1

Spatial interactions between grey wolves and Eurasian lynx in 2

Białowieża Primeval Forest, Poland 3

4

Krzysztof Schmidt*, Włodzimierz Jędrzejewski, Henryk Okarma1 and Rafał Kowalczyk 5

6

Mammal Research Institute, Polish Academy of Sciences, PL-17-230 Białowieża, Poland 7

8

* corresponding author: phone: +48 85 6827777, fax: +48 85 68 12 289; e-mail: 9

kschmidt@zbs.bialowieza.pl 10 11 1 present address: Institute of Nature Conservation, Polish Academy of Sciences, Mickiewicza 12

33 str., PL-31-120 Kraków, Poland 13

2

Abstract 1

Various species of large predators are reported to influence each other through interference or 2

exploitation competition that may affect demography and survival of the subordinate species. 3

We analyzed spatial relationships between grey wolf Canis lupus and Eurasian lynx Lynx lynx 4

in Białowieża Primeval Forest (BPF, eastern Poland) to determine how they partitioned the 5

space. The wolves (n=8) and lynx (n=14) were radio-tracked in 1991-1999. Three wolves and 6

seven lynx were radio-tracked simultaneously in 1994-1996. Territories of wolf packs and 7

home ranges of lynx overlapped considerably (76% of wolf territories and 50% of lynx home 8

ranges, on average). In three cases, their core areas were also overlapping. Wolf – lynx dyads 9

with overlapping home ranges were simultaneously located at distances from 0 to 28 km from 10

each other. We found neither avoidance nor attraction between wolves and lynx occupying 11

the same areas. We concluded that in BPF, the two large predators coexist due to 12

specialisation on different preferred prey and heterogeneous habitat. 13

14

Key words: Canis lupus, coexistence, exploitation competition, interference competition, 15

Lynx lynx, spatial overlap 16

17

Introduction 18

19

What enables various carnivorous mammals to coexist and how they interact with each other 20

when occurring sympatrically has been a subject of numerous studies. Rosenzweig (1966) 21

suggested that coexistence results from size differences between predator species. Felids and 22

canids seem to be particularly predisposed to “peaceful coexistence” (Major and Sherburne 23

1987) through separation of their ecological niches, which likely resulted from evolution of 24

their different social systems (Kleiman and Eisenberg 1973). 25

3

On the other hand, the competitive exclusion was recently suggested between tigers 1

Panthera tigris and wolves Canis lupus even despite of extreme size differences (Miquelle et 2

al. 2005). Cases of interspecific killing are common among predators, including felids and 3

canids (review in Palomares and Caro 1999). In some instances intraguild predation may have 4

serious consequences on demography and survival of the inferior species, as it has been 5

suggested for cheetahs Acinonyx jubatus (Laurenson 1995) and African wild dogs Lycaon 6

pictus (Creel and Creel 1996, Carbone et al. 1997). 7

Grey wolves and Eurasian lynx Lynx lynx are sympatric across most of their vast 8

geographical ranges, which stretch from Eastern Europe to the Far East of Asia (Bibikov 9

1985, Nowell and Jackson 1996). The spatial interactions between these species, however, 10

have not yet been studied. Anecdotal information on their interspecific relationships (based on 11

harvest data and incidental observations) are available from the Russian part of their ranges 12

and from Fennoscandia. According to those reports, coexistence of the two large carnivores is 13

often characterized by negative influences of wolf on lynx (Pulliainen 1965, Myrberget 1970, 14

Matyushkin 1985, Malafeev et al. 1986, Matyushkin and Vaisfeld 2003). Dynamics of wolf 15

and lynx population numbers reported there suggest that lynx may reach higher numbers only 16

when wolves are rare. Cases of wolves killing lynx were also reported (see Matyushkin and 17

Vaisfeld 2003 for review). On the other hand, it has also been suggested that the relationships 18

between the two species may vary with ecological circumstances (Matyushkin 1985). Indeed, 19

in some areas populations of both predators existed in large numbers without apparent 20

reciprocal influence (e.g. Upper Volga Region: Zheltukhin 1986) or even increased 21

simultaneously (West Siberia: Azarov and Shubin 2003; Eastern Poland and Western Belarus: 22

Jędrzejewska and Jędrzejewski 1998). Snow-tracking data from several study areas in Eurasia 23

(e.g. Upper Volga Region: Zheltukhin 1986; North-West Russia: Danilov et al. 2003; Far East 24

of Asia: Matyushkin et al. 2003) showed that lynx and wolves do not avoid using the same 25

4

sites and do not exhibit any particular interest in each other, which suggests a lack of 1

interference competition (sensu Case and Gilpin 1974). 2

Wolves and lynx also have similar diets across their common range (see Matyushkin 3

and Vaisfeld 2003 for review). Therefore, one can expect that the wolves may limit lynx 4

populations through exploitation of the common food resources (exploitation competition), as 5

suggested by Litvaitis and Harrison (1989) for sympatric bobcat Lynx rufus and coyote Canis 6

latrans in Maine (USA). 7

In Białowieża Primeval Forest (BPF, E Poland) the two predators coexist and no 8

negative relationship between their long-term dynamics was noted – the wolf and lynx 9

densities were found to be significantly correlated in time (Jędrzejewska and Jędrzejewski 10

