Download Spatial interactions between grey wolves and Eurasian lynx in

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Spatial interactions between grey wolves and Eurasian lynx in
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Białowieża Primeval Forest, Poland
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Krzysztof Schmidt*, Włodzimierz Jędrzejewski, Henryk Okarma1 and Rafał Kowalczyk
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Mammal Research Institute, Polish Academy of Sciences, PL-17-230 Białowieża, Poland
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* corresponding author: phone: +48 85 6827777, fax: +48 85 68 12 289; e-mail:
[email protected]
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33 str., PL-31-120 Kraków, Poland
present address: Institute of Nature Conservation, Polish Academy of Sciences, Mickiewicza
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Abstract
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Various species of large predators are reported to influence each other through interference or
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exploitation competition that may affect demography and survival of the subordinate species.
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We analyzed spatial relationships between grey wolf Canis lupus and Eurasian lynx Lynx lynx
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in Białowieża Primeval Forest (BPF, eastern Poland) to determine how they partitioned the
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space. The wolves (n=8) and lynx (n=14) were radio-tracked in 1991-1999. Three wolves and
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seven lynx were radio-tracked simultaneously in 1994-1996. Territories of wolf packs and
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home ranges of lynx overlapped considerably (76% of wolf territories and 50% of lynx home
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ranges, on average). In three cases, their core areas were also overlapping. Wolf – lynx dyads
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with overlapping home ranges were simultaneously located at distances from 0 to 28 km from
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each other. We found neither avoidance nor attraction between wolves and lynx occupying
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the same areas. We concluded that in BPF, the two large predators coexist due to
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specialisation on different preferred prey and heterogeneous habitat.
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Key words: Canis lupus, coexistence, exploitation competition, interference competition,
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Lynx lynx, spatial overlap
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Introduction
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What enables various carnivorous mammals to coexist and how they interact with each other
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when occurring sympatrically has been a subject of numerous studies. Rosenzweig (1966)
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suggested that coexistence results from size differences between predator species. Felids and
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canids seem to be particularly predisposed to “peaceful coexistence” (Major and Sherburne
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1987) through separation of their ecological niches, which likely resulted from evolution of
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their different social systems (Kleiman and Eisenberg 1973).
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On the other hand, the competitive exclusion was recently suggested between tigers
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Panthera tigris and wolves Canis lupus even despite of extreme size differences (Miquelle et
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al. 2005). Cases of interspecific killing are common among predators, including felids and
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canids (review in Palomares and Caro 1999). In some instances intraguild predation may have
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serious consequences on demography and survival of the inferior species, as it has been
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suggested for cheetahs Acinonyx jubatus (Laurenson 1995) and African wild dogs Lycaon
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pictus (Creel and Creel 1996, Carbone et al. 1997).
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Grey wolves and Eurasian lynx Lynx lynx are sympatric across most of their vast
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geographical ranges, which stretch from Eastern Europe to the Far East of Asia (Bibikov
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1985, Nowell and Jackson 1996). The spatial interactions between these species, however,
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have not yet been studied. Anecdotal information on their interspecific relationships (based on
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harvest data and incidental observations) are available from the Russian part of their ranges
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and from Fennoscandia. According to those reports, coexistence of the two large carnivores is
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often characterized by negative influences of wolf on lynx (Pulliainen 1965, Myrberget 1970,
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Matyushkin 1985, Malafeev et al. 1986, Matyushkin and Vaisfeld 2003). Dynamics of wolf
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and lynx population numbers reported there suggest that lynx may reach higher numbers only
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when wolves are rare. Cases of wolves killing lynx were also reported (see Matyushkin and
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Vaisfeld 2003 for review). On the other hand, it has also been suggested that the relationships
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between the two species may vary with ecological circumstances (Matyushkin 1985). Indeed,
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in some areas populations of both predators existed in large numbers without apparent
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reciprocal influence (e.g. Upper Volga Region: Zheltukhin 1986) or even increased
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simultaneously (West Siberia: Azarov and Shubin 2003; Eastern Poland and Western Belarus:
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Jędrzejewska and Jędrzejewski 1998). Snow-tracking data from several study areas in Eurasia
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(e.g. Upper Volga Region: Zheltukhin 1986; North-West Russia: Danilov et al. 2003; Far East
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of Asia: Matyushkin et al. 2003) showed that lynx and wolves do not avoid using the same
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sites and do not exhibit any particular interest in each other, which suggests a lack of
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interference competition (sensu Case and Gilpin 1974).
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Wolves and lynx also have similar diets across their common range (see Matyushkin
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and Vaisfeld 2003 for review). Therefore, one can expect that the wolves may limit lynx
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populations through exploitation of the common food resources (exploitation competition), as
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suggested by Litvaitis and Harrison (1989) for sympatric bobcat Lynx rufus and coyote Canis
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latrans in Maine (USA).
