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Mammal Review ISSN 0305-1838

REVIEW

Puma–livestock conflicts in the Americas: a review of the

evidence
Maria de las Mercedes GUERISOLI* GECOBI (Grupo de Genética y Ecología en Conservación y
Biodiversidad), División Mastozoología Museo Argentino de Ciencias Naturales “Bernardino
Rivadavia”, Angel Gallardo 490, Buenos Aires, 1405, Argentina and Departamento de Biología,
Bioquímica y Farmacia e Instituto de Ciencias Biológicas y Biomédicas del Sur, Universidad Nacional
del Sur - CONICET, San Juan 670, Bahía Blanca, 8000, Argentina and S.P.E.C.I.E.S, P.O. Box 7403,
Ventura, California, USA. Email: mariadelasmercedesguerisoli@gmail.com
Estela LUENGOS VIDAL Departamento de Biología, Bioquímica y Farmacia e Instituto de Ciencias
Biológicas y Biomédicas del Sur, Universidad Nacional del Sur - CONICET, San Juan 670, Bahía Blanca,
8000, Argentina. Email: eluengos@gmail.com
Nicolás CARUSO Departamento de Biología, Bioquímica y Farmacia e Instituto de Ciencias Biológicas y
Biomédicas del Sur, Universidad Nacional del Sur - CONICET, San Juan 670, Bahía Blanca, 8000,
Argentina. Email: nccaruso@gmail.com
Antony J. GIORDANO S.P.E.C.I.E.S, P.O. Box 7403, Ventura, California, USA. Email: species1@hotmail.com
Mauro LUCHERINI Departamento de Biología, Bioquímica y Farmacia e Instituto de Ciencias Biológicas
y Biomédicas del Sur, Universidad Nacional del Sur - CONICET, San Juan 670, Bahía Blanca, 8000,
Argentina. Email: lucherinima@yahoo.com

Keywords ABSTRACT
American continent, conflict mitigation,
cougar, human–wildlife, large Felidae, 1. Loss of livestock is one of the greatest sources of conflict between humans
livestock depredation, Puma concolor and large felids worldwide. The puma Puma concolor is the most widespread
apex predator in the Americas, and conflicts between this felid and humans
*Correspondence author
are common throughout its geographical range. In response to predation on
Received: 19 March 2019
livestock, humans persecute and hunt pumas.
Accepted: 21 July 2020 2. We identified the main environmental and anthropogenic variables that define
Editor: DR puma–livestock conflict areas in the Americas as 12 conflict predictor vari-
ables, and explored the techniques proposed to mitigate conflicts between
doi: 10.1111/mam.12224 the puma and livestock producers.
3. We conducted a systematic search and subsequent review of the scientific
literature and found 92 publications on puma–livestock conflicts. Through
single-variable analyses and generalised linear models (GLM), we identified
which of the 12 conflict predictors were most predictive of the occurrence
of predation.
4. The single-variable analyses showed that high livestock density (goat, sheep,
and cattle), low latitudes, low habitat steepness, low co-predator richness,
high distance to habitat (shrub), and high distance to roads characterised
areas with conflict. The binomial GLM indicated that areas with conflicts
were primarily located in the temperate southern hemisphere and characterised
by densities of livestock. The most frequently cited conflict mitigation tech-
niques were ‘improving livestock management’, ‘predator control’, and the
‘use of fencing’.
5. Although our knowledge about the puma and its relationships with human
communities has improved, there are wide geographical gaps, and many facets
of puma–livestock conflicts are still little understood. Scientists should work

Mammal Review (2020) Mammal Review (2020) © 2020 The Mammal Society and John Wiley & Sons Ltd. 1
Puma-livestock conflicts: a continental review M. M. Guerisoli et al.

with local stakeholders to generate reliable information regarding the ecologi-

cal and societal consequences of puma–livestock conflicts, and to develop
conflict mitigation techniques that could facilitate the coexistence of pumas
and humans.

RESUMEN EN ESPAÑOL
1. La predación de animales domésticos es una de las más importantes fuentes
de conflictos entre humanos y grandes felinos a nivel mundial. El puma Puma
concolor es el predador tope más ampliamente distribuido en las Américas, y
los conflictos entre este felino y los humanos resultan ser comunes a lo largo
de toda su distribución geográfica. Como respuesta a los eventos de predación
de animales domésticos, el hombre persigue y caza a esta especie.
2. Identificamos las principales variables ambientales y antrópicas, que describen
las áreas de conflicto entre el puma y la ganadería en el continente Americano,
como 12 variables predictoras del conflicto, y estudiamos las técnicas propuestas
para mitigar los conflictos entre el puma y los productores ganaderos.
3. Realizamos una búsqueda sistemática, y posterior revisión, de la literatura cientí-
fica y encontramos 92 publicaciones sobre conflictos puma-ganado. A través
del análisis individual de las variables y modelos lineales generalizados (GLM),
identificamos cuáles de las 12 variables caracterizaban las áreas de conflicto.
4. Los análisis individuales de las variables mostraron que la densidad de ganado
(caprino, ovino y bovino), la latitud, la pendiente del hábitat, la riqueza de
especies de copredadores, la distancia al hábitat (arbustales) y la distancia a
las carreteras caracterizaron las zonas con conflicto. El GLM binomial indicó
que las áreas con conflictos se ubicaron principalmente en zonas templadas
del hemisferio sur y se caracterizaron por altas densidades de ganado. Las
técnicas de mitigación citadas con mayor frecuencia fueron "mejorar el manejo
del ganado", "control de predadores" y "uso de corrales".
5. Si bien el conocimiento sobre el puma y sus relaciones con las comunidades
humanas ha sido caracterizado, existen amplias brechas geográficas, y muchas
facetas de los conflictos que aún se desconocen. Los científicos debemos
trabajar con los actores locales para generar información confiable sobre las
consecuencias ecológicas y sociales del conflicto entre el puma y el ganado,
y, así, desarrollar técnicas de mitigación que podrían facilitar la coexistencia
entre el felino y la producción.

INTRODUCTION predation of livestock (Treves & Karanth 2003). In an

Human–wildlife conflicts are defined as actions, by hu-
mans or wildlife, that have negative effects on the other;
these include actual threats posed by wildlife to human
life or economic resources, among others, and the per-
ception that wildlife threatens human safety, food, or
property (Nyhus 2016). Thus, these interactions may
negatively affect humans (e.g. via vehicle collisions;
Gunson et al. 2011), their resources (livestock depreda-
tion, crop raiding; Conover & Decker 1991, Inskip &
Zimmermann 2009), and/or wildlife (carnivore persecu-
tion; Mateo-Tomás et al. 2012). Large carnivores star
in human–wildlife conflicts worldwide, because of their
attempt to reduce depredation losses, people apply lethal
and/or non-lethal methods (van Eeden et al. 2018a, b).
Generally, non-lethal methods are more effective than
lethal methods in preventing carnivore predation of
livestock (Treves et al. 2016), and it is important to
address local cultural values and environmental condi-
tions in order to achieve long-term coexistence between
farmers and large carnivores (van Eeden et al. 2018a,
b, Ohrens et al. 2019a).
Large carnivores keep suffering from intense persecution
(Graham et al. 2005, Khorozyan et al. 2015), and conflicts
are enhanced by negative attitudes and perceptions of local
people towards carnivores (Mkonyi et al. 2017, Marchini

