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Acta Theriologica 54 (3): 207218, 2009.

PL ISSN 00017051

DOI: 10.4098/j.at.00017051.082.2008

Wolf predation on moose and roe deer: chase distances and outcome of encounters
Camilla WIKENROS, HDkan SAND, Petter WABAKKEN, Olof LIBERG and Hans C. PEDERSEN

Wikenros C., Sand H., Wabakken P., Liberg O. and Pedersen H. C. 2009. Wolf predation on moose and roe deer: chase distances and outcome of encounters. Acta Theriologica 54: 207218. We examined chase distances of gray wolves Canis lupus Linnaeus, 1758 hunting moose Alces alces and roe deer Capreolus capreolus, and recorded details of encounters between wolves and prey on the Scandinavian Peninsula, 19972003. In total, 252 wolf attacks on moose and 64 attacks on roe deer were registered during 4200 km of snow tracking in 28 wolf territories. Average chase distances were 76 m for moose and 237 m for roe deer, a difference likely due to variation in body size and vigilance between prey species. A model including prey species, outcome of the attack, and snow depth explained 1519% of the variation found in chase distances, with shorter chase distances associated with greater snow depth and with successful attacks on moose but not on roe deer. Wolf hunting success did not differ between prey species (moose 43%, roe deer 47%) but in 11% of the wolf attacks on moose at least one moose was injured but not killed, whereas no injured roe deer survived. Compared with most North American wolf studies chase distances were shorter, hunting success was greater, and fewer moose made a stand when attacked by wolves in our study. Differences in wolf encounters with moose and roe deer likely result from different anti-predator behaviour and predator-prey history between prey species.
Grims Wildlife Research Station, Swedish University of Agricultural Sciences, SE-730 91 Riddarhyttan, Sweden, e-mail: camilla.wikenros@ekol.slu.se (CW, HS, OL); Department of Forestry and Wildlife Research, Hedmark University College, Evenstad, NO-2480 Koppang, Norway (PW); Norwegian Institute for Nature Research, Tungasletta 2, NO-7485 Trondheim, Norway (HCP)

Key words: anti-predator behaviour, chase distance, hunting success, predatorprey history

Introduction
Adult ungulate species commonly use different anti-predator strategies to avoid attacks by predators such as birth synchrony (Estes 1966, Adams and Dale 1998), grouping (Hamilton 1971, Hebblewhite and Pletscher 2002), migra[207]

tion (Fryxell et al. 1988, Fryxell and Sinclair 1988, Nelson and Mech 1991), protective cover (Lima and Dill 1990), escape features such as standing in water (Mech 1966), increased vigilance (Hunter and Skinner 1998, Berger 1999), alarm vocalizations (Berger 1978) or physical traits such as size (Mech 1966). When attacked

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by a predator, ungulate prey species may either try to outrun the attacking predator, or make a stand to fend off the predator (Mech 1966, Kruuk 1972). To outrun a predator, vigilance and speed is important, while to stand at bay and fight a predator, body size and condition of potential prey relative to the predator is crucial (Mech 1966). Environmental factors such as snow characteristics (Peterson 1977), light conditions (Kruuk 1972), and habitat cover (Lima and Dill 1990) may also strongly affect prey vulnerability and the chances for escape by prey. Stalking, typical for hunting in felids, is sensitive to vegetation type and structure (Murray et al. 1995), whereas cursorial canids may be more successful when hunting in open habitats (Wells and Bekoff 1982), and may often exhaust larger prey by chasing them over long distances (Kruuk and Turner 1967, Kleiman and Eisenberg 1973). Even among prey species chased by canids, the use of protective cover may affect the risk of predation (Kunkel and Pletscher 2000, Dussault et al. 2005, Hebblewhite et al. 2005, Kauffman et al. 2007). In temperate regions, the vulnerability of prey to predation by wolves Canis lupus, Linnaeus 1758 has been shown to increase with greater snow depth (Peterson 1977, Nelson and Mech 1986, Huggard 1993, Jdrzejewski et al. 2002). The decision to escape or fight may be influenced by snow depth as it is more difficult for some prey species to outrun predators in deep snow (Paquet 1992). In North America, wolf chases on ungulates usually last one to two km (Mech 1966, Peterson 1977). In the classical studies of moose Alces alces and wolf encounters on Isle Royale, Mech (1966, 1970) showed that the best strategy for a moose to survive a wolf attack was to stand at bay and not try to outrun the attacker. Attempted escape as an unsuccessful strategy when attacked by wolves has also been reported for mule deer Odocoileus hemionus (Lingle and Pellis 2002), caribou Rangifer tarandus (Crisler 1956) bison Bison bison (Smith et al. 2000), elk Cervus elaphus (MacNulty et al. 2007), and musk ox Ovibos moschatus (Mech 1988). Although wolf predation behaviour when hunting ungulates has been studied in numerous areas (Mech 1966, Peterson et al. 1984,

