Spatial relationships between Iberian lynx and other carnivores
in an area of south-western Spain
F. PALOMARES, P. FERRERAS, J. M. FEDRIANI andM. DELIBES
Estación Biológica Dofiana(CSIC), Apdo. 1056, 41080 Sevilla, Spain
Summary
1. Spatial relationships between Iberian lynx and other carnivores were studied by
radio-tracking and/or track censusing in two adjacent areas of Doñana (south-western
Spain).
2. Both radio-tracking and track censusing showed that lynx were restricted to an
undisturbed area of Pistacia lentiscus shrubs called Matasgordas. Minimum and
maximum lynx density in Matasgordas were estimated as 055 and 075 in km2.
3. Egyptian mongoose tracks were mainly detected outside Matasgordas (83% of
tracks), European badger tracks were detected most often inside Matasgordas (76%
of tracks), and red fox tracks were frequently detected both outside (54%) and inside
(46%) Matasgordas. Surveys of tracks and faeces undertaken in other 14 areas where
P. lentiscus shrubs also dominated corroborated the census data obtained inside and
outside Matasgordas.
4. Trapping and radio-tracking of mongooses and common genets indicated that
both species avoided use of Matasgordas. They were almost exclusively trapped (24
out of 25 mongooses and all of 11 genets) and mainly radio-located (945% and
954% of times, for mongooses and genets, respectively) in the areas of P. lentiscus
shrubs situated outside Matasgordas. Their densities were estimated as 02 and 003
in km2 inside, and as 20 and 07 i. km2 outside Matasgordas for mongooses
and genets, respectively.
5. Lynx may kill mongooses, genets, and foxes; thus, the avoidance of Matasgordas
by smaller carnivores (mongooses and genets) could be related to the risk of lynx
predation. It is suggested that the decline of the lynx in the Dofiana area may have
caused the increase in the population size of smaller, previously rarer carnivores.
6. The true relationship between lynx and foxes remains unclear, and badgers were
apparently indifferent to lynx presence or absence.
Key-words: aggressive interactions, carnivore community, common genets, Dofiana,
Egyptian mongooses, Eurasian badgers, red foxes, space use.
Introduction
The study of coexisting species is important to conservation biology (Cornell & Lawton 1992), but frequently the factors influencing the distribution and
local abundance of species in nature remain unclear
(Schoener 1986; Simberloff & Dayan 1991). In this
study, the spatial relationships between five carnivore
species in a selected area of Doñana (south-western
Spain), were studied in relation to management and
conservation of the carnivore community.
Recent theoretical models on dynamics of metapopulations and species coexistence in patchy landscapes have suggested that removing patches of opti-
mal habitat and high heterogeneity of the landscape
can favour ‘weedy’ species (Nee & May 1992; Palmer
1992). These species can take advantage of the local,
stochastic extinctions of other species caused by small
habitat area, thus increasing their regional abundance.
This effect is especially obvious when the extinct species may prey upon weedy species (Polls, Myers & Holt
1989; Polis & Holt 1992). These predictions match the
case of Doflana carnivores since some of the smaller
predators (red foxes, Vulpes vulpes L., Egyptian mongooses, Herpestes ichneumon L., and common genets,
Gene tta genetta L.) have recently increased their populations (Valverde 1960; Rau, Beltrán & Delibes 1985;
Palomares & Delibes 1992a; M. Delibes, unpub5
lished), while the abundance and range of habitats
occupied by Felispardina Temminck (the Iberian lynx;
a predator of the above carnivores) have decreased
(Palomares et a!. 1991). This study addresses how
Iberian lynx presence affects the space use by the
remaining carnivores.
The study area and its carnivore community
The study was carried out in the Coto del Rey, a flat
area with sandy soils in south-western Spain, located
in the northern limit of Doñana National Park
(37°9’N 6°26’W; Fig. 1). The study area contains two
roughly homogeneous patches (about 4 and 3km2) of
Pistacia lentiscus L. shrubs with variable matorral of
Halimium halimfolium L. and scattered Quercus suber
L. trees. These patches are called Matasgordas (Fig.
1). The rest of the area is mainly afforested with pines,
Pinus pinea L., or eucalyptus, Eucalyptus sp., with
scattered undergrowth mainly of H. ha!imfolium.