2005). Their numbers were simultaneously shaped by temporarily varying hunting harvest. 11

The ecology of both species in BPF has been studied in detail (Jędrzejewski et al. 1996, 12

Okarma et al. 1997, Schmidt et. al. 1997, Okarma et al. 1998, Schmidt 1999, Jędrzejewski et 13

al. 2000, Theuerkauf et al. 2003), but their interspecific interactions have not been analyzed. 14

The feeding niches of the two carnivores partly overlap, as wolves primarily prey on red deer 15

Cervus elaphus and supplement their diet with wild boar Sus scrofa and roe deer Capreolus 16

capreolus (Jędrzejewski et al. 2000), whereas lynx feed mainly on roe deer with the addition 17

of female and young red deer (Okarma et al. 1997). Given the high densities of ungulates in 18

BPF (Jędrzejewski et al. 2002), exploitation competition is not very likely. 19

The aim of this study was to analyze fine scale spatial interactions between radio-20

tracked wolves and lynx inhabiting the Białowieża Primeval Forest. We investigated these 21

relationships at three spatial scales: (1) the study area level - utilization of the forest by both 22

species, (2) home range level – degree of overlap between lynx and wolf home ranges and (3) 23

individual level – dynamic interactions between dyads of lynx and wolf individuals located at 24

the same time. This way we intended to determine if wolves and lynx tended to be spatially 25

5

segregated and show any signs of exclusion of one by the other (interference competition) in 1

the BPF. 2

3

Material and methods 4

5

Study area 6

7

The study was conducted in Polish part of the Białowieża Primeval Forest (BPF), Eastern 8

Poland (52º30’-53º N, 23º30’-24º15’ E). The BPF is a temperate mixed lowland forest located 9

on the Polish-Belarussian border and characterized by a high percentage of natural stands 10

(Faliński 1986). The whole forest covers nearly 1500 km2 and its Polish part about 600 km2. 11

Most of the Polish part of the BPF (500 km2) is managed by State Forestry, while the rest is 12

protected as the Białowieża National Park (BNP, 100 km2) with a 50-km2 area of strict 13

reserve, where no human interference is allowed except for tourism and research. A network 14

of small reserves with partial protection is located within the managed part of the BPF 15

(Wesołowski 2005). The dominating tree species are: pine Pinus silvestris, spruce Picea 16

abies), oak Quercus robur, hornbeam Carpinus betulus, black alder Alnus glutinosa, ash 17

Fraxinus excelsior, lime Tilia cordata, birches Betula verrucosa and B. pubescens, aspen 18

Populus tremula and maple Acer platanoides. Despite of timber exploitation, the BPF 19

maintains a unique character in comparison to other European forest in terms of: (1) tree 20

diversity (with 26 tree and 55 shrub species forming a mosaic of diverse tree communities), 21

(2) a multi-storey profile of stands, (3) relatively large amount of dead wood and (4) 22

outstanding diversity of flora and fauna (Wesołowski 2005). The climate of BPF is temperate 23

with transitional character between Atlantic and continental ones with clearly marked warm 24

and cold periods (average temperatures during the study period were -3.9º C in January and 25

19.1º C in July; average annual precipitation was 622 mm; snow cover persisted for an 26

6

average of 96 days per year from November to March). Wolf and lynx are currently the only 1

large predators in BPF. The community of ungulate mammals consists of: red deer Cervus 2

elaphus, roe deer Capreolus capreolus, wild boar Sus scropha, moose Alces alces, and 3

European bison Bison bonasus. However, only three species – red deer, roe deer and wild 4

boar – play an important role in both predators’ diets (Jędrzejewski et al. 1993, Jędrzejewski 5

et al. 2000). 6

7

Radio-tracking 8

9

The study was a part of a larger research project on wolf and lynx ecology in BPF conducted 10

in 1991-1999 (Schmidt et al. 1997, Jędrzejewski et al. 2002). The principal data for this paper 11

originates from 1994-1996, when both wolf and lynx were studied by radio-tracking: 3 female 12

wolves belonging to 2 packs and 7 lynx (2 M, 5 F) (Appendix I). Five lynx provided 13

sufficient number of locations taken simultaneously with those of wolves for calculating 14

dynamic interactions between the species. Locations of wolves and lynx taken before and 15

after 1994-1996 were, however, used for general assessment of the forest use by both species 16

(9 wolves and 16 lynx). We acknowledge that the sample of studied animals is low and we 17

didn’t monitor whole the populations of both predators. However, the information obtained 18

during this study provided minimum estimates of their spatial overlap and existing potential 19

for interactions between them. Moreover, both wolves and lynx use large home ranges (173-20

294 km2 in wolves: Okarma et al. 1998 and 133-248 km2 in lynx: Schmidt et al. 1997), so that 21

the total area used by simultaneously radio-tracked wolves and lynx covered a representative 22

portion of the study area. We estimated that during the study, the population of wolves in 23

Polish part of BPF consisted of 3-4 packs of wolves containing 2-8 individuals each 24

