THE SOUTHWESTERN NATURALIST 59(2): 157–162 JUNE 2014

USE OF PREY BY SYMPATRIC BOBCAT (LYNX RUFUS) AND COYOTE

(CANIS LATRANS) IN THE IZTA-POPO NATIONAL PARK, MEXICO

JOSÉ ANTONIO MARTÍNEZ GARCÍA, GERMÁN D. MENDOZA MARTÍNEZ,* FERNANDO X. PLATA P., OCTAVIO CÉSAR ROSAS
ROSAS, LUÍS ANTONIO TARANGO ARÁMBULA, AND LOUIS C. BENDER

Universidad Autónoma Metropolitana, Unidad Xochimilco, 04960, México, Distrito Federal, México ( JAMG, GDMM, FXPP)
Colegio de Postgraduados, Carretera México-Texcoco km 36.5, 56230, Estado de México, México (OCRR, LATA)
Extension Animal Sciences and Natural Resources, New Mexico State University, Las Cruces, NM 88003 (LCB)
*Correspondent: gmendoza@correo.xoc.uam.mx

ABSTRACT—To investigate dietary differences in two coexisting competing predators, we determined prey
consumed by bobcats (Lynx rufus) and coyotes (Canis latrans) from scats in the Izta-Popo National Park,
central Mexico. Use of prey by bobcats and coyotes showed a Pianka’s index of overlap of 0.94, much higher
than in other studies of the diet of these two species. Diets of bobcats were comprised primarily of eastern
cottontail (Sylvilagus floridanus, 33.7%), rodents (22.7%), Mexican cottontail (S. cunicularius, 13.1%), the
endemic and endangered volcano rabbit (Romerolagus diazi, 12.5%), Merriam’s pocket gopher (Pappogeomys
merriani, 9.3%), and long-tailed partridge (Dendrortyx macroura, 1.4%). Prey used by coyotes included primarily
eastern cottontail (43.8%), rodents (15.7%), Mexican cottontail (12.0%), Merriam’s pocket gopher (6.4%),
and volcano rabbit (5.5%).

RESUMEN—Para investigar las diferencias de dieta de dos depredadores competidores coexistentes,

determinamos las dietas de linces (Lynx rufus) y coyotes (Canis latrans) a partir de excrementos en el Parque
Nacional Izta-Popo, en el centro de México. El uso de presas por linces y coyotes mostró un ı́ndice de Pianka
de traslape de 0.94, mucho más alto que en otros estudios de dieta de estas dos especies. La dieta de lince
consistió principalmente en el conejo de Florida (Sylvilagus floridanus; 33.7%), roedores (22.7%), conejo
montés (S. cunicularius; 13.1%), el conejo de los volcanes endémico y en peligro de extinción (Romerolagus
diazi, 12.5%), la tuza (Pappogeomys merriani; 9.3%) y la gallinita de monte (Dendrortyx macroura; 1.4%). La dieta
del coyote incluyó el conejo de Florida (43.8%), roedores (15.7%), el conejo montés (12.0%), la tuza (6.4%) y
el conejo de los volcanes (5.5%).

Similar-sized carnivores often differ in use of available
prey to facilitate coexistence (Witmer and DeCalesta,
1986; Major and Sherburne, 1987; Litvaitis and Harrison,
1989; White et al., 1995; Kitchen et al., 1999; Neale and
Sacks, 2001a; Azevedo et al., 2006). Dietary studies are,
thus, important for determining trophic relations among
predators and prey (Chuang and Lee, 1997). Intraspecific
competition can influence structure of the community
(Schoener, 1974, 1982), especially between predators that
are at the top of the food chain (Hairston et al., 1960;
Oksanen et al., 1981). The coyote (Canis latrans) is a
generalist carnivore distributed throughout Mexico and
has a broad diet composed of insects, vertebrates, fruits,
and vegetation (Andelt et al., 1987; Gese et al., 1988;
Neale et al., 1998). In contrast, the bobcat (Lynxs rufus) is
a strict carnivore that preys mainly on rodents and
lagomorphs (Aranda et al., 2002) and occasionally
livestock (Fritts and Sealander, 1978; Neale et al., 1998).
Bobcats and coyotes are sympatric throughout most of
their ranges, and, although competition may occur
between the species, it has been difficult to evaluate and
likely varies among areas and with availability of prey
(Palomares and Caro, 1999). High densities of coyotes
have been associated with low densities of bobcats (Henke
and Bryant, 1999; Neale and Sacks, 2001b), which could
indicate competition.
Bobcats and coyotes are sympatric in Izta-Popo
National Park, which is located in the Trans-Mexican
Volcanic Belt (Sierra Nevada), central Mexico. Prey for
these carnivores in this national park includes the volcano
rabbit (Romerolagus diazi; Cervantes and González, 1996)
and several other species of lagomorphs as well as rodents
and birds (Bojorges, 2004; Granados et al., 2004).
Because coyotes and bobcats are sympatric in Izta-Popo
National Park, our goal was to determine prey used by
each species to assess whether partitioning of the diet was
a potential ecological factor facilitating coexistence.
Additionally, we wanted to determine whether either
species preferentially preyed on the endangered volcano
rabbit and, thus, potentially contributed to the precarious
status of this species.