Messier 1994, Hayes and Harestad 2000, Jdrzejewski et al. 2000), relatively few reports have considered (1) actual distances covered during wolf attacks on different prey species ie chase distance, and (2) the outcome of encounters ie hunting success, and the proportion of ungulates injured but not killed during encounters with wolves. These aspects may provide further insight into the behavioural mechanisms important for the success of wolf attacks on their prey. The objective of this paper is to present data on chase distances of wolves on two important prey species during winter in Scandinavia, moose and roe deer Capreolus capreolus. We examine the length of chases in relation to wolf-related (pack size, individual pack effect, number and age of breeding wolves), prey-related (prey species, age class: possible for moose only and body condition) and a weather-related (relative snow depth) factor. We describe in detail the outcome of encounters between wolves and their two prey species in terms of hunting success, proportion of injured prey and prey defence behaviour, and discuss the results in the light of prey anti-predator behaviour and predator-prey history.

Study area
Sweden and Norway constitute the 837 000 km Scandinavian Peninsula, hereafter referred as Scandinavia. Boreal coniferous forest and alpine areas cover more than 75% of the peninsula. Wolf territories are dominated by forests consisting of Norway spruce Picea abies and Scots pine Pinus silvestris, but birch Betula pubescens, B. pendula and aspen Populus tremula are also available in various mixtures (Moen 1998, Swedish National Atlas 1991a). The study area is characterized by intensive forestry and mature stands are harvested by clear-cutting. This creates a mosaic of even-aged stands in the landscape together with lakes, bogs and mires. The terrain is hilly with various visibilities due to the forest age, but we lack a detailed description of habitat, topography and visibility at the time of data collec2 tion. Human population density averages 17 inhabitants/km , but large areas within the main wolf range contain less 2 than one inhabitant/km (Swedish National Atlas 1991b). The most important wild prey species for wolves are moose and roe deer (Sand et al. 2005). Badger Meles meles, beaver Castor fiber, and in Norway also red deer Cervus elaphus and potentially wild reindeer Rangifer tarandus are also available. Within the wolf breeding range, the winter densities of moose and roe deer range between 0.62.5 2 moose/km and 0.013.5 roe deer/km2, according to estimates
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from pellet group counts and aerial censuses (Sand et al. 2006a, Sand et al. unpublished data). In the mid-19th century the Scandinavian wolf population probably comprised more than 2000 individuals (Persson and Sand 1998). By the start of the 20th century few individuals remained due to intensive persecution and in 1966, when they became legally protected in Sweden, wolves were functionally extinct (Haglund 1968, Wabakken et al. 2001). In the late 1970s wolves re-appeared in south-central Scandinavia and since 1983 they have successfully reproduced every year, except for 1986 (Wabakken et al. 2001). During the winter of 20022003 the total population size was estimated at approximately 84100 wolves (including 9 pairs and 8 packs) and their distribution was concentrated in the boreal coniferous forest of south-central Scandinavia (Wabakken et al. 2003). The Scandinavian moose population has grown tremendously throughout the 20th century and has been exposed to an intensive management regime that replaced most natural mortality with human harvest (Lavsund and Sandegren 1989, Saether et al. 1996). The Scandinavian roe deer population was considered almost extinct in the 1920s, but recovered quickly and completely re-occupied south-central Scandinavia during the early 1980s (Liberg et al. 1994).