(Jędrzejewski et al. 2000). Lynx were estimated at 6 – 18 adult individuals (Jędrzejewski et. 25

al. 1996). 26

7

The wolves were captured both with “fladry” (a rope with strips of cloth that is used 1

to limit and drive wolves’ movements) and a net system (Okarma and Jędrzejewski 1997) or 2

foot snare traps (equipped with alarm-system) and fitted with VHF radio-collars made by 3

Telonics Inc., (Mesa, Arizona, USA), AVM Instrument Co. (Livermore, California, USA), 4

Telemetry Systems (Mequon, Wisconsin, USA), and Advanced Telemetry Systems (Isanti, 5

Minnesota, USA). Lynx were captured with snare traps set at fresh kill and marking sites or in 6

a box trap (equipped with alarm-system) and fitted with VHF radio-collars made by Wagener 7

Temetrieanlagen HF-NF Technik, (Köln, Germany) or AVM Instrument Co. The range of the 8

collars was 1 – 2 km. Both wolves and lynx were immobilized with a mixture of ketamine 9

hydrochloride (wolves: 3.4-4.5 mg/kg, lynx 5 mg/kg of body weight) and xylazine 10

hydrochloride (wolves: 5-6.5 mg/kg, lynx 6 mg/kg of body weight). 11

We located radio-collared animals 2-5 times per week by triangulation from the forest 12

roads. Additionally, each month we conducted sessions of continuous 24-h radio-tracking 13

lasting 2-7 days, during which we located animals every 30 min. During these sessions we 14

focused on one individual of a given species, although we recorded locations of the second 15

species if it was present nearby until the animals separated such that following two of them 16

simultaneously was too difficult. Locations were plotted on forest maps with 533x533 m 17

square grid (according to the forest division compartments marked in the terrain). Positions of 18

the animals were determined in the center of a square, in the middle of one of its sides or in 19

the corner between four adjacent squares. Their estimated locations could then differ from the 20

actual ones by a maximum of 373 m. We aimed at reducing the error by taking bearings from 21

points on the forest roads closest to animals (≥100 m). During continuous radio-tracking, 22

however, the observer usually stayed 500-1000 m from the focal animal to avoid disturbing it. 23

For a general assessment of space use by both species and their spatial relationships in 24

the study area we overlaid all locations of wolves (8 individuals with 28857 locations) and 25

8

lynx (14 individuals with 7521 locations) recorded between 1991-1999 on the forest map 1

using program Biotas™ v.1.03.1 Alpha (Ecological Software Solutions, Urnäsch, 2

Switzerland). To estimate an extent of overlap of both species’ occurrence in the entire study 3

area we calculated 95% minimum convex polygon ranges for pooled locations of all 4

individuals in the two species. Using 95% MCP, in this case, allowed us to maximise the 5

studied area used by both species as well as for excluding large portions of non-forested areas 6

that were not visited by them.We also calculated 75% fixed kernel ranges for the same data to 7

find out if there is an overlap of most frequently used areas by all radio-tracked lynx and 8

wolves. The fixed kernel ranges where estimated with a level of smoothing selected by least-9

squares cross-validation method and the chosen bandwidth was 750 m. 10

We also examined static and dynamic interactions (Macdonald et al. 1980) between 11

wolves and lynx. Static interactions were determined by calculating overlaps of lynx home 12

ranges with territories of wolves. We calculated an average overlap both between an 13

individual lynx home range and all available pooled wolf territories, between an individual 14

wolf pack territory and all available pooled lynx home ranges as well as between dyads of 15

lynx and wolves. We included all lynx radio-tracked simultaneously with wolves for the 16

pooled lynx home ranges, despite the low number of locations for some individuals (see 17

Appendix). The discrepancy in sample size, however, should not significantly affect the 18

results, because our aim was to estimate a minimum value of the overlap between wolf 19

territory and all detected home ranges of lynx inhabiting a common area. The home ranges 20

were estimated with 100% minimum convex polygon method using the program Biotas™ 21

v.1.03.1 Alpha, for all locations recorded during a simultaneous period of radio-tracking and 22

for a sub-sample of both species’ locations taken on the same day only (on average within 1 h, 23

max 12 h). This way we accounted for potential bias, resulting from a higher probability of 24

finding two animals with decreasing distance between them. The 100% MCP was used to 25

9

ensure detecting maximum potential overlapping area used by wolf and lynx dyads. To 1

determine core areas used by particular lynx and wolf individuals we calculated 50% fixed 2

kernel ranges (with the same smoothing parameters as for general – species’ level 3

assessment). 4

To determine if two predators ignore, avoid or attract each other, we examined their 5

dynamic interactions (Macdonald et al. 1980, Kenward 1992) using the program Ranges 6 v. 6

1.08 (R. Kenward, A. South and S. Walls, Anatrack Ltd. Dorset, UK). The program compares 7

observed and expected distances between locations of two animals. The expected distances 8

were estimated as a mean distance between each location of one animal and all locations of 9

another animal. The observed and expected distances were then compared with Jacobs’ index 10

