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Mammalia 2017; aop
Short Note
Alberto González-Gallina, Mircea G. Hidalgo-Mihart*, Freddy Pérez-Garduza,
Jesús A. Iglesias-Hernández, Adán Oliveras de Ita, Andrés Chacón-Hernández
and Octavio Vázquez-Zúñiga
Home range of a male jaguar spatially
associatedwith the landfill of the city of
PlayadelCarmen, Mexico
DOI 10.1515/mammalia-2016-0065
Received May 25, 2016; accepted December 13, 2016
Abstract: Understanding how jaguars (Panthera onca)
adapt to disturbed landscapes, especially urbanized
areas, can help us prevent adverse situations, thus reduc-
ing conflict, and perhaps even achieve coexistence with
these predators. Playa del Carmen, a city located in the
middle of a natural corridor linking two jaguar conserva-
tion units (JCUs; Sian Ka’an and Yum Balam), is facing
intense pressure from tourism-related city growth. From
January 2013 to March 2014, we tracked an adult male jag-
uar using a satellite collar and found that the presence of
the Playa del Carmen landfill had a clear influence on a
male jaguar’s home range and movements. We observed
that this particular jaguar had the smallest seasonal home
range and core areas reported in the literature (particu-
larly during the dry season in 2013, where the home range
was only 16.22 km2 and the core area was 2.5 km2) and also
that the seasonal core areas overlapped with the area cov-
ered by the landfill (with a number of important locations
within the landfill). Our results showed that male jaguars
are surviving in areas previously not considered as jaguar
habitat and are taking advantage of the human resources
within. We hope that these results encourage more jag-
uar studies to be carried out in areas disturbed by human
activities.
Keywords: food subsidy; landfill; Panthera onca; Yucatan
Peninsula.
Over the last 20years, it has been demonstrated that some
felids have the ability to live in human-dominated areas,
even suburbia. This has been observed for mountain lions
Puma concolor Linnaeus, 1771 (Riley et al. 2006), leop-
ards Panthera pardus Linnaeus, 1758 (Athreya etal. 2013)
and even tigers Panthera tigris Linnaeus, 1758 (Athreya
etal. 2014). The ecology of big cats in urban and subur-
ban areas is poorly known, especially for densely popu-
lated areas, where the potential for conflict is high. For
instance, jaguars (Panthera onca Linnaeus, 1758) are gen-
erally considered to be a species that avoids human set-
tlements (Colchero etal. 2011), to the point where habitat
modeling uses proximity to urban areas as a factor that
reduces the capacity of an area to be occupied by this
species (e.g. Rabinowitz and Zeller 2010, Rodríguez-Soto
et al. 2011). It is hard to say how jaguars adapt to urban-
ized areas as most studies on jaguar ecology and behav-
ior have been conducted in areas lacking a large human
population in the proximity or within natural protected
areas (e.g. de la Torre and Medellin 2011, Núñez-Pérez
2011, Avila etal. 2015). Jaguars in Mexico have lost most of
their distribution range to humans, with only around 16%
of the country’s land remaining as potentially suitable as
jaguar habitat (Rodríguez-Soto etal. 2013). One relevant
area for jaguar conservation is located in the northeast-
ern part of Quintana Roo state. Here, the tourism indus-
try has been the main driver of development (94.2% of
the GDP in 2006), affecting regional ecological patterns
(Pérez-Villegas and Carrascal 2000, Cascelli de Azevedo
and Murray 2007). Becoming the top tourist area of south-
ern Mexico has triggered high mean annual population
*Corresponding author: Mircea G. Hidalgo-Mihart, División
Académica de Ciencias Biológicas, Universidad Juárez Autónoma
deTabasco. km 0.5 Carretera Villahermosa-Cárdenas, Villahermosa,
Tabasco, 86039, Mexico, e-mail: mhidalgo@yahoo.com
Alberto González-Gallina: Instituto de Ecología A.C., Red de
Ambiente y Sustentabilidad. Carretera Antigua a Coatepec No. 351,
El Haya, Xalapa, Veracruz, 91070, México
Freddy Pérez-Garduza: División Académica de Ciencias Biológicas,
Universidad Juárez Autónoma de Tabasco. km 0.5 Carretera
Villahermosa-Cárdenas, Villahermosa, Tabasco, 86039, Mexico
Jesús A. Iglesias-Hernández, Adán Oliveras de Ita,
Andrés Chacón-Hernández and Octavio Vázquez-Zúñiga: Sistemas
Estratégicos para la Gestión Ambiental SEGA, S.A. de C.V. Río
Mixcoac No. 36 Int. 1001, Col Actipan, Delegación Benito Juárez,
Ciudad de México, 03230, México
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2 A. González-Gallina etal.: A jaguar’s spatial association with a landfill in Mexico
growths, particularly in the cities of Cancun and Playa del
Carmen. Both of these cities have had growth rates in the
last 20 years, reaching up to 20.5% a year (1995–2000),
with serious repercussions on the regional natural eco-
systems (Campos-Cámara 2007, Cascelli de Azevedo and
Murray 2007, Pérez-Villegas and Carrascal 2000). These
cities stand between two jaguar conservation units (JCUs;
Sian Ka’an and Yum Balam). The extensive areas that still
have preserved natural ecosystems and link both the JCUs
(Navarro-Serment et al. 2007, Chávez-Tovar and Zarza
2009, Rabinowitz and Zeller 2010, Rodríguez-Soto etal.
