Biological Conservation 159 (2013) 8087

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Biological Conservation
journal homepage: www.elsevier.com/locate/biocon

Spatial model of livestock predation by jaguar and puma in Mexico:

Conservation planning
Martha M. Zarco-Gonzlez a,b, Octavio Monroy-Vilchis a,b,, Jorge Alanz c
a

Sierra Nanchititla Biological Station, Faculty of Sciences, Autonomous University of the State of Mexico, Mexico
Instituto literario 100, Centro, Toluca, Estado de Mxico, CP 5000, Mexico
c
Faculty of Sciences, Autonomous University of Baja California, Carretera Tijuana-Ensenada Km 106, 22800 Ensenda, BC, Mexico
b

a r t i c l e

i n f o

Article history:
Received 15 May 2012
Received in revised form 23 October 2012
Accepted 4 November 2012
Available online 21 January 2013
Keywords:
Livestock
Predation
Ecological niche model
Puma
Jaguar
Mxico

a b s t r a c t
Predation on livestock is one of the main factors that cause the felids hunting, in particular for puma and
jaguar this conict with humans is severe. Most studies have assessed the predation impacts on livestock
production; however there is a spatial pattern in attacks occurrence that is feasible to analyze from ecological niche modeling. The objective of this research was to generate a risk model of livestock predation
by puma and jaguar in Mexico based on environmental and livestock management variables, which
allows identication of zones of risk in order to dene mitigation strategies at national level. We produced a geographic ensemble model of risk of predation from three algorithms for jaguar and ve for
puma. The variables most positively related with predation risk by jaguar were vegetative cover percentage, percentage of free grazing animals, and altitude, whereas arid vegetation has a negative inuence on
predation risk. In the case of puma the variables with highest contribution were livestock density, which
negatively inuences on the predation risk, in addition to forest and altitude, both with a positive relation. The ensemble models are an accurate approach to delineating the zones of predation risk by felids;
however at a regional scale the environmental characteristics that favor predation may be different. It is
recommended that researchers carry out studies for each biogeographic province that facilitate the identication of specic patterns and the denition of mitigation strategies most suitable for each one.
2012 Elsevier Ltd. All rights reserved.

1. Introduction
Populations of large carnivores are declining worldwide due to
the loss or fragmentation of their habitat, and by direct hunting of
the carnivores or their prey; currently, 17% of the species are threatened (IUCN, 2011). The hunting of felids takes place in response to
their supposed predation on livestock (Holmern et al., 2007; Kissui,
2008; Gusset et al., 2009); this problem affects 75% of felid species
worldwide, and in particular is severe for puma and jaguar (Inskip
and Zimmermann, 2009). In previous studies, data on felids hunted
in retaliation for predation have been reported, which range from
seven pumas killed per year on average in a ranch in Sao Paulo
to 150 pumas and jaguars in Alta Floresta, Brazil (Inskip and Zimmermann, 2009).
Hence, it is evident that a fundamental topic in large cats conservation is the analysis of the conicts with livestock holders. This
issue has mainly been approached by describing the impact of predation on livestock production and identifying factors related with
Corresponding author at: Instituto literario 100, Centro, Toluca, Estado de
Mxico, CP 5000, Mexico. Tel.: +1 722 296 55 53.
E-mail addresses: martha.zarco.g@gmail.com (M.M. Zarco-Gonzlez), tavomonroyvilchis@gmail.com, omv@uaemex.mx (O. Monroy-Vilchis).
0006-3207/$ - see front matter 2012 Elsevier Ltd. All rights reserved.
http://dx.doi.org/10.1016/j.biocon.2012.11.007