ing the route on a handheld GPS unit and then transferring coordinates to GIS, ArcView 3.3 (ESRI, Redlands, California). An attack was considered successful if a wolf-killed moose or roe deer was found along the chase route, and as a failure if no carcass was found. We identified species of ungulate prey remains by body size, hair, and skeletal remains. We collected mandibles from moose and used them to determine age by their ontogenetic development and classified age as calves (< 1 year old) or adults (> 1 year old) (Markgren 1969). For roe deer, age was determined by comparing tooth eruption of mandibles from wolf-killed roe deer with roe deer of known age (Cederlund and Liberg 1995). For the current analysis, age was classified into juvenile (< 1 year old) or adult (> 1 year old).

Snow depth
Snow (depth, density, texture) affects wolf-prey relationships mainly through restricting movements and may restrict the chances for successful escape of ungulate prey (Mech 1970, Peterson and Allen 1974, Peterson 1977, Huggard 1993). Compared to moose and deer, wolves have a foot loading (ie foot-load-on-track) that is several times lighter making wolves better supported in snow of a certain density (Peterson 1977, Mech and Peterson 2003). Therefore, we recorded the effect of snow by measuring how deeply each species sunk into the snow during the attack. To incorporate the predator-prey-specific constraints of snow, we analyzed the effect of snow depth by applying the difference in snow depth (cm) experienced by wolves and their prey during the attack, hereafter referred to as the relative snow depth. For example, if the prey sunk 40 cm and the wolves 20 cm in the snow during the attack the relative snow depth was 20 cm.

Material and methods


Chase distance
During the 19972003 winters, we snow-tracked both radio-collared (in 15 wolf territories that at least during one winter had a collared wolf; see Sand et al. 2006b for a detailed description of animal capture and collar use) and non-collared wolves (in 13 wolf territories) on foot, on skis and occasionally by snowmobile. The research project was evaluated and approved by The Swedish Agency of Animal Welfare (C 266/99) and The Norwegian Agency of Animal Welfare. Data recorded from snow-tracking included geographical location and length (km) of tracking route, name of territory, number of wolves (total and number of breeding wolves), determination of sex and social position (eg scentmarking behavior). During the pre-mating season, females were identified by the presence of vaginal blood in the urine on snow during estrus, whereas counts of newly-formed pairs and the number of breeding wolves within packs were based on their scent-marking behaviour (Mech 1970, Peters and Mech 1975). We used either a compass and map features or a handheld GPS unit (Garmin GPS 12/12XL) to plot wolf tracking routes onto 1:50000 topographic maps. During snow tracking, all wolf attacks on moose and roe deer were recorded. A hunting attack was defined as where lengthening stride patterns for both wolves and moose or roe deer indicated bounding gaits (fast running). When such tracks occurred together and local snow conditions indicated they had been made simultaneously, this was regarded as an attack (see also Murray et al. 1995 for a similar definition). The chase distance was defined as the stretch of fast running by the wolf/wolves and was measured by plotting the route on a topographic map, or by sav-

Outcome of wolf-prey encounters


When calculating hunting success and outcome of wolfprey encounters we used data only from territories with non-collared wolves. This was done to avoid a bias against successful attacks as we searched for carcasses using radio telemetry in territories with radio-collared wolves and therefore found a higher proportion of successful attacks in those territories. Each attack was described in terms of the number of moose or roe deer involved and the outcome of the attack (successful or failed). We classified prey as injured if blood, hair or skin were found along the bounding gaits, or if the prey had been pulled to ground by wolves. Prey defence behaviour was described as either (1) prey trying to outrun wolves (bounding gaits present), (2) making a stand (no bounding gaits but including a smaller area of frequent trampling of both wolves and moose) or (3) a combination of the two.