Smaller patches of P. lentiscus are also found both
interspersed with these forests and around some small
natural streams where Fraxinus sp. is the commonest
tree (Fig. 1). These smaller patches are fairly similar
to Matasgordas except that the overstorey, when
present, is constituted mainly of P. pinea or Fraxinus
spp. instead of Q. suber. Oryctolagus cuniculus L. (rabbits), are abundant in these patches (both Matasgordas and the smaller ones), but are rarer in other
habitats.
There are two intensities of human interference in
the study area. Matasgordas (inside the National
Park, Fig. 1) is under full protection, and people only
move occasionally and in low numbers. The remaining
area is subjected to: (i) religious pilgrimages (sometimes several thousand people); (ii) recreational activities including walking, riding, driving and picnicking;
and (iii) poaching of ungulates and rabbits, in which
carnivores are intentionally or accidentally killed
(Ferreras et a!. 1992; Palomares & Delibes 1992a).
Religious pilgrimages and intense recreational activities occur nearly every weekend and throughout the
summer. Poaching and driving are also frequent
throughout the year.
Nine carnivore species coexist in Doflana (Valverde
1960; Jaksic & Delibes 1987), but Lutra lutra (otter),
Mustela nivalis (weasel), Mustela putorious (polecat),
and Felis sylvestris L. (wild cat), are rare. The remaining species (Iberian lynx, Eurasian badgers, Meles
meles L., red foxes, Egyptian mongooses, and common genets) were considered in this study. The lynx
is the heaviest species (10—12 kg), followed by the
badger (7—8 kg), the fox (5—7 kg), the mongoose (3
kg) and the genet (2 kg).
Fig. 1. General sketch of the study area with the location of the transects for track censusing and the most relevant habitats.
Methods
.
Spatial relationships between lynx and other carnivores were studied by radio-tracking and/or track
censusing.
Twenty-five mongooses, eleven genets, and nine
lynx were captured with box-traps or padded foothold
traps, fitted with radio-collars containing tip switches
(Wildlife Materials, Inc., Carbondale, Illinois) (see
Delibes & Beltrán 1986; Palomares & Delibes 1992b),
and radio-tracked between June 1987 and November
1989 (mongooses and genets), and May 1991 and
March 1994 (lynx).
Two radio-tracking sampling strategies were performed: (i) one or two locations every day, and (ii)
periods of continuous tracking (between 12 and 48 h).
A total of 3861, 1385, and 682 locations were obtained
for mongooses, genets, and lynx, respectively.
Home ranges of mongooses and genets were estimated by the minimum convex polygon method, using
a maximum of two locations per day. Home range
overlap with the habitat used by lynx (i.e. Matasgordas; see below), the number of times that mongooses and genets were located inside this area, and
distances from capture points to both the nearest Matasgordas border and their furthest home range border
were obtained for each individual.
To see whether mongooses and genets used Matasgordas as expected by chance, the actual percentages
of home range overlap with Matasgordas were compared by Kolmogorov—Smirnov tests with the means
of five similar measurements of these proportions
obtained for each individual in randomly generated
home ranges of equal size and shape to actual home
ranges. Simulated home ranges were randomly
rotated about the initial capture point of each individual. Furthermore, results were not considered
acceptable if at least 70% of simulated home ranges
did not overlap with the total area (by the minimum
convex polygon) covered by all radio-tracked animals
(see Fig. 2b and c). These conditions imposed several
restrictions on the simulations, making the simulated
home ranges less likely to overlap with Matasgordas,
but counteracted the effect of uneven trapping effort
and unequal trapability of the animals across the study
area.
