478
Abstract: The annual and seasonal diet of the bobcat (Lynx rufus) was determined from analysis of 188 feces in the
Cape region of Baja California, Mexico, an arid zone with numerous subtropical elements in its flora and fauna. Bobcats
fed mainly on lagomorphs, which reached 74% of occurrence, followed by rodents (40%), reptiles (15%), and birds
(12%). No seasonal variations were observed. The results were consistent with those of studies elsewhere, indicating that
bobcats still rely upon lagomorphs for much of their food in southern latitudes. This supports the hypothesis that lynx
have evolved to prey on hares and rabbits. The prevalence of reptiles as prey of hohcats in our study area was the
highest ever reported. They were reported as bobcat prey in only I of 20 studies from north of latitude 40°, but in 14
of the 18 studies carried out south of this latitude. With regard to feeding on reptiles, the habits of bobcats in Baja
California Sur resemble those of other similar-sized felids in tropical areas, such as ocelots (Felis pardalis) and servals
(Leptailurus servo!).
Résumé : L’analyse de 188 tbces recueillies dans la region du Cap, en Basse Californie, Mexique, une zone aride
comptaot de nombreux éléments sub-tropicaux dans sa faune et sa fore, a permis de determiner les regimes alimentaires
annuel et saisonnier du Lynx roux (Lynx rufus) dans la region. Les lynx se nourrissent surtout de lagomorphes, jusqu’b
74% du régime, de rongeurs (40%), de reptiles (15%) et d’oiseaux (12%). Aucune variation saisonnière n’a été
enregistrée. Ces résultats concordent avee ceux obtenus en d’autres regions, ce qui semble indiquer que les lynx
comptent surtout sur les lagomorphes comme proies aux latitudes australes. Ces données appuient l’hypothèse selon
laquelle le lynx s’est adapté a chasser les lièvres et les lapins. La proportion de reptiles dans le régime alimentaire des
lynx de cette region est Ia plus haute jamais signalée. Des reptiles ont été mentionnés dans le régime alimentairc du
Lynx roux dans seulement l’une des 20 etudes connues au nord du 40° paralléle, mais daos 14 des 18 etudes connues
au sud de cette latitude. En cc qui conceroe Ia consommation de reptiles, les habitudes des lynx de la Basse Californie
se rapprochent de celles d’autres félins de taille semblable des regions tropicales, l’Ocelot (Felis pardalis) et le Serval
(Leptoilurus serval).
[Traduit par la Redaction]
Introduction
Conspecific populations of widely distributed species will
encounter geographical differences in the pressures that mold
their feeding behavior (Arnold 1981). Hence, we would expect
the regional diets of species having wide geographical distributions to reflect the availability of different prey resources.
The bobcat (Lynx rufus) is the most widely distributed
native cat in North America, ranging from southern Canada
(approximately 52°N) to southern Mexico (17°N) (Anderson
1987). Its food habits are well known in many parts of this
Received June 7, 1996. Accepted October 7, 1996.
M. Delibes’ and S.C. Zapata. EstaciOn Bioldgica dc Doiiana,
Consejo Superior de Investigaciones Cientificas (CS IC),
Apartado 1056, Scvilla 41080, Spain.
M.C. Blázquex. Estacidn BiolOgica dc DoAana, CSIC,
Apartado 1056, Sevilla 41080, Spain, and Ccntro de
tnvcstigaciones Bioldgicas dcl Norocste, Apartado 128,
La Paz 23000, Mexico.
R. Rodriguez-Estrella. Centro de lnvcstigacioncs BiolOgicas
dcl Norocstc, Apartado 128, La Paz 23000, Mexico, and
EstaciOn lliológica de DoOana, CSIC, Apartado 1056, Sevilla
41080, Spain.
Author to whom all correspondence should be addressed.
Can.
J. Zool. 74: 478—483 (1997)
range, and its diet is usually based on lagomorphs (hares and
rabbits), although there is some geographical variation, deer
and (or) small mammals being important prey in some areas
(McCord and Cardoza 1982; Maehr and Brady 1986). Up to
nt)w, no information has existed on the food habits of this
felid in the southernmost part of its range, which is included
in or neighboring the Neotropical region.
