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Transcript of Ulves jagt på elg og rådyr i Sverige og Norge og byttedyrenes forsvarsadfærd. Videnskabelig...
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7/28/2019 Ulves jagt p elg og rdyr i Sverige og Norge og byttedyrenes forsvarsadfrd. Videnskabelig artikel 2009.
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Introduction
Adult ungulate species commonly use differ-
ent anti-predator strategies to avoid attacks by
predators such as birth synchrony (Estes 1966,
Adams and Dale 1998) , grouping (Ha milton
1971, Hebblewhite and Pletscher 2002), migra-
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 vigi-
lance (Hunter and Skinner 1998, Berger 1999),
alarm vocalizations (Berger 1978) or physical
traits such as size (Mech 1966). When attacked
[207]
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. Wolfpredation 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, 1758hunting moose Alces alces and roe deer Capreolus capreolus, and recordeddetails of encounters between wolves and prey on the Scandinavian Penin-sula, 19972003. In total, 252 wolf attacks on moose and 64 attacks on roedeer 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, adifference likely due to variation in body size and vigilance between preyspecies. A model including prey species, outcome of the attack, and snow depthexplained 1519% of the variation found in chase distances, with shorter chase
distances associated with greater snow depth and with successful attacks onmoose but not on roe deer. Wolf hunting success did not differ between preyspecies (moose 43%, roe deer 47%) but in 11% of the wolf attacks on moose atleast one moose was injured but not killed, whereas no injured roe deersurvived. Compared with most North American wolf studies chase distanceswere shorter, hunting success was greater, and fewer moose made a standwhen attacked by wolves in our study. Differences in wolf encounters withmoose and roe deer likely result from different anti-predator behaviour andpredator-prey history between prey species.
Grims Wildlife Research Station, Swedish University of Agricultural Sciences, SE-730 91Riddarhyttan, Sweden, e-mail: [email protected] (CW, HS, OL); Department ofForestry 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, predator-prey history
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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 con-
ditions (Kruuk 1972), and habitat cover (Lima
and Dill 1990) may also strongly affect prey vul-
nerability and the chances for escape by prey.
Stalking, typical for hunting in felids, is sen-
sitive to vegetation type and structure (Murray
et al. 1995), whereas cursorial canids may bemore 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 vulnerabil-
ity of prey to predation by wolves Canis lupus,
Linnaeus 1758 has been shown to increase with
greater snow depth (Peterson 1977, Nelson andMech 1986, Huggard 1993, Jdrzejewski et al.
2002). The decision to escape or fight may be in-
fluenced 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 at-
tacked 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).
Alt hough wolf pre dat ion behavio ur when
hunting ungulates has been studied in numer-
ous areas (Mech 1966, Peterson et al. 1984,
Messier 1994, Hayes and Harestad 2000, Jd-
rzejewski 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 encoun-
ters ie hunting success, and the proportion of
ungulates injured but not killed during encoun-
ters with wolves. These aspects may provide fur-
ther 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 examinethe length of chases in relation to wolf-related
(pack size, individual pack effect, number and
age of breeding wolves), prey-related (prey spe-
cies, 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-pred-
ator behaviour and predator-prey history.
Study area
Sweden and Norway constitute the 837 000 km2
Scan-
dinavian 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 collec-
tion. Human population density averages 17 inhabitants/km2,
but large areas within the main wolf range contain less
than one inhabitant/km2
(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
moose/km2
and 0.013.5 roe deer/km2, according to estimates
208 C. Wikenros et al.
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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 popula-
tion probably comprised more than 2000 individuals (Perssonand Sand 1998). By the start of the 20th century few indi-
viduals 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 Scan-
dinavia 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 esti-
mated 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 popula-
tion 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).
Material and methods
Chase distance
During the 19972003 winters, we snow-tracked both
radio-collared (in 15 wolf territories that at least during onewinter had a collared wolf; see Sand et al. 2006b for a de-
tailed 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 geo-
graphical 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 scent-
marking 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, Petersand Mech 1975). We used either a compass and map fea-
tures 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 indi-
cated they had been made simultaneously, this was re-
garded 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 mea-
sured by plotting the route on a topographic map, or by sav-
ing the route on a handheld GPS unit and then transferring
coordinates to GIS, ArcView 3.3 (ESRI, Redlands, Califor-
nia). An attack was considered successful if a wolf-killed
moose or roe deer was found along the chase route, and as afailure if no carcass was found. We identified species of un-
gulate prey remains by body size, hair, and skeletal re-
mains. 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 com-
paring 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 rela-
tionships 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). There-
fore, 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 rel-ative snow depth was 20 cm.
Outcome of wolf-prey encounters
When calculating hunting success and outcome of wolf-
prey 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 boundinggaits, 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
Wolf chase distances and prey encounters 209
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of captured adults (Gipson et al. 2000), and (3) construction
of a population pedigree based on DNA analyses for non-col-
lared 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.