Three track censuses undertaken in July 1991, January 1992 and April 1994 were used to complement
and control the radio-tracking data. These censuses
were also used to detect fox, badger and rabbit relative
densities. Censuses in July 1991 and January 1992
were undertaken early in the morning along sandy
fire-break roads during three consecutive days, and
for distances of 56 km in Matasgordas and 88 km in
the remaining area (Fig. 1). Tracks were recorded
separately for each 0-4 km section of transect. About
4 h were needed to complete each census. The ground
was cleaned of old tracks a day before each census,
and again cleaned each day after the track count until
the census ended. For logistic problems the census in
April 1994 could not be undertaken every day during
three consecutive days, so tracks were counted 3 days
later than the last rainfall. The overall distance (56
km) could be sampled in Matasgordas, but only
60 km in the remaining area. Only carnivore tracks
were recorded in this census. Obviously data collection
in the last census was not as accurate as in the two
first ones; nevertheless, results were included as data
were valuable for the purpose of two-areas comparisons.
To add support to the data collected in the Coto
del Rey area, results of tracks and faeces searching of
the involved carnivore over the Dofiana area
(2750 km2) will also be presented. Data collection took
place during studies of lynx density and distribution
during the winters of 1986/87 (Palomares et a!. 1991)
and 1992/93 (Delibes, Ferreras & Fredriani 1993).
Details of the methods and the general study area can
be found in Palomares et a!. (1991). To remove a
possible habitat effect on the results, only sampling
undertaken in areas where P. lentiscus shrubs predominated (the plant predominating in our main
study area) are included. Results are expressed as
number of signs found per hour of sampling for a
given species.
Results
RADIO-TRACKING OF LYNX
Due to the previous observed absence of lynx outside
Matasgordas (Palomares et a!. 1991), trapping for
this species was only undertaken inside this area. Six
individuals (two adult females, one adult male, one
subadult female, one subadult male and one young
male) were trapped in the western portion of Matasgordas and three (one adult female, one young male
and one young female) in the eastern one.
Lynx were mainly restricted to Matasgordas
(Fig. 2a). They were located outside Matasgordas only
on 15% (SD = 97, n = 9 lynx) of occasions.
Locations to the north outside Matasgordas (the area
used by mongooses and genets; see below) only represented 8% (SD = 68, n = 9 lynx). Most locations
outside Matasgordas were close to the Matasgordas
border (see Fig. 2a). One subadult female and one
subadult male dispersed, quickly traversing the adjacent area to the north of Matasgordas.
RADIO-TRACKING OF MONGOOSES AND
GENETS
Twenty-four mongooses out of 25, and all of 11 genets
were captured outside Matasgordas; the other mongoose was captured on the northern edge of Matasgordas (Table 1). Comparing trapping success when
it was carried out simultaneously in both areas (for
125 days), 29 traps day’ per animal were needed to
Fig. 2. Areas used by radio-tracked lynx (a), mongooses (b),
and genets (c), between 1987 and 1994 in Coto del Rey,
Dofiana area. Every radio-tracking location is drawn for
each species.
were 19 and 37 times larger, respectively, than the
mean distances from the capture point to the nearest
Matasgordas border (Table 1), indicating that results
were probably not influenced by mongooses and
genets living too far away from Matasgordas. In fact,
only in the case of three mongooses and one genet was
the distance from the capture point to Matasgordas
border larger than the distance to the furthest home
range border (Table 1). However, these individuals
also situated their home range limits close to Matasgordas (always closer than 039km; Table 1).
Results of the random simulations of home ranges
over the capture points confirmed the above observations (Table 1). Simulated home ranges included a
significantly higher proportion of Matasgordas than
observed ones in both mongooses (D = 071,
P = 0004,
n = 17)
and
genets
(D = 071,
P = 00562, n = 7) (In both cases individuals with
home ranges unable to reach Matasgordas border (see
above) were excluded from the comparisons).