the predation levels (Patterson et al., 2004; Kolowski and Holekamp, 2006; Azevedo and Murray, 2007; Van et al., 2007; Palmeira
et al., 2008; Iftikhar et al., 2009). Other authors have observed differences in the environmental characteristics of sites with and
without predation (Jackson et al., 1996), and this may be an indicator of conditions inherent to the sites that inuence on the risk
(Stahl et al., 2002). Among the proposed environmental variables
that may be related with the frequency of attacks are proximity
to forest zones (Mazzolli et al., 2002; Stahl et al., 2002; Azevedo
and Murray, 2007; Palmeira et al., 2008), the vegetation type (Rosas-Rosas et al., 2010), altitude (Lui et al., 2006), topography (Stahl
et al., 2002; Michalski et al., 2006; Kissling et al., 2009), density of
livestock and wild prey (Treves et al., 2004; Bagchi and Mishra,
2006; Kolowski and Holekamp, 2006), distance to protected areas,
human settlements, roads, and water sources (Lui et al., 2006; Van
et al., 2007; Gusset et al., 2009; Rosas-Rosas et al., 2010). An approach that may be useful in decreasing the severity of the conict
is to anticipate its spatial location and to propose preventive actions in specic areas, optimizing economic and human resources.
A commonly used tool to plan strategies of wildlife management, which relates ecological variables and spatial processes,
are ecological niche models (Gibson et al., 2007; Zarco-Gonzlez
et al., 2012), the modeling provides estimates of occurrence

M.M. Zarco-Gonzlez et al. / Biological Conservation 159 (2013) 8087

probability of these processes in the study area. This technique has

been applied to study the relations between environmental parameters and richness of species (Arajo and Williams, 2000; Ferrier
et al., 2002; Scotts and Drielsma, 2003; Mac Nally and Fleishman,
2004), invasive potential of exotic species (Peterson, 2003; Goolsby, 2004), and species distributions (Bakkenes et al., 2002; Hugall
et al., 2002; Arajo et al., 2004; Peterson et al., 2004; Skov and
Svenning, 2004; Thomas et al., 2004; Thuiller et al., 2005; Rodrguez-Soto et al., 2011).
By applying this methodology to the study of human-wildlife
conict, from the location and characterization of the sites where
predation on livestock is present, it is possible to spatially predict
the zones of predation risk, as well as to identify the variables that
propitiate this interaction. This enables planning actions to manage livestock and predators in specic zones to minimize the conicts, optimize livestock production, and reduce the hunting of
wild felids. The objective of this study was to generate a risk model
of livestock predation by puma and jaguar in Mexico based on
environmental and livestock management variables. The model
further allows identifying and prioritizing of zones of felid predation risk in order to develop mitigation strategies at national level.

2. Materials and methods

2.1. Study area
Mexico is located in northern America, its territorial extension
is 1,953,162 km2 (Fig. 1). Its extreme coordinates are: 14320 2700
south, 32430 0600 north; 86420 3600 east and 118270 2400 west (INEGI, 2011). Mexico possesses a varied topography, more than 65%

81

of the country is above 1000 m above sea level and nearly 47% of
the surface has slopes over 27% (UNAM, 1990; Mittermeier and
Mittermeier, 1992). In conservation importance, Mexico hosts between 8% and 12% of the total of the species on the planet (Mittermeier and Mittermeier, 1992), including six feline species, of which
puma (Puma concolor) and jaguar (Panthera onca) are the largest.
The records of attacks on livestock by puma and jaguar in Mexico were collected in three ways: (1) databases of the Mexican government, Secretariat of Environment and Natural Resources
(SEMARNAT), Pronatura (civil association) and technical reports
(Nez, 2007a); (2) review of scientic literature: theses (BuenoCabrera, 2004; Villordo-Galvn, 2009), books (Brown and Lpez,
2001; Rosas-Rosas and Lpez-Soto, 2002; Caso, 2007; Cruz et al.,
2007; Leyequin and Balvanera, 2007; Lira and Ramos-Fernndez,
2007; Navarro et al., 2007; Nez, 2007b), and papers (Rosas-Rosas et al., 2008; Chvez and Zarza, 2009; Zarco-Gonzlez et al.,
2012); (3) and eld work in different states of the country (states
of Mexico, Chihuahua, Yucatn, Campeche, Baja California and
Guerrero). Approximately 78% of predation data were recorded at
the site of attack with GPS devices and 22% were inferred using
Google Earth and maps and descriptions presented in the original
sources. A database of predation events was produced with the records specifying date, predator species, place of attack, and geographic coordinates (Fig. 1).
Considering the reports of other studies in relation to the inuence of the environmental factors on the predation risk (Stahl et al.,
2002; Treves et al., 2004; Bagchi and Mishra, 2006; Kolowski and
Holekamp, 2006; Lui et al., 2006; Michalski et al., 2006; Van
et al., 2007; Gusset et al., 2009; Kissling et al., 2009; Rosas-Rosas
et al., 2010), the variables used to characterize the sites of attack
were grouped in three types: landscape, livestock management,