2 Mammal Review (2020) © 2020 The Mammal Society and John Wiley & Sons Ltd.
M. M. Guerisoli et al.
& Macdonald 2018). This is also true for jaguars Panthera
onca and pumas Puma concolor (e.g. de Azevedo & Murray
2007, Thornton & Quinn 2009, Zarco-González et al. 2013),
the only large felids in the Americas. Due to its extensive
geographic range and broad ecological niche, the puma is
the top predator in a variety of ecosystems of the American
continent (Nielsen et al. 2015). Throughout their range
(Fig. 1), pumas are persecuted and hunted by humans,
mainly because of livestock predation (e.g. Polisar et al.
2003, Peebles et al. 2013, Llanos et al. 2014, Guerisoli et
al. 2017). Pumas attack mostly cattle Bos taurus (Conforti
& de Azevedo 2003, Polisar et al. 2003), sheep Ovis aries
(Elbroch & Wittmer 2012, Guerisoli et al. 2017) and goats
Capra hircus (Mazzolli et al. 2002); predation by pumas
on horses Equus caballus, domestic Camelidae, and swine
Sus scrofa domesticus has been recorded more rarely. Faecal
analyses detected livestock remains in puma diet in 38%
of studies from Mexico to Brazil (Laundré & Hernández
2010). Although livestock remains typically occurred in
<10% of puma scats, higher percentages have been docu-
mented locally (Murphy & Ruth 2010). Monetary losses
resulting from pumas killing livestock may be important
(Palmeira et al. 2008, Zarco-González et al. 2012, Lucherini
et al. 2018) and may trigger intense persecution of this
species (Yáñez et al. 1986, Iriarte et al. 1990, Mazzolli et
al. 2002, Polisar et al. 2003). Only a few studies have quan-
tified retaliatory or preventative hunting, but the available
Fig. 1. Map of the continental Western Hemisphere showing the geographic range of the puma Puma concolor and the geographical locations of the
92 studies on puma–livestock conflict that were found in the systematic search and included in the analysis. The regions used in the analysis (North,
Central, and South America) are indicated.
Mammal Review (2020) © 2020 The Mammal Society and John Wiley & Sons Ltd. 
Puma-livestock conflicts: a continental review
information suggests that lethal control may heavily reduce
puma numbers and lead to their local ecological extinction
(Novaro et al. 2000). For example, in the Brazilian Amazonia,
110–150 individual pumas and jaguars were hunted annu-
ally due to livestock losses in an area of 34200 km2 (Michalsky
et al. 2006). In the Argentinean Espinal, 1.1–1.4 pumas
were killed/year/100 km2 in an area of 27000 km2 in retali-
ation to livestock depredation (Guerisoli et al. 2017).
Thus, identifying common patterns in conflicts between
pumas and livestock producers is fundamental for under-
standing this phenomenon, designing more efficient pre-
vention and mitigation techniques (Graham et al. 2005,
Miller 2015), and, ultimately, improving conditions for
the coexistence of humans and pumas (Castaño-Uribe et
al. 2016). A number of environmental variables may affect
the occurrence and frequency of puma attacks on livestock
locally, including vegetation type, elevation, topography,
density of livestock, distance to forest, land protection
status, human settlements, roads, water sources, and wild
prey availability (Mazzolli et al. 2002, Palmeira et al. 2008,
Rosas-Rosas et al. 2008, Zarco-González et al. 2013, Burgas
et al. 2014). In many locations, poor livestock husbandry
and management facilitate puma attacks on livestock
(Polisar et al. 2003, Inskip & Zimmermann 2009, Zarco-
González et al. 2018).
Although conflicts over livestock between humans and
pumas seem to be widespread and potentially threaten
3
Puma-livestock conflicts: a continental review
the conservation of pumas in both natural and anthro-
pogenic ecosystems, there is still no range-wide review on
this subject. We propose to: (1) assess the research efforts
dedicated to the study of puma depredation on livestock
throughout the Americas; (2) describe the environmental
and anthropogenic variables that characterise the presence
of conflict in a given area, and identify the main factors
that correlate with depredation; and (3) explore the tech-
niques chosen and applied to mitigate conflicts between
puma and livestock producers.
METHODS
Sourcing information and assembling data
We conducted a systematic search and review of the sci-
entific literature (Pullin & Stewart 2006) in English and
Spanish that addressed predation by pumas in the Americas
(Appendix S1a). Because several common names (i.e. puma,
cougar, mountain lion, panther) are used for Puma concolor
(Murphy & Ruth 2010), we included all of them in our
search and combined them (in both English and Spanish)
with the following search terms: ‘predation’, ‘livestock’,
‘diet’, ‘prey’, ‘attack’. We did not use the term ’conflict’
in our search because it is a broad term including preda-
tion on livestock, predation of pets, road kills, and attacks
on humans (see the International Union for Conservation
of Nature Species Survival Commission’s Human-Wildlife
Conflict Task Force: http://www.hwctf.org/). All scientific
literature published by December 2017 was searched using
Web of Science and Google Scholar. Following Inskip and
Zimmerman (2009), we also employed a ‘snowball’ sam-
pling approach (Goodman 1961) to identify and include
additional relevant literature matching our criteria and
listed in the reference section of articles and chapters
identified during our primary searches. We intentionally
excluded from our analyses unpublished reports, grey lit-
erature, and MSc and PhD theses.
Characterisation and geographical
distribution of the literature
Because puma distribution is expansive in widely varying
socio-cultural and economic contexts, we divided the
American continent into three broad-scale regions: North
America (defined here as Canada and the USA), Central
America (from Mexico to Panama), and South America
(all remaining countries; Fig. 1). To clarify whether re-
search effort on puma–livestock conflict was homogenously
distributed throughout the geographical range of the puma,
we compared the number of publications found in the
systematic search per region (observed value; Nielsen et
al. 2015) with the number expected per region, calculated
4 Mammal Review (2020) © 2020 The Mammal Society and John Wiley & Sons Ltd.
M. M. Guerisoli et al.
from the total number and the area of the puma’s range
in each region, by using a chi-square test with the func-
tion chisq.test of R (R Core Team 2017). To prevent bias
in the quantification of the total area occupied by pumas
in each region, we summed only the areas of those coun-
tries where studies occurred. In this analysis, we used the
publication as the sampling unit.
Characterisation and geographical
distribution of the presence and absence of
conflict
We defined puma–livestock conflict as ‘present’ when the
authors of a study, regardless of the source of informa-
tion, reported depredation of livestock by puma, and as
‘absent’ when a study did not report any livestock attacks
by pumas. We then analysed each study in which conflict
was present to extract information on the techniques em-
ployed to collect data and on all factors affecting the
presence of conflict. Whereas puma diet data derived from
kill-site inspection were considered for this review, we
excluded from the analyses the literature based on puma
scat analysis, because the presence of a prey item in faecal
samples could be due to opportunistic scavenging and
not, necessarily, to active predation (Bauer et al. 2005).
In order to understand whether conflict areas presented
a geographic pattern, we used the publication as the sam-
pling unit, and related the number of publications reporting
conflicts in each of the three regions (observed value)
with the total number of published papers found in our
systematic search per region (expected value). We then
applied a chi-square test, to test whether conflicts were
homogeneously distributed throughout the puma’s geo-
graphic range.
Protected areas and presence of conflict
Human–wildlife conflict tends to be minimal or absent
inside Protected Areas (PAs; Borg et al. 2016, Borón et
al. 2016). In order to test this for puma–livestock conflict,
we identified in the literature those studies that were car-
ried out inside PAs. We excluded from this analysis the
literature in which: the study area (StA) was an entire
state or country, the StA included both protected and
unprotected areas, and the study was carried out in a
PA, but conflict occurred in areas with livestock adjacent
to the PA. To quantify the size of the area covered by
PAs, we used the World Database on Protected Areas
(IUCN & UNEP-WCMC 2015). Finally, since sample sizes
were small, we used the fisher.test function of R (R Core
Team 2017) and carried out a Fisher test (Fisher 1922)
to test, through a contingency table, whether the frequency
of puma–livestock conflicts in the PAs was greater than
M. M. Guerisoli et al.
expected, based on the proportion of land area that was
protected within the geographic range of the puma. In
this analysis, we used the publication as the sampling
unit.
Livestock as prey: preferences and densities
To understand which livestock species were taken as prey
by pumas, we extracted, directly from the literature, which
livestock species were present in each StA, and, when
several species co-occurred, which was the livestock species
most commonly taken as prey by pumas. Additionally,
we noted the specified age of the animal or animals at-
tacked. We excluded from this analysis horses, domestic
Camelidae, and swine, because they were infrequently taken
in the studies we reviewed.
To estimate average livestock densities within the puma’s
geographic range, we extracted the densities of livestock
species from a continent-scale map (Table 1; FAO 2010)
overlaid with the puma’s distribution map. In this analysis,
we used the publication as the sampling unit.
Conflict predictors
Based on prior information associated with livestock pre-
dation by large felids, and particularly pumas (Mazzolli
et al. 2002, Rosas-Rosas et al. 2008, Zarco-González et al.
2013), we defined and considered a set of conflict predic-
tors and associated predictions (Table 1) that could be
used to compare areas where conflicts occurred with areas
where conflict was absent.
For this analysis, we used the StA as the sampling unit.
To map the locations covered by the literature, we ex-
tracted the geographical coordinates of the central point
of each StA directly from the text (if provided) or, when
coordinates were not provided, through online maps based
on the name of the location reported. In order to include
as many studies as possible, we assigned a mean central
set of coordinates to the literature referring to a large
region (e.g. a national park or reserve). When a given
publication reported data from several StAs located at least
approximately 100 km apart, geographical coordinates were
assigned to each one of those areas. Because large land-
scapes typically include ample variations in environmental
conditions, StAs ≥30000 km2 were not considered in sub-
sequent analyses. Additionally, in order to avoid overes-
timation of the presence or absence of conflict, when
literature sources sampled the same StA, we considered
only one of the sources.
For each StA, we extracted the values of the variables
(Table 1) that we used to predict the occurrence of puma–
livestock conflicts, by building a buffer area around the
StA location. We applied two buffer radii: 5 km (BA5),
Mammal Review (2020) © 2020 The Mammal Society and John Wiley & Sons Ltd. 
Puma-livestock conflicts: a continental review
thus creating an area of 78.5 km2 (i.e. approximately the
size of an average puma home range; Franklin et al. 1999,
Grigione et al. 2002, Elbroch & Wittmer 2012); and 10 km
(BA10), corresponding to an area of 314 km2 (i.e. an
area potentially large enough to contain a resident male,
some resident females, and transients; Quigley & Hornocker
2010). We used Geographic Information System software
(QGIS 3.0.2) to extract the values of the conflict predic-
tors in each buffer area (Table 1).
Statistical analysis of conflict predictors
We used Mann–Whitney U-tests to understand which of
the predictive variables were significantly associated with
conflict. One-tailed tests were performed because we ex-
pected a priori a directional relationship of each variable
under examination with the presence of conflicts. We used
the wilcox.test function of R (R Core Team 2017).
To understand what combination of conflict predictors
was related to the presence of conflict, we used generalised
linear models (GLM; Zuur et al. 2009). The Mann–Whitney
U-test results, indicating which variables differed signifi-
cantly between areas where conflict was present and absent,
and the visual inspection of the predictors’ box plots as-
sisted our selection of the predictors to include in the
GLM analysis. We first performed a Kendall correlation
analysis (Zar 1999) to identify and eliminate highly cor-
related variables (those with r ≥ .5) and thus reduce po-
tential problems due to multicollinearity among predictors
(Zuur et al. 2009). We then fit a set of different binomial
GLMs using the presence or absence of conflict as the
response variable (with ‘logit’ as the link function). Using
the function dredge (package ‘MuMIn’, R Core Team 2017),
we created a list of all possible models assuming only
additive effects for predictors. We ranked all models using
Akaike Information Criteria adjusted for finite samples
(AICc) and used ΔAICc and relative weights to evaluate
the relative importance of each model in the final set
(Burnham & Anderson 2002). We identified those models
with ΔAICc < 2 to be competitive with the top-ranked
model (Burnham & Anderson 2002, Caruso et al. 2016).
Finally, we analysed the 95% confidence intervals (CI) of
β coefficients (i.e. regression coefficients) of individual
covariates to evaluate their significance (i.e. when CI ex-
cluded zero; Zeller et al. 2011, Caruso et al. 2016).
Conflict mitigation
The use of mitigation techniques is fundamental to efforts
to promote and enhance coexistence between predators
and livestock production. In order to understand which
strategies can be more effective in mitigating puma–live-
stock conflicts, we extracted from each literature all the
5
Puma-livestock conflicts: a continental review M. M. Guerisoli et al.