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 158 The Southwestern Naturalist
Table 1. Elevation and length of transects, primary plant-associations along transects in Izta-Popo National Park, central Mexico.
Site Elevation (m)
a
Altzomoni 1 4,248–3,813
Tlamacas 1b 3,524–3,707
Tlamacas 2b 3,685–3,898
Tlamacas 3b 3,621–3,957
Zoquiapan 1a 3,418–3,445
Zoquiapan 2a 3,425–3,434
Zoquiapan 3a 3,406–3,457
a
With human activity.
b
Without human activity.
MATERIALS AND METHODS—Our study area was located in Izta-
Popo National Park and Zoquiapan, Mexico (18854’39 00 –
19833’00 00 N, 98831’11 00 –98848’10 00 W) which encompasses ca.
45,097 ha. The study area included the volcanoes Iztaccı́huatl
and Popocatépetl, which are the second and third highest
elevations in Mexico at 5,280 and 5,482 m, respectively (Arriaga
et al., 2000a). Climate was temperate, and the main vegetative
communities were oak (Quercus) and pine (Pinus) forests. Pine
forest was present at 2,500–4,000 m, and dominant species
included P. hartwegii, P. montezumae, P. pseudostrobus, P. rudis, P.
leiophylla, and P. teocote (Table 1; Arriaga et al., 2000a, 2000b;
Hernández and Granados, 2006).
Potential prey in the study area included white-tailed deer
(Odocoileus virginianus mexicanus), common opossum (Didelphis
marsupialis), nine-banded armadillo (Dasypus novemcinctus),
Mexican cottontail (Sylvilagus cunicularius), eastern cottontail
(S. floridanus), and volcano rabbit (R. diazi; Chapman et al.,
1980; Cervantes et al., 2005). Additionally, 13 species of rodents,
including Peromyscus melanotis (Castro et al., 2005b), P. aztecus
(Ramı́rez et al., 2005a), P. difficillis (Chávez and Ceballos, 2005),
P. maniculatus (Ramı́rez et al., 2005d), Reithrodontomys chrysopsis
(Lira and Gaona, 2005), R. sumichrasti (Ramı́rez et al., 2005c), R.
megalotis (Sánchez and Oliva, 2005), Neotomodon alstoni (Chávez,
2005), Neotoma mexicana (Zarza and Ceballos, 2005), Liomys
irroratus (Espinosa and Chávez, 2005), Dipodomys phillipsi (Oliva,
2005), Sigmodon leucotis (Ramı́rez et al., 2005b), and Pappogeomys
merriami (Villa and Valencia, 1991), were present. Four species
of insectivores also occurred on the site, including Sorex vagrans
orizabae (Velázquez et al., 2001), S. saussurei (Castro, 2005), S.
oreopolus ventralis (Castro, 2005a), and Cryptotis alticola (Ceballos
and Carréon, 2005). Multiple species of birds, primarily Buteo
jamaicensis, B. lineatus, Falco peregrinus, Cypseloides niger, Hirundo
rustica, and Corvus corax, were present, including potential prey
such as Dendrortyx macroura, Grallaria guatimelensis, Columbina
inca, and Zenaida macroura. Some of these species were endemic
to the study area (Chavez and Trigo, 1996; Chávez, 1999;
Bojorges, 2004).
Other carnivores present included Urocyon cinereoargenteus
nigrirostris, Bassariscus astatus astatus, Procyon lotor hernandezii, and
Nasua nasua molaris (Ramı́rez et al., 1982; Ramı́rez and
Müdespacher, 1987; Chávez and Trigo, 1996; Granados et al.,
2004; Trites and Roy, 2005). Endemic species recorded in the
study area included Romerolagus diazi, Pappogeomys merriami,
Neotomodon alstoni, Reithrodontomys chrysopsis, Thomomys umbrinus
volcanius, T. gambrinus peregrinus, Sorex vagrans orizabae, and
Peromyscus aztecas hylocetes (Ceballos and Galindo, 1984; Chávez
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vol. 59, no. 2
Length (m) Plant-association
7,400 Zacatón
4,900 Pine-Zacatón
4,900 Pine-Zacatón
4,900 Pine-Zacatón
1,900 Pine-Oyamel-Zacatón
1,900 Pine-Oyamel-Zacatón
1,900 Pine-Oyamel-Zacatón
and Trigo, 1996; Velázquez and Romero, 1999; Velázquez et al.,
2001).
We collected scats monthly during the dry (December 2004–
May 2005) and wet seasons (June–November 2005). We
randomly placed seven transects of different lengths in three
areas of the National Park (Tlamacas, Zoquiapan, and Altzomo-
ni). Transects were placed on semi-abandoned trails and roads
that had sign of wildlife (lagomorphs, rodents, and birds). In
Tlamacas (Puebla and Mexico states), we placed three transects
each 4,900 m long at an elevation of 4,122 m. In Zoquiapan in
the state of Puebla, we placed three transects each 1,900 m long
at an elevation of 3,752 m. One transect was placed in
Altzomoni, an area located between the states of Mexico and
Puebla at an elevation between 3,712 and 3,983 m, and was 7,400
m in length (Table 1).
We identified scats of bobcats and coyotes by associated tracks
and other sign and odor, color, and shape of scat (Aranda,
2000). We placed scats in plastic or paper bags labeled with the
name of the transect, location, date, and species.
Scats were analyzed at the Animal Nutrition Laboratory of
the Universidad Autónoma Metropolitana, México Distrito
Federal, México. We washed scats through sieves of different
sizes (5, 10, and 20 mm2) and separated skin, hair, and bones,
removing traces of vegetation. We identified remains using
reference guides for hairs, bones, and skulls (Glass, 1981;
Monroy and Rubio, 2003). We identified larger prey (i.e.,
lagomorphs and larger) to species and smaller prey (e.g.,
rodents, insectivores, and birds) to class. We calculated
frequency of occurrence and biomass of each item in scats
(Ackerman et al., 1984) and used the model of Corbett (1989)
to estimate biomass consumed by coyotes and the model of
Ackerman et al. (1984) to estimate biomass consumed by
bobcats.
We compared frequency of prey in scats using Fisher’s exact
test (Zar, 1996) and partitioned experiment-wise error rate by
the number of comparisons to determine significant differences
by species or class of prey in diets. We also used Fisher’s exact
test to compare the frequency of the volcano rabbit in diets to
that of all other species of prey in scats. We estimated overlap of
diet betweenPcoyotes P and bobcats using Pianka’s index (Pianka,
P 2 1/2
1973): 0 = pi qi/( pi2 qi ) , where pi and qi represent
proportions of prey in the diet of both species, respectively.
Pianka’s index varies between 0 (no overlap) and 1 (total
overlap).
Quality of habitat was ranked for prey at each site according
to the density of grasses per square meter and human activity
based on a visual estimate. Density of grasses was measured with
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 June 2014 Garcı́a et al.—Prey of sympatric bobcat and coyote 159