Age determination of wolves


We determined the age of breeding adult wolves by a combination of three methods: (1) known age of captured animals classified as pups at first capture by the presence of a growth zone in the tibia (Rausch 1967), (2) tooth wear

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of captured adults (Gipson et al. 2000), and (3) construction of a population pedigree based on DNA analyses for non-collared wolves (Liberg et al. 2005). We used the population pedigree in combination with a complete set of data on the chronological order and geographical location of breeding packs in Scandinavia to identify the year of birth and, thus, current age for each wolf (Wabakken et al. 2001, Liberg et al. 2005). Samples for genetic analyses were derived from either blood or tissue from captured or dead wolves, or from blood on snow (females in heat) and scats collected during snow tracking of wolves. If the different methods estimated different ages for the same animal, we ranked the methods in order of accuracy as follows: capture of pups > pedigree construction of birth year > tooth wear of adult wolves. If results indicated that the birth year could have two or three alternatives, we used the lower or median age, respectively.

Results
Chase distances

Analyses and statistical procedures


We used the total dataset (n = 316) and the students t-test to analyze differences in chase distances between kills of adult and calf moose (the sample size did not allow for a comparison between roe deer age classes). As data of snow depth and number of wolves was not always recorded in the total dataset, we used a reduced dataset (n = 212) and an ANCOVA to discriminate between the effects of outcome of wolf attacks, prey species, number of wolves, number of breeding wolves, and snow depth on the length of chases. In order to test whether the main variation in chase distances explained by model variables was found within or between wolf territories (temporal pseudo replication) we included wolf territory (n = 24) as a random variable in the model. The body condition of temperate ungulates tends to deteriorate towards the end of the winter (Mech and Peterson 2003) so we also tested for the effect of body conditions by including time during the season of the wolf attack (by including the number of days from 1 November) in the model. Age and sex of the breeding wolves were not known in all cases even in the reduced dataset (n = 212), so we tested for the potential effect of these factors in a further reduced dataset (n = 174). In this dataset the ages of the oldest of the two breeding wolves in each pack were grouped into five classes (1, 2, 3, 4, and 5 years and older) and used as a continuous variable together with a factorial variable denoting the number of breeding wolves present (1 or 2), and the sex of the breeding wolf (1 = male, 0 = female) present. Test for interaction (age number of breeding wolves) between these two variables allowed an evaluation of the potential effect of sex, in the case of only one breeding wolf present, and age of the breeding wolf/wolves on the dependent variable. As chase distances were not normally distributed, numbers were transformed by ln(x) + 1 but results are presented as back-transformed data in meters. Variables included in the model were considered significant at an alpha level of 0.05 and we applied the model building strategy of stepwise forward inclusion of independent variables. Due to the difficulty of biological interpretation of data we did not include multiple second-order terms in addition to single factors. Akaikes information criteria AIC were also calculated and showed the same model selection results among models as the one based on traditional significance levels. Analyses were performed with SPSS version 11.5 for Windows.

A total of 316 chase distances (average 13 chases per pack, range 197, all years pooled) by wolves were registered including both successful and failed attacks on moose (n = 252) and roe deer (n = 64), during approximately 4200 km of snow tracking for 28 packs (pack size 111 wolves). Successful chase distances ranged from 0 m (one roe deer and nine moose killed while bedded) to 1.7 km and 2.3 km for moose and roe deer, respectively. The longest failed chases were 5 km for moose and 13.7 km for roe deer, but most chase distances were shorter than 400 meters (90% and 69% for moose and roe deer, respectively). Multiple kill of prey was registered only on one occasion where two roe deer were killed during one attack by two adult wolves. There were no significant differences in chase distances between kills of adult (mean = 82 m) and calf (mean = 53 m) moose (t = 1.070, df = 1, nadult = 15, ncalf = 84, p = 0.29).
Factors affecting chase distances

Inclusion of single and multiple variables in an ANCOVA showed that prey species was the single most important factor, explaining 12% of the total variance (F = 24.13, p < 0.001, n = 212), with an average of 68% shorter chase distances for moose as compared to roe deer (Fig. 1). The outcome of an attack (successful or failed) and the effect of snow slightly increased the predictive power of the model (Table 1). Successful attacks were shorter than failed attacks and chase distances decreased with an increase in the relative snow depth (Figs 1 and 2). On average ( SD) wolves sunk 17 12 cm, moose 26 13 cm and roe deer 21 14 cm during wolf attacks, and snow depth ranged between 1 and 80 cm. Consequently, according to the model, an increase in the relative snow depth from 10 to 40 cm, resulted in an average 46% reduction in chase distance from 140 to 75 m (controlling the effects of outcome and prey species). In this model, including only first-order terms, neither the number of wolves nor the number of breeding wolves in-