If mongooses and genets actually avoided Matasgordas for reasons other than habitat selection, a
differential use of the portion of the largest P. lentiscus
and Fraxinus sp patch that separates both halves of
Matasgordas (see Fig. 1), should also be expected. To
test this, the numbers of locations of mongooses and
genets inside this portion (just below the limit of
National Park on Fig. 1) and in the rest of the patch,
were compared with the numbers expected according
to their areas by a chi-square test. Inside the portion
separating Matasgordas (22% of the total patch),
both mongooses and genets were located significantly
less often (57%, n = 1963; and 09%, n = 666) often
than expected (x2 = 304 and 148, respectively,
d.f. = 1, P < 00001 in both cases; also see Fig. 2).
No mongoose and genet situated its core area in this
portion of the patch, while they frequently did so in
the rest of the patch (Palomares & Delibes 1994).
TRACK CENSUSING
capture 20 individuals (16 mongooses and four genets)
outside Matasgordas, against 440 traps day’ per animal to capture the sole mongoose inside (x2 = 4266,
P = 00001; Exact test of Wells & King 1980).
Overall, mongooses and genets were rarely located
inside Matasgordas (mean = 55% and 46% of
locations, for n = 24 and 10 individuals, respectively;
Table 2, Fig. 2b, c), and they included only a small
portion of this area inside their home ranges (mean
= 89% and 86%, respectively; Table 1). Most individuals of both species situated their home ranges at
the limits of Matasgordas or very close to the Matasgordas border (Table 1). Only an adult male mongoose, an adult female mongoose, and a young male
genet regularly used Matasgordas (Table 1).
Mean distances from the capture point to the furthest home range border for mongooses and genets
Only lynx, mongooses, foxes, badgers, and rabbits
were detected by track counts (Table 2). Foxes were
the carnivore detected most frequently (86% of all
carnivore tracks), followed by badgers, mongooses
and lynx. All lynx tracks were inside Matasgordas; on
the other hand, only 17% of mongoose tracks were
there (Table 2), in spite of the fact that the transect
outside Matasgordas did not include the best areas
for mongooses (see below and Figs 1 and 2b). Badgers
were more frequently detected inside Matasgordas.
Foxes were more frequently detected outside Matasgordas in the summer census, but inside it in the
winter and spring ones. Rabbits were detected nearly
43 times more frequently inside Matasgordas than
outside (Table 2). The Mann—Whitney test showed
that differences were statistically significant for all
species except foxes (Table 2).
The observations over the Doñana area agreed with
Table 1. Distances (km) from the capture point to the nearest Matasgordas border (DM) and to the furthest home range
border (DHRM), numbers of locations inside Matasgordas, and percentages of home ranges (observed and simulated) inside
Matasgordas for each mongoose and genet capture and/or radio-tracked in Coto del Rey. In cases where actual home ranges
did not include any portion of Matasgordas, the distances (km) between their nearest borders are given in parentheses.
M = male, F = female, A = adult, Y = young
% loca. in
Animal
DM
DHRM
Matas. (n)
% home range in
Matasgordas
Observed (Dist.)
Simulated
Mongooses
MY1
MA2
MA3
MA4
MY5
MA7
MA8
MY9
MA1O
FA1
FY2
FA3
FY4
FA6
FA8
FA9
FA1O
FY11
100
176
143
143
095
1•57
110
133
000
071
081
062
148
119
114
052
143
143
410
2-71
133
a
a
200
a
142
319
157
b
129
238
319
086
405
405
167
0(124)
0-6(338)
0(21)
0(4)
0(1)
09 (574)
0(2)
0(200)
705 (78)
31 (32)
b
0 (36)
0(29)
0(336)
0(150)
0.4 (271)
6.9 (659)
(122)
FY12
1’67
190
0(285)
FY13
FA14
162
114
a
238
0(3)
0(56)
FA15
114
105
0(153)
FA16
FA17
FA18
143
133
133
381
200
276
498(247)
0(87)
0(53)
Genets
MA1
MY2
MA3
MA4
MY5
MY6
MA7
FA1
FA2
FY3
FY4
100
157
105
162
181
029
152
167
029
029
157
410
a
381
419
b
529
705
3.6 (247)
0(2)
0(128)
0(179)
b
258 (150)
0(128)
0(288)
0 (212)
167 (6)
0(49)
a
b
333
452
a
148
2
0(0.01)
a
a
2
a
0(0.48)
73
2
b
0 (033)
1
0(0.14)
(038)
8
31
13
4
0
a
a
a
1
54
5
b
18
2
24
0
11
11
1
1
a
0(1.05)20
0(0.39)
a
0
55
1
0
22
4
6
a
1
26
a
21
7
25
b
49
1
10
0(0.28)
a
0(0.38)
b
15
21
21
14
a
0
Home ranges in these individuals could not be suitably estimated.
