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Interspecific and intraspecific social interactions among brown bears and wolves in an enclosure?

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We investigated bear-bear, wolf-wolf, and bear-wolf interactions in a 2-ha enclosure, which was occupied by 13 brown bears (Ursus arctos) and a wolf (Canis lupus) pack. The hierarchy of bears and wolves was determined by behavioral observation and rank order analysis. All behavioral interactions between bears and wolves were systematically videotaped for 1 month and analyzed. Bears and wolves had a hierarchical organization similar to that found in nature, although we found a relatively low position in the rank order of blind bears. Social interactions between wolves appeared to be very serious at times. Bear-wolf interactions were mostly playful, but were sometimes agonistic. Young bears were more often victims of wolf harassment than other bears and were sometimes seriously bitten. However, no deaths were recorded, contrary to descriptions from natural bear-wolf interactions. We concluded that bears and wolves could be kept safely together in a single enclosure with apparently few consequences. Interactions we observed were comparable to interactions between bears and wolves under natural conditions and concerned mainly competition over food or a den (wolf or bear). The size of the food (large prey carcasses in the wild vs. chicken in the enclosure) probably plays a role in the severity and the eventual solution of the bear-wolf conflict.
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INTERSPECIFIC
AND
INTRASPECIFIC SOCIAL
INTERACTIONS
AMONG
BROWN
BEARS
AND
WOLVES
IN
AN
ENCLOSURE
PAUL KOENE, Ethology Group, Department of Animal Sciences, Wageningen University, PO Box 338, 6700 AH Wageningen, the
Netherlands,
email:
paul.koene@etho.vh.wau.nl
JENTINA
ARDESCH,
Ethology
Group, Department
of Animal
Sciences, Wageningen
University,
PO Box 338, 6700 AH
Wageningen,
the Netherlands
ANNETTE LUDRIKS, Ethology Group, Department of Animal Sciences, Wageningen University, PO Box 338, 6700 AH Wageningen,
the Netherlands
EGBERT URFF, Ethology Group, Department of Animal Sciences, Wageningen University, PO Box 338, 6700 AH Wageningen, the
Netherlands,
email:
egberrt.urff@
etho.vh.wau.nl
LUDGER
WENZELIDES,
Ethology
Group,
Department
of Animal
Sciences, Wageningen
University,
PO Box
338, 6700 AH
Wageningen,
the Netherlands
VERENA WITTENBERG,
Ethology Group, Department of Animal Sciences, Wageningen University, PO Box 338, 6700 AH
Wageningen, the Netherlands
Abstract: We investigated
bear-bear, wolf-wolf, and bear-wolf interactions
in a 2-ha enclosure, which was occupied by 13 brown bears (Ursus
arctos) and a wolf (Canis lupus) pack. The hierarchy
of bears and wolves was determined
by behavioral observation
and rank order
analysis. All
behavioral
interactions between bears and wolves were systematically videotaped
for 1 month and analyzed.
Bears and wolves had a hierarchical
organization
similar
to that found in nature,
although
we found a relatively
low position
in the rank
order of blind bears.
Social interactions
between
wolves
appeared
to be very
serious at times. Bear-wolf
interactions were
mostly
playful,
but were sometimes
agonistic.
Young
bears
were
more
often victims
of wolf harassment than other bears and were sometimes
seriously
bitten.
However,
no
deaths
were
recorded,
contrary
to descriptions
from natural bear-wolf interactions. We concluded that bears and wolves could be kept safely together
in a single enclosure with apparently
few
consequences.
Interactions
we observed were comparable
to interactions between bears
and wolves under
natural conditions
and
concerned
mainly
competition
over food or a den (wolf or bear).
The size of the food (large
prey
carcasses in the wild vs. chicken
in the enclosure)
probably
plays a role
in the
severity
and the eventual solution of the bear-wolf
conflict.
Ursus 13:85-93 (2002)
Key words: bear-wolf interactions,
brown
bear,
Canis lupus,
enclosure,
Ursus arctos, wolf
Bear research in enclosures has advantages
and disad-
vantages
over bear research in natural conditions.