Table 2. Biomass estimated relative to number of consumed individuals by bobcat and coyote, in the Izta-Popo National Park,
Zoquiapan, central Mexico.

No. of Frequency of Relative biomass Relative no. of

Predator Prey prey items occurrence (%) consumed (%) individuals consumed (%)

Bobcat Sylvilagus cunicularius 41 15.24 33.00 13.13

Sylvilagus floridanus 62 23.05 49.18 33.73
Romerolagus diazi 39 15.50 7.28 12.48
Pappogeomys merrani 29 10.78 8.28 9.28
Other rodents 71 26.39 1.32 22.72
Birds 4 1.49 0.54 1.28
Not identifified 23 8.55 0.36 7.36
Coyote S. cunicularius 13 14.61 28.75 11.96
S. floridanus 28 31.46 61.02 43.79
R. diazi 6 6.74 3.07 5.51
P. merrani 7 7.87 5.49 6.43
Other rodents 17 19.20 0.87 15.71
Birds 0 0.00 0.00 0.00
Not identifified 18 22.22 0.77 16.55

five quadrants of 5 m2 in each site (Martinez et al., 2012). We
defined human activity by the presence of traces of humans
(trash, footprints, and feces). We classified habitat as excellent
when density was >2 grasses/m2 without human activity, good
when 2 grasses/m2 with human activity, fair when <2 grasses/m2
without human activity, and poor when <2 grasses/m2 with
human activity.
RESULTS—We collected 126 scats of bobcats and 70 scats
of coyotes. Frequency of prey differed in diets of bobcats
and coyotes (Fisher’s exact P = 0.012); scats of bobcats
contained fewer (8.6%) unidentified remains of prey
than did scats of coyotes (22.2%; P = 0.003). Although
only marginally different (P = 0.056), frequency of
volcano rabbits was greater in scats of bobcats than in
those of coyotes (15.5 versus 6.7%). Frequency of all
other parts of prey was similar (experiment-wise P >
0.05). Use of prey by bobcats and coyotes showed an
index of overlap of 0.94.
By class of prey, bobcats preyed primarily on lago-
morphs (59%), with approximately similar use of eastern
cottontails, Mexican cottontails, and volcano rabbits.
Similarly, lagomorphs (62%), primarily eastern cotton-
tails (44%; Table 2), were most frequent in diets of
coyotes.
Altzomoni had the highest density of grasses (2.6 –
2.8) and low human-activity. The site at El Papayo had the
lowest density of grasses (1.4 – 0.0) and high human-
activity.
DISCUSSION—Overall, use of prey by bobcats and coyotes
was extremely similar. The dietary overlap (0.94) was
much greater than that in previous studies (Fedriani et
al., 1999, 2000), although this result was confounded by
inclusion of vegetative matter in dietary analyses of
coyotes in other studies. The primary difference in use
of prey between bobcats and coyotes was the presence of
birds in the diet of bobcats (Table 2). Birds were absent in
scats of coyotes, although low frequency of occurrence
precluded statistical differences.
Dietary overlap between bobcats and coyotes was only
0.49 in central Florida (Thornton et al., 2004). Compared
to our results, this was due to greater use of larger species
by coyotes. Coyotes consumed a wide variety of larger prey
including white-tailed deer (Odocoileus viginianus), feral
pigs (Sus scrofa), and cattle (Bos taurus). In contrast,
bobcats used only one of these species of large prey,
white-tailed deer (Thornton et al., 2004). Similarly, diets
of bobcats and coyotes in California also showed less
overlap (0.69; Fedriani et al., 1999, 2000) than we
observed, with diets of coyotes differing mainly by greater
use of plants as food. Other studies have similarly shown
varying degrees of dietary overlap between bobcats and
coyotes (Huegel and Rongstad, 1986; Major and Sher-
burne, 1987; Kitchen et al., 1999). Most of these studies
differed from ours in that they included plants as food in
their analyses of the diet of coyotes. The coyote, an
omnivore, would be expected to have a greater intake of
fruits and other vegetation than would bobcats, an
obligate carnivore.
Lagomorphs were the most common prey in the diets
of bobcats and coyotes in our study, probably because
they were the most available and profitable prey present
in the study area (Griffiths, 1975). The average mass for
the Mexican cottontail (2.0 kg) and the eastern cottontail
(1.2 kg) combined offered comparable energetic han-
dling costs for coyotes and bobcats and likely resulted in
both predators obtaining more net energetic gains from
these larger lagomorphs than from volcano rabbits (0.5
kg) or the similarly-sized pocket gopher (0.7 kg;
Cervantes et al., 1990; Villa and Valencia, 1991). Energetic
gains suggest that volcano rabbits and pocket gophers
were less used as prey due to their smaller size and lower
energetic value for the predator. However, bobcats