The cat species of northern areas seem to be quite specialized in their diet, and Holarctic lynx species (genus Lynx,
following Wilson and Recdcr 1993) are thought to have
evolved to prey on Iagomorphs (Kurten 1968). ‘l’ropical felids,
by contrast, usually take all the vertebrate taxa they can handle,
especially reptiles (Emmons 1987; Kitchcner 1991).
This paper reports the annual and seasonal diet of bobcats
in the Cape region of Baja California (Mexico), an area with
numerous suhtropical elements in its vegetation and fauna
(Goldman and Moore 1946). We expected that lagomorphs
were important in the diet of Baja California bobcats (as they,
like other lynx species, have evolved to capture lagomorphs),
but also that reptiles would be more important here than elsewhere (owing to subtropical conditions in the area). To try
to detect whether there is a latitudinal gradient in the impor
tance of lagomorphs and reptiles as bobcat prey, we review
38 papers concerning the Ibod habits of this species in differ
cnt areas.
© 1997
NRC
Canada
Delibes el al.
Bobcats in Baja California belong to the subspecies Lynx
rufus peninsularis and are reported to be the sma Nest members
of the species (Thomas 1898; Samson 1979). Given the strong
relationship between predator and prey sizes (Rosenzweig
1966), especially in feuds (Leyhausen 1965), we expected
this small size to influence the kind of prey taken.
Study area
Fieldwork was conducted in an area of about 10 km2 in and
around El Comitán, owned by the Centro de Investigaciones
Biológicas del Noroeste. It is a coastal lowland located 17 km
north of La Paz City (24°17’N, 110°20’W). Soils are sandy
loam, while the xerophytic sarcocaulescent scrub vegetation
(Leon de Ia Luz et a!. 1996) is dominated by cardon cacti
(Pachycereus pringlel), dagger cacti (Stenocereus gu,n,nosus),
mesquites (Prosopis articulara), palo verde trees (ercidiu,n
microphyllurn), Adam’s trees (Fouquieria thguem), plum
(Cyriocarpa edulis), copales (Bursero spp.), lomboys
(Jairopha cinerea), and chollas (Opunüa cholla). This kind
of vegetation (“cardonal”) is very characteristic of the best
preserved habitats in Baja California Sur. Mean annual rainfall
is 150 mm, with precipiuition concentrated during the summer
months, from July to September. Mean annual temperature
is 23.9°C.
Methods
188) during 1994 by walking periodically along sandy paths and dry riverbeds. At least two females
with kittens and some males were known to be living in the area
during the study period. All of the area was covered regularly, so
We collected bobcat feces (n
we considered the collected feces to be a representative sample of
those produced, although some were found in easily recognizable
clusters ( toilets” or fecal marking locations; Bailey 1972). Feces
were separated, labeled, and dried at 60°C until they reached a
constant dry mass. If possible, prey items were identified to species
by comparing hairs, teeth, feathers, scales, and bones with a reference collection. Results are presented as frequency of occurrence
(number of occurrences of each prey type x 100 divided by the
number of fecal samples; this implies that the sum of frequencies
may be above 100). This method does not accurately reflect the
mass of ingested material (e.g., Weaver 1993), but it is usually considered to give a good representation of food habits (Corbett 1989).
Samples were divided into four, according to season: winter
(January—March), spring (April—June), summer (July—September),
and autumn (October—December). Numbers of occurrences were
compared among samples through a contingency-table analysis
(G test; Sokal and Rohlf 1981).
Results
Prey items in bobcat feces from Baja California Sur included
at least 9 species of mammals, 7 birds, 5 reptiles, and 1 arthro
pod (Table 1). The most important prey category was lagomorphs, which occurred in 73.9% of the feces. Rodents were
the second most important component of the diet, as they
occurred in 40. 1 % of the feces. Reptiles ranked third in
importance (15.4% in occurrence), followed by birds (12.2%).
Scorpions were also found.
In the study area the predominani prey of the bobcat were
the jackrahbit and cottontail rabbit, consumed in about the
same proportions. The pocket mouse was the most frequently
479
consumed rodent, followed by the white-tooted mouse and
desert wood rat. The California quail was the most frequent
bird in the analyzed feces. Among reptiles, the spiny-tailed
iguana was the predominant prey species. It ranked fourth in
occurrence, exceeded only by the two leporids and the pocket
mouse (Table 1).