2013) will eventually narrow due to human development
thus becoming corridors. Playa del Carmen is amidst the
corridor connecting Sian Ka’an and Yum Balam JCUs.
Jaguars have been reported in the outskirts of the city
(Navarro-Serment etal. 2007), and have had conflicts with
the settlers, because of jaguars preying on pet dogs (Canis
familiaris Linnaeus, 1758; Remolina-Suárez 2014). Given
the rapid growth of Playa (from 10,531 inhabitants in 1990
to 135,512 in 2005 and increasing; Campos-Cámara 2007,
Secretaría de Desarrollo Urbano y Vivienda de Quintana
Roo 2010), it is possible that interactions between humans
and jaguars will increase, particularly far from the coast
where new settlements are being built in areas previously
covered by jungles (Cascelli de Azevedo and Murray 2007,
Remolina-Suárez 2014). Management activity, given the
lack of more complete knowledge on carnivores living in
human-dominated areas has been to relocate animals to a
nearby protected area (Athreya etal. 2007). In most cases,
this means relocating the problem as well. However, evi-
dence shows these animals as part of natural populations
sharing the space with densely populated human settle-
ments (Athreya etal. 2013). Given this situation, it is nec-
essary to understand how jaguars use the areas close to
human settlements and the influence on their behavior
by human-generated food resources for wildlife. Little is
known about how food sources of anthropogenic origin
(without including livestock) influence the movement and
spatial patterns of felids (Newsome etal. 2014a).
The study area is the vicinity of Playa del Carmen city,
in Quintana Roo state, Mexico (Figure 1). Because of the
city’s growth rate, it has spread in a disorderly fashion over
the surrounding landscape such that on the outskirts of
the town, wild habitat now intermingles with urban areas,
making it possible to encounter a wide variety of wildlife,
even jaguars (Remolina-Suárez 2014). The original vegeta-
tion in the area was tropical evergreen forest (Rzedowski
2006), but due to regular hurricanes and slash-and-burn
agricultural fires, most of the area is now covered by a gra-
dient of succession stages. Playa del Carmen’s city land-
fill (UTM 16N WGS84 483789, 2292051) covers 40hectares
and receives 400 tons of solid municipal waste daily. The
landfill is located outside the city, embedded in a natural
habitat matrix. As the landfill is inside an area with
natural vegetation, it is visited by a wide array of wild
species, including mammals such as raccoons (Procyon
lotor Linnaeus 1758), coatis (Nasua narica Linnaeus 1766),
grey foxes (Urocyon cinereoargenteus Schreber, 1775) and
feral dogs, as well as birds including vultures (Cathartes
aura Linnaeus 1758 and Coragyps atratus Bechstein 1793)
and herons (Bubulcus ibis Linnaeus 1758; pers. obs.).
A male jaguar (4years old and weighting 50 kg) was
captured using an Aldrich snare trap (Aldrich Snare Co.,
Clallam Bay, WA, USA) (Logan etal. 1999) on January 26,
2013, 12 km away from the outskirts of Playa del Carmen
and 7 km away from the city’s landfill. The capture, man-
agement and collaring of the jaguar was carried out under
the collection permit SGPA/DGVS/9611/12 (15 October
2012), granted to Mircea Gabriel Hidalgo Mihart by the
Dirección General de Vida Silvestre-SEMARNAT-México,
following the capture and management guidelines of the
American Society of Mammalogists (Sikes etal. 2011). The
jaguar was tracked using a satellite collar [Vectronic GPS
Plus Pro (VECTRONIC Aerospace GmbH, Berlin, Germany)
with Globalstar system and a drop-off device] during
2013–2014. The collar was programmed to take and send
a GPS location every 6h and the drop-off mechanism set
for releasing the collar 1 year and 2months after activation
(from 26 January 2013 to 13 March 2014). The collar worked
properly on the jaguar for the 411days. VHF telemetry was
used to locate the collar after drop-off. Once recovered,
the collar had recorded 61.84% successful GPS locations,
i.e. 1525 of 2466 tries). The jaguar’s location data were
distributed throughout three seasons as follows: 466 for
the 2013 dry season (January–May 2013), 551 for the 2013
rainy season (June–October 2013), and 508 for the 2014
dry season (November 2013–March 2014).