Fig. 1. Mexico location and records of predation by jaguar and puma.

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M.M. Zarco-Gonzlez et al. / Biological Conservation 159 (2013) 8087

and anthropogenic; topographic variables were altitude (USGS/

NASA, 2007) and slope. The vegetative associations of the National
Forest Inventory were also included (SEMARNAP, 2001) and
grouped into seven types: forest (conifers, oaks and riverside vegetation), dry forest, rainforests, arid vegetation, underwater vegetation, and agriculture. The percentage of tree cover was
obtained from Global Land Cover Facility (De Fries et al., 2000).
In relation to livestock management, layers of livestock density
and percentage in free grazing were generated by interpolating
the values of each municipality from the 2007 agricultural, livestock, and forest census (INEGI, 2009). Human perturbation, considering the human population density (Salvatore et al., 2005)
and distance to paved roads (CONABIO, 2008), was included as
an anthropogenic variable. Each of these layers was processed in
a raster format, with a 1-km2 resolution; to unify them, pixels with
no data and water sources were excluded. Although the information represented in each variable is different, the correlation between them was calculated using BioMapper4 (Hirzel, 2008).
Variables retained in the models have a correlation of less than 0.5.
For reducing the spatial correlation between records, they were
ltered to obtain only one datum per pixel. The records thus obtained were randomly divided into two groups for each species:
75% to calibrate the prediction models and 25% to validate the
models (Guisan and Zimmermann, 2000). It is assumed that this
sample of the original data contains independent observations that
can be used in a statistical test (Arajo et al., 2005).
To obtain the different models of predation risk for each species,
we used the software Open Modeller version 1.1.0 (http://
www.openmodeller.sf.net), including the algorithms: Articial
Neural Network (Gevrey et al., 2003; Pearson et al., 2004), Environmental Distance (Hirzel and Arlettaz, 2003), GARP (Genetic Algorithm for Rule-set Production; in both versions: single run and
with best subsets, Stockwell and Peters, 1999) and SVM (Support
Vector Machines; Cristianini and Scholkopf, 2002; Huang et al.,
2002; Guo et al., 2005; Drake and Bossenbroek, 2009). We also
used BioMapper4 (Hirzel, 2008) and Maximum Entropy Species
Distribution Modeling v3.3.3e (Phillips et al., 2006), to obtain the
algorithms of ENFA (Hirzel et al., 2002) and Maxent (Phillips
et al., 2006), respectively, the latter also to get the inuence of each
variable in the predation risk.
Once the models were obtained, their individual performance
was evaluated from the area under the curve (AUC) ROC (Receiver
Operating Characteristic, Hanley and McNeil, 1982). The AUC values between 0.5 and 0.7 are considered low (model with poor performance); between 0.7 and 0.9 are moderate; and >0.9 are high
performance (Manel et al., 2001). We used the ROC module of
the Idrisi Andes software (Clark Labs, 2006) in order to calculate
this value. We obtained two evaluations for each model, one based
on the calibration data (internal) and other with the validation
data (external).
Comparing the efcacy of different modeling methods we have
noticed that in spite of some being more effective for prediction
(Elith et al., 2006; Tsoar et al., 2007), none proved to be the best
in all cases (Hernandez et al., 2008). To overcome this variability,
a solution is to develop models using multiple algorithms that
identify common areas of consistent prediction (Anderson et al.,
2003; Arajo et al., 2006). The consensual area between predictions incorporates the uncertainties of modeling and produces
more reliable estimations (Hartley et al., 2006). This methodology
has been applied in the prediction of the distribution of species of
plants (Marmion et al., 2009), invasive arthropod species (RouraPascual et al., 2008), reptiles (Domnguez-Vega et al., 2012) and
carnivorous mammals (Rodrguez-Soto et al., 2011). Consensus approaches are particularly useful in cases where there are not adequate data to evaluate which model makes the most accurate
predictions, as is the case with species invading new areas, project-