Table 1. List of conflict predictors and predictions (in italics) on puma–livestock conflict, and description of the data extraction processes used. Sources
of information (references cited only in this Table) are presented in Appendix S1(b). Around the central point of each study area (StA), data were ob-
tained for two buffer areas (BA5, with a 5 km buffer radius, and BA10, with a 10 km radius) where appropriate.

Conflict predictor Predictions and reasoning Data extraction process for each study area

Livestock density Higher densities of livestock will characterise areas with conflict.
Reason: for puma–livestock conflict to occur, livestock needs to be
present. High density of livestock has been found to result in
conflicts between farmers and large felids (Woodroffe & Frank
2005, Peebles et al. 2013, Hoogesteijn et al. 2016a), including
pumas (Reyna-Sáenz et al. 2019).
Livestock richness A greater richness of livestock favours the occurrence of conflict.
Reason: the availability of a greater variety of livestock, differing in
sizes and habits, may favour predation by pumas.
Livestock diversity A greater diversity/richness of livestock will occur in areas with conflict. We used the mean numbers of cattle, sheep and goats
Reason: the availability of a greater variety of livestock, differing in
sizes and habits, may favour predation by pumas.
Human density The presence of puma–livestock conflict increases with increasing
human population density.
Reason: greater human density is usually associated with reductions in
wild prey numbers, which, in turn, may force carnivores to prey on
livestock (Halfpenny et al. 1991, Treves et al. 2002).
Habitat The presence of puma–livestock conflict is greater in areas where the To estimate, for BA5 and BA10, the intensity of habitat
fragmentation fragmentation of natural habitats is stronger.
Reason: heterogeneous, unfragmented natural habitat is an important
determinant of the persistence of wild carnivore prey (Crooks 2002),
thus habitat fragmentation may indirectly favour predation on
livestock (Mazzolli et al. 2002, Michalsky et al. 2006).
Habitat steepness Puma attacks on livestock are less likely in areas where habitat is
steeper
Reason: puma wild prey (guanaco Lama guanicoe, vicuña Vicugna
vicugna, bighorn sheep Ovis canadiensis) occupy high-elevation
areas and tend to avoid livestock (Pedrana et al. 2010, Schröder et
al. 2013, Borgnia et al. 2008); most domestic livestock are kept on
flat or gently sloping terrain. Additionally, pumas typically use gentle
slopes for hunting (Dickson et al. 2005, Dickson & Beier 2007).
Thus, we expected that areas with conflicts would be characterised
by low habitat steepness.
Co-predator Predation on livestock is more likely to occur in areas where pumas
richness co-occur with other large predators.
Reason: resource partitioning can be present where large carnivores
co-occur (Alexander et al. 2006). Black bears Ursus americanus,
grizzly bears Ursus arctos horribilis, wolves Canis lupus and jaguars
Panthera onca are dominant over pumas (Elbroch & Kusler 2018).
Thus, where these large carnivores occur, pumas have more limited
access to their preferred wild prey, and are more likely to switch to
livestock.
We used a continent-scale map of livestock density
(head/km2) with spatial resolution of 1 km2 (FAO
2010). We considered only cattle, goats, and sheep,
and excluded equines, swine and domestic
camelids, which are infrequently taken by pumas.
For each StA, we computed the mean value of
livestock density of the pixels within BA5 and BA10.
We used the mean numbers of cattle, sheep and goats
within BA5 and BA10 to estimate richness, we
counted the number of livestock species present
within BA5 and BA10 using the livestock continent-
scale map.
within BA5 and BA10 to calculate the Shannon
Diversity Index (Shannon & Weaver 1949). The value
of the index equals 0 when only one domestic
species is present and reaches its maximum when
the diversity of livestock is the highest. We used the
diversity function of R (R Core Team 2017).
We calculated mean human density inside BA5 and
BA10 from the global population density map (FAO
2015), where pixel values indicate the number of
persons/km2.
fragmentation, we used the magnitude given in the
habitat fragmentation map (Hoekstra et al. 2010),
which considers agriculture, urban infrastructure,
roads, and railroads as fragmenting features in the
landscape. Lower values indicate more fragmented
areas.
To estimate habitat steepness in each StA, we
calculated the difference between the maximum
and the minimum elevation for BA5 and BA10 using
Shuttle Land Elevation Data (Farr et al. 2007).
Although ruggedness, rather than steepness, has
been associated with puma habitat use (Burdett et
al. 2010, Dickson et al. 2013), most of the study
areas had low values of the Terrain Ruggedness
Index (Riley et al. 1999); thus we selected steepness
for our analyses.
We used the number of co-predator species in each
StA. Data on presence/absence of each co-predator
were from the Terrestrial Mammal Distribution Map
(IUCN 2014). The species considered were large
carnivores that prey on cattle: black bear, grizzly
bear, wolf and jaguar. The spectacled bear
Tremarctos ornatus was not included because its
range overlapped with a very small number of StA.