The jaguar’s home range and core areas were cal-
culated by season, using all of the data obtained from
the collar. Home range, understood as the outer limit of
the animal’s movement during its daily activities (Burt
1943), was calculated to determine the space used by the
jaguar during his movements over a given period of time.
We calculated core areas because they represent areas
of more intense use and allow us to identify sites where
resources appear to be clustered and are important to
the animal, providing a better understanding of the spe-
cific life requirements of the animal rather than simply
delimiting the peripheral areas (Harris etal. 1990, Powell
2000). Home range calculations were made using the 95%
minimum convex polygon (MCP; Mohr 1947) and at 95%
with the adaptive Kernel method (Kernel; Worton 1989).
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A. González-Gallina etal.: A jaguar’s spatial association with a landfill in Mexico 3
Also, core areas were calculated using the 50% MCP and
the Kernel estimators (Worton 1989). All home range cal-
culations were made using the Home Range Tools exten-
sion for ArcGis (Rodgers et al. 2007). Seasonal home
ranges in square kilometers for the dry season of 2013, the
rainy season of 2013 and the dry season of 2014with MCP
(95%) were 163.3, 123.9 and 106.08 km2 and with Kernel
(95%; Figure 2) were 16.22, 97.46 and 82.38 km2, respec-
tively. In the case of core areas the seasonal home ranges
for the dry season of 2013, the rainy season of 2013 and
the dry season of 2014with MCP (50%) were 11.91, 31.64,
38.15 km2 and with Kernel (50%; Figure 2) were 2.5, 2.55
and 2.48 km2, respectively. Seasonal differences in the
home range sizes and core areas were calculated with a
χ2-test contrasting observed sizes against the expected
values, being the average of the home range size and core
area for the three seasons (Siegel and Castellan 1995). The
seasonal home range sizes varied from 16.22 to 97.6 km2
with significant differences between seasons (χ2 = 307.5,
d.f. 2, p > 0.001). The average core area size was 2.5 km2,
with little variation reflecting our finding of no seasonal
differences (χ2 = 0.001, d.f. 2, p < 0.99). The seasonal home
range overlap was obtained with Kernohan etal.’s (2001)
index HRi,j = Ai/Aij (and HRj,i) calculated with the areas of
Kernel (95% and 50%). The spatial overlap index showed
values of HRi,j between 0.17 (dry season of 2014 against dry
season of 2013) and 1.75 (dry season of 2013 against rainy
season of 2013). Core areas (Kernel 50%) were even more
stable, with seasonal overlap between 0.97 and 1.
Our results showed that there was association of the
jaguar’s range with the Playa del Carmen landfill, as the
landfill polygon formed part of the animal’s core areas
during all three seasons (Figure 2). Throughout the study
period, 21.3% of the jaguar locations (325) were within
the first 500 m from the landfill polygon, including 51
inside the landfill (26 during the 2013 dry season, 18 in
Figure 1: Location of the study area in the northwest of the city of Playa del Carmen in the municipality of Solidaridad in the state of
Quintana Roo, Mexico and south of the Merida-Cancun highway.
The area has an elevation between 5 and 10m above sea level. Climate is warm sub-humid with average annual temperatures ranging from
26°C to 33°C. Mean annual rainfall is 1300mm and normally falls from June to October (Instituto Nacional de Estadística y Geografía 2013).
Main vegetation is a gradient of succession stages of evergreen forest. In the figure, the position of the Playa del Carmen city landfill is
highlighted.