(D) (Jacobs 1974): 11

OE

OE

dd

ddD

+

−= 12

where dE is expected distance and dO is observed distance between pairs of simultaneous 13

locations of a wolf-lynx dyad with overlapping home ranges. D ranges from -1 in case of 14

mutual avoidance, to 0 if the animals move independently, to +1 if they are attracted to each 15

other. 16

17

Results 18

19

Wolves and lynx utilized the study area in nearly the same extent (Fig. 1). Although the eight 20

wolves of four packs were more intensively radio-tracked than the 14 lynx, the minimum 21

convex polygons with 95% of pooled locations of all individuals in the two species 22

overlapped in 88%. Most frequently used areas by both predators (75% fixed kernel) also 23

showed a substantial spatial overlap: wolves on 18% of lynx core area, and lynx on 32% of 24

wolves’ core area. It is striking that both wolves and lynx were utilizing mostly the main solid 25

10

forest body, leaving it only occasionally. However, wolves seemed to wander out of the forest 1

slightly more readily than lynx (note westernmost locations of wolves on Fig. 1). 2

Home ranges of lynx individuals and territories of wolf packs overlapped each other 3

extensively (Fig. 2). Territory of each of the studied wolf packs was on average, covered by 4

home ranges of radio-collared lynx by 76% (Table 1). Home range of each lynx was covered 5

by wolf territories by 50%, on average. Smaller overlap (mean 12-14%) was recorded 6

between core areas (Table 1). These measures of overlap are certainly underestimated, since 7

in both years shown in Fig. 2, there were other (non-collared) wolf packs and lynx individuals 8

recorded by snow-tracking in the nearest vicinities of the territories/home ranges of radio-9

collared animals (comp. Jędrzejewski et al. 1996, Okarma et al. 1998). 10

When found at approximately the same time, particular dyads of wolves and lynx with 11

overlapping home ranges were located at an average distance from 4.4 to 11.0 km apart 12

(Table 2). It is noteworthy that occasionally they were found at the same location as the 13

minimum recorded distance was 0 or 0.3 km in some dyads. According to analysis of 14

dynamic interactions (based on Jacobs’ index, D) conducted for pairs of simultaneous 15

locations of wolves and lynx, the animals showed neither avoidance nor attraction to each 16

other (Table 2). A slight attraction was recorded in the case of a dyad L4-W3, which also 17

showed the most extensive overlap of their home ranges. 18

A male lynx L4 and a pack of wolves with female W3 were radio-tracked 19

simultaneously on three separate occasions, in which the animals were found at very short 20

(less than 1 km) distances from each other for longer periods (Fig. 3). On 23 May 1995, the 21

lynx approached the site where the wolf pack stayed and remained there in close proximity to 22

wolves (0.25 – 0.8 km) for 5 h. The lynx left that area in the evening and headed in the 23

opposite direction to the one he arrived. On 22 June 1995, both lynx and wolf were at 24

approximately the same location (maximum 0.25 km from each other) for 1 h. Afterwards, 25

11

they split, however, they still remained close by (0.25 – 1.4 km) in an area of about 0.8 km2 1

for the subsequent 8 h, before the lynx departed. On another occasion, 23/24 October 1995, 2

the female wolf with her pack and the lynx moved along nearly the same route over 13 km for 3

22 h. During this time, the animals rested four times, each time in different place, at a distance 4

of 0.5 – 1.5 km from each other. After the resting bouts that took from 0.5 to 2.5 h, they 5

moved alternately following each other, heading approximately in the same direction until 6

early morning, when they went separate ways (Fig. 3). 7

8

Discussion 9

10

The competition between two species of predators may be mitigated by various factors, such 11

as differences in size (Rosenzweig 1966) and social organization (Kleiman and Eisenberg 12

1973). Therefore, it is difficult to predict the effect of interactions between given species at 13

given site. Grey wolf inhabiting BPF is nearly twice as heavy as Eurasian lynx (on average, 14

45 and 35 kg, for adult male and female wolves, respectively: Jędrzejewski et al. 2007; 22 15

and 17 kg for male and female lynx, respectively: Schmidt et al. 1997). Size difference may 16

lead to interference competition through interspecific killing, as suggested by Palomares and 17

Caro (1999). On the other hand it may diminish exploitive competition through partitioning of 18

prey resources (Scognamillo et al. 2003). Different life habits of canids and felids – sociality 19

and coursing hunting behaviour of wolves versus solitary life and stalking hunting common in 20

felids – may farther ease resource partitioning, as found by Husseman et al. (2003) for 21

sympatric wolves and cougars Puma concolor. 22

Our study showed that the wolves and the lynx did not exclude each other spatially in 23

the Białowieża Primeval Forest. They behaved neutrally, neither avoiding nor attracting each 24

other, which suggests that they used the available area completely independently. Cases of 25

12

extensively overlapping core areas and parallel travelling recorded during this study are also 1

indicating that individuals of the two species do not particularly tend to separate from each 2

other while using the same area. The patterns of their activity were also greatly overlapping 3

(compare Schmidt 1999 and Theuerkauf et al. 2003), because wolves were active throughout 4

the day with peaks at dawn and dusk, while in lynx the most active period covered the whole 5

night including dawn and dusk. Moreover, except one record by Gavrin and Donaurov (1954) 6

of lynx remains in the wolf scats, we did not obtain any recent or other past reports on lynx 7

being killed by wolves or vice versa in BPF. 8

How is the “peaceful coexistence” (Major and Sherburne 1987) possible between the 9

wolf and the lynx in Białowieża Forest? The separation of ecological niches in community of 10

carnivores often lies in different diets (Sunquist et al. 1989, Jędrzejewska and Jędrzejewski 11