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800 700

Successful Failed

600 500 400 300 200 100 0 Moose Roe deer

Fig. 1. Chase distances (mean SD) in successful and failed wolf attacks on moose (nsuccessful = 68, nfailed = 92) and roe deer (nsuccessful = 30, nfailed = 22), in Scandinavia during winters of 19972003.

volved in the attack were significant predictors of chase distances (Table 1). In addition, it was shown that two second-order terms were significantly related to chase distances for wolves on moose and roe deer. Chase distances differed between moose and roe deer but were also dependent on the outcome of the attack. Thus, successful attacks were shorter than failed attacks for moose (66 versus 123 m) whereas the reverse was true for roe deer (273 versus 212 m) (Fig. 1). Chase distances also decreased with an increase in the number of wolves involved in successful attacks but increased with an increase in the number of wolves involved in failed attacks. The multiple model including the three significant predictor variables (prey species, outcome, snow depth) explained 15% of the total

Chase distance (m)

Table 1. Single factors, multiple factors and interaction terms used in an ANCOVA analysis to estimate effects on the length of chase distances during wolf attacks (n = 212) on moose and roe deer, in Scandinavia during winters of 19972003. Variable(s) in model Constant Outcome Prey species Snow depth n wolves n breeding wolves Prey species + Outcome Snow depth n wolves n breeding wolves Prey species + outcome + Snow depth n wolves n breeding wolves Prey species + outcome + snow depth + Outcome prey species Outcome snow depth Outcome n wolves Outcome n breeding wolves Prey species snow depth Prey species n wolves Prey species n breeding wolves Snow depth n wolves Snow depth n breeding wolves n wolves n breeding wolves df F R
2

Power

AIC 356.89 356.02 335.84 347.01 357.82 355.87

1 1 1 1 1

2.87 24.13 12.11 1.07 3.02

0.01 0.12 0.06 0.01 0.01

0.09 0.00 0.00 0.30 0.08

0.38 1.00 0.95 0.18 0.41

1 1 1 1

6.11 5.83 0.04 0.84

0.13 0.13 0.10 0.11

0.01 0.02 0.84 0.36

0.69 0.67 0.06 0.15

331.74 332.00 337.80 336.99

1 1 1

5.94 0.33 0.69

0.15 0.13 0.13

0.02 0.56 0.41

0.68 0.09 0.13

327.76 333.40 333.04

1 1 1 1 1 1 1 1 1 1

5.17 0.01 4.92 0.00 0.38 1.66 2.71 3.23 0.52 0.08

0.17 0.15 0.19 0.16 0.15 0.17 0.17 0.17 0.16 0.15

0.02 0.93 0.01 0.96 0.54 0.19 0.07 0.07 0.47 0.78

0.62 0.05 0.80 0.05 0.10 0.35 0.53 0.43 0.11 0.06

324.54 329.76 321.87 331.21 329.37 328.37 326.25 326.48 329.23 329.68

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variation found in chase distances. Including the interaction term with the strongest predictive power (outcome*number of wolves, p = 0.01) slightly increased the total variation explained to 19% whereas inclusion of the second strongest interaction term (outcome*prey species, p = 0.02) explained 17% of the total variation (Table 1). Consequently, most of the variation in chase distances remained unexplained even when second-order terms were included. Inclusion of wolf territory (n = 24) in the multiple model in addition to the above significant variables explained some of the remaining variation (F = 1.69, p = 2 0.03, R = 0.034), but this did not influence the effect size of the model parameters for prey spe-

cies or outcome. However, it did result in a non-significant effect of snow depth (F = 1.80, p = 0.18), indicating that the effect of snow on chase distances was mainly found at the interterritory level. Inclusion of the time during winter (date) of the wolf attack did not increase the predictability of the multiple model (F = 0.11, p = 0.74). Finally, in a reduced dataset (n = 174) neither age of breeding wolves (n = 174, F = 0.40, p = 0.53), nor the number of breeding wolves or sex of the remaining breeding wolf (in cases of only one breeding animal) (F = 1.98, p = 0.14) were significant predictors of chase distances in addition to prey species, outcome and snow depth. The latter three predictor variables also