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In Białowieża Primeval Forest (BPF, E Poland) the two predators coexist and no
negative relationship between their long-term dynamics was noted – the wolf and lynx
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densities were found to be significantly correlated in time (Jędrzejewska and Jędrzejewski
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2005). Their numbers were simultaneously shaped by temporarily varying hunting harvest.
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The ecology of both species in BPF has been studied in detail (Jędrzejewski et al. 1996,
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Okarma et al. 1997, Schmidt et. al. 1997, Okarma et al. 1998, Schmidt 1999, Jędrzejewski et
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al. 2000, Theuerkauf et al. 2003), but their interspecific interactions have not been analyzed.
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The feeding niches of the two carnivores partly overlap, as wolves primarily prey on red deer
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Cervus elaphus and supplement their diet with wild boar Sus scrofa and roe deer Capreolus
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capreolus (Jędrzejewski et al. 2000), whereas lynx feed mainly on roe deer with the addition
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of female and young red deer (Okarma et al. 1997). Given the high densities of ungulates in
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BPF (Jędrzejewski et al. 2002), exploitation competition is not very likely.
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The aim of this study was to analyze fine scale spatial interactions between radio-
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tracked wolves and lynx inhabiting the Białowieża Primeval Forest. We investigated these
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relationships at three spatial scales: (1) the study area level - utilization of the forest by both
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species, (2) home range level – degree of overlap between lynx and wolf home ranges and (3)
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individual level – dynamic interactions between dyads of lynx and wolf individuals located at
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the same time. This way we intended to determine if wolves and lynx tended to be spatially
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segregated and show any signs of exclusion of one by the other (interference competition) in
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the BPF.
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Material and methods
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Study area
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The study was conducted in Polish part of the Białowieża Primeval Forest (BPF), Eastern
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Poland (52º30’-53º N, 23º30’-24º15’ E). The BPF is a temperate mixed lowland forest located
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on the Polish-Belarussian border and characterized by a high percentage of natural stands
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(Faliński 1986). The whole forest covers nearly 1500 km2 and its Polish part about 600 km2.
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Most of the Polish part of the BPF (500 km2) is managed by State Forestry, while the rest is
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protected as the Białowieża National Park (BNP, 100 km2) with a 50-km2 area of strict
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reserve, where no human interference is allowed except for tourism and research. A network
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of small reserves with partial protection is located within the managed part of the BPF
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(Wesołowski 2005). The dominating tree species are: pine Pinus silvestris, spruce Picea
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abies), oak Quercus robur, hornbeam Carpinus betulus, black alder Alnus glutinosa, ash
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Fraxinus excelsior, lime Tilia cordata, birches Betula verrucosa and B. pubescens, aspen
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Populus tremula and maple Acer platanoides. Despite of timber exploitation, the BPF
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maintains a unique character in comparison to other European forest in terms of: (1) tree
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diversity (with 26 tree and 55 shrub species forming a mosaic of diverse tree communities),
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(2) a multi-storey profile of stands, (3) relatively large amount of dead wood and (4)
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outstanding diversity of flora and fauna (Wesołowski 2005). The climate of BPF is temperate
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with transitional character between Atlantic and continental ones with clearly marked warm
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and cold periods (average temperatures during the study period were -3.9º C in January and
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19.1º C in July; average annual precipitation was 622 mm; snow cover persisted for an
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average of 96 days per year from November to March). Wolf and lynx are currently the only
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large predators in BPF. The community of ungulate mammals consists of: red deer Cervus
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elaphus, roe deer Capreolus capreolus, wild boar Sus scropha, moose Alces alces, and
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European bison Bison bonasus. However, only three species – red deer, roe deer and wild
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boar – play an important role in both predators’ diets (Jędrzejewski et al. 1993, Jędrzejewski
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et al. 2000).
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Radio-tracking
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The study was a part of a larger research project on wolf and lynx ecology in BPF conducted
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in 1991-1999 (Schmidt et al. 1997, Jędrzejewski et al. 2002). The principal data for this paper
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originates from 1994-1996, when both wolf and lynx were studied by radio-tracking: 3 female
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wolves belonging to 2 packs and 7 lynx (2 M, 5 F) (Appendix I). Five lynx provided
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sufficient number of locations taken simultaneously with those of wolves for calculating
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dynamic interactions between the species. Locations of wolves and lynx taken before and
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after 1994-1996 were, however, used for general assessment of the forest use by both species
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(9 wolves and 16 lynx). We acknowledge that the sample of studied animals is low and we
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didn’t monitor whole the populations of both predators. However, the information obtained
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during this study provided minimum estimates of their spatial overlap and existing potential
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for interactions between them. Moreover, both wolves and lynx use large home ranges (173-
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294 km2 in wolves: Okarma et al. 1998 and 133-248 km2 in lynx: Schmidt et al. 1997), so that
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the total area used by simultaneously radio-tracked wolves and lynx covered a representative
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portion of the study area. We estimated that during the study, the population of wolves in
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Polish part of BPF consisted of 3-4 packs of wolves containing 2-8 individuals each
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(Jędrzejewski et al. 2000). Lynx were estimated at 6 – 18 adult individuals (Jędrzejewski et.