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 160 The Southwestern Naturalist
consumed marginally more volcano rabbits than did
coyotes (15.5 versus 6.7%; P = 0.056). Because the
volcano rabbit is listed as threatened in CITES Appendix I
(United Nations Environment Programme, 2012) and
endangered in Mexico (Secretarı́a de Medio Ambiente y
Recursos Naturales, 2010), intensive management of
bobcats is one tool managers might consider to aid
recovery of the volcano rabbit.
In terms of classes of prey used, our results were
comparable with other studies in North America (Delibes
and Hiraldo, 1987), e.g., lagomorphs were the most
common item in diets of bobcats (59% by frequency).
Studies in Mexico had similar results; in northwestern
Mexico, diets of bobcats consisted of 68% lagomorphs
(36% S. auduboni, 32% Lepus), and, in the Ajusco
Mountain Range, their diets consisted of 69% lago-
morphs (42% eastern cottontail, 15% Mexican cottontail,
12% volcano rabbit; Aranda et al., 2002). In contrast, diets
of coyotes were more diverse, especially given the
inclusion of plants as food. For example, along the
Pacific Coast of Mexico, coyotes consumed mainly
rodents and insects during the dry season and included
fruits (mainly papaya and mango) during the wet season
(Huegel and Rongstad, 1986; Andelt et al., 1987; Hidalgo-
Mihart et al., 2001; Guerrero et al., 2004). Use of prey by
coyotes in our study was similar to that in studies
conducted in the Sierra del Ajusco, Mexico, where diets
were comprised of 30% lagomorphs, 24% rodents, and
22% livestock (Aranda et al., 1995). The low use of
lagomorphs when compared to our results was likely a
consequence of higher availability of larger prey, i.e.,
livestock (primarily domestic sheep), in the Sierra del
Ajusco. In areas where larger prey are common, they
increase in the diet of coyotes. For example, white-tailed
deer were the most common food of coyotes in Michigan,
followed by lagomorphs (Ozoga and Harger, 1966).
In conclusion, use of prey by bobcats and coyotes was
similar, and both species preyed upon the endangered
volcano rabbit. Although coyotes and bobcats preyed on
volcano rabbits, our study did not determine whether
either species was having a harmful effect on the
population of volcano rabbits; any decision to initiate
control of predators would require further study.
LITERATURE CITED
ACKERMAN, B. B., F. G. LINDZEY, AND T. P. HEMKER. 1984. Cougar
food habits in southern Utah. Journal of Wildlife Manage-
ment 48:147–155.
ANDELT, W. F., J. G. KIE, F. F. KNOWLTON, AND C. CARDWELL. 1987.
Variation in coyote diets associated with season and
successional changes in vegetation. Journal of Wildlife
Management 51:273–277.
ARANDA, M. 2000. Huellas y otros rastros de los mamı́feros
grandes y medianos de México. Instituto de Ecologı́a A.C. y
Comisión Nacional para el Conocimiento y uso de la
Biodiversidad, México.
ARANDA, M., N. LÓPEZ-RIVERA, AND L. LÓPEZ DE BUEN. 1995. Hábitos
//titan/production/s/swna/live_jobs/swna-59-02/swna-59-02-01/layouts/swna-59-02-01.3d  7 November 2014  12:52 pm  Allen Press, Inc.
vol. 59, no. 2
alimentarios del coyote (Canis latrans) en la Sierra del Ajusco,
México. Acta Zoológica Mexicana (nueva serie) 65:89–99.
ARANDA, M., O. ROSAS, J. J. RIOS, AND N. GARCÍA. 2002. Análisis
comparativo de la alimentación del gato montés (Lynx rufus)
en dos diferentes ambientes de México. Acta Zoológica
Mexicana (nueva serie) 87:99–109.