ing future distributions under environmental change scenarios,

and in very data-poor regions (Hernandez et al., 2008). Because
of uncertainties in individual models, an ensemble model of predation risk for each felid species was produced.
In the ensemble model, we included the models with AUC values P0.75 in the internal evaluation. We used the weighted average, as it has been demonstrated that it has the highest predictive
performance of all the consensus methods (Marmion et al., 2009),
AUC value was also calculated to evaluate the ensemble model performance. To facilitate their interpretation, the original prediction
values (0100) were reclassied considering the mean probability
value of the records used to calibrate the models, so that the zones
with a prediction value greater than or equal to this mean were
considered those with high risk and those with values lower were
considered low risk (Liu et al., 2005).

3. Theory
Machine learning methods are algorithms that are used to learn
the mapping function or classication rules inductively, directly
from the training data (Franklin, 2009).
They include articial neural networks, genetic algorithms,
maximum entropy and support vector machines. These methods
tend to perform well given complex classication problems, however, interpreting the modeled relationships between predictors
and response is not always straightforward in some machine learning implementations.
ANNs consist of many processing elements (articial neurons)
that are interconnected to form a network. ANNs are trained
by repeatedly passing large numbers of known examples of the
problem under consideration through the network. By repeatedly
adjusting the connections between processing elements the difference between the network predictions and the known examples
can be minimized (Pearson et al., 2004). Neural networks can identify non-linear responses to environmental variables, as well as
incorporating multiple types of variables, including qualitative
and quantitative. A disadvantage of this method is that it does
not identify the relative contribution of the different variables
(Gevrey et al., 2003). The details of the classier developed using
neural networks are not obvious or interpretable, it is considered
a black box approach to classication, and so the main advantage
of this method is that it sometimes achieves much higher classication accuracy than other methods in high-dimensional, complex
classication problems (Franklin, 2009).
The GARP software implements an ensemble method that generates a population of prediction rules based on several different
types of models, and then uses a genetic algorithm to select among
them and develop a nal rule set to make predictions. Further, because the output from GARP is stochastic, it is typical to run the
model multiple times, and then average a subset of the best models
(Stockwell and Peters, 1999). GARP is not able to model responses
to categorical predictors and tends to over predict the extent of
species distributions, that is, to have higher commission errors
(Franklin, 2009).
In Maxent the multivariate distribution of suitable habitat conditions in environmental feature-space is estimated using the best
approximation of an unknown distribution with maximum entropy (the most dispersed) subject to known constraints. The advantages of this method are that it can use both categorical and
continuous data, as well as incorporate interactions between different variables (Phillips et al., 2006), additionally Maxent can
work well with very small samples that have relatively widespread
geographic distributions (Hernandez et al., 2008).
SVM use a functional relationship known as a kernel to map
data onto a new hyperspace in which complicated patterns can

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M.M. Zarco-Gonzlez et al. / Biological Conservation 159 (2013) 8087