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al. 1996).
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The wolves were captured both with “fladry” (a rope with strips of cloth that is used
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to limit and drive wolves’ movements) and a net system (Okarma and Jędrzejewski 1997) or
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foot snare traps (equipped with alarm-system) and fitted with VHF radio-collars made by
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Telonics Inc., (Mesa, Arizona, USA), AVM Instrument Co. (Livermore, California, USA),
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Telemetry Systems (Mequon, Wisconsin, USA), and Advanced Telemetry Systems (Isanti,
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Minnesota, USA). Lynx were captured with snare traps set at fresh kill and marking sites or in
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a box trap (equipped with alarm-system) and fitted with VHF radio-collars made by Wagener
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Temetrieanlagen HF-NF Technik, (Köln, Germany) or AVM Instrument Co. The range of the
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collars was 1 – 2 km. Both wolves and lynx were immobilized with a mixture of ketamine
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hydrochloride (wolves: 3.4-4.5 mg/kg, lynx 5 mg/kg of body weight) and xylazine
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hydrochloride (wolves: 5-6.5 mg/kg, lynx 6 mg/kg of body weight).
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We located radio-collared animals 2-5 times per week by triangulation from the forest
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roads. Additionally, each month we conducted sessions of continuous 24-h radio-tracking
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lasting 2-7 days, during which we located animals every 30 min. During these sessions we
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focused on one individual of a given species, although we recorded locations of the second
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species if it was present nearby until the animals separated such that following two of them
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simultaneously was too difficult. Locations were plotted on forest maps with 533x533 m
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square grid (according to the forest division compartments marked in the terrain). Positions of
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the animals were determined in the center of a square, in the middle of one of its sides or in
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the corner between four adjacent squares. Their estimated locations could then differ from the
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actual ones by a maximum of 373 m. We aimed at reducing the error by taking bearings from
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points on the forest roads closest to animals (≥100 m). During continuous radio-tracking,
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however, the observer usually stayed 500-1000 m from the focal animal to avoid disturbing it.
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For a general assessment of space use by both species and their spatial relationships in
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the study area we overlaid all locations of wolves (8 individuals with 28857 locations) and
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lynx (14 individuals with 7521 locations) recorded between 1991-1999 on the forest map
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using program Biotas™ v.1.03.1 Alpha (Ecological Software Solutions, Urnäsch,
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Switzerland). To estimate an extent of overlap of both species’ occurrence in the entire study
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area we calculated 95% minimum convex polygon ranges for pooled locations of all
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individuals in the two species. Using 95% MCP, in this case, allowed us to maximise the
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studied area used by both species as well as for excluding large portions of non-forested areas
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that were not visited by them.We also calculated 75% fixed kernel ranges for the same data to
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find out if there is an overlap of most frequently used areas by all radio-tracked lynx and
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wolves. The fixed kernel ranges where estimated with a level of smoothing selected by least-
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squares cross-validation method and the chosen bandwidth was 750 m.
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We also examined static and dynamic interactions (Macdonald et al. 1980) between
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wolves and lynx. Static interactions were determined by calculating overlaps of lynx home
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ranges with territories of wolves. We calculated an average overlap both between an
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individual lynx home range and all available pooled wolf territories, between an individual
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wolf pack territory and all available pooled lynx home ranges as well as between dyads of
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lynx and wolves. We included all lynx radio-tracked simultaneously with wolves for the
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pooled lynx home ranges, despite the low number of locations for some individuals (see
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Appendix). The discrepancy in sample size, however, should not significantly affect the
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results, because our aim was to estimate a minimum value of the overlap between wolf
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territory and all detected home ranges of lynx inhabiting a common area. The home ranges
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were estimated with 100% minimum convex polygon method using the program Biotas™
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v.1.03.1 Alpha, for all locations recorded during a simultaneous period of radio-tracking and
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for a sub-sample of both species’ locations taken on the same day only (on average within 1 h,
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max 12 h). This way we accounted for potential bias, resulting from a higher probability of
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finding two animals with decreasing distance between them. The 100% MCP was used to
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ensure detecting maximum potential overlapping area used by wolf and lynx dyads. To
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determine core areas used by particular lynx and wolf individuals we calculated 50% fixed
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kernel ranges (with the same smoothing parameters as for general – species’ level
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assessment).
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To determine if two predators ignore, avoid or attract each other, we examined their
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dynamic interactions (Macdonald et al. 1980, Kenward 1992) using the program Ranges 6 v.