ARRIAGA, L., J. M. ESPINOZA, C. AGUILAR, E. MARTÍNEZ, L. GÓMEZ, AND
E. LOA. 2000a. Sierra Nevada RTP-107. Pages 427–429 en
Regiones terrestres prioritarias de México. Comisión Nacio-
nal para el Conocimiento y Uso de la Biodiversidad, México.
ARRIAGA L., J. M. ESPINOZA, C. AGUILAR, E. MARTÍNEZ, L. GÓMEZ, AND
E. LOA. 2000b. Ajusco-Chichinautzin RTP-108. Pages 430–432
en Regiones terrestres prioritarias de México. Comisión
Nacional para el Conocimiento y Uso de la Biodiversidad,
México.
AZEVEDO, F. C. C. , V. LESTER, W. GORSUCH, S. LARIVIÈRE, A. J.
WIRSING, AND D. L. MURRAY. 2006. Dietary breadth and overlap
among five sympatric prairie Carnivores. Journal of Zoology
Society of London 269:127–135.
BOJORGES, B. J. C. 2004. Riqueza de aves de la región noreste de la
Sierra Nevada. Estado de México. Acta Zoológica Mexicana
(nueva serie) 20:15–29.
CASTRO, A. I. 2005. Musaraña (Sorox saussurei, Merriam, 1892).
Pages 150–151 en Los mamı́feros silvestres de México. Fondo
de Cultura Económica, Comisión Nacional para el Conoci-
miento y Uso de la Biodiversidad, México.
CASTRO, C. A., C. M. MARTÍNEZ, U. AGUILERA, AND P. J. RAMÍREZ
2005a. Musaraña (Sorex oreopolus, Merriam, 1892). Page 148
en Los mamı́feros silvestres de México. Fondo de Cultura
Económica, Comisión Nacional para el Conocimiento y Uso
de la Biodiversidad, México.
CASTRO, C. A., C. M. MARTÍNEZ, U. AGUILERA, AND P. J. RAMÍREZ.
2005b. Ratón (Peromyscus melanotis, J. A. Allen y J. A.
Chapman, 1897). Pages 754–756 en Los mamı́feros silvestres
de México. Fondo de Cultura Económica, Comisión Nacio-
nal para el Conocimiento y Uso de la Biodiversidad, México.
CEBALLOS, G. G., AND A. G. CARREÓN. 2005. Musaraña (Cryptotis
alticola, Merriam, 1895). Page 126–127 en Los mamı́feros
silvestres de México. Fondo de Cultura Económica, Comisión
Nacional para el Conocimiento y Uso de la Biodiversidad,
México.
CEBALLOS, G. G., AND L. C. GALINDO. 1984. Mamı́feros silvestres de
la cuenca de México. Ed. Limusa S. A. de C. V., México
Distrito Federal.
CERVANTES, A. F., AND F. X. GONZÁLEZ. 1996. I. Los conejos y libres
silvestres de México. Pages 17–25 en Ecologı́a y conservación
del conejo zacatuche y su hábitat. Universidad Nacional
Autónoma de México Fondo de Cultura Económica México.
CERVANTES, A. F., P. DELGADO, AND A. L. COLMENARES. 2005. Conejo.
Sylvilagus cunicularius (Watherhouse, 1848). Pages 842–843
en Los mamı́feros silvestres de México. Fondo de Cultura
Económica, Comisión Nacional para el Conocimiento y Uso
de la Biodiversidad, México.
CERVANTES, A. F., C. LORENZO, AND R. S. HOFFMANN. 1990.
Romerolagus diazi. Mammalian Species 360:1–7.
CHAPMAN, J. A., J. G. HOCKMAN, AND C. M. M. OJEDA. 1980.
Sylvilagus floridanus. Mammalian Species 136:1–8.
CHÁVEZ, C. J. M., AND B. N. TRIGO. 1996. Programa de manejo
para el Parque Nacional Izta-Popo. Universidad Autónoma
Metropolitana Unidad Xochimilco, México Distrito Federal.
CHÁVEZ, L. G. 1999. Long-tailed tree-quail (Dendrortyx macroura):
current knowledge and research needs. Pages 15–26 in
Page 160
 June 2014 Garcı́a et al.—Prey of sympatric bobcat and coyote
Conservation of quail in the Neotropics (J. C. Eitniear, J. T.
Baccus, S. L. Dingle, and J. P. Carroll, editors). Center for the
Study of Tropical Birds, San Antonio, Texas.
CHÁVEZ, T. C. 2005. Ratón de los volcanes (Neotomodon alstoni;
Merriam, 1898). Pages 699–701 en Los mamı́feros silvestres
de México. Fondo de Cultura Económica, Comisión Nacio-