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 out-
come of wolf attacks, prey species, number of wolves, num-
ber 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 Peter-son 2003) so we also tested for the effect of body conditions
by including time during the season of the wolf attack (by in-
cluding 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 con-
tinuous 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 vari-
able. As chase distances were not normally distributed, num-
bers 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 diffi-
culty 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.
Results
Chase distances
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, re-
spectively). 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 predic-
tive power of the model (Table 1). Successful at-
tacks were shorter than failed attacks and chase
distances decreased with an increase in the rela-
tive 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. Conse-
quently, according to the model, an increase in
the relative snow depth from 10 to 40 cm, re-
sulted in an average 46% reduction in chase dis-
tance from 140 to 75 m (controlling the effects of
outcome and prey species). In this model, includ-
ing only first-order terms, neither the number of
wolves nor the number of breeding wolves in-
210 C. Wikenros et al.
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volved in the attack were significant predictors
of chase distances (Table 1).
In addition, it was shown that two second-or-
der terms were significantly related to chase dis-
tances 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 de-
creased with an increase in the number of
wolves involved in successful attacks but in-
creased with an increase in the number of
wolves involved in failed attacks.The multiple model including the three sig-
nificant predictor variables (prey species, out-
come, snow depth) explained 15% of the total
Wolf chase distances and prey encounters 211
Table 1. Single factors, multiple factors and interaction terms used in an ANCOVA analysis to estimateeffects on the length of chase distances during wolf attacks ( n = 212) on moose and roe deer, in Scandina-
via during winters of 19972003.
Variable(s) in model df F R2
p Power AIC
Constant 356.89
Outcome 1 2.87 0.01 0.09 0.38 356.02
Prey species 1 24.13 0.12 0.00 1.00 335.84
Snow depth 1 12.11 0.06 0.00 0.95 347.01
n wolves 1 1.07 0.01 0.30 0.18 357.82
n breeding wolves 1 3.02 0.01 0.08 0.41 355.87
Prey species +
Outcome 1 6.11 0.13 0.01 0.69 331.74
Snow depth 1 5.83 0.13 0.02 0.67 332.00
n wolves 1 0.04 0.10 0.84 0.06 337.80
n breeding wolves 1 0.84 0.11 0.36 0.15 336.99
Prey species + outcome +
Snow depth 1 5.94 0.15 0.02 0.68 327.76
n wolves 1 0.33 0.13 0.56 0.09 333.40
n breeding wolves 1 0.69 0.13 0.41 0.13 333.04
Prey species + outcome + snow depth +
Outcome prey species 1 5.17 0.17 0.02 0.62 324.54
Outcome snow depth 1 0.01 0.15 0.93 0.05 329.76
Outcome n wolves 1 4.92 0.19 0.01 0.80 321.87
Outcome n breeding wolves 1 0.00 0.16 0.96 0.05 331.21
Prey species snow depth 1 0.38 0.15 0.54 0.10 329.37
Prey species n wolves 1 1.66 0.17 0.19 0.35 328.37
Prey species n breeding wolves 1 2.71 0.17 0.07 0.53 326.25
Snow depth n wolves 1 3.23 0.17 0.07 0.43 326.48
Snow depth n breeding wolves 1 0.52 0.16 0.47 0.11 329.23
n wolves n breeding wolves 1 0.08 0.15 0.78 0.06 329.68
0
100
200
300
400
500
600
700
800
Moose Roe deer
Successful
Failed
Chasedistance(m)
Fig. 1. Chase distances (mean SD) in successful and failedwolf attacks on moose (nsuccessful = 68, nfailed = 92) and roedeer (nsuccessful = 30, nfailed = 22), in Scandinavia during win-ters of 19972003.
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variation found in chase distances. Including