be more simply represented. Functionally, SVMs seek to nd an

optimal separating hyperplane with the maximal margin between
the training points for presence and absence data in multidimensional space (Cristianini and Scholkopf, 2002; Huang et al., 2002;
Guo et al., 2005; Drake and Bossenbroek, 2009). SVM are able to
handle categorical and non-linear data and not make assumptions
about the probability of data density (Guo et al., 2005). However, it
is difcult to tell if it is possible to interpret its classication rules
from these models in order to verify its ecological meaning or
validity (in terms of the importance of predictors and the form of
their relationship with the response; Franklin, 2009).
Distance-based methods, like Environmental Distance and
ENFA, have limitations, they work best when organisms are using
optimal habitat, are well-sampled in environmental space, and
when habitat variables are not dynamic (Franklin, 2009). Additionally, these methods are not well-suited for categorical predictors;
they equally weight predictors, and assume linear relationships between environmental variables and habitat suitability.
Environmental distance denes the suitability area in the environmental space of each species record. The geometric media or
the distance of each record to rest of them are calculated, in this
way, a high density of records in the environment space imply a
high habitat suitability (Hirzel and Arlettaz, 2003).
ENFA is based on a multivariate description of species occurrence locations. It estimates species niche more explicitly based
on the magnitude of the difference between the species mean
and the entire range of environmental conditions observed. This
model estimates marginality and specialization (Hirzel et al.,
2002). ENFA is prone to high commission error or low specicity
in comparison with other methods (Franklin, 2009).

Table 2
AUC values of the models for jaguar.
Algorithm

Maxenta
GARP (with best subsets)a
SVMa
ENFA
GARP (single run)
ANN
Maxent (open modeller)
Environmental distance
ENFA (open modeller)
a

AUC
External evaluation

Internal evaluation

0.799
0.784
0.767
0.728
0.728
0.725
0.704
0.626
0.500

0.919
0.792
0.837
0.743
0.714
0.730
0.669
0.923
0.520

Algorithms included in the ensemble model.

Table 3
AUC values of the models for puma.
Algorithm

Maxenta
Environmental distancea
ENFAa
GARP (with best subsets)a
GARP (single run)a
ANN
SVM
Maxent (open modeller)
ENFA (open modeller)
a

AUC
External evaluation

Internal evaluation

0.944
0.875
0.866
0.817
0.767
0.720
0.617
0.503
0.500

0.934
0.961
0.828
0.802
0.790
0.730
0.591
0.505
0.510

Algorithms included in the ensemble model.

Table 4
Importance of variables in the models generated with Maxent for each felid species
(only the most important are shown).

4. Results
We obtained 241 records of felid attacks on livestock in Mexico,
and after data reduction we retained 222 records; 152 predation
events by jaguar and 70 by puma. The retained records are from
1990 to 2010, and are distributed in 19 states (Table 1).
For jaguar, three algorithms were considered suitable according
to the internal evaluation: Maxent, GARP-with best subsets and
SVM (Table 2). For puma ve algorithms were considered: Maxent,
Environmental distance, ENFA, GARP-with best subsets and GARPsingle run (Table 3). For both species the algorithm with the best
performance was Maxent.
According to ENFA, for data from jaguars, we obtained a global
marginality value of 0.793, which indicates that the observed conditions in the sites of attack are different from the average environmental conditions and the range in which predation occurs is very
restricted in relation to the global characteristics. Maxent indicates
that the variables with highest contribution to the model are tree
cover percentage, percentage of animals in free grazing areas,
and altitude. These variables are positively related with the risk

Table 1
Number of data obtained of each source and year of collect.
Source

Number of
data

Year of collect

Secretariat of Environment and

Natural Resources (SEMARNAT)
Pronatura (civil association) and
technical reports
Theses
Books
Papers
Field work

80

20092010

10

20002007

13
36
26
76

20002006
19912006
19982004
20062010

Variable

Altitude
Forest
Arid vegetation
Cover percentage
Livestock density
Percentage of free gazing livestock

Maxent (contribution percentage)