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1.08 (R. Kenward, A. South and S. Walls, Anatrack Ltd. Dorset, UK). The program compares
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observed and expected distances between locations of two animals. The expected distances
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were estimated as a mean distance between each location of one animal and all locations of
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another animal. The observed and expected distances were then compared with Jacobs’ index
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(D) (Jacobs 1974):
D=
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d E − dO
d E + dO
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where dE is expected distance and dO is observed distance between pairs of simultaneous
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locations of a wolf-lynx dyad with overlapping home ranges. D ranges from -1 in case of
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mutual avoidance, to 0 if the animals move independently, to +1 if they are attracted to each
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other.
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Results
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Wolves and lynx utilized the study area in nearly the same extent (Fig. 1). Although the eight
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wolves of four packs were more intensively radio-tracked than the 14 lynx, the minimum
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convex polygons with 95% of pooled locations of all individuals in the two species
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overlapped in 88%. Most frequently used areas by both predators (75% fixed kernel) also
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showed a substantial spatial overlap: wolves on 18% of lynx core area, and lynx on 32% of
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wolves’ core area. It is striking that both wolves and lynx were utilizing mostly the main solid
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forest body, leaving it only occasionally. However, wolves seemed to wander out of the forest
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slightly more readily than lynx (note westernmost locations of wolves on Fig. 1).
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Home ranges of lynx individuals and territories of wolf packs overlapped each other
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extensively (Fig. 2). Territory of each of the studied wolf packs was on average, covered by
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home ranges of radio-collared lynx by 76% (Table 1). Home range of each lynx was covered
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by wolf territories by 50%, on average. Smaller overlap (mean 12-14%) was recorded
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between core areas (Table 1). These measures of overlap are certainly underestimated, since
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in both years shown in Fig. 2, there were other (non-collared) wolf packs and lynx individuals
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recorded by snow-tracking in the nearest vicinities of the territories/home ranges of radio-
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collared animals (comp. Jędrzejewski et al. 1996, Okarma et al. 1998).
When found at approximately the same time, particular dyads of wolves and lynx with
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overlapping home ranges were located at an average distance from 4.4 to 11.0 km apart
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(Table 2). It is noteworthy that occasionally they were found at the same location as the
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minimum recorded distance was 0 or 0.3 km in some dyads. According to analysis of
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dynamic interactions (based on Jacobs’ index, D) conducted for pairs of simultaneous
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locations of wolves and lynx, the animals showed neither avoidance nor attraction to each
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other (Table 2). A slight attraction was recorded in the case of a dyad L4-W3, which also
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showed the most extensive overlap of their home ranges.
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A male lynx L4 and a pack of wolves with female W3 were radio-tracked
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simultaneously on three separate occasions, in which the animals were found at very short
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(less than 1 km) distances from each other for longer periods (Fig. 3). On 23 May 1995, the
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lynx approached the site where the wolf pack stayed and remained there in close proximity to
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wolves (0.25 – 0.8 km) for 5 h. The lynx left that area in the evening and headed in the
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opposite direction to the one he arrived. On 22 June 1995, both lynx and wolf were at
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approximately the same location (maximum 0.25 km from each other) for 1 h. Afterwards,
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they split, however, they still remained close by (0.25 – 1.4 km) in an area of about 0.8 km2
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for the subsequent 8 h, before the lynx departed. On another occasion, 23/24 October 1995,
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the female wolf with her pack and the lynx moved along nearly the same route over 13 km for
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22 h. During this time, the animals rested four times, each time in different place, at a distance
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of 0.5 – 1.5 km from each other. After the resting bouts that took from 0.5 to 2.5 h, they
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moved alternately following each other, heading approximately in the same direction until
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early morning, when they went separate ways (Fig. 3).
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Discussion
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The competition between two species of predators may be mitigated by various factors, such
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as differences in size (Rosenzweig 1966) and social organization (Kleiman and Eisenberg
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1973). Therefore, it is difficult to predict the effect of interactions between given species at
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given site. Grey wolf inhabiting BPF is nearly twice as heavy as Eurasian lynx (on average,
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45 and 35 kg, for adult male and female wolves, respectively: Jędrzejewski et al. 2007; 22
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and 17 kg for male and female lynx, respectively: Schmidt et al. 1997). Size difference may
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lead to interference competition through interspecific killing, as suggested by Palomares and
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Caro (1999). On the other hand it may diminish exploitive competition through partitioning of
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prey resources (Scognamillo et al. 2003). Different life habits of canids and felids – sociality
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and coursing hunting behaviour of wolves versus solitary life and stalking hunting common in
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felids – may farther ease resource partitioning, as found by Husseman et al. (2003) for
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sympatric wolves and cougars Puma concolor.
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Our study showed that the wolves and the lynx did not exclude each other spatially in
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the Białowieża Primeval Forest. They behaved neutrally, neither avoiding nor attracting each
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other, which suggests that they used the available area completely independently. Cases of
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extensively overlapping core areas and parallel travelling recorded during this study are also
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indicating that individuals of the two species do not particularly tend to separate from each
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other while using the same area. The patterns of their activity were also greatly overlapping
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(compare Schmidt 1999 and Theuerkauf et al. 2003), because wolves were active throughout
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the day with peaks at dawn and dusk, while in lynx the most active period covered the whole
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night including dawn and dusk. Moreover, except one record by Gavrin and Donaurov (1954)
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of lynx remains in the wolf scats, we did not obtain any recent or other past reports on lynx
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being killed by wolves or vice versa in BPF.