1998, Karanth and Sunquist 2000, Ray and Sunquist 2001). However, in sympatric bobcats 12

and coyotes, no interference competition was even found in situations, where their diets 13

overlapped almost completely (Major and Sherburne 1987). In BPF, red deer and roe deer are 14

essential prey species of wolves and lynx. Lynx highly prefers roe deer, taking them more 15

often than expected from their share in the ungulate community (Okarma et al. 1997). Red 16

deer, on the other hand, were taken more often than expected from their abundance by wolves 17

(Jędrzejewski et al. 2000). The diet of both predators, however, overlapped partly due to less 18

frequent predation of lynx on red deer (22% of prey killed) and wolves on roe deer (17% of 19

prey killed). Although wolves appeared to be strongly dependent on red deer in BPF, these 20

canids are also well suited to kill and sustain themselves on the smaller species, e.g. roe deer. 21

In areas where red deer are absent or very rare the roe deer may become a staple food of 22

wolves (Valdmann et al. 2005). Nevertheless, there is probably quite wide range of densities 23

at which the wolves focus their predation on red deer. In the Western Carpathian Mountains 24

wolves were found to prey predominantly on red deer, despite its relatively low (21%) share 25

13

in ungulates community (Nowak et al. 2005). Thus, it seems that possibility for exploitive 1

competition between wolves and lynx is low. It would be, however very likely only in case of 2

a severe alteration of red and roe deer proportions in the living ungulate community (e.g. due 3

to hunting harvest). For example, the coyotes occurring sympatrically with Canada lynx Lynx 4

canadensis, were suggested to influence lynx densities through heavy exploiting the 5

snowshoe hares Lepus americanus (Buskirk et al. 2000). 6

Yet another way of avoiding competition are differences in hunting behaviour between 7

wolves and lynx. As a solitary predator, the lynx is able to conceal itself as well as its prey, 8

making it less detectable for wolves, as suggested by Stander et al. (1997) for African 9

leopards Panthera pardus. Lynx usually drag their kills into thick vegetation and cover it with 10

available material (leaves, snow etc.) (Jędrzejewski et al. 1993). In BPF, wolves were rarely 11

recorded scavenging at kills made by lynx (Jędrzejewski et al. 1993, Selva et al. 2005). Lynx 12

were also never found feeding on kills made by wolves (Selva et al. 2005). 13

According to Matyushkin (1985), direct competition between wolf and lynx is 14

possible, but the relationships between these two species vary depending on the circumstances 15

in which they share resources. Similarly, based on interactions among African wild dogs and 16

larger carnivores, Creel (2001) proposed that the effect of competition on carnivores’ 17

population dynamics may be significantly modified by several factors, including habitat 18

characteristics. Although the Białowieża Forest is characterized by lack of topographic 19

heterogeneity that would ensure escape avenues, it offers heterogeneous habitat, which may 20

facilitate the coexistence of these two large carnivores. The lynx were found to select specific 21

habitat features (fallen logs, uprooted trees and dense thickets) when hunting and resting 22

(Podgórski et al. 2008). In a study on sympatric wolves and cougars, Husseman et al. (2003) 23

showed that the two carnivores used the habitat in different ways when hunting. Ambushing 24

cougars focused on small patches facilitating stalking, whereas coursing wolves pursued their 25

14

prey irrespective of specific habitat type. Wolves, in contrast to lynx, were also occasionally 1

found outside of the forest during this study. Therefore, wolves may probably do well in less 2

diverse habitat, while lynx may need heterogeneous environment to escape competition with 3

stronger predator. 4

In conclusion, differences in morphology, diet, habitat use and hunting behaviour all 5

contribute to allow grey wolf and Eurasian lynx to coexist wherever local habitat and 6

available prey resources fulfil both predators’ requirements. 7

8

Acknowledgements: 9

We thank S. Śnieżko, R. Kozak, I. Ruczyński, M. Chudziński, P. Wasiak, J. Theuerkauf, S. 10

Rouys and R. Gula and numerous students and volunteers, who helped during the field work. 11

Permissions to capture and radio-collar wolves were issued by Ministry of Forestry and 12

Nature Protection and the Director of Białowieża National Park. We are grateful to M. W. 13

Hayward for correcting English and his valuable comments on the manuscript. We also 14

appreciate comments by two anonymous referees that contributed to clarity of the final 15

version of this paper. This study was financed by the budget of Mammal Research Institute 16

PAS and the Polish State Committee for Scientific Research (grants 6P20503405, 17

6P20401905 and 6P04F02612). 18

19

20

References: 21

22

Azarov VI, Shubin NG (2003) Western Siberia. In: Matyushkin EN, Vaisfeld MA (eds) The 23

lynx. Regional features of ecology, use and protection. Nauka, Moscow, pp 249-282 (in 24

Russian with English summary) 25

15

Bibikov DI (ed.) (1985) The Wolf. History, systematics, morphology, ecology. Nauka, 1