No significant seasonal changes in prey composition were
detected: bobcats fed primarily on rabbits and secondarily on
rodents in all seasons (Table 1). This lack of seasonal change
in diet suggests that prey abundance and vulnerability do not
vary greatly during the year, which is characteristic of tropical environments.
Discussion
Lagomorphs occurred in most of the seats, with little variation between seasons, indicating that bobcats still rely upon
them for much of their food near the southern limit of their
range. This is consistent with the results of most studies of
bobcat diets, irrespective of latitude. Lagomorphs occurred
in more than 30% of the samples (stomachs and (or) intestine
and feces) in 30 (83%) of 36 studies of bobcat diets in areas
ranging from 48°N to 24°N (Table 2). Lagomorph remains
were present in 49% or more of the samples in almost onehalf of these studies. In two cases only, they reached less
than 10% occurrence, in each case being replaced by another
mammal species, the mountain beaver (Aplodontia rufa; 74%
frequency of occurrence versus 7% for lagomorphs in 247
feces from Oregon; Witmer and deCalesta 1986) and whitetailed deer (79% of occurrence in 43 feces collected during
the winter in Massachussetts, with only traces of leporids,
being found; McCord 1974).
These results support Kurten‘s (1968) hypothesis concerning the evolution of the feeding specialization of lynx species
on lagomorphs. Nevertheless, rodents seem to be more important than rabbits as food of bobcats in the southern Appalachians and western United States, and deer are also a notable
prey of bobcats in the northeastern United States (Machr and
Brady 1986).
Besides the fact that bobcats depend upon lagomorphs
even at subtropical latitudes, the most distinctive feature in
the diet of bobcats in Baja California is the high frequency of
occurrence of reptiles, which appeared in more than 15% of
the feces. The reptiles included at least two species of iguanas
and three species of snakes, one of them venomous (Table 1).
Both of the iguanas are relatively large, reaching 300 g in the
case of the spiny-tailed iguana. These were consumed also in
winter, in spite of the fact that they have a short hibernation
period in the area (Blázqucz and Ortcga-Ruhio 1996).
Predation on reptiles by bobcats has been considered
unusual elsewhere. In the review by Anderson (1987), reptiles
were not even referred to as an occasional prey of the species, while in the reviews by McCord and Cardoza (1982)
and Rolley (1987), it is stated that bobcats cat mammals and
some birds, although in certain circumstances they can take
almost anything available, including reptiles.
Approximately 40% of the studies reviewed reported one
or more reptiles as prey of the bobcat (Table 2), including
snakes (51.4% of all identified reptiles; most were nonvenomous) aiid lizards (48.6%). The frequency of occurrence
Cc)
t997 NRC Canada
480
Can. J. Zool, Vol. 75, 1997
Table 1. Seasonal diet (numbers of prey items) of bobcats from El Comitán, Baja California Sur, through analysis
of feces.
Frequency of occurrence
Total
(n
Mammals
Lagomorphs
Cottontail (Sylvilagus sp.)
Black-tailed jackrabbit (Lepus californicus)
Unknown lagomorph
Total lagomorphs
Rodents
Antelope squirrel (Animosperniophilus leucurus)
Pocket gopher (Thoniornys sp.) Pocket
mouse (Chaetodipus sp.) Kangaroo rat
(Dipodomys merriami) White-footed
mouse (Peromyscus sp.) Desert wood
rat (Neotoma lepida) Unknown rodent
Total rodents
Others
Domestic cat (Fells catus)
Total mammals
Birds
American kestrel (Falco spa rverius
California quail (Callipepla calfornica)
White-winged dove (Zenaida asiatica)
Ground dove (Columbina passe rina)
Common flicker (Colaptes auratus)
Flycatcher (Myiarchus sp.)
Thrasher (Toxostoma sp.)