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How is the “peaceful coexistence” (Major and Sherburne 1987) possible between the
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wolf and the lynx in Białowieża Forest? The separation of ecological niches in community of
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carnivores often lies in different diets (Sunquist et al. 1989, Jędrzejewska and Jędrzejewski
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1998, Karanth and Sunquist 2000, Ray and Sunquist 2001). However, in sympatric bobcats
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and coyotes, no interference competition was even found in situations, where their diets
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overlapped almost completely (Major and Sherburne 1987). In BPF, red deer and roe deer are
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essential prey species of wolves and lynx. Lynx highly prefers roe deer, taking them more
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often than expected from their share in the ungulate community (Okarma et al. 1997). Red
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deer, on the other hand, were taken more often than expected from their abundance by wolves
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(Jędrzejewski et al. 2000). The diet of both predators, however, overlapped partly due to less
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frequent predation of lynx on red deer (22% of prey killed) and wolves on roe deer (17% of
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prey killed). Although wolves appeared to be strongly dependent on red deer in BPF, these
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canids are also well suited to kill and sustain themselves on the smaller species, e.g. roe deer.
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In areas where red deer are absent or very rare the roe deer may become a staple food of
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wolves (Valdmann et al. 2005). Nevertheless, there is probably quite wide range of densities
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at which the wolves focus their predation on red deer. In the Western Carpathian Mountains
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wolves were found to prey predominantly on red deer, despite its relatively low (21%) share
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in ungulates community (Nowak et al. 2005). Thus, it seems that possibility for exploitive
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competition between wolves and lynx is low. It would be, however very likely only in case of
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a severe alteration of red and roe deer proportions in the living ungulate community (e.g. due
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to hunting harvest). For example, the coyotes occurring sympatrically with Canada lynx Lynx
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canadensis, were suggested to influence lynx densities through heavy exploiting the
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snowshoe hares Lepus americanus (Buskirk et al. 2000).
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Yet another way of avoiding competition are differences in hunting behaviour between
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wolves and lynx. As a solitary predator, the lynx is able to conceal itself as well as its prey,
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making it less detectable for wolves, as suggested by Stander et al. (1997) for African
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leopards Panthera pardus. Lynx usually drag their kills into thick vegetation and cover it with
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available material (leaves, snow etc.) (Jędrzejewski et al. 1993). In BPF, wolves were rarely
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recorded scavenging at kills made by lynx (Jędrzejewski et al. 1993, Selva et al. 2005). Lynx
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were also never found feeding on kills made by wolves (Selva et al. 2005).
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According to Matyushkin (1985), direct competition between wolf and lynx is
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possible, but the relationships between these two species vary depending on the circumstances
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in which they share resources. Similarly, based on interactions among African wild dogs and
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larger carnivores, Creel (2001) proposed that the effect of competition on carnivores’
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population dynamics may be significantly modified by several factors, including habitat
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characteristics. Although the Białowieża Forest is characterized by lack of topographic
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heterogeneity that would ensure escape avenues, it offers heterogeneous habitat, which may
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facilitate the coexistence of these two large carnivores. The lynx were found to select specific
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habitat features (fallen logs, uprooted trees and dense thickets) when hunting and resting
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(Podgórski et al. 2008). In a study on sympatric wolves and cougars, Husseman et al. (2003)
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showed that the two carnivores used the habitat in different ways when hunting. Ambushing
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cougars focused on small patches facilitating stalking, whereas coursing wolves pursued their
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prey irrespective of specific habitat type. Wolves, in contrast to lynx, were also occasionally
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found outside of the forest during this study. Therefore, wolves may probably do well in less
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diverse habitat, while lynx may need heterogeneous environment to escape competition with
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stronger predator.
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In conclusion, differences in morphology, diet, habitat use and hunting behaviour all
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contribute to allow grey wolf and Eurasian lynx to coexist wherever local habitat and
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available prey resources fulfil both predators’ requirements.
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Acknowledgements:
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We thank S. Śnieżko, R. Kozak, I. Ruczyński, M. Chudziński, P. Wasiak, J. Theuerkauf, S.
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Rouys and R. Gula and numerous students and volunteers, who helped during the field work.
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Permissions to capture and radio-collar wolves were issued by Ministry of Forestry and
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Nature Protection and the Director of Białowieża National Park. We are grateful to M. W.
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Hayward for correcting English and his valuable comments on the manuscript. We also
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appreciate comments by two anonymous referees that contributed to clarity of the final
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version of this paper. This study was financed by the budget of Mammal Research Institute
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PAS and the Polish State Committee for Scientific Research (grants 6P20503405,
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6P20401905 and 6P04F02612).