(Continues)

6 Mammal Review (2020) © 2020 The Mammal Society and John Wiley & Sons Ltd.
M. M. Guerisoli et al. Puma-livestock conflicts: a continental review

Table 1. Continued

Conflict predictor Predictions and reasoning Data extraction process for each study area
Distance to Predation on livestock is more likely to occur near areas with We calculated the linear distance (km) from each StA
habitat (trees vegetation such as trees and shrubs. to the nearest patch of trees or shrubs using the
and shrubs) Reason: puma occurrence is typically associated with areas with Global Land Cover SHARE map (FAO 2014).
semi-closed/closed vegetation (Scognamillo et al.2003, Guerisoli et
al. 2019), so ranches in proximity to these habitats are more
exposed to predation by pumas (Michalsky et al. 2006, de Azevedo
& Murray 2007, Inskip & Zimmermann 2009).
Distance to roads Proximity to roads favours the occurrence of conflict. We used the World Street Map (ESRI 2009) to
Reason: in natural landscapes, roads represent a major source of calculate the linear distance (km) from each StA to
anthropogenic disturbance (e.g. resulting in road-kills). In rangeland the nearest road. This map includes highways, major
areas, roads facilitate access for hunters and, indirectly, lead to a roads and minor roads.
decrease in wild prey populations. Thus, the proximity to roads may
indirectly be associated with greater predation on livestock by
carnivores (Treves et al. 2004, Miller et al. 2016).
Distance to Conflict is more frequent in areas far from human settlements. We calculated the linear distance (km) from each StA
human Reason: with the potential exception of some specific areas of the USA to the nearest human settlement (cities and towns)
settlement (e.g. southern California), large human settlements are incompatible using the World Topographic Map (ESRI 2013).
with the presence of dense puma populations, so puma–livestock
conflicts are unlikely to occur in the proximity of towns and cities
(Mazzolli et al. 2002, Inskip & Zimmermann 2009).
Latitude Conflicts are less frequent in the temperate regions of the American We used the angle (in degrees) from each StA to the
continent (at high latitudes) than in the tropics (low latitudes). Equator to indicate latitude.
Reason: in high-latitude regions, pumas find a greater availability of
their preferred wild prey, medium-sized ungulates, e.g. in North
America mule deer Odocoileus hemionus, white-tailed deer
Odocoileus virginianus, and bighorn sheep; in South America
guanaco (Anderson 1983, Murphy & Ruth 2010, Lucherini et al.
unpubl. data).

information on mitigation techniques that had been ap-
plied or proposed, for comparison with information from
most recent publications on human–carnivore conflict
mitigation.
RESULTS
Characterisation and geographic distribution
of the literature
The database created, based on our literature search and
selection process for the analyses, produced 92 original
literature sources, published between 1984 and 2017, de-
scribing studies that took place in North, Central, and
South America (Fig. 1). Among the sampling techniques
used, interviews and community/ranch surveys were the
most common method (n = 57), followed by field surveys
of predation (e.g. kill-site inspections; n = 40), radio-
telemetry (n = 18), and mortality records of pumas (n = 4).
In several papers, a combination of sampling techniques
was used (e.g. interviews and field surveys; Appendix S1).
The literature sources were associated with 14 countries,
corresponding to 61% of countries where pumas occur
and to 94% of the area covered by the puma’s geographic
range. We found no literature on puma–livestock conflict
from Belize, Honduras, El Salvador, Nicaragua, Guyana,
Suriname, French Guiana, Ecuador, and Paraguay (Fig. 1).
Most of the studies took place in South America (52%,
n = 48), followed by North America (32%, n = 29) and
Central America (16%, n = 15). Based on the area of
the puma’s geographic range in North America and Central
America, the observed number of studies from these re-
gions was greater than expected, while the opposite was
true for South America (χ2 = 10, df = 2, P < 0.005;
Fig. 2).
Characterisation and geographical
distribution of presence and absence of
conflict
Puma–livestock conflict was reported in 82% (n = 75) of
the literature included in our database, but only 42 studies
(56%) mentioned a factor, or factors, potentially conducive
to conflict. Proximity to dense vegetation (48%) and poor
or absent livestock husbandry (55%) were the most fre-
quently cited factors that were believed to lead to conflict.