2250 2000 1750 1500 1250 1000 750 500

(a)

Failed Successful

Chase distance (m)

250 0 0 5 10 15 20 25 30 35 40 45 50

2000 1750 1500 1250 1000 750 500 250 0

(b)

10

15

20

25

30

35

40

45

50

Snow depth (cm)


Fig. 2. Chase distances (m) in relation to snow depth (measured as the difference between the depth prey and wolves had sunk into the snow) during wolf attacks on (a) moose (nsuccessful = 68, nfailed = 91) and (b) roe deer (nsuccessful = 30, nfailed = 22), in Scandinavia during winters of 19972003. One wolf attack on moose, with chase distance 5000 m at 40 cm difference in snow depth, is not shown in Fig. 2a.

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remained significant in this reduced dataset and with approximately the same size of the model parameter estimates as in the larger dataset (n = 212).
Outcome of wolf-prey encounters

Detailed reconstruction of the outcome of wolf and moose encounters was available for 40 wolf attacks involving 66 moose (Fig. 3) in 19 wolf territories. Four of the 66 moose (6%) were confirmed to have made a stand, two of them (age unknown) directly when confronted with the wolves and two (cow and calf) after a 1 km long chase. All four survived unharmed. Fortytwo moose (64%) escaped unharmed by fleeing immediately and seven more (11%) after being injured by the wolves, as we found blood, hair or skin along the flight track. Two of the injured moose were pulled to the ground one and six times, during chases that lasted 2 km and 5 km, respectively, but both moose managed to get up

and outrun the wolves. We do not know if those moose later died due to the wounds they sustained during the attack. Seventeen of the 66 moose (26%) were killed, two of them (one adult, one calf) while bedded, and 15 after being chased and caught by the wolves. Nine of these were killed during first contact (all calves) and the other six (two adults, four calves) after repeated contacts with wolves during the flight. Three of the six moose that were killed after repeated contacts were also pulled to the ground while trying to escape the wolves. A total of 32 wolf attacks on roe deer were distributed over 14 wolf territories. Fifteen (47%) of these attacks were successful. In 23 of the 32 attacks the number of roe deer attacked was also registered resulting in a total of 36 roe deer. Eight of the 36 roe deer attacked were killed, resulting in a hunting success of 22% per individual roe deer attacked. Detailed description of the outcome of wolf attacks on roe deer was available for ten attacks (Fig. 4) involving 14 individuals. No

66 moose attacked by wolves 2 made a stand and survived 40 escaped unharmed

62 chased by wolves

22 physical contact with wolves 2 stopped and made a stand 9 killed at first contact (9 calves)

13 injured (loss of hair/blood/skin)

5 escaped injured

5 pulled to ground

3 killed ( 1 adult, 2 calves)

2 escaped injured

3 killed (1 adult, 2 calves)

Fig. 3. Outcome of encounters between moose and wolves in 40 attacks in Scandinavia during winters of 19972003. Dashed arrows indicate moose that survived wolf attacks and filled arrows indicate killed moose.

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14 roe deer attacked by wolves 0 made a stand 8 escaped unharmed 13 chased by wolves 1 killed while bedded ( adult)

5 physical contact with wolves 0 stopped and made a stand 4 killed at first contact (2 adult, 2 age unknown)

1 injured (loss of hair/blood/skin)

0 pulled to ground

1 killed (age unknown)

Fig. 4. Outcome of encounters between roe deer and wolves in 10 attacks in Scandinavia during winters of 19972003. Dashed arrows indicate roe deer that survived wolf attacks and filled arrows indicate killed roe deer.

roe deer made a stand when wolves approached. Eight roe deer outran the wolves and escaped unharmed. One roe deer was killed while bedded, four were killed at first contact during flight, and one was killed after repeated injuries during flight. No roe deer injured during an attack survived. Wolf hunting success on roe deer was similar to that of moose, both when calculated as the proportion of successful attacks (moose: 43%, roe 2 deer: 47%, c = 0.14, df = 1, p = 0.71) and as the proportion of prey animals killed out of all prey animals involved in the attacks (moose: 26%, roe 2 deer: 22%, c = 0.16, df = 1, p = 0.69).

