction in food sources and altered local
ecological relationships.

Feral dogs represent an underrecognized public health challenge in rabies-endemic areas.

In Arequipa, 12.4% of periurban residents report being bitten by dogs annually—the highest
rate recorded in Latin America (De la Puente-León et al., 2020) . Of those bitten, 84% say the
dog was unfamiliar, and 73% do not seek medical care. At the time that study was
conducted, feral dog populations were unrecognized in the area and the proportion of dog
bites associated to feral dogs was not estimated. Even if recognized, in those same areas
the proportion of unrestricted owned dogs is also very high making the estimation very
challenging (Castillo-Neyra et al., 2020) . Local media frequently report attacks, even fatal, on
vulnerable populations, such as children and intoxicated individuals. Additionally, sightings
and conflicts involving feral dogs have been reported in multiple districts, including Cayma,
Cerro Colorado, Yura, and Mariano Melgar
(Diario Correo, 2022; Diario La República, 2019a, 2019b, 2021)
, indicating that this is not an isolated issue.
Concerningly, current dog rabies vaccination programs focus solely on owned dogs and
surveillance activities are restricted to areas with human settlements, excluding feral dog
populations
(Norma Técnica de Salud Para La Prevención y Control de Rabia Humana En El Perú, 2017; Raynor et a
. Because these dog populations are not recognized and
quantified, they are not accounted in the calculations of rabies vaccine coverage causing
overestimation of the actual vaccine coverage of dogs at risk (e.g., free-roaming dogs). Our
data suggest untapped opportunities to work with feral dogs. Periurban dwellers are already
in conflict with these dog populations. This offers a critical opportunity to engage them for
passive disease surveillance and pathogen monitoring (Douangngeun et al., 2017) .
Importantly, the COVID-19 pandemic demonstrated how quickly environmental changes can
shift the dynamics of feral dog populations. This reinforces the need for comprehensive
public policies—particularly around solid waste management—not just as a sanitation issue,
but as a vital component of zoonotic disease prevention and human-wildlife coexistence
(Sangkachai et al., 2024; Vourc’h G et al., 2022) .

The persistence of rabies in Arequipa, the ecological adaptability of feral dogs, and their
increasing interactions with humans and wildlife call for an urgent, integrated response.
Effective interventions must include targeted surveillance, inclusion of feral dogs in rabies
control efforts, and robust environmental management to reduce the conditions that allow
these populations to thrive.

Our study had some challenges. The sensory abilities and behavioral nature of feral dogs
allow them to detect human presence from a distance, potentially causing them to flee or
leave their shelters before detection. Due to this limitation, we supplemented our
observations with indirect evidence of dog presence, including feces, tracks, scratch marks,
and nearby food and water sources. These indirect signs varied in reliability; for instance,
fresh feces and tracks are considered more dependable indicators, particularly under the
climatic conditions of Arequipa, where such evidence rapidly degrades. While indirect
evidence helps infer site use by feral dogs, it does not provide precise information about the
timing of their presence, making direct observations more conclusive. Similarly, the
identification of deceased dogs within the study area did not allow for confirmation of their
feral status, as they could have been unowned roaming dogs, community dogs, or even
owned dogs with outdoor access. Consequently, the actual mortality of feral dogs may be
overestimated. However, this potential overestimation would have affected both pre-
pandemic and pandemic-era data equally. During our visits, we noticed that some caves had
been destroyed which could cause overestimation of occupancy. We adjust for that by using
the number of caves at each visit as an offset. Although our study implies interaction of feral
dogs and humans and other animals base on proximity, it did not permit evaluation of
interspecific interactions, such as with owned dogs or native wildlife like foxes, before and
during the pandemic.

The presence of feral dog increases the vulnerability of local human communities, especially
in a region that already reports the highest rates of dog bites in Latin America, has extremely
low socio-economic status, and lacks access to healthcare, including PEP. Based on our
findings, environmental management emerges as a key factor in modulating the interaction
between human settlements and feral dog populations; improved practices around animal
husbandry and solid waste management could offer an efficacious and sustainable approach
to reduce the problems associated with feral dogs. However, it remains unclear whether the
 observed reduction in occupied caves during the pandemic is due to mortality or migration to
other areas. If migration is occurring, it raises concerns about the potential spread of
pathogens such as the rabies virus into rabies-free zones, either urban or the wilds, as well
as the ecological impacts of interspecies interactions and predation on native wildlife by feral
dogs. Given the complexity of the periurban ecology, any environment-based intervention
intended to reduce the carrying capacity for feral dogs should be accompanied by monitoring
of intended and also unintended consequences. It is critical to integrate these newly
identified dog subpopulations in the dog rabies control program to move towards the goal of
eliminating dog-mediated human rabies by 2030 (WHO et al., 2018) .