Mammal Review (2020) © 2020 The Mammal Society and John Wiley & Sons Ltd.  7
Puma-livestock conflicts: a continental review
Additional important factors included scarcity of wild prey
(19%), proximity to water sources (19%), and high hu-
man densities (4.8%). Conflicts were reported more fre-
quently in studies originating from South and Central
America (92% and 93%, respectively); in North America,
conflict occurred in 59% of studies, but the differences
were not significant (χ2 = 4.3, df = 2, P = 0.1).
Protected areas and presence of conflict
Approximately 27% of the geographical range of the puma
has some form of protected land status (e.g. national park,
reserve). South America has the greatest percentage of
protected land area within the puma’s geographic range
(30%; n = 3792 PAs), followed closely by Central America
(23%; n = 1329), and lastly, North America (18%;
n = 7287). The percentage of studies mentioning puma–
livestock conflicts occurring inside PAs (37%, n = 26,
P < 0.05) was greater than expected based on the per-
centage of puma range comprised by PAs.
Overall, breeding livestock within PA boundaries was
unrelated to conflict occurrence, as conflicts were simi-
larly frequent inside (80%, n = 16) and outside (91%,
n = 40) PAs (P = 0.4). However, all the studies refer-
encing conflict in North America (n = 6) were located
outside PAs, whereas most studies inside PAs from Central
America and South America reported conflict (Fig. 3).
Livestock as prey: preferences and densities
Cattle were taken as prey in 52% of all StAs with con-
flict (n = 75), sheep in 37% and goats in 25%. To
understand which of the three livestock species was most
20000
18000
16000
14000
Area, in 1000 km2
12000
10000
8000
6000
4000
2000
0 0
North America
Fig. 2. Regional comparison between the percentage of papers found in the systematic search on puma–livestock conflicts (right axis), and the size of
the puma’s geographic range in each region (left axis).
8 Mammal Review (2020) © 2020 The Mammal Society and John Wiley & Sons Ltd.
M. M. Guerisoli et al.
commonly taken as prey by pumas, we extracted data
from the StAs with conflicts where the three species
co-occurred (n = 9). Sheep were the species most fre-
quently attacked in 56% of these areas, while goats were
the species most intensely taken as prey in 33% and
cattle in 22%.
Only 17 of 40 studies where predation on cattle oc-
curred specified the age of the animal attacked; these cases
reported that pumas mainly took calves (usually
<12 months old). Eight of 47 studies with predation re-
cords of sheep and goat specified that both juveniles and
adults were puma prey; none of the others provided details
on the age of individuals attacked.
Individual analyses of conflict predictors
Central America and South America had higher average
livestock densities within the puma distribution range than
North America. Cattle were the most widespread livestock
species throughout the three regions (Appendix S2). On
average, and for both BA5 and BA10, the total livestock
density was higher in those StA with puma–livestock conflict
than in areas where no conflict records were published
(Table 2). Livestock diversity and richness did not differ
between areas where conflict was present and absent
(Table 2).
Human population density tended to be greater around
the StAs where conflict was reported than in areas where
no conflict with pumas occurred (Table 2), but this vari-
ation was not significant (either for BA5 and BA10). Habitat
fragmentation did not differ between StAs where conflicts
occurred and those where no conflicts were reported in
the literature (Table 2). Habitat steepness: conflict areas
100
90
Size of the puma’s geographic
% of papers found in systematic search
range 80
Literature
70
60
n = 48
50
40
n = 29
30
n = 15 20
10
Central America South America
M. M. Guerisoli et al.
100
90
80
70
% of study areas
60
50
40
30
20
10
0
North Central
America America
Protected
Fig. 3. Percentages of studies found in the systematic search in which puma–livestock conflict was present and absent, within Protected Areas (left)
and outside Protected Areas (right), in each of the three regions in the Americas.
were less steep than areas without conflict; these results
were similar for BA5 and BA10 (Table 2).
In 48% of all StAs, pumas co-occurred with at least
one other species of large carnivore. In contrast to what
we expected, the median number of co-predator species
was significantly lower in conflict areas (Table 2). Although
the median distance to the closest patch of trees tended
to be lower in areas where conflict was reported than in
areas with no conflict, this difference was not significant
(Table 2). The median distance to the closest patch of
shrubs was greater for areas with conflict than for areas
where no conflict was reported (Table 2).
The median distance to roads in the areas with conflict
was significantly greater than in those areas where no
conflict occurred, for both BA5 and BA10 (Table 2). The
median distance to human settlements was similar in areas
with conflict and those with no conflict (Table 2). The
latitudes of the StAs ranged from: −50.96° to 52.07°. On
average, latitude differed between areas with and without
conflict (Table 2). StAs with conflicts were located closer
to the Equator, while more publications from areas reg-
istering no conflicts were located in the Northern
Hemisphere (Appendix S3).
Combined analysis of conflict predictors
From Table 2 and from box plots of the conflict predic-
tors (Appendix S3), we selected the variables which were
related to the presence and absence of conflict. The cor-
relation matrix showed that a correlation existed between
‘lat’ (latitude) and ‘copred’ (co-predators), for both BA5
and BA10 (Appendix S4). Thus, the explanatory variables
included in the binomial GLM were as follows: ‘dtot’
(livestock total density), ‘step’ (habitat steepness), ‘dshrub’
Mammal Review (2020) © 2020 The Mammal Society and John Wiley & Sons Ltd. 
Puma-livestock conflicts: a continental review
Conflict presence
Conflict absence
South North Central South
America America America America
No protected
(distance to shrub), ‘droad’ (distance to road), and ‘lat’
(latitude). Four of the top-ranked models showed
ΔAICc < 2, for BA5 and BA10 (Table 3). We followed
the principle of parsimony sensu Midlane et al. (2014)
and selected the model with the lowest number of vari-
ables (i.e. the model with ‘dtot’ and ‘lat’ as predictors,
for both BA5 and BA10) as the final model. This model
strongly supported a positive relationship between livestock
density and the occurrence of conflict (BA5: βdtot = 1.2,
CI95%:0.04–3.4; BA10: βdtot = 1.7, CI95%:0.1–4.6), and a
negative relationship between the latitude of the StAs and
the occurrence of conflict (BA5: βdtot = −1.1, CI95%:-2.08
to −0.4; BA10: βdtot = −0.3, CI95%:-1.7 to −0.1; Table 3).
Areas in which puma–livestock conflict occurred were
likely to have high livestock density and low latitude.
Conflict mitigation
Mitigation techniques were applied and/or proposed in
81% of the articles reporting conflicts (n = 75). Between
one and five techniques were cited in all studies surveyed
(considering also two papers including only analysis of
the application of mitigation techniques, n = 94) for a
total of 123 mentions overall. The most frequently cited
mitigation techniques (Fig. 4) were as follows: (1) improv-
ing livestock management (29% of cases and 42% of areas),
(2) predator control (18% and 28%, respectively), and
(3) the use of fencing (18% and 28%, respectively).
DISCUSSION
Puma–livestock conflict was studied in only eight of the 13
countries of South America, and the number of articles was
lower than expected based on the extent of the puma’s
9
Puma-livestock conflicts: a continental review M. M. Guerisoli et al.

Table 2. Values of conflict predictors in the study areas (StAs) with the presence and absence of puma–livestock conflict, shown as median (interquar-
tile range; minimum-maximum, n = sample size). Distances were measured from the central coordinates assigned to each StA. All other values were
obtained for two buffer areas (BA5, with a 5 km buffer radius, and BA10, with a 10 km radius). For each predictor, we also present the Mann–Whitney
U-test statistics.

Buffer
area Conflict predictor Presence Absence Mann Whitney U test

BA5 Livestock density (head/km2)*
Livestock richness (number of livestock
species)
Livestock diversity (Shannon index)
Human density (inhabitants/km2)
Habitat fragmentation index (low =
more fragmented)
Habitat steepness (m)*
Co-predator richness (number of
species)*
BA10 Livestock density*
Livestock richness
Livestock diversity
Human density
Habitat fragmentation index
Habitat steepness*
Co-predator richness*
Distances Distance to shrubs (km)*
Distance to trees (km)
Distance to roads (km)*
Distance to human settlement (km)
Latitude (degrees N or S)*
14.5 (43.06; 0–123.7, n = 68)
3 (1; 1–3, n = 52)
0.3 (0.58; 0–0.95, n = 52)
5.3 (32.48; 0–841.5, n = 68)
2.5 (2.32; 0.33–84.95, n = 68)
256 (951.5; 2–2186, n = 68)
0 (1; 0–3, n = 68)
145.5 (298.83; 0–942.3, n = 68)
3 (0; 0–3, n = 68)
0.2 (0.55; 0–1, n = 68)
6 (37.41; 0–994.58, n = 68)
15.53 (30; 2.4–91.4, n = 68)
254 (951.5; 2–2178, n = 68)
0.4 (1; 0–3, n = 68)
3 (10.9; 0–167, n = 68)
0 (1.7; 0–14.33, n = 68)
16.1 (52.54; 0–564, n = 68)
27.8 (42.80; 0–537.4, n = 68)
−12.3°(34.44°; −50.96°−52.07°,
n = 68)
4.5 (8.39; 0–33.2, n = 12) U = 265, P < 0.05
2.5 (1; 2–3, n = 8) U = 196, P = 0.4
0.3 (0.23; 0.1–0.6, n = 8) U = 201.5, P = 0.4
4.5 (18.22; 0–149.2, n = 12) U = 357.5, P = 0.2
6.2 (5.34; 1.47–25, n = 12) U = 204, P = 0.3
849 (499.5; 24–1401, n = 12) U = 268.5, P < 0.05
1 (2; 0–3, n = 12) U = 229, P < 0.05
24.5 (19.39; 0–268, n = 12) U = 180.5, P < 0.05
3 (0.5; 0–3, n = 12) U = 399.5, P = 0.4
0.4 (0.51; 0–1, n = 12) U = 305, P = 0.08
3.6 (10.75; 1–481.46, n = 12) U = 351, P = 0.2
16.2 (32.6; 3.1–92.7, n = 12) U = 298, P = 0.4
849 (499.5; 86–1401, n = 12) U = 261, P < 0.05
1.3 (2; 0–3, n = 12) U = 229, P < 0.05
0.7 (5.8; 0–37, n = 12) U = 291.5, P < 0.05
0.4 (4.6; 0–84, n = 12) U = 319.5, P = 0.06
11.6 (26.22; 0–89.4, n = 12) U = 343.5, P < 0.05
51.6 (47.12; 0–112.4, n = 12) U = 321.5, P = 0.1
39.1° (19.97°; −29.3°−51.16°, U = 177, P < 0.05
n = 12)