Moscow (in Russian) 2

Buskirk SW, Ruggiero LF, Krebs CJ (2000) Habitat fragmentation and interspecific 3

competition: implications for lynx conservation. In: Ruggiero LF, Aubry KB, Buskirk SW, 4

Koehler GM, Krebs CJ, McKelvey KS, Squires JR (eds) Ecology and conservation of lynx 5

in the United States. University Press of Colorado and USDA, Rocky Mountains Research 6

Station, pp 83-100 7

Carbone C, Du Toit JT, Gordon IJ (1997) Feeding success in African wild dogs: does 8

kleptoparasitism by spotted hyenas influence hunting group size? J Anim Ecol 66: 318-326 9

Case TJ, Gilpin ME (1974) Interference competition and niche theory. Proc Natl Acad Sci 10

USA 71: 3073-3077 11

Creel S (2001) Four factors modifying the effect of competition on carnivore population 12

dynamics as illustrated by African wild dogs. Cons Biol 15: 271-274 13

Creel S, Creel NM (1996) Limitation of African wild dogs by competition with larger 14

carnivores. Cons Biol 10: 526-538 15

Danilov PI, Rusakov OS, Tumanov IL, Belkin VV, Makarova OA (2003) The north-west of 16

Russia. In: Matyushkin EN, Vaisfeld MA (eds) The lynx. Regional features of ecology, use 17

and protection. Nauka, Moscow, pp 31-52 (in Russian with English summary) 18

Faliński JB (1986) Vegetation dynamics in temperate lowland primeval forests. Ecological 19

studies in Białowieża Forest. Dr W. Junk Publishers, Dordrecht 20

Gavrin VF, Donaurov SS (1954) The wolf in Białowieża Primeval Forest. Zool Zhurn 33: 21

904-924 [in Russian] 22

Jacobs J (1974) Quantitative measurements of food selection. A modification of the forage 23

ratio and Ivlev’s electivity index. Oecologia 14: 413-417 24

16

Husseman JS, Murray DL, Power G, Mack C, Wenger CR, Quigley H (2003) Assessing 1

differential prey selection patterns between two sympatric large carnivores. Oikos 101: 2

591-601 3

Jędrzejewska B, Jędrzejewski W (1998) Predation in vertebrate communities. The Białowieża 4

Primeval Forest as a case study. Springer, Berlin Heidelberg New York. 5

Jędrzejewska B, Jędrzejewski W (2005) Large carnivores and ungulates in European 6

temperate forest ecosystems: Bottom-up and top-down control. In: Ray JC, Redford KH, 7

Steneck RS, Berger J (eds) Large carnivores and the conservation of biodiversity. Island 8

Press, Washington-Covelo-London, pp 230-246 9

Jędrzejewski W, Schmidt K, Miłkowski L, Jędrzejewska B, Okarma H. (1993) Foraging by 10

lynx and its role in ungulate mortality: the local (Białowieża Forest) and the Palearctic 11

viewpoints. Acta Theriol 38: 385-403 12

Jędrzejewski W, Jędrzejewska B, Okarma H, Schmidt K, Bunevich AN, Miłkowski L (1996) 13

Population dynamics (1869-1994), demography and home ranges of the lynx in Białowieża 14

Primeval Forest (Poland and Belarus). Ecography 19: 122-138 15

Jędrzejewski W, Jędrzejewska B, Okarma H, Schmidt K, Zub K, Musiani M (2000) Prey 16

selection and predation by wolf in Białowieża Primeval Forest (Poland). J Mammal 81: 17

197-212 18

Jędrzejewski W, Schmidt K, Theuerkauf J, Jedrzejewska B, Selva N, Zub K, Szymura L 19

(2002) Kill rates and predation by wolves on ungulate populations in Białowieża Primeval 20

Forest (Poland). Ecology 83: 1341-1356 21

Jędrzejewski W, Schmidt K, Theuerkauf J, Jędrzejewska B, Kowalczyk R (2007) Territory 22

size of wolves Canis lupus: linking local (Białowieża Primeval Forest, Poland) and 23

Holarctic-scale patterns. Ecography 30: 66-76 24

17

Karanth KU, Sunquist ME (2000) Behavioural correlates of predation by tiger (Panthera 1

tigris), leopard (Panthera pardus) and dhole (Cuon alpinus) in Nagarahole, India. J Zool 2

(Lond) 250: 255-265 3

Kenward RE (1992) Quantity versus quality: programmed collection and analysis of radio-4

tracking data. In: Priede IG, Swift SM (eds). Wildlife telemetry: remote monitoring and 5

tracking of animals. Ellis Horwood, Chichester, pp 231-246 6

Kleiman DG, Eisenberg JF (1973) Comparisons of canid and felid social systems from an 7

evolutionary perspective. Anim Behav 21: 637-659 8

Litvaitis JA, Harrison DJ (1989) Bobcat-coyote niche relationships during a period of coyote 9

population increase. Can J Zool 67: 1180-1188 10

Laurenson MK (1995) Implications of high offspring mortality for cheetah population 11

dynamics. In: Sinclair ARE, Arcese P (eds) Serengeti II: dynamics, management, and 12

conservation of an ecosystem. The University of Chicago Press, Chicago and London, pp 13