Funding

MDL was a doctoral student studying an Epidemiological Research Doctorate at Universidad

Peruana Cayetano Heredia under FONDECYT/CIENCIACTIVA scholarship EF033-235-
2015 and supported by training grant D43TW007393 awarded by the Fogarty International
Center of the US National Institutes of Health. RCN was supported by NIH-NIAID grants
K01AI139284 and R01AI168291. RCN, EWD, and VPZ were supported by NIH-Fogarty
International Center grant D43TW012741.

Acknowledgments

We extend our gratitude to the officials and staff of GERESA-Arequipa, the Arequipa-
Caylloma Health Network, and the INS Arequipa Rabies Reference Regional Laboratory for
their work and support in fighting rabies. We are also grateful to the families who welcomed
us into their homes and allowed us to learn from their experiences and realities. Finally, we
acknowledge the support of the U.S. Centers for Disease Control and Prevention (CDC) and
Mission Rabies, who provided access to the World Veterinary Service (WVS) mobile
application to collect part of our data.

Conflict of interest statement

The views expressed in this publication do not necessarily reflect the views of the National
Center for Epidemiology, Prevention, and Disease Control Center (CDC, Peru).

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Supplement 1. Form for Recording Information on Cave Area Visits in the WVS Application
Nro Pregunta Tipo de pregunta Respuestas

PREGUNTAS GENERALES

1 Zona DropDown List

APSIL

San Luis zona A

San Luis zona D

El Roble

Otra zona

Completar 1a si marcó "Otra zona" en la pregunta 1.

1a Especificar la zona Text

2 Fecha Datepicker

3 Hora Timepicker

4 Tipo de entrada

Características del sitio

Cueva

Perro muerto

Crías de perros

Perro vivo
 Jauría

Observaciones o hallazgos
adicionales

PERRO VIVO

Si marcó "PERRO VIVO" en la pregunta 4, completar las preguntas 5-14.

5 Foto 1 del perro vivo Image

6 Foto 2 del perro vivo Image

7 Tamaño del perro Radio Button

Pequeño

Mediano

Grande

8 Edad en años Numeric

9 Edad en meses Numeric

10 Condición corporal DropDown List

1 (caquéctico)

2 (bajo peso)

3 (peso ideal)

4 (sobrepeso)

5 (obeso)

No puedo determinarlo

11 Calidad del pelaje DropDown List

Regular

Brilloso

No puedo determinarlo

Descuidado

12 Raza aparente DropDown List

Pura

Características distintivas
 Mestizo o criollo

No puedo determinarlo

Completar 12a si marcó "PURA" en la pregunta 12.

12a Especificar raza DropDown List

Beagle

Collie

Boxer

Bull Terrier

Bulldog

Caniche/Poodle

Carlino/Pug

Chihuahua

Chow Chow

Cocker

Dálmata

Dobermann

Dogo

Golden Retriever

Labrador

Pastor Alemán

Pekinés

Pitbull

Rottweiler

Samoyedo

San Bernardo

Husky Siberiano

Otro

Completar 12b si marcó "OTRO" en la pregunta 12a.

12b Especificar otra raza Textbox

 Completar 12c si marcó CARACTERÍSTICAS DISTINTIVAS en la pregunta
12.

12c Especificar raza DropDown List

Beagle

Collie

Boxer

Bull Terrier

Bulldog

Caniche/Poodle

Carlino/Pug

Chihuahua

Chow Chow

Cocker

Dálmata

Dobermann

Dogo

Golden Retriever

Labrador

Pastor Aleán

Pekinés

Pitbull

Rottweiler

Samoyedo

San Bernardo

Husky Siberiano

Otro

Completar 12d si marcó "OTRO" en pregunta 12c.

12d Especificar otra raza Textbox

13 ¿El perro está asociado a alguna casa? Radio Button

 Sí

No

14 Observaciones adicionales Textbox

JAURÍA

Si marcó "JAURÍA" en la pregunta 4, completar de la pregunta 15-20.

15 Foto 1 de la jauría Image

16 Foto 2 de la jauría Image

17 ¿Cuántos perros hay? Numeric

18 ¿Dónde están los perros? Radio Button

Con una casa

Deambulando

Otro

Completar 18a si marcó "OTRO" en la pregunta 18.

18a Especificar Textbox

19 ¿Qué están haciendo los perros? DropDown List

Caminando

Buscando comida

Durmiendo

Peleando

Cuidando casa

Otro

Completar 19a si marcó "OTRO" en la pregunta 19.

19a Especificar Textbox

20 Observaciones adicionales Textbox

 CRÍAS DE PERROS

Si marcó "CRÍAS DE PERROS" en la pregunta 4, completar de la pregunta

21-27.

21 Foto 1 de camada Image

22 Foto 2 de camada Image

23 Número de cachorros Numeric

24 Edad de los cachorros Radio Button

0-10 días

11 días - 3 semanas

3-8 semanas

> 8 semanas

25 ¿Están en una cueva? Radio Button

Sí

No

26 ¿La mamá está cerca? Radio Button

Sí

No

27 Observaciones adicionales Textbox

PERRO MUERTO

Si marcó "PERRO MUERTO" en la pregunta 4, completar de la pregunta

28-37.