* Conflict predictors with significant P-values in the Mann Whitney U Test.

geographic range. This is contrary to what we found for
publications in the rest of the continent. Several factors may
underlie these differences. Differential availability of economic
resources for wildlife research is likely to play a major role
(dos Santos et al. 2020), but this gap may also reflect the
value ascribed to wildlife in North American society. In the
19th and 20th Centuries, in Canada and the USA, wildlife
conservation began to play an important role in society,
which led people to consider wildlife as a resource in the
public domain and science as a fundamental tool to regulate
the protection of wildlife (the ‘North American Model of
Wildlife Conservation’; Organ et al. 2012).
We found that interviewing local ranchers was the main
approach used to collect data on puma–livestock conflict.
This methodology is particularly suitable in developing
countries, where direct data collection (i.e. ranch monitor-
ing, radio-tracking data) is generally limited by costs.
Interviews also enable researchers to study the human
dimension of conflict.
These limitations, however, do not affect the results of
this review, as we based our analyses exclusively on the
presence or absence and not on the magnitude of conflict.
The evidence shows that puma–livestock conflicts are re-
current throughout most of South America, including in
many PAs. Because PAs are intended to conserve wildlife,
ecosystem services, and associated cultural values (Day et
al. 2012), we anticipated that conflict within these areas
would be minimal. Nonetheless, when livestock are bred
within their borders, the presence of PAs does not reduce
the occurrence of conflict. The relationship between PAs
and conflict differed among regions: in North America,
no conflict was found in PAs, indicating that depredation
is concentrated in rural areas, where ranching is predomi-
nant or wildlife protection is less strict. The situation was
different in Central and South America, where the presence
of conflicts was also common within parks and reserves.
This contrast could be explained by several factors: the
extent or category of PAs, the density of livestock within
and adjacent to PAs, the abundance of wild prey, the
level and effectiveness of protection that PAs provide to
predator populations, or the fact that PAs were created
on lands that belonged to local communities and early
settlers. Although we used the most comprehensive global
database on terrestrial PAs, we acknowledge that some of
the StAs that are not listed as PAs in some countries,
especially in North America, may be managed similarly
to areas classified as PAs in the rest of the continent.
Conflicts between livestock producers and large carnivores

10 Mammal Review (2020) © 2020 The Mammal Society and John Wiley & Sons Ltd.
 M. M. Guerisoli et al. Puma-livestock conflicts: a continental review

Table 3. Parameters of fitted models and parameters of the model with the lowest number of variables for the presence of puma–livestock conflict in
the Americas for BA5 and BA10. Only models with ΔAICc < 2 are reported. Variable response: presence/absence of conflict. Int.: intersection; df: de-
grees of freedom; LogLik: log likelihood; AICc: value of the Akaike information criterion adjusted for small samples; ΔAICc: difference between the
AICc values; CI: confidence intervals (2.5% and 97.5%); Z-value: statistical test Wald; dtot: livestock total density; lat: latitude; dshrub: distance to
shrub; droad: distance to road; step: habitat steepness.

BA5 Models Int. dtot lat dshrub droad step df logLik AICc ΔAICc AIC weight

1 2.5 1.2 −1.1 – – – 3 −25.2 56.7 0 0.2

2 2.6 1.2 −1.1 0.7 – – 4 −24.6 57.9 1.1 0.1
3 2.7 1.3 −1.1 – 0.6 – 4 −24.7 58 1.2 0.1
4 2.6 1.2 −1.1 – – −0.3 4 −24.9 58.4 1.6 0.09
Estimate 2.5 1.2 −1.1
Std. Error 0.6 0.8 0.4
Z value 4.2 1.5 −2.8
2.5% 1.6 0.04 −2.08
97.5% 4.2 3.4 −0.4

BA10 AIC
Models Int. dtot lat dshrub droad step df logLik AICc ΔAICc weight

1 2.8 1.7 −0.8 – – – 3 −24.9 56.2 0 0.2

2 3.03 2.06 −0.7 – 0.7 – 4 −24.4 57.4 1.2 0.1
3 2.8 1.6 −0.8 0.6 – – 4 −24.5 57.5 1.3 0.1
4 2.8 1.6 −0.8 – – −0.3 4 −24.5 57.7 1.4 0.09
Estimate 2.8 1.7 −0.8
Std. Error 0.7 1.1 0.3
Z value 3.6 1.6 −2.2
2.5% 1.7 0.1 −1.7
97.5% 4.9 4.6 −0.1

40
LM: improve livestock management
35 FC: fencing
PC: predator control
WP: wild prey (increase or protect)
30 GA: use guard animals (mostly dogs)
Number of mentions

EA: education/awareness campaigns

25 WB: water buffaloes Bubalus bubalis/‘criollos’ livestock
CM: compensation schemes
20 EC: ecotourism
RL: relocate problem individuals
DT: deterrents
15

10

0
LM=36 FC=22 PC=22 WP=10 GA=10 EA=9 WB=5 CM=3 EC=3 RL=2 DT=1
Fig. 4. Numbers of mentions of techniques to mitigate puma–livestock conflicts, as cited in the literature sourced. In some cases, more than one
method was mentioned in a single publication. Water buffaloes and ‘criollos’ cattle (WB) are considered to be guard animals, but they are used
particularly in areas of the Pantanal and Llanos (Paraguay, Venezuela, and Brazil). They were therefore separated from other guard animals (GA).