385 -399 14

Macdonald DW, Ball FG, Hough NG (1980) The evaluation of home range size and 15

configuration using radio-tracking data. In: Amlaner CJ, Macdonald DW (eds) A 16

handbook on biotelemetry and radio tracking. Pergamon Press, Oxford, pp 405-424 17

Major JT, Sherburne JA (1987) Interspecific relationships of coyotes, bobcats and red foxes 18

in western Maine. J Wildl Manage 51: 606-616 19

Malafeev YM, Kryazhimskii FW, Dobrinskii LN (1986) Analiz populacii rysi Srednego 20

Urala. (Analysis of lynx population in the Middle Ural). Akademya Nauk SSSR, 21

Sverdlovsk, (in Russian) 22

Matyushkin EN (1985) Vzaimootnoshenya s drugimi khishchnymi mlekopitayushchimi 23

(Relationships with other predatory mammals). In: Bibikov DI (ed.) The Wolf. History, 24

systematics, morphology, ecology. Moscow, Nauka, pp 355-370 (in Russian) 25

18

Matyushkin EN, Vaisfeld MA (eds) (2003) The lynx. Regional features of ecology, use and 1

protection. Nauka, Moscow (in Russian with English summary) 2

Matyushkin EN, Podolskii SA, Tkachenko KN (2003) The south of Far East. In: Matyushkin 3

EN, Vaisfeld MA (eds). The lynx. Regional features of ecology, use and protection. 4

Nauka, Moscow, pp 423-472 (in Russian with English summary) 5

Miquelle DG, Stephens PA, Smirnov EN, Goodrich JM, Zaumyslova OJ, Myslenkov AE 6

(2005) Tigers and wolves in the Russian Far East: Competitive exclusion, functional 7

redundancy, and conservation implications. In: Pages in Ray JC, Redford KH, Steneck 8

RS, and Berger J, (eds). Large carnivores and the conservation of biodiversity. Island 9

Press, Washington DC, pp 179-207 10

Myrberget S (1970) The Norwegian population of wolverine, Gulo gulo (L.), and lynx, Lynx 11

lynx (L.). Medd Stat Viltunders 2: 1-35 12

Nowak S, Mysłajek RW, Jędrzejewska B (2005) Patterns of wolf Canis lupus predation on 13

wild and domestic ungulates in the Western Carpathian Mountains (S Poland). Acta 14

Theriol 50: 263-276 15

Nowell K, Jackson P (1996) Wild cats. Status survey and conservation action plan. IUCN, 16

Gland. 17

Okarma H, Jędrzejewski W, Schmidt K, Kowalczyk R, Jędrzejewska B (1997) Predation of 18

Eurasian lynx on roe deer and red deer in Białowieża Primeval Forest, Poland. Acta 19

Theriol 42: 203-224 20

Okarma H, Jędrzejewski W, Schmidt K, Śnieżko S, Bunevich AN, Jędrzejewska B (1998) 21

Home ranges of wolves in Białowieża Primeval Forest, Poland, compared with other 22

Eurasian populations. J Mammal 79: 842-852 23

Okarma H, Jędrzejewski W (1997) Livetrapping wolves with nets. Wildl Soc Bull 25: 78-82 24

19

Palomares F, Caro TM (1999) Interspecific killing among mammalian carnivores. Am Nat: 1

153: 492-508 2

Podgórski T, Schmidt K, Kowalczyk R, Gulczyńska A (2008) Microhabitat selection by 3

Eurasian lynx and its implications for species conservation. Acta Theriol 53: 97-110. 4

Pulliainen E (1965) Studies on the wolf (Canis lupus L.) in Finland. Ann Zool Fenn 2: 215-5

259 6

Ray JC, Sunquist ME (2001) Trophic relations in a community of African rainforest 7

carnivores. Oecologia 127: 395-408 8

Rosenzweig ML (1966) Community structure in sympatric Carnivora. J Mammal 47: 602-612 9

Schmidt K, Jędrzejewski W, Okarma H (1997) Spatial organization and social relations in the 10

Eurasian lynx population in Białowieża Primeval Forest, Poland. Acta Theriol 42: 289-312 11

Schmidt K (1999) Variation in daily activity of the free living Eurasian lynx in Białowieża 12

Primeval Forest, Poland. J Zool (Lond.) 249: 417-425 13

Scognamillo D, Maxit IE, Sunquist ME, Polisar J (2003) Coexistence of jaguar (Panthera 14

onca) and puma (Puma concolor) in a mosaic landscape in the Venezuelan llanos. J Zool 15

(Lond.) 259: 269-279 16

Selva N, Jędrzejewska B, Jędrzejewski W, Wajrak A (2005) Factors affecting carcass use by a 17

guild of scavengers in European temperate woodland. Can J Zool 83: 1590-1601 18

Stander PE, Haden PJ, Kaqece //, Ghau // (1997) The ecology of asociality in Namibian 19

leopards. J Zool ( Lond) 242: 343-364 20

Sunquist ME, Sunquist F, Daneke DE (1989) Ecological separation in a venezuelan llanos 21

carnivore community. In: Redford KH, Eisenberg JF (eds). Advances in neotropical 22

mammalogy. Sandhill Crane Press, Gainesville, Florida, 197-232 23

20

Theuerkauf J, Jędrzejewski W, Schmidt K, Okarma H, Ruczyński I, Śnieżko S, Gula R (2003) 1