Unknown bird
Total birds
Reptiles
Desert iguana (Dipsosaurus dorsalis)
Spiny-tailed iguana (Ctenosaura hemilopha)
Unknown lizard
Whipsnake (Masticophis flagellum)
Gopher snake (Pituophis melanoleucus)
Speckled rattlesnake (Crotalus mitchelli)
Unknown snake
Total reptiles
Arachnida
Scorpion
Note:
vi
=
Winter
= 74)
188) (n
Spring
(n
=
Summer
= 24)
Autumn
(n = 41)
49) (a
G
p
28.2
26.0
19.7
73.9
76.9
67.3
70.8
78.0
1.919 0.7
39.0
1.3
95.9
38.7
0
97.9
41.4
0
91.6
41.4
0
100
1.778 0.7
8.1
22.4
4.2
12.5
4.519 0.2
2.1
15.4
21.0
12.6
16.7
9.7
2.802 0.4
1.1
0
0
4.8
1.6
1.0
19.1
3.2
4.7
4.7
5.8
40.1
0.5
96.8
—
1.787 0.7
0.5
4.2
1.1
1.1
1.1
1.6
0.5
2.1
12.2
2.1
5.3
1.6
1.1
2.1
1.1
0
is the number of feces examined.
of reptiles ranged between 0 and 15.4% (this study), hut
rarely exceeded 2%. A definite north to south trend was not
detected, but reptiles were reported as bobcat prey in only I
of 20 studies carried out north of 40N, but in 14 of 18 studies
carried out south of this latitude (Table 2).
North to south differences in predation upon reptiles could
he related to changes in reptile abundance and the distributi()n of reptile sizes. A high incidence of reptiles in the diet
of predators has been reported in desert areas at low latitudes. In a pioneer study on the ecology of Saharan birds,
Valverde (1957) stated that “in hot climates, vegetarian reptiles play the same role as prey of carnivores as rodents in
cooler climates.” Likewise, Hernández et al. (1994) reported
the importance of reptiles and insects in the diet of coyotes
(canis lutrans) living in the arid Sonoran desert, and other
authors (e.g., Bothma et al. 1984) have also reported earniV
CO
S feeding on reptiles in other deserts. This is possibly
due to the high species divcrsity and abundance of reptiles in
these areas, because of the increased solar radiation (Schall
and Pianka 1978) and the high production effIciency of small
ectotherms under these conditions (Turner Cf al. 1976).
Moreover, reptile abundance and diversity (and possibly
total reptile biomass as well) increase toward the equator,
independently of aridity (Schall and Pianka 1978; Zug 1993;
© 1997 NRC Canads
Delibes et al.
481
Table 2. Frequency of occurrence of Iagomorphs and reptiles in the bobcat diet in different states or provinces in North
America, ordered approximately from north to south.
No. of
Geographic area
stomachs
Washington State
Washington State
Minnesota
Minnesota
North Dakota
Maine
Maine
Maine
Maine
Idaho
Idaho
Nova Scotia
Nova Scotia
Vermont
Oregon
Oregon
New Hampshire
New York
Massachussetts
Massachussetts
Utah
Virginia
Tennessee
Tennessee
California
California
Oklahoma
Oklahoma
Arkansas
Arkansas
Arizona
Arizona
Alabama
Texas
Florida
Florida
Durango
Baja California
76
404
50
73
74
88
170
No. of feces
and intestinal
samples
Percentage with
latitude
lagomorphs
reptiles
(°N)
Source
1)
250
43
81
124
40
27
37
40
50
22
51
71
72
66
36
71
54
31
30
7
49
60
52
2
60
55
34
35
176
51
32
29
Brittell et al. 1979
Knick et al. 1984
Rollings 1945
Berg 1979
Trevor et al. 1989
Westfall 1956
Litvaitis et al. 1986a
Litvaitis et al. l986b
Dibello et al. 1990
Baitey 1979
Koehler and Hornocker 1991
Matlack and Evans 1992
Mills 1984
Hamilton and Hunter 1939
Toweill and Anthony 1988
Witmer and deCalesta 1986
Litvaitis et al. 1984
Pollack 1951
Potlack 1951
McCord 1974
Gashwiler et a!. 1960
Progulske 1955
Buttrey 1979
Story et al. 1982
Sperry (in Young 1958)
Leach and Frazier 1953
Sperry (in Young 1958)
Rolley and Warde 1985
Fritts and Sealander 1978
Tumlison and McDaniel 1990
Sperry (in Young 1958)
Jones and Smith 1979
Miller and Speake 1978
Beasom and Moore 1977
Fickett 1971
Maehr and Brady 1986
Delibes and Hiraldo 1987
This study
462
452
223
160
662
113
140
47
494
247
388
208
53
107
15
1202
166
57
145
150
115
252
137
125
75
413
Approx.