Mammal Review (2020) © 2020 The Mammal Society and John Wiley & Sons Ltd.  11
Puma-livestock conflicts: a continental review
in PAs and their buffer areas were observed elsewhere,
including in Mexico (Anaya-Zamora et al. 2017), Brazil
(Schultz et al. 2014), India (Karanth et al. 2012), and
Bhutan (Wang & Macdonald 2006), and can be attributed
to the large spatial requirements of large carnivores, which
can mean that they must look for food outside the borders
of PAs (Woodroffe & Ginsberg 1998).
Individual analyses of conflict predictors
Our exploratory analyses showed that several variables
differ between conflict areas and areas without conflicts,
such as anthropogenic pressure, habitat, and geographic
location. The consistency between the two buffer areas
we used suggests that the spatial scale did not affect our
results and that they may relate to landscape-scale
variations.
Contrary to our prediction, conflict areas were farther
from roads than areas where conflict was absent. Perhaps
this is related to the fact that we did not include second-
ary roads or rough tracks, which are associated with in-
creased human penetration (Hoogesteijn & Hoogesteijn
2010) and which are used more than expected by pumas
in Mexico (Ávila-Nájera et al. 2018). Large-scale maps that
include these types of more relevant roads are not easily
available. In Mexico, the distance to roads was positively
associated with greater vulnerability of cattle to predation
by jaguars; however, this effect was weaker that the effects
of other variables, including vegetation cover and proximity
to riparian environments (Rosas-Rosas et al. 2010).
In general, a greater density of livestock means a greater
availability of these alternative prey. Thus, it is not sur-
prising that conflicts occurred in regions where livestock
were more abundant.
In concordance with other studies (Michalsky et al.
2006, Anaya-Zamora et al. 2017), we found evidence that
pumas prefer to prey on small livestock, primarily sheep,
and, secondarily, on goats and young calves. Nonetheless,
we found that cattle are the livestock species most fre-
quently involved in conflicts, perhaps because sheep and
goats are much less abundant than cattle in the Americas
(Appendix S2).
Although conflict areas were also characterised by being
far from shrub habitats, the median distance was 3 km
(Table 2). Pumas are associated with semi-closed habitats
(Maehr & Cox 1995, Maehr 1996, Kerkhoff et al. 2000,
Guerisoli et al. 2019); they are stealth or ambush preda-
tors (e.g. Hornocker & Negri 2010) so the success of their
hunting strategy is maximised in habitats that offer both
cover for stalking and open spaces to pursue and capture
prey. On the contrary, proximity to shrub was positively
related to predation of livestock by wolves Canis lupus in
Mongolia (Davie et al. 2014), and, in the Nepalese Himalaya,
12
M. M. Guerisoli et al.
large carnivores were more likely to kill livestock in shrub-
land than elsewhere (Jackson et al. 1996). Still, proximity
to semi-closed habitat in areas with livestock grazing could
lead to increased puma predation of domestic species.
We found that areas with conflict had lower habitat
steepness than areas without conflict. This is probably
driven by the greater occurrence of large-scale livestock
breeding in less steep terrain, as this terrain favours hu-
man control, accessibility, and management. We found
that conflict areas, throughout the continent, were regions
where pumas were the only large predators, or co-existed
with, at most, one other large carnivore species (Table 2).
This occurs in almost all of Latin America, where pumas
either are the unique top predators or co-exist with jag-
uars. The abundance and availability of large native prey
is usually lower in these regions than in other areas
(Laudré & Hernández 2010). Although misidentification
of puma faecal samples used in dietary studies may bias
findings towards small prey (Elbroch & Wittmer 2013),
the available evidence suggests that in most of Patagonia
and the southern Andes – a vast area where the puma
is the only apex carnivore – this feline preys mainly on
exotic species, probably because large native prey are
nearly extinct or are outcompeted by domestic species
(e.g. guanacos Lama guanicoe; Novaro et al. 2000, Walker
& Novaro 2010, Buenavista & Palomares 2018, M.G.
unpublished data). In other areas of Latin America (e.g.
Brazil and Mexico), where jaguars and pumas live sym-
patrically, both felids are responsible for predation of
livestock (Conforti & de Azevedo 2003, de Azevedo 2008,
Rosas-Rosas et al. 2008, Amador-Alcalá et al. 2013).
Conversely, in Canada and the USA, pumas have access
to thriving populations of wild prey (Bowyer et al. 2019),
and, in some areas, form part of comparatively rich
carnivore guilds, where they are subordinate to larger
predators (bears Ursus americanus, Ursus arctos) or gre-
garious predators (wolves). The scenario we propose to
explain the association between the co-predator richness
and the occurrence of conflict is consistent with the
strong positive correlation between co-predators and
latitude, which we found in the analysis aimed at ex-
cluding redundant predictive variables for our GLM.
Combined analysis of conflict predictors
The final multivariate model to predict puma–livestock
conflict included only two variables, livestock density and
distance from the Equator, indicating that areas with con-
flicts between pumas and livestock are likely to have high
densities of livestock and be located in the Southern
Hemisphere. We believe that it is likely that latitude and
livestock density are not only mutually associated, but
also related to the other variables that appear to differ
Mammal Review (2020) © 2020 The Mammal Society and John Wiley & Sons Ltd.
M. M. Guerisoli et al.
between conflict areas and areas without conflict, and that
the variables interact to explain the occurrence of puma–
livestock conflicts throughout the Americas. The importance
of livestock density as a predictor of the distribution and
occurrence of human-felid conflict is not surprising and
is recognised (Miller 2015, Hoogesteijn et al. 2016a, Reyna-
Sáenz et al. 2019). Domestic species tend to compact the
soil, preventing seed germination and affecting the structural
diversity of the vegetation (Kauffman & Krueger 1984).
They also reduce the availability of pastures and vegeta-
tion cover for wild ungulates, which can lead to competi-
tion in foraging (Lee et al. 1998). Thus, high livestock
density may favour conditions leading to puma–livestock
conflicts both directly and indirectly.
Central and South America are the regions of the con-
tinent with the highest livestock densities (Appendix S2),
but are also the regions where anthropogenic pressure on
wildlife is the strongest and intense habitat conversion
and overexploitation is still occurring (Rodríguez 2001,
Ripple et al. 2016, Taubert et al. 2018). We could not
measure the effects of the ecological, cultural, and eco-
nomic differences between these two regions and North
America, and, as previously noted, we believe that these
differences largely affect the effectiveness of wildlife con-
servation strategies (Fragoso et al. 2004). Central and South
America are important worldwide mammalian hotspots
of species richness (Ceballos & Ehrlich 2006), and support
a great variety of medium-sized to large wild ungulates
(several cervid species, two camelids and three peccaries).
However, the populations of these species are frequently
depleted, either directly by sport hunting or poaching, or
indirectly by habitat alterations (e.g. Robinson & Redford
1991, Wilkie et al. 2011). In contrast, in Canada and the
USA, many wildlife populations are intensively managed
as recreational resources (Bowyer et al. 2019). As a con-
sequence, in North America, pumas still find relatively
abundant populations of the ungulates considered to be
their preferred prey (e.g. elk Cervus elaphus, mule deer
Odocoileus hemionus, white-tailed deer Odocoileus virgin-
ianus, pronghorn Antilocapra americana, and bighorn sheep
Ovis canadensis; Anderson 1983, Murphy & Ruth 2010).
Livestock, for pumas, may represent more easily accessible
prey than native prey of equal size (Linnell et al. 1999,
Palmeira et al. 2008), and when large to medium-sized
wild species are scarce or less available, predation on al-
ternative species (e.g. livestock) is likely to occur (Khorozyan
et al. 2015).
Conflict mitigation and management
implications
In 1240, De Bracton wisely stated: “An ounce of preven-
tion is worth a pound of cure”. Several approaches can
Mammal Review (2020) © 2020 The Mammal Society and John Wiley & Sons Ltd. 
Puma-livestock conflicts: a continental review
be used to address human–carnivore conflicts, including
the following: fully engaging key stakeholders; testing and
adapting predation reduction methods and social conflict
mitigation strategies; and working closely with the affected
local communities (Zarco-González et al. 2018, Ohrens et
al. 2019a). Several recent reviews concurred in concluding
that there are wide gaps in the scientific evidence on the
effectiveness of techniques to reduce livestock depredation
(Eklund et al. 2017, van Eeden et al. 2018b, Treves et al.
2019). Concurrently, the list of potential strategies men-
tioned in the literature we reviewed is comprehensive, but
most of the studies suggested mitigation techniques based
exclusively on researchers’ opinions or on simple observa-
tions of the practices adopted in the StAs. Three tools
(control of predator populations, improvement of livestock
husbandry and use or improvement of fences) made up
over 64% of the mentions of mitigation techniques. Other
techniques that are adopted to mitigate human-carnivore
conflicts, such as economic compensation, relocation of
problematic individuals, and the use of deterrents (van
Eeden et al. 2018a), were rarely cited.
Lethal control of carnivores is widely applied, both le-
gally (Treves & Karanth 2003) and illegally (Liberg et al.
2011, Guerisoli et al. 2017). The limited evidence available