1900 m for failed attacks, during two successive winters, and 2100 m during both winters for successful attacks (Kolenosky 1972). Mech (1966) found that wolves on Isle Royale most often (78%) gave up after less then 800 m when chasing moose, with the longest chase being 4.8 km. Paquet (1989) recorded average distances of 886 m, 159 and 115 m for successful chases on moose, deer and elk, respectively, in Manitoba, Canada, whereas Hebblewhite et al . (2005) showed that the average chase distance by wolves on elk in Alberta, Canada was 260 m and ranged between 10 and 1700 m.
Causes of variable chase distances

Discussion
Chase distances

Chase distances of Scandinavian wolves on moose and roe deer showed large variation but were mostly shorter than 400 m. This is in the lower range of distances reported in most North American wolf studies including white-tailed deer Odocoileus virginianus with 600 m and

Defense strategies to predator attacks are likely to differ between moose and roe deer with the larger moose being able to confront the attacking predator, an option not open to roe deer (roe deer/moose size ratio 1:15 of adult and 1:6 of calf moose). Roe deer are therefore totally dependent on their vigilance, their initial acceleration speed, and their endurance to successfully escape wolves. Mech and Korb (1978) reported

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that one white-tailed deer fled for more than 21 kilometres without being killed, and in our study wolves chased one roe deer for almost 14 km before giving up the attack. Once caught up by wolves, small deer species like white-tailed deer and roe deer seem to be doomed, which is supported by our study where all roe deer that were in physical contact with wolves were killed. Considering that moose and roe deer are radically different in meat yield, and that hunting success was shown to be approximately equal for both prey species, longer chase distances might be expected for moose compared to roe deer. However, all moose attacked are not vulnerable to wolf predation (Mech 1970, Sand et al. 2005), and because of the higher risk involved in attacking moose compared to smaller deer (Weaver et al. 1992, Mech and Peterson 2003), wolves may continuously evaluate the vulnerability of individual moose during attacks and terminate the attack on certain individuals.
Snow depth

body mass to foot area), in this case resulting in shorter chase distances with increased snow depth.
Other factors important for chase distances

Several studies have shown that snow depth affects the vulnerability of ungulates to predation (Peterson 1977, Nelson and Mech 1986, Jdrzejewski et al. 1992, 2002, Huggard 1993), a finding consistent with our study. More snow generally resulted in shorter chases for both prey species irrespective of outcome of the attack, but we lacked data on roe deer in snow deeper than 20 cm. Moose vulnerability to predation by wolves has been shown to increase at snow depths of more then 75 cm (Peterson 1977). During our study winters snow depths exceeded this depth in only one wolf territory. Therefore, it is likely that the constraint of snow will be much more marked at snow depth greater than what was found for majority of wolf-prey encounters in this study. In our analyses, snow depth was measured as the difference between the depth prey and wolves sunk into the snow during the attack, which enabled us to incorporate the predator-prey-specific constraints of snow. Despite the relatively limited variation in snow depth, our results conform to earlier findings that wolves have an advantage over prey in snow due to their lower foot-load (ratio of

Our results showed that the number of wolves involved in an attack was negatively related to chase distances, but only for successful attacks. Inspection of data showed that this relationship was a statistical artifact that was related to differences in chase distances between prey species. As chase distances on roe deer were longer than on moose, and we had a maximum of three wolves involved in attacks on roe deer, this resulted in a negative relationship between chase distances and number of wolves during successful attacks. Consequently, this study provided no support for the assertion that the number of wolves involved in the attack was an important factor affecting chase distances on moose and roe deer. Although several factors significantly explained variation in chase distances, most of the variation (8183%) remained unexplained and may be attributed to several predator-, preyand environmentally-related factors. For example, age and sex of breeding wolves have been shown to affect hunting success on Scandinavian moose (Sand et al. 2006b). Therefore, we assumed that individual characteristics also would be important affecting chase distances if wolves of greater age and experience learn how to kill prey more efficiently. In contrast, this study (using largely the same dataset as in Sand et al. 2006b) found no support for our assumption that the age of breeding wolves was an important factor affecting chase distances. Nor did we found evidence that the time during winter, a proxy for general condition of prey, explained further variation in chase distances. One factor expected to have implications for the type and outcome of interactions between predators and prey is the physical features of the landscape. Landscape heterogeneity is likely to affect the visibility and influence the distance at which prey and predators may reach visual contact (Mills et al. 2004, Kauffman et al. 2007). However, our study area in Scandinavia is domi-