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References:
22
23
Azarov VI, Shubin NG (2003) Western Siberia. In: Matyushkin EN, Vaisfeld MA (eds) The
24
lynx. Regional features of ecology, use and protection. Nauka, Moscow, pp 249-282 (in
25
Russian with English summary)
14
1
2
3
Bibikov DI (ed.) (1985) The Wolf. History, systematics, morphology, ecology. Nauka,
Moscow (in Russian)
Buskirk SW, Ruggiero LF, Krebs CJ (2000) Habitat fragmentation and interspecific
4
competition: implications for lynx conservation. In: Ruggiero LF, Aubry KB, Buskirk SW,
5
Koehler GM, Krebs CJ, McKelvey KS, Squires JR (eds) Ecology and conservation of lynx
6
in the United States. University Press of Colorado and USDA, Rocky Mountains Research
7
Station, pp 83-100
8
9
10
11
12
13
14
15
16
Carbone C, Du Toit JT, Gordon IJ (1997) Feeding success in African wild dogs: does
kleptoparasitism by spotted hyenas influence hunting group size? J Anim Ecol 66: 318-326
Case TJ, Gilpin ME (1974) Interference competition and niche theory. Proc Natl Acad Sci
USA 71: 3073-3077
Creel S (2001) Four factors modifying the effect of competition on carnivore population
dynamics as illustrated by African wild dogs. Cons Biol 15: 271-274
Creel S, Creel NM (1996) Limitation of African wild dogs by competition with larger
carnivores. Cons Biol 10: 526-538
Danilov PI, Rusakov OS, Tumanov IL, Belkin VV, Makarova OA (2003) The north-west of
17
Russia. In: Matyushkin EN, Vaisfeld MA (eds) The lynx. Regional features of ecology, use
18
and protection. Nauka, Moscow, pp 31-52 (in Russian with English summary)
19
20
21
22
23
24
Faliński JB (1986) Vegetation dynamics in temperate lowland primeval forests. Ecological
studies in Białowieża Forest. Dr W. Junk Publishers, Dordrecht
Gavrin VF, Donaurov SS (1954) The wolf in Białowieża Primeval Forest. Zool Zhurn 33:
904-924 [in Russian]
Jacobs J (1974) Quantitative measurements of food selection. A modification of the forage
ratio and Ivlev’s electivity index. Oecologia 14: 413-417
15
1
Husseman JS, Murray DL, Power G, Mack C, Wenger CR, Quigley H (2003) Assessing
2
differential prey selection patterns between two sympatric large carnivores. Oikos 101:
3
591-601
4
5
6
Jędrzejewska B, Jędrzejewski W (1998) Predation in vertebrate communities. The Białowieża
Primeval Forest as a case study. Springer, Berlin Heidelberg New York.
Jędrzejewska B, Jędrzejewski W (2005) Large carnivores and ungulates in European
7
temperate forest ecosystems: Bottom-up and top-down control. In: Ray JC, Redford KH,
8
Steneck RS, Berger J (eds) Large carnivores and the conservation of biodiversity. Island
9
Press, Washington-Covelo-London, pp 230-246
10
Jędrzejewski W, Schmidt K, Miłkowski L, Jędrzejewska B, Okarma H. (1993) Foraging by
11
lynx and its role in ungulate mortality: the local (Białowieża Forest) and the Palearctic
12
viewpoints. Acta Theriol 38: 385-403
13
Jędrzejewski W, Jędrzejewska B, Okarma H, Schmidt K, Bunevich AN, Miłkowski L (1996)
14
Population dynamics (1869-1994), demography and home ranges of the lynx in Białowieża
15
Primeval Forest (Poland and Belarus). Ecography 19: 122-138
16
Jędrzejewski W, Jędrzejewska B, Okarma H, Schmidt K, Zub K, Musiani M (2000) Prey
17
selection and predation by wolf in Białowieża Primeval Forest (Poland). J Mammal 81:
18
197-212
19
Jędrzejewski W, Schmidt K, Theuerkauf J, Jedrzejewska B, Selva N, Zub K, Szymura L
20
(2002) Kill rates and predation by wolves on ungulate populations in Białowieża Primeval
21
Forest (Poland). Ecology 83: 1341-1356
22
Jędrzejewski W, Schmidt K, Theuerkauf J, Jędrzejewska B, Kowalczyk R (2007) Territory
23
size of wolves Canis lupus: linking local (Białowieża Primeval Forest, Poland) and
24
Holarctic-scale patterns. Ecography 30: 66-76
16
1