for pumas suggests that hunting-related mortality may
disrupt this felid’s population dynamics (e.g. by altering
patterns of dispersal of young individuals or females with
cubs, or by modifying home range size areas) and social
structure (e.g. sex or age structure), and may lead to
increased puma–livestock conflicts (Robinson et al. 2008,
Keehener et al. 2015, Teichman et al. 2016). Predator
control as a conflict mitigation technique is based on the
assumption that a decrease in carnivore abundance would
lead to a decrease in losses caused by predation. This
interpretation represents a simplification of complex preda-
tor–prey dynamics, and several studies have shown that
increasing efforts to control predator populations do not
produce reductions in livestock predation (Graham et al.
2005, McManus et al. 2015, Treves et al. 2016). Peebles
et al. (2013) concluded that heavy hunting of pumas in
Washington, USA, largely amplified the odds of complaints
and depredations. Besides increasing conflict, lethal control
is generally not selective towards the conflictive individual
(Treves & Naughton-Treves 2005). Therefore, evidence
that the elimination of carnivores helps to reduce livestock
predation must be considered equivocal at best (Berger
2006, McManus et al. 2015, Lennox et al. 2018), and
ecologically and politically misguided at worst.
Studies assessing the effectiveness of mitigation tech-
niques to reduce predation by pumas are still scarce and
have a geographical bias, but they show that the effective-
ness of interventions to protect livestock from puma pre-
dation is highly variable (Khorozyan & Waltert 2019).
13
Puma-livestock conflicts: a continental review
Our results support the recent reviews by Eklund et al.
(2017) and Moreira-Arce et al. (2018), both of which
showed that livestock management is one of the most
promising methods to limit depredation and that improv-
ing husbandry and management techniques is considered
a suitable tool for successful mitigation of conflicts with
pumas (Breck et al. 2011, Soto-Shoender & Giuliano 2011,
Zarco-González et al. 2018, Reyna-Sáenz et al. 2019).
However, other tools are also probably practical for de-
creasing puma depredation (Khorozyan & Waltert 2019).
In the Brazilian Pantanal, the use of electric fences de-
creased losses caused by jaguars and pumas (Hoogesteijn
et al. 2016b). In northern Chile, light deterrents discour-
aged pumas from preying on alpacas Vicugna pacos and
llamas (Lama glama; Ohrens et al. 2019b). In Mexico,
Zarco-González and Monroy-Vilchis (2014) found that
visual and auditory repellents were useful to protect goats
and cattle from large felids. A recent literature review
suggested that the use of guardian animals could be the
most appropriate mitigation technique, because it reduced
losses of livestock (especially sheep and goats), and had
minimal negative effects on carnivore populations (van
Eeden et al. 2018a). We observed that this method was
one of the least frequently mentioned mitigation techniques
for pumas in the Americas, possibly because of the cost
of purchasing and subsequently maintaining the pure-bred
guard dogs that are used (Moral et al. 2016). Mixed-breed
guard dogs proved to be a successful, cheaper alternative
for protecting goat herds in northern Argentinean Patagonia
(González et al. 2012), and may have great potential, also
because they are more adaptable to certain environments
than the dog breeds traditionally used. Nevertheless, im-
properly managed guard dogs may become semi-feral and
may have negative effects on wildlife (Moral et al. 2016,
Allen et al. 2019).
The success of mitigation techniques can be condi-
tional, since strategies may work only under certain
conditions, and ineffective mitigation techniques can
intensify levels of conflict (Inskip & Zimmermann 2009,
Ohrens et al. 2019b). Economic constraints may limit
the range of applicable interventions in developing coun-
tries (Khorozyan & Waltert 2019). Each ranch presents
a unique scenario, depending on the livestock species
and management, neighbouring properties, habitat type,
wild prey availability, ranch size, and its owner’s income
and educational level. Thus, successful mitigation tech-
niques have to be tailored to the local scenarios, and
the livestock owner’s budget, available time, and pre-
disposition must be considered. Given the potential
variety of scenarios where puma–livestock conflicts can
occur, this felid’s ecological flexibility, and the available
information, we conclude that scientists and managers
wishing to address conflict need to work along with
14
M. M. Guerisoli et al.
livestock producers to consider a diverse array of strate-
gies which account for both the ecological and human
dimensions (Hulet et al. 1987, Ohrens et al. 2019a).
Limitations of our analyses
We presented an exploratory analysis of the presence
of puma–livestock conflict at the continental scale. A
variety of ecosystems, countries, cultures, and socio-
economic variables characterise the puma’s geographic
range, so we tried to be inclusive in conflict predictor
selection and in the analysis. Despite this, we recognise
some methodology limitations. A major weakness in our
work is that we were unable to include a variable in-
dicative of the abundance of wild prey populations. Prey
availability is commonly identified as important in de-
termining levels of conflict between humans and preda-
tors (Graham et al. 2005). Although some of the literature
reviewed (19% of papers) mentioned that scarcity of
wild prey influenced the occurrence of puma predation
of livestock, the importance of this factor is still unclear.
We strongly recommend that authors of future studies
exploring the factors affecting the presence and, espe-
cially, the intensity of conflicts between large cats and
livestock, should consider the effect of wild prey popula-
tions. In particular in Central and South America, there
is little information on wild prey abundance and avail-
ability and its effects on jaguar and puma populations.
Poor livestock husbandry or poorly implemented ranch
management strategies can affect the intensity of dep-
redation by pumas (Mazzolli et al. 2002, Polisar et al.
2003), so livestock husbandry is an additional predictor
that could have improved our understanding of the
dynamics of puma–livestock conflicts. Regrettably, in-
formation on these strategies used in the different StAs
was usually poor and difficult to standardise.
An additional limitation of our approach is that, by
using the central geographical coordinates of the StA to
obtain the values of the predictive variables, we may have
failed to reflect the detailed characteristics of sites where
predation took place. The inclusion of this geographically
explicit information – which is typically not reported by
authors – in the supplementary material of studies analys-
ing carnivore predation would improve the quality of
future meta-analyses.
CONCLUSIONS
Unlike most other species of Felidae, pumas are charac-
terised by great ecological plasticity (e.g. Hornocker &
Negri 2010, Zanin et al. 2015), which has allowed them
to be successful apex predators and to persist throughout
most of their range. Paradoxically, this also has made
Mammal Review (2020) © 2020 The Mammal Society and John Wiley & Sons Ltd.
M. M. Guerisoli et al.
them more likely to engage in conflicts with humans, and
to suffer retaliatory killings.
Pumas exert top-down effects on food webs, and their
disappearance may disrupt the natural functioning of
ecosystems (Terborgh et al. 1999, Ripple et al. 2014).
However, when planning the conservation of a conflic-
tive species, its effects on local human communities must
be acknowledged. The information available on the con-
sequences of puma depredation on regional economies
is still poor. However, under certain circumstances, puma
attacks may have dramatic effects on the livelihoods of
local people (e.g. Michalsky et al. 2006, Palmeira &
Barrella 2007), particularly in developing countries and
on small ranches (Guerisoli et al. 2017, Lucherini et al.
2018). Although the number of publications on puma–
livestock conflict is increasing, we found that information
on the applicability and effectiveness of potential tech-
niques for conflict mitigation is still extremely poor in
quality and quantity. It is time to take a further step
in puma–livestock conflict research, and increase research
efforts aimed at identifying tools capable of reducing
the negative impacts of conflicts for both humans and
pumas. The development and testing of locally effective
mitigation techniques is the next frontier in human–
wildlife conflicts.
ACKNOWLEDGEMENTS
We acknowledge the effort made by the anonymous re-
viewers. The comments and suggestions made by them
improved this manuscript. Additionally, we want to ac-
knowledge the follow-up and revisions made by the Editor-
in-Chief, Dr Danilo Russo, and the Managing Editor, Dr
Nancy Jennings.
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SUPPORTING INFORMATION
Additional supporting information may be found in the
online version of this article at the publisher’s web-site.
Appendix S1. (a) List of the literature reviewed, including
the country and specific area where each study was con-
ducted and the main methods adopted. (b) List of the
information sources used for the extraction of the conflict
predictors described in Table 1.
Appendix S2. Livestock densities (mean ± standard devia-
tion; head/km2) within the puma’s geographic range, per
region.
Appendix S3. Box-plots for each potential predictor of
puma-livestock conflicts included in our analysis, for
buffer areas around study areas, of two different sizes:
BA5 (with a radius of 5 km) and BA10 (with a radius
of 10 km).
Appendix S4. Kendall correlation matrix showing all the
predictor variables of puma-livestock conflict considered
in the modelling analyses with two buffer sizes: BA5 (with
a radius of 5 km) and BA10 (with a radius of 10 km).
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