28 Foto 1 de perro muerto Image

29 Foto 2 de perro muerto Image

30 Estado de descomposición DropDown List

Definiciones de los estados de descomposición: Fresco (sin decoloración

de piel o actividad de insectos), Temprana (decoloración gris o verdosa
de la piel, hinchazón, pérdida de pelo), Avanzada (descomposición
húmeda de los tejidos, alta actividad de insectos, exposición de huesos
de menos de la mitad del cuerpo, momificación), Casi extrema (huesos
con algunos fluidos o tejido cubriendo menos de la mitad del cuerpo,
huesos secos) y Extrema (esqueleto)
 Fresco

Temprana

Avanzada

Extrema

Casi extrema

31 ¿Su cadáver ha sido aparentemente descartado por personas? Radio Button

Sí

No

32 ¿Su cabeza ha sido removida? Radio Button

Sí

No

33 ¿Hemos colectado muestra de cerebro? Radio Button

Sí

No

33a Código de la muestra colectada Textbox

34 ¿Hemos marcado al perro? Radio Button

Sí

No

34a ¿Con qué hemos marcado al perro? Textbox

35 Escriba el código de la última cueva visitada Textbox

¿A cuántos metros aproximadamente se encuentra de la última cueva

36 visitada? Numeric

37 Observaciones adicionales Textbox

CUEVA

Si marcó "CUEVA" en la pregunta 4, completar de la pregunta 38-46.

38 Número de cueva

39 ¿Esta cueva existe? Multi Select

Sí, existe
 No, está derrumbada

No existe

40 Foto 1 de la cueva Image

41 Foto 2 de la cueva Image

42 ¿Hay evidencia de que los perros usen la cueva? Radio Button

Sí

No

No estoy seguro

Completar 42a si marcó "NO ESTOY SEGURO" en la pregunta 42.

42a Especificar por qué no estás seguro Textbox

43 Marca la posible evidencia de que los perros usen la cueva Multi Select

Huellas

Arañazos

Heces frescas

Heces secas

Camada

Perro(s) en cueva

Perros cerca a la cueva

Restos de animales

Fuente de comida

No hay evidencia

Otro

Fuente de agua

Completar 43a si marcó "OTRO" en la pregunta 43.

43a Especificar otra evidencia Textbox

44 Descripción de los restos animales Multi Select

Huesos

Pelo

Carcasa fresca
 Oveja o cabra

Pollo

Perro

Otro

Completar 44a si marcó "OTRO" en la pregunta 44.

44a Especificar otro resto de animal Textbox

En la pregunta siguiente solo marcar la opción "Perro" si el perro está

siendo consumido como comida de otros canes. De lo contrario (si es un
perro muerto que no es fuente de comida), no marcar esa opción.

45 Especie animal probable Multi Select

Pollo

Oveja

Cabra

Perro

Gato

Cuy

Otro

No puedo determinarlo

Completar 45a si marcó "OTRO" en la pregunta 45.

45a Especificar animal probable Textbox

46 Observaciones adicionales Textbox

CARACTERÍSTICAS DEL SITIO

Si marcó "CARACTERÍSTICAS DEL SITIO" en la pregunta 4, completar de la

pregunta 47-51.

47 ¿Qué has observado de los siguientes en este sitio? Multi Select

Fuente de agua

Basura

Recipiente de agua

Recipiente de comida
48 Número de perros Numeric

49 Número de jaurías Numeric

50 ¿Cuál es el tipo de suelo? DropDown List

Arenoso

Rocoso

Arcilloso

Agrícola

Otro

Completar 50a si marcó "OTRO" en la pregunta 50.

50a Especificar tipo de suelo Textbox

51 Observaciones adicionales Textbox

HALLAZGOS ADICIONALES

Si marcó "HALLAZGOS ADICIONALES" en la pregunta 4, completar de la

pregunta 47-51.

52 Foto 1 Image

53 Describir Textbox

Supplement 2.

Model Without offset With offset

Estimate (95% CI) -0.76 (-1.09 - -0.45) -0.90 (-1.16 - -0.66)
Standard Error 0.162 0.128
p value 2.34E-06 1.38E-12
AIC 844.4 790.16
BIC 852.61 798.37

Comparison between two generalized linear models (GLMs): one simple model and another
including the number of visited caves as an offset. Both models use the number of caves
with general evidence as the response variable and the presence of restrictions as the
explanatory variable. The AIC comparison shows that the model with the offset provides a
substantially better fit, with a difference of 50 AIC units in favor of the offset model.
According to this model, the log of the number of caves with general evidence is 0.90 units
lower under restriction conditions compared to before the restrictions.
Similar results were observed when using the number of caves with direct evidence and the
number of caves with indirect evidence as response variables.