nal para el Conocimiento y Uso de la Biodiversidad, México.
CHÁVEZ, T. J. C., AND G. CEBALLOS. 2005. Ratón (Peromyscus
difficillis; J. A. Allen, 1891). Pages 729–730 en Los mamı́feros
silvestres de México. Fondo de Cultura Económica, Comisión
Nacional para el Conocimiento y Uso de la Biodiversidad,
México.
CHUANG, S. A., AND L. L. LEE. 1997. Food habits of three carnivore
species (Viverriicula indica, Herpestes urva, and Melogale
moschata) in fushan forest, northern Taiwan. Journal of
Zoology 243:71–79.
CORBETT, L. K. 1989. Assessing the diet of dingoes from feces: a
comparison of three methods. Journal of Wildlife Manage-
ment 53:343–346.
DELIBES, M., AND F. HIRALDO. 1987. Food habits of the bobcat in
two habitats of the southern Chihuahuan desert. Southwest-
ern Naturalist 32:457–461.
ESPINOSA, L. A., AND T. C. CHÁVEZ. 2005. Ratón espinoso (Liomys
irroratus; Gray, 1868). Pages 628–629 en Los mamı́feros
silvestres de México. Fondo de Cultura Económica, Comisión
Nacional para el Conocimiento y Uso de la Biodiversidad,
México.
FEDRIANI, J. M., F. PALOMARES, AND M. DELIBES. 1999. Niche
relations among three sympatric Mediterranean carnivores.
Oecologia 121:138–148.
FEDRIANI, J. M., T. K. FULLER, R. M. SAUVAJOT, AND E. C. YORK. 2000.
Competition and intraguild predation among three sympat-
ric carnivores. Oecologia 125:258–270.
FRITTS, S. H., AND J. A. SEALANDER. 1978. Diets of bobcat in
Arkansas with special reference to age and sex differences.
Journal of Wildlife Management 42:533–539.
GESE, E. M., O. J. RONGSTAD, AND W. R. MITON. 1988. Relationship
between coyote group size and diet in southeastern Colo-
rado. Journal of Wildlife Management 52:647–563.
GLASS, P. B. 1981. A key to the skulls of North American
mammals. Oklahoma State University, Stillwater.
GRANADOS, S. D., R. G. F. LÓPEZ, G. M. A. HERNÁNDEZ, AND A.
SÁNCHEZ-GONZÁLEZ. 2004. Ecologı́a de la fauna silvestre de la
Sierra Nevada y la Sierra del Ajusco. Revista Chapingo, Serie
Ciencias Forestales y del Ambiente 10:111–117.
GRIFFITHS, D. 1975. Prey availability and the food of predators.
Ecology 56:1209–1214.
GUERRERO, S., M. H. BADII, S. S. ZALAPA, AND J. A. ARCE. 2004.
Variación espacio-temporal en la dieta del coyote en la costa
norte de Jalisco. México. Acta Zoológica Mexicana (nueva
serie) 20:145–157.
HAIRSTON, N. G., F. E. SMITH, AND L. B. SLOBODKIN. 1960.
Community structure, population control, and competition.
American Naturalist 94:421–425.
HENKE, S. E., AND F. C. BRYANT. 1999. Effects of coyote removal on
the faunal community in western Texas. Journal of Wildlife
Management 63:1066–1081.
HERNÁNDEZ, G. M. A., AND S. D. GRANADOS. 2006. El Parque
Nacional Iztaccı́huatl-Popocatépetl-Zoquiapan y el impacto
ecológico-social de su deterioro. Revista Chapingo, Serie
Ciencias Forestales y del Ambiente 12:101–109.
HIDALGO-MIHART, M. G., L. CANTÚ-SALAZAR, C. A. LÓPEZ-GONZÁLEZ,
//titan/production/s/swna/live_jobs/swna-59-02/swna-59-02-01/layouts/swna-59-02-01.3d  7 November 2014  12:52 pm  Allen Press, Inc.
161
E. MARTÍNEZ-MEYER, AND A. GONZÁLEZ-ROMERO. 2001. Coyote
(Canis latrans) food habits in a tropical deciduous forest of
western Mexico. American Midland Naturalist 146:210–216.
HUEGEL, C. N., AND O. J. RONGSTAD. 1986. Winter foraging
patterns and consumption rates of northern Wisconsin
coyotes. American Midland Naturalist 113:203–207.
KITCHEN, A. M., E. M. GESE, AND E. R. SCHAUSTER. 1999. Resource