Karanth KU, Sunquist ME (2000) Behavioural correlates of predation by tiger (Panthera
2
tigris), leopard (Panthera pardus) and dhole (Cuon alpinus) in Nagarahole, India. J Zool
3
(Lond) 250: 255-265
4
Kenward RE (1992) Quantity versus quality: programmed collection and analysis of radio-
5
tracking data. In: Priede IG, Swift SM (eds). Wildlife telemetry: remote monitoring and
6
tracking of animals. Ellis Horwood, Chichester, pp 231-246
7
8
9
10
11
Kleiman DG, Eisenberg JF (1973) Comparisons of canid and felid social systems from an
evolutionary perspective. Anim Behav 21: 637-659
Litvaitis JA, Harrison DJ (1989) Bobcat-coyote niche relationships during a period of coyote
population increase. Can J Zool 67: 1180-1188
Laurenson MK (1995) Implications of high offspring mortality for cheetah population
12
dynamics. In: Sinclair ARE, Arcese P (eds) Serengeti II: dynamics, management, and
13
conservation of an ecosystem. The University of Chicago Press, Chicago and London, pp
14
385 -399
15
Macdonald DW, Ball FG, Hough NG (1980) The evaluation of home range size and
16
configuration using radio-tracking data. In: Amlaner CJ, Macdonald DW (eds) A
17
handbook on biotelemetry and radio tracking. Pergamon Press, Oxford, pp 405-424
18
19
20
Major JT, Sherburne JA (1987) Interspecific relationships of coyotes, bobcats and red foxes
in western Maine. J Wildl Manage 51: 606-616
Malafeev YM, Kryazhimskii FW, Dobrinskii LN (1986) Analiz populacii rysi Srednego
21
Urala. (Analysis of lynx population in the Middle Ural). Akademya Nauk SSSR,
22
Sverdlovsk, (in Russian)
23
Matyushkin EN (1985) Vzaimootnoshenya s drugimi khishchnymi mlekopitayushchimi
24
(Relationships with other predatory mammals). In: Bibikov DI (ed.) The Wolf. History,
25
systematics, morphology, ecology. Moscow, Nauka, pp 355-370 (in Russian)
17
1
2
3
Matyushkin EN, Vaisfeld MA (eds) (2003) The lynx. Regional features of ecology, use and
protection. Nauka, Moscow (in Russian with English summary)
Matyushkin EN, Podolskii SA, Tkachenko KN (2003) The south of Far East. In: Matyushkin
4
EN, Vaisfeld MA (eds). The lynx. Regional features of ecology, use and protection.
5
Nauka, Moscow, pp 423-472 (in Russian with English summary)
6
Miquelle DG, Stephens PA, Smirnov EN, Goodrich JM, Zaumyslova OJ, Myslenkov AE
7
(2005) Tigers and wolves in the Russian Far East: Competitive exclusion, functional
8
redundancy, and conservation implications. In: Pages in Ray JC, Redford KH, Steneck
9
RS, and Berger J, (eds). Large carnivores and the conservation of biodiversity. Island
10
11
12
Press, Washington DC, pp 179-207
Myrberget S (1970) The Norwegian population of wolverine, Gulo gulo (L.), and lynx, Lynx
lynx (L.). Medd Stat Viltunders 2: 1-35
13
Nowak S, Mysłajek RW, Jędrzejewska B (2005) Patterns of wolf Canis lupus predation on
14
wild and domestic ungulates in the Western Carpathian Mountains (S Poland). Acta
15
Theriol 50: 263-276
16
17
18
Nowell K, Jackson P (1996) Wild cats. Status survey and conservation action plan. IUCN,
Gland.
Okarma H, Jędrzejewski W, Schmidt K, Kowalczyk R, Jędrzejewska B (1997) Predation of
19
Eurasian lynx on roe deer and red deer in Białowieża Primeval Forest, Poland. Acta
20
Theriol 42: 203-224
21
Okarma H, Jędrzejewski W, Schmidt K, Śnieżko S, Bunevich AN, Jędrzejewska B (1998)
22
Home ranges of wolves in Białowieża Primeval Forest, Poland, compared with other
23
Eurasian populations. J Mammal 79: 842-852
24
Okarma H, Jędrzejewski W (1997) Livetrapping wolves with nets. Wildl Soc Bull 25: 78-82
18
1
2
Palomares F, Caro TM (1999) Interspecific killing among mammalian carnivores. Am Nat:
153: 492-508
3
Podgórski T, Schmidt K, Kowalczyk R, Gulczyńska A (2008) Microhabitat selection by
4
Eurasian lynx and its implications for species conservation. Acta Theriol 53: 97-110.