Jaguar

Puma

10.3
3.7
8.5
47
4.5
12.1

12.3
34.1
0.5
6.7
35.2
6.6

of attack by jaguar, whereas arid vegetation has a negative correlation with predation risk (Table 4).
In the case of puma, ENFA indicated a marginality global value
of 1.157, these results imply that the characteristics of predation
sites by puma are very specic, even higher than those of jaguar,
and the general variation in these characteristics is low. Maxent
showed that the variables with highest contribution were livestock
density, which negatively inuences on the predation risk by
puma, in addition to forest and altitude, both with a positive relation (Table 4).
The ensemble model for jaguar had an AUC value of 0.8. The
zones of highest predation risk by this species (probability > 64)
are in the western Sierra Madre Occidental, at the border with
Sonorense province, the Costa del Pacco, western and center of
Cuenca del Balsas, eastern Sierra Madre del Sur, eastern Soconusco,
the northern and southern limits of Altos de Chiapas and Golfo de
Mxico, mainly northern and limits with Sierra Madre del Sur,
Yucatn, Petn and southern Sierra Madre Oriental (Fig. 2).
The ensemble model for puma presented an AUC value of 0.9. In
this case the risk zones (probability > 69) are concentrated on
southern California, western and central Sierra Madre Occidental,

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M.M. Zarco-Gonzlez et al. / Biological Conservation 159 (2013) 8087

Fig. 2. Ensemble model of risk predation by jaguar in Mexico. (1) Sierra Madre Occidental, (2) Sonorense, (3) Costa del Pacco, (4) Cuenca del Balsas, (5) Sierra Madre del Sur,
(6) Soconusco, (7) Altos de Chiapas, (8) Golfo de Mxico, (9) Yucatn, (10) Petn y, (11) Sierra Madre Oriental.

Fig. 3. Ensemble model of risk predation by puma in Mexico. (1) California, (2) Sierra Madre Occidental, (3) Sierra Madre Oriental, (4) Eje Volcnico Transmexicano, (5)
Cuenca del Balsas, (6) Sierra Madre del Sur, (7) Soconusco y, (8) Altos de Chiapas.

northern Sierra Madre Oriental, the southern limit of the Eje Volcnico Transmexicano, the center of Cuenca del Balsas, Sierra Madre
del Sur, Soconusco and the center of Altos de Chiapas (Fig. 3).

5. Discussion
The conict of livestock predation by the two large cats of Mexico has negatively impacted on the populations of both jaguar and
puma. In the north of the country, the illegal hunting of jaguar is
perhaps the main threat for its conservation (Rosas-Rosas et al.,
2008). However, in the case of puma the severity of the conict
has not been assessed, but it is known that in some areas predation
on livestock is the most frequent reason for their hunting by hu-

mans (Zarco-Gonzlez et al., 2012). Despite this situation, systematic data on the topic are scarce, and most of the published studies
are focused on jaguar (Rosas-Rosas et al., 2008, 2010; Chvez and
Zarza, 2009; Villordo-Galvn, 2009). Fewer studies have been carried out on puma (Bueno-Cabrera, 2004; Zarco-Gonzlez et al.,
2012). As a consequence, there are no data where livestock predation occurs, and much less on the impact on livestock production
or on the felids conservation, so the present risk model shows a
spatial panorama of the predation problem in Mexico.
One of the most recent actions to address this issue was carried
out in 2009 by the National Livestock Confederation and the Secretariat of Agriculture, Livestock, Rural Development, Fishing and Alimentation (SAGARPA). They established a program providing
livestock insurance for death due to predators. The general objec-