These individuals were not radio-tracked.
the results obtained in Coto del Rey (Table 3). Mongoose signs were more often found in areas where
lynx were not detected than in areas where lynx were
detected (Z = 226, P 00024, Mann—Whitney test).
No significant difference was found between areas
where lynx were and were not detected for foxes or
badgers (Z = 173 and 066, P = 00842 and 0506O,
respectively).
=
ESTIMATED DENSITY INSIDE AND OUTSIDE
MATASGORDAS
Only the density of radio-tracked species (mongooses,
genets, and lynx) could be reliably estimated.
From home range overlap, mimimum mongoose
density outside Matasgordas during the study was
estimated as 20 in km2 (see Palomares & Delibes
1992a, for details). Since only two mongooses used
Matasgordas regularly (both during the second study
year) and they were located in this area on 50% and
71 % of occasions, minimum mongoose density in
Matasgordas was estimated as 02 md. km 2 Mongoose density in this area during the first study year
was probably lower since no radio-tracked individual
was located in Matasgordas more than 7% of
occasions.
Using a similar method, minimum genet density
outside Matasgordas was estimated to be 07 md.
Table 2. Number of tracks km in 3 days of sampling* for carnivores and rabbits inside (In; 56 km) and outside (Out; 88 km
in July and January censuses, and 60 km in April census) Matasgordas. Results of Mann—Whitney tests for testing differences
between inside and outside Matasgordas for data from July and January censuses and using 04 km sections as the sampling
units are also given. Number total of tracks detected for each species are given in parentheses
between lynx and
small carnivores
Lynx
36)
(n
=
Mongoose
Fox
(n
(n
=
52)
=
Badger
1025)
(n
=
99)
Rabbit
(n 8819)
=
July 1991
In
4.3
Out
0’O
04
27
168
61
2646
39.9
18
75.7
January 1992
In
Out
11
00
07
438
46
8880
19
275
13
1930
April 1994
In
Out
11
00
0•0
61
98
14
07
b
b
11
666
121
11527
5•5
772
Z = 006
P = 095O
38
Z = 390
P < 0.001
2686
Z = 497
P < 0.001
Totals
In
Out
08
65
00
Z = 433
P < 0.001
Z
P
219
= 0O29
=
* Ground was cleaned and tracks counted during three consecutive days in the censuses of July 1991 and January 1992;
tracks were counted 3 days later than the last rainfall in the census of April 1994 (see Methods for more details).
t Rabbit tracks were not recorded.
Only 04 km sections crossing or close to P.lentiscus patches (the habitat used by mongooses; see Palomares & Delibes
1993b and Figs 1 and 2b) in the area outside Matasgordas were considered for the test.
Table 3. Signs (832% tracks and 168% faeces) found per
hour of sampling from lynx, mongooses, foxes and badgers
in places over the Doflana area where P. lentiscus shrubs
predominate
Signs (h-)
Lynx
Mongooses Foxes
Badgers
With lynx signs
Mean
431
SE
141
033
7’31
206
7
311
1’14
7
No lynx sign
Mean
SE
283
086
321
088
081
6
7
4
n
—
056
other young lynx were also using the area. Thus, minimum lynx density in Matasgordas was estimated as
075 md, km2. During 1991, only two adult females
seemed to be simultaneously using Matasgordas, and
probably two males and two young as well; thus, lynx
density was estimated as 055 md. km2. Lynx presence
outside Matasgordas varied depending on distance to
Matasgordas border (see Fig. 2a), but considering the
area where any lynx was radio-located, lynx density
to the north of Matasgordas (i.e. where mongoose
and genet density was estimated) was 005 i. km2
during the year of highest lynx use of this area.