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nated by a mixture of relatively small and dense coniferous forest stands in variable age classes and tree species mixtures. Therefore, in general we would predict much shorter detection distances than in other predator-prey systems having a larger proportion of open landscapes, for example Yellowstone, USA (Boyce et al. 2003). Interestingly, chase distances on elk was reported to average 980 m in this system (Kauffman et al. 2007). Although landscape features of our study system offers a relatively low degree of visibility, we suggest that between-patch variability in visibility is likely responsible for a significant portion of the unexplained variation in chase distances found. Regrettably, this parameter was not recorded during field work, nor available at adequate resolution from GIS-data. In addition, individual characteristics like age, sex and body condition of prey animals involved in failed attacks may also contribute to some of the unexplained variation in chase distances found.

main mortality factor for moose during the last century in Scandinavia (Sand et al. 2006a). We suggest that differences in chase distances between prey species are explained by different anti-predator behaviour partly due to variation in size and vigilance in moose and roe deer. Compared to moose, roe deer may be a less predator-naive prey species since their major predator, the Eurasian lynx, reoccupied south-central Scandinavia 3040 years before the return of wolves to this region (Liberg 1998). In contrast to the cursorial wolf, the lynx is a stalking predator likely forcing roe deer to adopt a more vigilant behaviour, as compared to moose, in response to the return of the main predator.
Acknowledgements: We thank a number of people who helped with snow tracking of wolves, especially C. Aronson, B. Broman, B. Dahlen, J. Dahlen, H-E. Eriksson, G. Jansson, J. Johansson, K. Johansson, M. Johansson, P. Johansson, T. Mellgren, S. Nordgren, D. Palm, M. Rapp, H. Rrnning, M. Sandstrm, O. K. Steinset, T. Holm Strrmseth, A. StDhl and R. Wiklund. M. Apollonio, J. W. Laundr, R. Peterson and one anonymous reviewer provided valuable comments on an earlier draft of this paper. This study was supported by the Swedish Environmental Protection Agency, the Association for Hunting and Wildlife Management, the World Wildlife Fund for Nature (WWF Sweden), the Swedish University of Agricultural Sciences, the Norwegian Directorate for Nature Management, the Norwegian Research Council, the Norwegian Institute for Nature Research, Hedmark University College, the County Governor of Hedmark, Borregaard Skoger AS, Glommen Skogeierforening, Olle and Signhild Engkvists Stiftelse, Carl Tryggers Stiftelse, the Swedish Carnivore Association, Stor-Elvdal, Trysil, Cmot and Csnes Municipalities.

Outcome of wolf-prey encounters

Adult moose were generally more difficult to kill for wolves once caught up than calves that in turn were more difficult to kill than roe deer. This was likely a result of size difference and the capability of defence of the different prey. In contrast, we found no difference in chase distances for successful attacks between calf and adult moose. The proportion of injured moose, 11%, was similar of that reported from Denali, Alaska (13%, Mech et al. 1998) but higher than reported from Isle Royale, USA (1%, Mech 1966). Hunting success on moose was greater than that reported from North America (Mech 1966, Mech et al. 1998) but our data indicated that only a small proportion of the moose attacked by wolves made a stand while, for example on Isle Royale, up to 40% of the moose showed this behaviour when confronted with wolves (Mech 1970, Peterson 1977). The relatively high hunting success rate, short chase distances, and a low proportion of moose that made a stand are likely to be a result of moose being a predator-naive prey to wolves, a result of predator-prey history and man as the

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