M.M. Zarco-Gonzlez et al. / Biological Conservation 159 (2013) 8087

tive of the fund is to protect the patrimony of Mexican livestock

holders and the operative continuity of their production units. This
program is intended to address at a regional level the problem of
predation, however, the compensation to affected livestock holders
has not been done efciently. In this sense, the accurate identication of the risk zones will enable the directing of nancial and human resources to the zones most vulnerable to predation.
Most of the studies in Mexico have analyzed the impact on felid
predation on bovines (Rosas-Rosas et al., 2008, 2010; Chvez and
Zarza, 2009; Villordo-Galvn, 2009), the interest in this livestock
is related to the higher economic value in relation to the smaller
domestic species (SAGARPA, 2006). Additionally, the reports of attacks on small livestock are less frequent. These conditions have favored that the most data on predation are related to jaguar, as that
jaguars more frequently consume large domestic prey (bovines,
Hoogesteijn, 2001). Hence, the impact of predation by puma is
probably underestimated and eventually so would be the risk area
predicted in this study. To overcome these limitations is necessary
to evaluate with greater precision the cases of attacks by puma,
particularly on sheep and goats. On the other hand, considering
the priority areas for jaguar conservation proposed by RodrguezSoto et al. (2011), it is observed that they coincide with the zones
of highest predation risk, in this sense it is necessary to invest resources on the application of strategies to reduce the human-jaguar conict.
According to the specialization values for both species is clear
that the attributes of the zones with attacks are particular, even
different from those that characterize the general species habitat.
Chvez and Zarza (2009) found that vegetation was the variable
that best explains the jaguars potential distribution in Yucatan
Peninsula, conversely, the human-jaguar conict model generated
in the same study shows that the most of predation events takes
place around human settlements. This reafrms the fact that sites
in which the attacks on livestock occur have particular characteristics that make their inclusion into a predictive model feasible.
Maxent indicates that the percentage of cover and the presence
of arid vegetation are positively and negatively related, respectively, with predation risk by jaguar. In particular, tree cover inuences on the success probability of attack by jaguar, which is a
stalk-and-ambush predator; on the contrary, scarce cover, a characteristic of arid vegetation, explains the negative relation with
the risk. The zones with highest predation risk by jaguar are those
with a tree cover percentage over 70%. In a study carried out at local level, it is also mentioned that the predation sites by jaguar are
characterized by having dense vegetation (Rosas-Rosas et al.,
2010), particularly oaks, thorny shrubs, xeric bushes and mesquites. However, Michalski et al. (2006) suggest that a tendency
exists in the spatial variation in predation events, such that most
events occur in ranches embedded in large forested zones. This
must be taken cautiously, as it has been observed that in these
zones frequency of attacks is negatively related to the abundance
of wild prey (Stahl et al., 2002; Polisar et al., 2003; Woodroffe
et al., 2005; Kolowski and Holekamp, 2006).
Maxent also indicates that the percentage of free-grazing livestock is related with the risk, being higher when this variable is
above 20%. The lack of shelter for livestock is a situation that in
many studies has been considered the main cause of predation
by felids (Jackson et al., 1996; Mazzolli et al., 2002; Wang and Macdonald, 2006; Van et al., 2007; Inskip and Zimmermann, 2009). The
spatial exposition of livestock to predators, lack of vigilance as well
as a decient sanitary and feeding management favor a higher incidence of diseases that weakens the animals. Likewise, the birth of
offspring in sites of risk and the lack of forage in dry season are
both factors that propitiate predation.
The zones identied with the highest predation risk by jaguar
and where the percentage of free-grazing animals is the highest