158
km2 (Palomares & Delibes 1994). As only one
young genet used Matasgordas, estimated density inside this area was OO3 md. km2.
In 1993, three adult female lynx simultaneously
radio-tracked were located, on average, 84% of
occasions inside Matasgordas. Additionally, one
adult male was radio-located 73% of occasions inside
the western portion of Matasgordas, and two subadults (one in the western portion and another in
the eastern) were radio-located on average 77% of
occasions inside Matasgordas in the same period. One
untagged adult male was repeatedly observed in the
eastern portion of Matasgordas and it is likely that
AGGRESSIVE INTERACTIONS BETWEEN LYNX
AND OTHER CARNIVORES
Valverde (1957) reported two foxes and one mongoose
killed by lynx. An adult radio-tracked male genet was
killed by a lynx. Another young radio-tracked ‘female
mongoose might have been killed by a lynx, though
tracks were not clear enough to obtain a reliable conclusion. Additionally, five genets, a domestic cat, and
a young fox have been found killed by lynx in the
Doñana National Park since 1973.
Two adult radio-tracked mongooses (a male and a
female) were chased by a lynx on 14 April 1989; both
animals managed to escape, but none had returned to
the area by 9 and 19 June 1989, when their radiotracking stopped. An adult radio-tracked male lynx
was observed chasing a mongoose in 1993. In 1992 a
.
fight between a lynx and a fox was observed; the lynx
was biting the fox’s neck.
No evidence of aggressive interactions exists
between other carnivores in Dofiana.
Discussion
Whereas Iberian lynx used Matasgordas, mongooses
and genets used the other smaller P. lentiscus patches
outside the National Park. Foxes and badgers used
both Matasgordas and the remaining area.
Previous information suggests that lynx are very
susceptible to human disturbance and rabbit density
(Delibes 1980; Palomares et a!. 1991; Ferreras et a!.
1992), and the results of this study confirm it. They
used the relatively small but continuous patches of
suitable habitat with high rabbit density under total
protection, but did not use the discontinuous and
smaller ones (although apparently suitable for them
as well) outside the National Park. It remains unclear
which factor was more decisive (human influence or
high patchiness of suitable habitats), and further
research is required on this issue.
Human interference and presence of lynx are
different bewteen Matasgordas and the remaining P.
lentiscus patches in Coto del Rey; also, subtle differences in habitat could be important. One or more
of these differences could explain the avoidance of
Matasgordas by mongooses and genets. The absence
of human interference inside Matasgordas should not
be a reason for mongooses and genets avoiding this
area, and this aspect does not deserve further discussion.
Habitats selected by mongooses and genets should
provide adequate shelter when resting, and both protection from predation and abundant prey when foraging. Both mongooses and genets mainly selected P.
lentiscus patches with Fraxinus sp. both for activity
and resting, followed by other P. lentiscus patches
outside Matasgordas (Palomares & Delibes 1994). For
resting, mongooses use rabbit warrens and thickets,
and genets use thickets and tree hollows (Palomares
& Delibes 1994).