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4 A. González-Gallina etal.: A jaguar’s spatial association with a landfill in Mexico
the 2013 rainy season and 7 in the 2014 dry season). To
test if the landfill polygon had an effect on the observed
jaguar locations, we calculated the seasonal distance from
each jaguar location to the landfill polygon and compared
it with the distance obtained from an equal number of
random points within the jaguar seasonal MCP at 95%
(we used MCP because it is not biased by the frequency of
use of specific areas and allowing us to identify areas that
are potentially used by the jaguar). Once we obtained the
seasonal observed and random location distances from
the landfill, we divided the distances in 500-m intervals
to construct an observed and random location distance
frequency table (Figure 3). We compared the observed
and random tables using the nonparametric Kolmogorov-
Smirnov test (K-S, Siegel and Castellan 1995). The K-S test
showed that in all seasons the observed distance frequen-
cies from the landfill were different from the random gen-
erated ones (for 2013 dry season K-S = 198.391, p < 0.001;
for 2013 rainy season K-S = 21.521, p < 0.001; 2014 dry
season K-S = 24.571, p < 0.001). Finally, the results showed
that 58.33% of the occasions that the jaguar was in the
landfill polygon were during the night (between 18:00
and 06:00h; 35 locations) and the rest of the occasions
(41.67%; 15 locations) were during the day (between 06:00
and 18:00h).
Male jaguars’ seasonal home ranges reported in lit-
erature in Mexico varied from 63 km2 (Núñez-Pérez 2006)
to 380 km2 (Chávez-Tovar 2009). This Playa del Carmen
jaguar, despite having a seasonal home range up to 97
km2, had a home range of only 16.22 km2 during the 2013
dry season. This, to our knowledge, is the smallest home
range described for a jaguar in the literature to date along
its distribution. It is quite possible that this jaguar behaves
like other carnivores in the presence of the concentrated
resources offered by the landfill (food) reducing his home
range. The increase in home range in response to the rainy
season has been documented for jaguars in areas where
seasonality is marked, probably as a consequence of the
greater mobility of potential prey as water sources are
no longer spatially concentrated (e.g. Núñez-Pérez 2006,
Figure 2: Seasonal locations, home ranges and core areas of the studied jaguar relative to the Playa del Carmen city landfill, Quintana Roo,
Mexico.
Home ranges and core areas were generated by the adaptive Kernel method at 95% and 50%, respectively. (A) Dry season 2013 (January–
May 2013), (B) rainy season 2013 (June–October 2013) and (C) dry season 2014 (November 2013–March 2014).
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A. González-Gallina etal.: A jaguar’s spatial association with a landfill in Mexico 5
Cavalcanti and Gese 2009, Chávez-Tovar 2009). There
is evidence that predators alter their activity patterns
and home range sizes in response to resource subsidies
(Newsome etal. 2014a). Home range size decreases in a
way consistent with the presence of a resource subsidy,
such as a landfill, because it offers a predictable and
concentrated food source (Blanchard and Knight 1991,
Hidalgo-Mihart et al. 2004, Bino et al. 2010). Whenever
the predators use an anthropogenic food subsidy they
generally avoid human presence (Gehrt etal. 2009, Valeix
et al. 2012). This is likely occurring with the jaguar we
tracked because most of his locations within the landfill
polygon were recorded during the night (38) when human
activity is minimal, if any at all. However, it is interest-
ing to observe that 24% (12) of the locations within the
polygon were recorded during the day (from 06:00 to
13:00h), perhaps indicating that the jaguar was searching
for diurnal prey despite the risk involved.
200
A
B
C
180
160
140
120
100
80
60
40
20
0
120
Jaguar
Random points
100
80
Number of GPS locations
Distance (m) from nearest landfill edge
60
40
20
0
120
100
80
60
40
20
0
0–500
500–1000
1500–2000
1000–1500
2000–2500
2500–3000
3000–3500
3500–4000
4000–4500
4500–5000
5000–5500
5500–6000
6000–6500
6500–7000
7000–7500
7500–8000
8000–8500
8500–9500
9000–9500
9500–10,000
>10,000
Figure 3: Comparison of the observed and randomly generated distances of the jaguar’s GPS locations relative to the Playa del Carmen
landfill polygon during the three studied seasons.
(A) Dry season 2013 (January–May 2013), (B) rainy season 2013 (June–October 2013) and (C) dry season 2014 (November 2013–March 2014).
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6 A. González-Gallina etal.: A jaguar’s spatial association with a landfill in Mexico
The jaguar probably became associated with this
landfill because he identified it as a reliable food source.