partitioning between coyotes and swift foxes: space, time, and
diet. Canadian Journal of Zoology 77:1645–1656.
LIRA, I. E., AND S. GAONA, 2005. Ratón (Reithrodontomys chrysopsis;
Merriam, 1900). Page 780 en Los mamı́feros silvestres de
México. Fondo de Cultura Económica, Comisión Nacional
para el Conocimiento y Uso de la Biodiversidad, México.
LITVAITIS, J. A., AND D. J. HARRISON. 1989. Bobcat–coyote niche
relationships during a period of coyote population increase.
Canadian Journal of Zoology 67:1180–1188.
MAJOR, J. T., AND J. A. SHERBURNE. 1987. Interspecific relationships
of coyotes, bobcats, and red foxes in western Maine. Journal
of Wildlife Management 51:606–616.
MARTINEZ, J. A., G. G. Mendoza, J. L. Alcantara, L. A. Tarango, T.
Sanchez, R. Rodriguez, and P. A. Hernandez. 2012.
Composición de la dieta y capacidad nutricional de carga
del hábitat del conejo de los volcanes (Romerolagus diazi) en
México. Revista Chapingo Serie Ciencias Forestales y del
Ambiente 18:423–434.
MONROY, V. O., AND R. R. RUBIO. 2003. Guı́a de identificación de
mamı́feros terrestres del Estado de México, a través del pelo
de guardia. México. Universidad Autónoma del Estado de
México, México.
NEALE, J. C., B. N. SACKS, M. M. JAEGER, AND D. R. MCCULLOUGH.
1998. A comparison of bobcat and coyote predation on lambs
in north-coastal California. Journal of Wildlife Management
62:700–706.
NEALE, J. C. C., AND B. N. SACKS. 2001a. Food habits and space use
of gray foxes in relation to sympatric coyotes and bobcats.
Canadian Journal of Zoology 79:1794–1800.
NEALE, J. C. C., AND B. N. SACKS. 2001b. Resource utilization and
interspecific relations of sympatric bobcats and coyotes.
Oikos 94:236–249.
OKSANEN, L., S. D. FRETWELL, J. ARRUDA, AND P. NIEMELÄ. 1981.
Exploitation ecosystems in gradients of primary productivity.
American Naturalist 118:240–262.
OLIVA, G. 2005. Rata canguro (Dipodomys phillipsi; Gray, 1841).
Pages 620–621 en Los mamı́feros silvestres de México. Fondo
de Cultura Económica, Comisión Nacional para el Conoci-
miento y Uso de la Biodiversidad, México.
OZOGA, J. J., AND E. M. HARGER. 1966. Winter activities and
feeding habits of northern Michigan coyotes. Journal of
Wildlife Management 30:809–818.
PALOMARES, F., AND T. M. CARO. 1999. Interspecific killing among
mammalian carnivores. American Naturalist 153:492–508.
PIANKA, E. R. 1973. The structure of lizard communities. Annual
Review of Ecology and Systematics 4:53–74.
RAMÍREZ, P. J., AND C. MÜDESPACHER. 1987. Estado actual y
perspectivas del conocimiento de los mamı́feros de México.
Ciencia 38:49–67.
RAMÍREZ, P. J., C. AGUILAR, AND C. A. CASTRO. 2005a. Ratón
(Peromyscus aztecus, Saussure, 1860). Pages 721–722 en Los
mamı́feros silvestres de México. Fondo de Cultura Econó-
mica, Comisión Nacional para el Conocimiento y Uso de la
Biodiversidad, México.
RAMÍREZ, P. J., T. J. C. CHÁVEZ, AND G. OLIVA. 2005b. Rata
Page 161
 162 The Southwestern Naturalist
algodonera (Sigmodon leucotis, Bailey, 1902). Pages 801–803
en Los mamı́feros silvestres de México. Fondo de Cultura
Económica, Comisión Nacional para el Conocimiento y Uso
de la Biodiversidad, México.
RAMÍREZ, P. J., W. R. LÓPEZ, C. MÜDESPACHER, AND I. J. LIRA. 1982.
Catálogo de los mamı́feros terrestres nativos de México.
Editorial Trillas-Universidad Autónoma Metropolitana Uni-
dad Iztapalapa, México, Distrito Federal.
RAMÍREZ, P. J., P. R. QUIJANO, U. AGUILERA, AND C. A. CASTRO. 2005c.