Percentage with
7
1
38
34
22
49
25
72
74
1
2
0
48
48
48
48
47
45
45
45
45
45
45
45
44
44
43
43
43
42
41
41
38
37
36
36
35
35
34
34
34
34
33
33
33
30
2
1
15
27
26
24
?
39
39
61
176
218
540
188
for North America see Currie 1991). This tendency is reflected
in an analysis of the food of feral cas (Fells catus) on a
global scale (Fitzgerald 1988). That author found a significant negative correlation bctween latitude and incidence of
predation on reptiles: below 35°N, reptiles were usually found
in more than 20% of cat stomachs and intestines, but above
40°N they were found in no more than 10%. Also, bobcatsized tropical Cats, such as ocelols (Fells pardalis) in South
America and servals (Leplailurus serval) in Africa, prey on
snakes and lizards regularly (Emmons 1987; Gccrtscma 1985).
The relationship between low latitude and high incidence
of reptiles in the diets of carnivores could be related also to
differences in the availability of medium-sized and large reptile species. On average, reptiles reach larger sizes at low
0
0
0
0
0
0
0
0
0
0
0
0
0
3
0
0
0
0
0
2
2
0
1
1
5
2
0
2
0
27
latitudes with a greater number of hours of sunlight (Andrews
1982); hence, they would appear to be more rewarding in
energetic terms to carnivores. The large spiny-tailed iguanas
preyed upon in our study area belong to a subtropical species
whose range is restricted to below 29°N.
Some minor features in the diet of bobcats in Baja California
Sur arc also interesting. Several authors (Bailey 1972 and
later works) have reported selectivity for cottontails in comparison with jackrabbits. Unfortunately, the availability of
both species is unknown in our study area, but we saw jackrabbits and their sign much more often than cottontails and
their sign, although both were consumed in approximately
equal proportions. Thus, there could be also selectivity for
cottontails in Baja California.
© 1997
NRC Canada
Can. J. Zool. Vol. 75, 1997
482
The rather high frequency of occurrence of small rodents,
such as the pocket mouse, and the lack of ungulate remains
in the feces could be related to the small size of bobcats in
Baja California. In the southern and southwestern United
States and northern Mexico, rodents are important prey for
bobcats (e.g., Beasom and Moore 1977; Miller and Speake
1978; Jones and Smith 1979; Delibes and Hiraldo 1987), but
medium-sized species (above 100 g mass), such as cotton rats
(Sigmodon spp.), wood rats (Neotorna spp.), and to a lesser
degree squirrels (Sciuridae), are consumed most often, whereas
the smaller pocket mice and white-footed mice (under 30 g)
are rarely captured. Maybe these small rodents are energetically rewarding for the small bobcats in the study area.
On the other hand, we did not find ungulates in the analyzed
feces. Usually, larger individual bobcats (adult males) take
deer more often than smaller ones (females and juveniles;
Litvaitis et al. 1986a). Hence, the small Baja California bobcats could be limited in their ability to subdue these large
prey. However, the scarcity of wild ungulates (mule deer,
Odocoileus hemionus) in our study area makes it impossible
to resolve this issue.
Although the bobcat is considered one of the most studied
of all the wild felids (Kitchener 1991), more research in the
south of its range would no doubt be very useful for a more
comprehensive understanding of its ecology.
Acknowledgements
This research was partially supported by the Junta de Andalucia, Spain, through a travel grant to M.D., by the Centro
de Envestigaciones BiolOgicas del Noroeste, Mexico, and by
the CSIC, Spain. M.C.B. had a fellowship granted by CSIC
Consejo Nacional de Ciencia y TdcnologIa (C2l0/393) and
R.R.E. had a predoctoral fellowship from the CSIC. Special
thanks are extended to P. Ferreras, J. Svenson, A. Travaini,
J. Weaver, R.A. Every, and an anonymous reviewer for
useful comments on the manuscript, to Avclino Cota and
Andrbs Sanchez for their assistance in the field, and to
Javier Juste for providing literature.
—
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