Daily patterns and duration of wolf activity in the Białowieża Forest (Poland). J Mammal 2

84: 243-253 3

Valdmann H, Andersone-Lilley Z, Koppa O, Ozolins J, Bagrade G (2005) Winter diets of 4

wolf Canis lupus and lynx Lynx lynx in Estonia and Latvia. Acta Theriol 50: 521-527 5

Wesołowski T (2005) Virtual conservation: how the European Union is turning a blind eye to 6

its vanishing primeval forests. Cons Biol 19: 1349–1358 7

Zheltukhin AS (1986) Biocenotic relationships of the European lynx (Lynx lynx) in the 8

southern taiga of the upper Volga. Zool Zhurn 65: 259-271 (in Russian with English 9

summary). 10

21

Table 1. Spatial overlap among home ranges of 7 lynx and territories of 3 wolf packs radio-1

tracked simultaneously in the Białowieża Primeval Forest in 1994-1996. The overlaps were 2

calculated for combinations of particular wolf pack territories with all lynx home ranges and 3

for particular lynx home ranges with all wolf packs monitored concurrently. See Appendix for 4

number of locations of individual lynx and wolves. 5

Percentage overlap (percent of territory/home range

of the first species overlapped by all radio-tracked

individuals of the second species)

Pair of species,

territory/home range estimate

N cases Mean ± SE (min – max)

Wolf pack – lynx, MCP 100% 3 76 ± 7 (68 – 89)

Wolf pack – lynx, Kernel 50% 3 12 ± 12 (0 – 36)

Lynx – wolf packs, MCP 100% 7 50 ± 12 (13 – 99)

Lynx – wolf packs, Kernel 50% 7 14 ± 10 (0 – 70)

6

7

8

9

10

11

12

13

14

15

16

17

22

1

Table 2. Static (home range percentage overlap) and dynamic (Jacobs’ indices and average 2

distances between individuals) interactions between individual dyads of wolves and lynx 3

radio-tracked in the Białowieża Primeval Forest in 1994-1996, based on simultaneous 4

locations (on average within 1 h) only. n – number of simultaneous locations. 5

Percentage overlap Distance between individuals (km) Dyads

(n) of wolf

territory

of lynx

home

range

Mean ± SE (min-max)

Jacobs’ index

(D)

W1 – L1

(159)

3 5 6.9±0.2 2.3-18.4 0.03

W1 – L2

(424)

40 40 5.5±0.1 0.3-16.0 0.03

W2 – L3

(111)

46 29 11.0±0.5 0.4-23.8 0.02

W3 – L3

(225)

31 19 7.1±0.4 0.3-27.8 0.04

W3 – L4

(237)

63 52 4.4±0.3 0-18.3 0.36

6

23

Figure captions: 1

2

Figure 1. Distribution of all radio-locations of lynx (n=7521 of 14 radio-tracked individuals in 3

1991-1996) and wolves (n=28857 of 8 individuals belonging to 4 packs radio-tracked in 1994 4

-1999) in the Białowieża Primeval Forest and most frequently used areas (75% fixed kernel) 5

estimated for all locations pooled for each species. Question marks denote area where lynx 6

were not trapped for radio-collaring. 7

8

Figure 2. Territories (100% MCP) and core areas (50% fixed kernel) of wolf packs (n=3) and 9

home ranges of lynx (n=7) radio-tracked in the Białowieża Primeval Forest in 1994-1996. 10

11

Figure 3. Examples of movement routes of a pack of wolves (with the female W3) and male 12

lynx (L4) recorded during simultaneous, continuous radio-tracking of both carnivores in BPF. 13

24

Figure 1.

25

Figure 2.

26

Figure 3.

27

Appendix. Grey wolves and Eurasian lynx radio-tracked in Białowieża Primeval Forest in

1994-1996. * this period includes a 4-month gap in tracking (22 Sep 1995 – 2 Feb 1996)

due to collar failure. Numbers following names of wolves are numbers of individuals in the

packs.

Animal

name

ID

code

Sex Age Time of simultaneous radio-

tracking

Number of

locations

Wolves:

Szara (6-7) W1 F Subadult 31 Mar 1994 – 31 Jan 1995 1650

Bura (5-8) W2 F Adult 22 Mar 1995 – 1 Oct 1996* 1229

Ruda (5-7) W3 F Adult 26 Mar 1995 – 22 Oct 1996 1547

Lynx:

Tamara L1 F Subadult 30 Mar 1994 – 10 Aug 1994 614

Sonia L2 F Adult 31 Mar 1994 – 31 Jan 1995 902

Trofim L3 M Adult 22 Mar 1995 – 14 Oct 1996 1200

Iwan

Zoja

Olga

Diana

L4

L5

L6

L7

M

F

F

F

Adult

Subadult

Adult

Adult

29 Mar 1995 – 1 Apr 1996

5 Apr 1994 – 24 Aug 1994

22 Mar 1994 – 7 Sep 1994

27 Mar 1995 – 3 Aug 1995

923

8

99

72