Disad-
vantages include the very small area, compared with
natural home ranges,
and the absence
of escape possibili-
ties. These factors can increase social stress even in ani-
mals kept in larger
enclosures.
Advantages
of relatively
large
enclosures lie in the
possibilities
of intensive
research
concerning
animal
behavior,
welfare,
and
health.
Bear behavior varies considerably
among individuals
(Fagen and
Fagen 1996, Koene 1998). This can be dem-
onstrated
to the public
in a bear
enclosure and
can be used
to illustrate the individual
distinctiveness
and
personality
of bears
(Fagen
and
Fagen 1996),
together
with their
cog-
nitive abilities. Thus, enclosures
are important
in public
education. Furthermore,
behavioral and veterinary
re-
search in enclosures can add to knowledge acquired
in
nature,
as for example, testing
radiotelemetry
equipment
in relation to activity.
Also, the
study
of behavioral
changes
relative to environmental
conditions
can be related
to the
welfare
of the animals;
for example,
a decrease
in stereo-
typic behavior or an increase in play behavior
may indi-
cate a better relationship between a bear and its
environment or increased well being (Koene 1998). In
this paper we present an analysis of social interactions
between brown
bears
and wolves, coexisting in the Bear
Forest enclosure in the Ouwehand Zoo in Rhenen, the
Netherlands.
We
addressed
the following questions
in this paper:
(1)
How are
bears
and
wolves socially organized
in the 'Bear
Forest' enclosure
of the Ouwehand
Zoo, in Rhenen, the
Netherlands?
(2) Do bears and wolves interact?
(3) How
does the behavior
of bears and wolves in the enclosure
compare
to free-ranging
animals? and (4) What are the
management
implications
for similar
situations in captiv-
ity?
STUDY AREA
Bear Forest, a refuge for bears, was created in the
Ouwehand
Zoo in Rhenen, the Netherlands,
in 1993 by
the International
Bear Foundation
(Koene 1998). The 2-
ha area is enclosed by a 3.5-m high double fence (3 m
apart)
electrified on top with a 1-m electric fence inside
the double fence. The area is bisected into northern
and
southern
sections
by a public
walkway
through
the enclo-
sure.
Tunnels
under
the
walkway
connect the
northern
and
southern
sections. The refuge houses former
zoo bears,
circus
bears,
Turkish
dancing
bears,
and
bears
from
Bosnia
relocated
because of the war.
In 1994, 3 wolves from
Bel-
gium and Germany
were moved to the Bear Forest in
Rhenen along with 7 wolves from the Ouwehand
Zoo.
Since then interspecific
and intraspecific
behavioral in-
teractions
among bears
and wolves have been noted, but
not systematically investigated.
86 Ursus 13:2002
Thirteen
European
brown bears
(6 male:7
female)
of
different
ages (range
3-23 years)
resided in
the enclosure
(Table
1).
Three of the
bears were blind due to previous
mistreatment. Years of birth were
known
for all bears ex-
cept
the blind
bears
and a veterinarian
estimated their
ap-
proximate age
in 1993.
We
designated
3 age
classes:
young
(0-4 years),
adult
(5-10 years)
and
old
(>10
years
of age).
Bears
were
individually
recognized
by
their
posture,
color,
and facial details.
In addition
to the
13
bears,
7 wolves
lived
in the
enclo-
sure
at
the time
of observation
(Table
1).
Each
wolf
was
distinguished by
facial and
body
characteristics.
All
male
animals
except
Cazar
were castrated and all
females
were
sterilized. The bears and
wolves
lived outdoors
all year
round;
the
bears had
unimpeded
access to 8 dens,
while
the wolves
dug
their own den.
Normally
the bears and
wolves were
fed 3 times
daily
at
irregular
hours.
During
the first
feeding,
only dog pellets
were
given.
The second
and third
feeding
consisted
mainly
of apples,
bread,
cab-
bage,
and carrots
for the bears and
rats, chicken,
ducks,
fishes,
or
offal
for
the
wolves.
There were
2 feeding places,
and the caretakers
distributed
the food
from
the
walkway.