Rabbit warrens are very common in Matasgordas,
and though thickets and tree hollows are less abundant than in P. lentiscus patches with Fraxinus sp.,
they are more abundant than in the other P. lent iscus
patches in the study area. Furthermore, rabbits (the
staple prey of mongooses in the area; Palomares &
Delibes 1991a) are more abundant in Matasgordas
than in the other patches of P. lentiscus (Palomares et
a!. 1995), and data on foraging movements indicate
that Matasgordas is just as suitable for foraging by
mongooses as the other two types of P. lentiscus
patches (see Palomares & Delibes l993b). Although
there is no specific information on foraging movements of genets in each type of P. lentiscus patch,
there is no reason to think that their staple prey in
Dofiana (small mammals and small birds; Palomares
& Delibes 1991 b) are less abundant in Matasgordas
(see Camacho & Moreno 1989; Garcia, Calderén &
Castroviejo 1989). Therefore, subtle differences in
habitat appear unlikely to be the reason for the avoidance by small carnivores of Matasgordas. This
hypothesis is supported by the differential use that
both mongooses and genets made of the portion of
P. lentiscus with Fraxinus sp. (the best habitat for
mongooses and genets) separating both patches of
Matasgordas (i.e. where habitat was identical and lynx
were common). It appears more reasonable to interpret the results as a behavioural response of small
carnivores avoiding lynx predation risk (intraguild
predation, sensu Polis et al. 1989) inside Matasgordas,
because lynx may kill them. The results obtained during the tracks and faeces searching over the Doflana
area in places where P. lent iscus shrubs predominated
also seem to confirm the last hypothesis.
Track counts in other parts of the National Park
have shown that lynx and mongooses coexist to a
higher degree than in Matasgordas (Delibes et a!.
1992). Lynx density in that area was lower than in
Matasgordas; hence, spatial relationships between
lynx and smaller carnivores may be greatly influenced
by levels of lynx density. This possibility should be
further investigated. Some level of spatial segregation
has previously been reported between other carnivores, especially canids (e.g. Voigt & Earle 1983; Sargeant, Allen & Hastings 1987).
If the space use of smaller carnivores is influenced
by lynx presence, then the decrease in the density
of the Iberian lynx may have allowed the smaller,
previously rarer carnivores
from the comments of
Valverde (1960), mongooses and especially genets
were rarer before to increase their numbers over the
Doflana area.
The results for foxes and badgers were different
from those found for smaller carnivores. As expected,
badgers were detected more often inside Matasgordas
because no aggressive interactions were detected
between badgers and lynx, and rabbits are important
food for badgers in Doflana (Martin-Franquelo
1984).
Data on fox tracks inside and outside Matasgordas
were different in each sampling, but their numbers
were apparently very high in the three periods and
both areas. High rates of immigration from surrounding areas of Matasgordas might compensate for
population losses by lynx predation, and the so-called
‘mass effect’ (sensu Shmida & Wilson 1985; Auerbach
& Shmida 1987) might prevent low densities of foxes
where lynx are abundant due to dispersal. If the mass
effect is important, it might be expected that foxes
using Matasgordas were subordinate individuals (see
van Home
1983; Pulliam & Danielson 1991).
However, other alternative non-exclusive hypotheses
might be that foxes living inside Matasgordas are big
enough to avoid lynx, or that they are able to avoid
encounters with lynx.
—
Suitable patches for lynx and both mongooses and
genets may be different in size, prey type and density.
Planning to increase lynx distribution and density
through managing habitat composition and quality,
should include suitable patches at least as large as
those of Matasgordas. Small patches of less favourable habitat for lynx (e.g. very dense vegetation where
mongooses and genets may avoid predation and maintain a viable population) should also be interspersed
in order to favour the biodiversity of the carnivore
community.
Finally, though lynx feed almost exclusively on rabbits (Delibes 1980; Beltrán & Delibes 1991), high densities of this carnivore may favour higher densities of
rabbits by controlling populations of smaller carnivores which also feed on rabbits. The effect of lynx
presence on numbers of rabbits was studied through
simulations and field samplings (Palomares et al.
1995). Results suggest that a policy which deals with
the protection and improvement of lynx populations,
might also improve small game species’ densities.
Acknowledgements
The research was supported by DGICYT (project
PB87—0405). F.P. and P.F. had postdoctoral and predoctoral grants, respectively, of Consejo Superior de
Investigaciones Cientificas. H. Okarma, C. P. Doncaster, L. Halliwell and three anonymous referees provided useful comments and suggestions which
improved an early version of this manuscript. N. Bustamante, L. Halliwell and A. Green kindly reviewed
the English version.
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Received 19 March 1993; revision received 24 August 1994