5
Pulliainen E (1965) Studies on the wolf (Canis lupus L.) in Finland. Ann Zool Fenn 2: 215-
6
7
8
259
Ray JC, Sunquist ME (2001) Trophic relations in a community of African rainforest
carnivores. Oecologia 127: 395-408
9
Rosenzweig ML (1966) Community structure in sympatric Carnivora. J Mammal 47: 602-612
10
Schmidt K, Jędrzejewski W, Okarma H (1997) Spatial organization and social relations in the
11
Eurasian lynx population in Białowieża Primeval Forest, Poland. Acta Theriol 42: 289-312
12
13
14
Schmidt K (1999) Variation in daily activity of the free living Eurasian lynx in Białowieża
Primeval Forest, Poland. J Zool (Lond.) 249: 417-425
Scognamillo D, Maxit IE, Sunquist ME, Polisar J (2003) Coexistence of jaguar (Panthera
15
onca) and puma (Puma concolor) in a mosaic landscape in the Venezuelan llanos. J Zool
16
(Lond.) 259: 269-279
17
Selva N, Jędrzejewska B, Jędrzejewski W, Wajrak A (2005) Factors affecting carcass use by a
18
guild of scavengers in European temperate woodland. Can J Zool 83: 1590-1601
19
Stander PE, Haden PJ, Kaqece //, Ghau // (1997) The ecology of asociality in Namibian
20
leopards. J Zool ( Lond) 242: 343-364
21
Sunquist ME, Sunquist F, Daneke DE (1989) Ecological separation in a venezuelan llanos
22
carnivore community. In: Redford KH, Eisenberg JF (eds). Advances in neotropical
23
mammalogy. Sandhill Crane Press, Gainesville, Florida, 197-232
19
1
Theuerkauf J, Jędrzejewski W, Schmidt K, Okarma H, Ruczyński I, Śnieżko S, Gula R (2003)
2
Daily patterns and duration of wolf activity in the Białowieża Forest (Poland). J Mammal
3
84: 243-253
4
5
6
7
8
9
10
Valdmann H, Andersone-Lilley Z, Koppa O, Ozolins J, Bagrade G (2005) Winter diets of
wolf Canis lupus and lynx Lynx lynx in Estonia and Latvia. Acta Theriol 50: 521-527
Wesołowski T (2005) Virtual conservation: how the European Union is turning a blind eye to
its vanishing primeval forests. Cons Biol 19: 1349–1358
Zheltukhin AS (1986) Biocenotic relationships of the European lynx (Lynx lynx) in the
southern taiga of the upper Volga. Zool Zhurn 65: 259-271 (in Russian with English
summary).
20
1
Table 1. Spatial overlap among home ranges of 7 lynx and territories of 3 wolf packs radio-
2
tracked simultaneously in the Białowieża Primeval Forest in 1994-1996. The overlaps were
3
calculated for combinations of particular wolf pack territories with all lynx home ranges and
4
for particular lynx home ranges with all wolf packs monitored concurrently. See Appendix for
5
number of locations of individual lynx and wolves.
Pair of species,
territory/home range estimate
Percentage overlap (percent of territory/home range
of the first species overlapped by all radio-tracked
individuals of the second species)
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
21
1
2
Table 2. Static (home range percentage overlap) and dynamic (Jacobs’ indices and average
3
distances between individuals) interactions between individual dyads of wolves and lynx
4
radio-tracked in the Białowieża Primeval Forest in 1994-1996, based on simultaneous
5
locations (on average within 1 h) only. n – number of simultaneous locations.
Dyads
(n)
Percentage overlap
of wolf
of lynx
territory
home
Distance between individuals (km)
Jacobs’ index
Mean ± SE
(min-max)
(D)
range
W1 – L1
3
5
6.9±0.2
2.3-18.4
0.03
40
40
5.5±0.1
0.3-16.0
0.03
46
29
11.0±0.5
0.4-23.8
0.02
31
19
7.1±0.4
0.3-27.8
0.04
63
52
4.4±0.3
0-18.3
0.36
(159)
W1 – L2
(424)
W2 – L3
(111)
W3 – L3
(225)
W3 – L4
(237)
6
22
1
Figure captions:
2
3
Figure 1. Distribution of all radio-locations of lynx (n=7521 of 14 radio-tracked individuals in
4
1991-1996) and wolves (n=28857 of 8 individuals belonging to 4 packs radio-tracked in 1994
5
-1999) in the Białowieża Primeval Forest and most frequently used areas (75% fixed kernel)
6
estimated for all locations pooled for each species. Question marks denote area where lynx
7
were not trapped for radio-collaring.
8
9
10
Figure 2. Territories (100% MCP) and core areas (50% fixed kernel) of wolf packs (n=3) and
home ranges of lynx (n=7) radio-tracked in the Białowieża Primeval Forest in 1994-1996.
11
12
Figure 3. Examples of movement routes of a pack of wolves (with the female W3) and male
13
lynx (L4) recorded during simultaneous, continuous radio-tracking of both carnivores in BPF.
23
Figure 1.
24
Figure 2.
25
Figure 3.
26
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
ID
name
code
Sex
Age
Time of simultaneous radio-
Number of
tracking
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
L4
M
Adult
29 Mar 1995 – 1 Apr 1996
923
Zoja
L5
F
Subadult
5 Apr 1994 – 24 Aug 1994
8
Olga
L6
F
Adult
22 Mar 1994 – 7 Sep 1994
99
Diana
L7
F
Adult
27 Mar 1995 – 3 Aug 1995
72
27