In general, jaguars are considered to be opportunistic
predators that vary their diet according to prey density
and ease of capture (Crawshaw and Quigley 2002, de
Oliveira 2002, Polisar etal. 2003, Hernández-San Martín
etal. 2015). Jaguars have a great dietary plasticity, includ-
ing over 85 documented species. The main prey items
being terrestrial diurnal animals with a body mass > 1 kg,
though other mammals, birds and reptiles are also impor-
tant components of the jaguar diet (Da Silveira etal. 2010,
Foster etal. 2010). In natural preserved habitats of similar
Mayan jungles in Belize, jaguars depend on medium-sized
prey such as armadillos (Dasypus novemcinctus) and large
prey ( > 10 kg). In disturbed areas, on the other hand,
the jaguar’s diet consists more of small wild prey com-
plemented with domestic species such as cattle and to a
lesser degree, dogs (0.3%, Foster etal. 2010). Jaguars even
occasionally feed on carrion (López-González and Loren-
zana-Piña 2002), but like most felids, they have a clear
preference for live prey (Newsome etal. 2014a). Thus, a
jaguar that constantly roams a landfill is more likely to be
attracted by the species that visit the place and feed on
the waste, such as dogs, raccoons, coatis or vultures, all
of which are regularly observed inside the landfill and its
surroundings (most natural prey items) rather than by the
human waste itself, although it can not be ruled out that
once in a while jaguars feed on disposed meat. Changes
in the diet of carnivores in the presence of anthropogenic
subsidies have been demonstrated (Dahle etal. 1998, Yirga
etal. 2012, Newsome etal. 2014b); such a change was seen
in this particular jaguar that seemed to have learned to
exploit the black vultures (Coragyps atratus) that gathered
in great numbers around the landfill (González-Gallina,
pers. com.). Jaguars have been reported to prey on domes-
tic animals on numerous occasions (e.g. Hoogesteijn etal.
2002, Navarro-Serment et al. 2007, Cavalcanti and Gese
2010, Amit etal. 2013); however, in the study area, there
were virtually no cattle, so the jaguar was likely comple-
menting his diet with dogs. Despite the few documented
cases (e.g. Belize and México; Foster et al. 2010, Cruz
et al. 2011), these predation incidents were reported in
an informal manner in many parts of Central and South
America (Butler etal. 2014). In the surroundings of Playa
del Carmen, this behavior had already led to a jaguar’s
relocation (Remolina-Suárez 2014). Public policies regard-
ing environmental education and conflict prevention
should be applied in critical areas of high probability
of conflicts, such as the outskirts of Playa del Carmen
in order to reduce conflicts to a minimum in particular
issues related to dog predation and human safety. Also,
interacting with the fauna attracted to the landfill – espe-
cially dogs – puts jaguars at risk of contracting diseases
such as canine parvovirus, canine distemper virus (CDV),
rabies, Sarcoptes, etc. Knobel etal. 2014 for a review). For
instance, CDV is capable of producing severe infections
in captive jaguars (Appel etal. 1994), and the presence of
dogs has been associated with high degrees of seropositiv-
ity in wild jaguars (Furtado etal. 2013). There are records
of dogs living on the periphery of the Playa del Carmen
area exhibiting the symptoms of CDV (Remolina-Suárez
2014), but despite this our jaguar was in good physical
condition and had no evident signs of injury or severe
parasitosis.
We hope that this note serves to encourage more
jaguar studies to be carried out in areas disturbed by
human activities, and to promote conservation actions
in the corridors that join the currently protected areas, in
addition to generating new spaces for the conservation of
this species, even in disturbed areas that still have a high
environmental value.
Acknowledgments: This study was made possible by the
support and funds granted by Sistemas Estratégicos para
la Gestión Ambiental SEGA S. A. de C.V in charge of the
biological survey for the highway project “Ramales Cedral-
Tintal, Tintal-Playa del Carmen con una longitud de 54 km
en el estado de Quintana Roo, México”. The project was
sponsored by Ingenieros Civiles Asociados, Infraestruc-
ture Division (ICAi) through the Mayab Consortium. We
thank the División Académica de Ciencias Biológicas de
la Universidad Juárez Autónoma de Tabasco for logisti-
cal support in the realization of this project. We specially
thank all the landowners of the ejidos of Agua Azul, Benito
Juárez, Cedral, Laguna de Costa Rica, Guadalupe Victoria,
Héroes de Nacozari and El Tintal for allowing us to do our
research in their land. We are grateful to those who helped
with capturing the jaguar: A. Rivera, F. Zavala, E. López, A.
de la Torre de Lara, S. Carrillo, M. Tobler and D. Simpson;
and to the veterinary team: I. Cassaigne, S. Ortiz Amador,
S. Ilescas, O. Lofer and B. Portillo. We thank CONACYT
for graduate studies scholarship number 335814/232663
awarded to A. González Gallina who is studying at the
Instituto de Ecología A.C.
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