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. 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