Ratón (Reithrodontomys sumichrasti, Saussure, 1861). Pages
789–790 en Los mamı́feros silvestres de México. Fondo de
Cultura Económica, Comisión Nacional para el Conocimien-
to y Uso de la Biodiversidad, México.
RAMÍREZ, P. J., A. SÁNCHEZ, U. AGUILERA, AND C. A. CASTRO. 2005d.
Ratón (Peromyscus maniculatus, Wagner, 1845). Pages 748–750
en Los mamı́feros silvestres de México. Fondo de Cultura
Económica, Comisión Nacional para el Conocimiento y Uso
de la Biodiversidad, México.
SÁNCHEZ, O., AND G. OLIVA. 2005. Ratón (Reithrodontomys megalotis;
Baird, 1858). Pages 784–785 en Los mamı́feros silvestres de
México. Fondo de Cultura Económica, Comisión Nacional
para el Conocimiento y Uso de la Biodiversidad, México.
SCHOENER, T. W. 1974. Some methods for calculating competi-
tion coefficients from resource-utilization spectra. American
Naturalist 108:332–340.
SCHOENER, T. W. 1982. The controversy over interspecific
competition. American Scientist 70:586–595.
SECRETARÍA DE MEDIO AMBIENTE Y RECURSOS NATURALES. 2010. Oficial
Mexicana NOM-059-SEMARNAT-2010, protección ambiental-
especies nativas de México de flora y fauna silvestres-
categorı́as de riesgo y especificaciones para su inclusion o
cambio-lista de especies en riesgo. Segunda sección. Secre-
tarı́a de Medio Ambiente y Recursos Naturales, Diario
Oficial, 30 de diciembre de 2010, México, Distrito Federal,
México.
THORNTON, D. N., M. E. SUNQUIST, AND M. B. MAIN. 2004.
Ecological separation within newly sympatric populations of
//titan/production/s/swna/live_jobs/swna-59-02/swna-59-02-01/layouts/swna-59-02-01.3d  7 November 2014  12:52 pm  Allen Press, Inc.
vol. 59, no. 2
coyotes and bobcats in South–Central Florida. Journal of
Mammalogy 85:973–982.
TRITES, A. W., AND R. ROY. 2005. Dietary analysis from fecal
samples: how many scats are enough? Journal of Mammalogy
86:704–712.
UNITED NATIONS ENVIRONMENT PROGRAMME. 2012. CITES Appen-
dice I. United Nations Environment Programme, Geneva,
Switzerland.
VELÁZQUEZ, A., AND F. J. ROMERO. 1999. Biodiversidad de la región
de montaña del sur de la cuenca de México. Universidad
Autónoma Metropolitana Unidad Xochimilco-Secretaria del
Medio Ambiente, Gobierno del Distrito Federal.
VELÁZQUEZ, A. F. J. ROMERO, H. RANGEL-CORDERO, AND G. W. HEIL.
2001. Effects of landscape changes on mammalian assem-
blages at Izta-Popo Volcanoes, Mexico. Biodiversity and
Conservation 10:1059–1075.
VILLA, C. B., AND M. J. VALENCIA. 1991. Actividad reproductiva de
la tuza Pappogeomys merriami Merriami (Rodentia: Geomyidae)
de Chalco, México. Anales del Instituto de Biologı́a. Serie
Zoologı́a 62:235–247.
WHITE, P. J., K. RALLS, AND C. A. VANDERBILT-WHITE. 1995. Overlap
in habitat and food use between coyotes and San Joaquin kit
foxes. Southwestern Naturalist 40:342–349.
WITMER, G. W., AND D. S. DECALESTA. 1986. Resource use by
unexploited sympatric bobcats and coyotes in Oregon.
Canadian Journal of Zoology 64:2333–2338.
ZAR, J. H. 1999. Biostatistical analysis. Prentice Hall, Upper
Saddle River, New Jersey.
ZARZA, H., AND G. CEBALLOS. 2005. Rata magueyera (Neotoma
mexicana; Baird, 1855). Pages 620–621 en Los mamı́feros
silvestres de México. Fondo de Cultura Económica, Comisión
Nacional para el Conocimiento y Uso de la Biodiversidad,
México.
Submitted 21 February 2013. Accepted .
Acceptance recommended by Associate Editor, Troy A. Ladine, 13 June
2013.
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