85

at national level correspond to pennsula de Yucatn, the states

of Sonora, Durango, San Luis Potos, Nuevo Len and Tamaulipas,
the central part of the Costa del Pacco, Veracruz and Tabasco (INEGI, 2009). Pennsula de Yucatn holds one of the most important
jaguar populations, in this region the population of cattle reaches
898,393 animals, of which more than 60% are in free-grazing production system (Chvez and Zarza, 2009). Tamaulipas, Durango
and San Luis Potos belong to the arid and semi-arid productive region, characterized by deteriorated rangelands, whereas the Golfo
de Mxico and the Peninsula de Yucatn are comprised in the tropical humid region that provides 33% of the national beef production; however the reproductive parameters are low due to the
lack of management (SAGARPA, 2006). Particularly, in Veracruz
most of the operations are characterized by resorting to free grazing (Vilaboa and Daz, 2009).
Altitude explained about 10% of model variation, and there are
two ranges (from 0 to 300 and from 2700 to 3000 m above sea level) in which the risk of attack by jaguar increases; the rst related
to the cases that took place in the Costa del Pacco, the Golfo de
Mxico and the Pennsula de Yucatn; and the second, to the Cuenca del Balsas and Sierra Madre Oriental, where it has been registered the presence of jaguar at higher altitudes that its typical
distribution (Monroy-Vilchis et al., 2008).
Maxent indicates that the predation risk by puma increases
from 25% of forest vegetation (conifer forest, oaks and riverside
vegetation), the use of this vegetation by puma, in particular
pine-oak forest, was previously reported in central Mexico (Monroy-Vilchis et al., 2009). These results differ from Polisar et al.
(2003), who mention that in Venezuelan llanos predation by puma
seems to be less limited by the proximity of forest in comparison
with jaguar; however, in Brazil, Palmeira et al. (2008) found that
50% of the pastures with predation events by puma were close to
forest.
As for altitude, from 2300 m above sea level the risk of attack by
puma is considered high. The relation of sites of attack by puma
with this variable is evident in the model, in which the zones with
the highest risk virtually are present in all Mexican sierras, which
are the areas with highest altitude in the country. It is also possible
that the predation risk in these zones is determined by the proximity to cliffs, as it has been observed related to it in regional level
studies (Bueno-Cabrera, 2004; Zarco-Gonzlez et al., 2012). In a
similar way, in a study performed in Patagonia, Kissling et al.
(2009) found that at paddock level, topography had a notable effect
on the risk of attacks by puma.
There is an important negative relation between livestock density and predation risk by puma, even with a contribution percentage higher than before mentioned variables. The highest values of
livestock density are present in central Mexico, mainly in the
States of Mexico, Hidalgo, Puebla and Quertaro. The management
of livestock in this zone, which is one of the most important in livestock production at national level (INEGI, 2008), is different to the
rest of the country, farms in this region are intensive, many of them
dedicated to fattening and export cattle (SAGARPA, 2006). These
farms generally use fences and are away from forested areas,
which reduce the risk of predation by puma.
In comparing the variables that determine the risk of predation
for each species of felid, it is clear the puma prey more frequently
in mountain environments. The puma has a less elusive behavior,
so it often hunts near human settlements, unlike the jaguar which
usually hunts animals that are grazing freely in preserved environments with a high percentage of forest cover. These conditions occur mainly in the Southeast zone of the country, and reect the
primarily neotropical distribution of the jaguar.
It is noteworthy that the use of algorithms for ecological niche
modeling to identifying zones of predation risk is an innovative approach to this topic. The AUC of the models obtained by the differ-

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M.M. Zarco-Gonzlez et al. / Biological Conservation 159 (2013) 8087

ent algorithms were generally acceptable; the greatest accuracy

that was obtained by Maxent has been mentioned in other studies,
which shown that it produces good results with species of wide
distribution and few records (Hernandez et al., 2008). In this case,
AUC value from Maxent for puma model was even higher than the
ensemble model, which has been observed in other studies (Marmion et al., 2009). However, some authors have mentioned that the
evaluation of a model basically depends on the study objectives
and its usefulness, more than only on statistical results (Guisan
and Zimmermann, 2000). One of the disadvantages of Maxent is
that the prediction area is generally smaller contrasted to the results from other algorithms. Considering the approach of this study
and the intended application for the risk map, it is not convenient
to use a model that may predict a reduced area; in this case it is
better to incorporate uncertainty or disagreement areas between
algorithms in order to prevent underestimating the potential risk
area, which was achieved through the ensemble model.
The ensemble models are accurate approaching of the zones of
predation risk by felids; however at a regional scale the environmental characteristics that favor predation may be different. Then
it is recommended to carry out studies for each biogeographic province that facilitate the identication of specic patterns and the definition of mitigation strategies more suitable for each of them.
6. Conclusion
We generated ensemble risk models of livestock predation by
puma and jaguar in Mexico from ecological niche models, identifying the zones of risk and the environmental and livestock management variables that contributed to this risk. However, although the
initial results are potentially useful, the characteristics of specic
areas are very variable, ne-tuned models could be produced by
applying them to regional scales.
Acknowledgments
We thank to Mexican people for funding this study through
PROMEP (Project 103/10/0942), CONACYT (Project 101254), and
with a scholarship to MMZG (212618). Fernando Corts give us
some records. Two reviews anonymous strengthen the manuscript.
Luis Cejudo helps us with the translation of the text and C. Gienger
review the english version.
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