the interaction term with the strongest predic-
tive 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 sec-
ond-order terms were included. Inclusion of wolf
territory (n = 24) in the multiple model in addi-
tion to the above significant variables explained
some of the remaining variation (F = 1.69, p =
0.03, R2
= 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 inter-
territory level. Inclusion of the time during win-
ter (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
212 C. Wikenros et al.
0
250
500
750
1000
1250
1500
1750
2000
2250
0 5 10 15 20 25 30 35 40 45 50
Successful
Failed
0
250
500
750
1000
1250
1500
1750
2000
0 5 10 15 20 25 30 35 40 45 50
Snow depth (cm)
(a)
(b)
Chase
distance
(m)
Fig. 2. Chase distances (m) in relation to snow depth (measured as the difference between the depth prey and wolves hadsunk into the snow) during wolf attacks on (a) moose (nsuccessful = 68, nfailed = 91) and (b) roe deer (nsuccessful = 30, nfailed = 22), inScandinavia during winters of 19972003. One wolf attack on moose, with chase distance 5000 m at 40 cm difference in snowdepth, 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. Forty-two 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 sus-
tained 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 dis-
tributed over 14 wolf territories. Fifteen (47%) of
these attacks were successful. In 23 of the 32 at-tacks 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, re-
sulting in a hunting success of 22% per individual
roe deer attacked. Detailed description of the out-
come of wolf attacks on roe deer was available for
ten attacks (Fig. 4) involving 14 individuals. No
Wolf chase distances and prey encounters 213
62
chased by wolves
22
physical contact with wolves
66
moose attacked by wolves
13
injured (loss of hair/blood/skin)
5
pulled to ground
3
killed
( 1 adult, 2 calves)
40
escaped
unharmed
3
killed
(1 adult, 2 calves)
5
escaped
injured
2
escaped
injured
2
made a standand survived
9
killed at first contact
(9 calves)
2
stopped and made
a stand
Fig. 3. Outcome of encounters between moose and wolves in 40 attacks in Scandinavia during winters of 19972003. Dashedarrows indicate moose that survived wolf attacks and filled arrows indicate killed moose.
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roe deer made a stand when wolves approached.
Eight roe deer outran the wolves and escaped un-
harmed. 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 sur-
vived. Wolf hunting success on roe deer was simi-
lar to that of moose, both when calculated as the
proportion of successful attacks (moose: 43%, roe
deer: 47%, c2
= 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
deer: 22%, c2
= 0.16, df = 1, p = 0.69).
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
1900 m for failed attacks, during two successive
winters, and 2100 m during both winters for suc-
cessful attacks (Kolenosky 1972). Mech (1966)found that wolves on Isle Royale most often
(78%) gave up after less then 800 m when chas-
ing 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 a l. (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
Defense strategies to predator attacks are
likely to differ between moose and roe deer with
the larger moose being able to confront the at-
tacking 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 de-
pendent on their vigilance, their initial accelera-
tion speed, and their endurance to successfully
escape wolves. Mech and Korb (1978) reported
214 C. Wikenros et al.
1
killed while bedded
(adult)13
chased by wolves
5
physical contact with wolves
14
roe deer attacked by wolves
1
injured (loss of hair/blood/skin)
0
pulled to ground
1
killed
(age unknown)
8
escaped
unharmed
0
made a stand
4
killed at first contact
(2 adult, 2 age unknown)
0
stopped and made
a stand
Fig. 4. Outcome of encounters between roe deer and wolves in 10 attacks in Scandinavia during winters of 19972003. Dashedarrows indicate roe deer that survived wolf attacks and filled arrows indicate killed roe deer.
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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 be-
fore 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 sup-
ported by our study where all roe deer that were
in physical contact with wolves were killed.
Considering that moose and roe deer are rad-
ically 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 at-
tacking 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
Several studies have shown that snow depth
affects the vulnerability of ungulates to preda-
tion (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 at-
tack, but we lacked data on roe deer in snow
deeper than 20 cm. Moose vulnerability to pre-
dation 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 en-
counters 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 incorp-
orate 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
body mass to foot area), in this case resulting in
shorter chase distances with increased snow
depth.
Other factors important for chase distances
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 dif-
ferences in chase distances between prey spe-
cies. 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 re-sulted in a negative relationship between chase
distances and number of wolves during success-
ful 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 signif icantly ex-
plained variation in chase distances, most of the
variation (8183%) remained unexplained and
may be attributed to several predator-, prey-
and environmentally-related factors. For exam-ple, age and sex of breeding wolves have been
shown to affect hunting success on Scandina-
vian moose (Sand et al. 2006b). Therefore, we as-
sumed 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 (us-
ing 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 fac-
tor 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-
Wolf chase distances and prey encounters 215
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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 dis-
tances than in other predator-prey systems hav-
ing 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 visibil-
ity, 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 parameterwas not recorded during field work, nor avail-
able 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.
Outcome of wolf-prey encounters
Adult moose were generally more difficult to
kill for wolves once caught up than calves that inturn 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 dis-
tances 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
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. Com-
pared to moose, roe deer may be a less preda-
tor-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 be-
haviour, as compared to moose, in response to
the return of the main predator.Acknowledgements: We thank a number of people who helpedwith 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. StDhland R. Wiklund. M. Apollonio, J. W. Laundr, R. Petersonand one anonymous reviewer provided valuable commentson an earlier draft of this paper. This study was supportedby the Swedish Environmental Protection Agency, the Asso-ciation for Hunting and Wildlife Management, the WorldWildlife Fund for Nature (WWF Sweden), the Swedish Uni-
versity of Agricultural Sciences, the Norwegian Directoratefor 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, Olleand Signhild Engkvists Stiftelse, Carl Tryggers Stiftelse,the Swedish Carnivore Association, Stor-Elvdal, Trysil, Cmotand Csnes Municipalities.
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