METHODS
Bear-bear
Interactions
We observed
bears
at
feeding
times
during
13-21
May
1997.
Observations
began
0.5 hr before and
ended 1.5
hr
after
food
was offered.
We
(2 observers)
recorded
agonis-
tic encounters,
which we defined as any
situation
where
>2 bears
interacted
with each
other
in such a way as to
disrupt
their ongoing patterns
of moving or feeding
Table
1. Brown
bears and
wolves
residing
in
the Bear
Forest,
Ouwehand
Zoo, Rhenen,
the Netherlands,
for a behavioral
study
conducted
in 1997.
Species Name Label Sex Age(yr) Age class
Bear Battir Ba M 23 old
Geert Ge M 22 old
Koroglf Ko M >1Ob old
Mackenzie Mk M 7 adult
Bora Bo M 5-10b adult
Axel Ax M 4 young
Nelly Ne F 23 old
Fiona Fi F 5-10 adult
Mascha Ma F 8 adult
Niki Ni F 5 adult
Tory To F 4 young
Wolke Wo F 4 young
Bjorna Bj F 3 young
Wolf Cazar Ca M 7 adult
Traan Tr M 4 adult
Streep St M 4 adult
Poot Po M 4 adult
Brenda Br F 4 adult
Noeska No F 4 adult
Svlvia Sv F 4 adult
a Wolves were not categorized
by age class.
b Blind bear;
age estimated
by a veterinarian.
(Stonorov
and
Stokes
1972).
A bear
was labeled domi-
nant
(DOM)
when it
approached,
charged,
and
maintained
a frontal orientation to another
bear. A bear was
labeled
subordinate
(SUB)
when it ran or walked
away,
backed
up,
and had a lateral
orientation
to another bear. We
cal-
culated 2 measures of rank order. The
DS (dominance-
subordinate)
rank
order
was based
on reordering
the
dominance-subordinate
matrix
(Martin
and
Bateson
1993)
using
the MatMan
program
(De Vries
1994).
The
DV-
rank
order
was
based
on
the
individual
dominance
value
(DV;
Lehner
1996)
of each animal
(No. DOM encoun-
ters/total
[No.
DOM
+ No. SUB]
encounters).
Wolf-wolf
Interactions
We observed
wolves
during
3-28 February
1997.
We
normally
observed
from
0800-1700, Monday-Friday,
using
a Latin
square design.
Total observation time was
144
hrs. We
collected
data
by
behavior
sampling
(Martin
and
Bateson
1993)
on a portable
recorder
with
The
Ob-
server
software
(Noldus
1991).
We
distinguished
22 be-
havioral elements that
occurred
during
social interactions
based
on the wolf sociogram
from
Zimen
(1982).
Actor,
behavior,
and receiver
were
recorded on
tape.
Active sub-
mission
appeared
to be the best
indication
of rank order
according
to our
analysis
and Van
Hooff
and
Wensing
(1987).
Thus,
we based
rank
order
only on this behav-
ioral element.
Active submission
is a
behavioral
complex
in which the
actor
actively
seeks
contact
with a recipient
by
approaching
it in a crouched manner
with
curved back
and
bent
legs, the tail
curled
down,
often
wagging,
with
ears
folded back.
From this
position,
the animal tries to
contact
the
recipient
by licking
its nose
(Van
Hooff and
Wensing
1987,
Derix
1994).
When
constituent
elements
occurred
separately,
we recorded
them
as separate
cat-
egories.
We also
calculated
the same
2 measures
of rank
order
for
the wolves.
Bear-wolf
Interactions
We observed
bear-wolf
interactions
on 3-21 February
1997
around the
feeding
hours
(1000-1400),
when most
interactions
occurred
(Koene
1998).
We
videotaped
in-
teractions
when
(1) a bear
or wolf
approached
a member
of the other
species,
or (2) bears
or wolves
started
run-
ning
without an
apparent
obvious
trigger.
We recorded
the following
data
from
the videotaped
sequences:
date,
time,
and
context
(bear
den,
wolf den,
food,
unknown).
Concerning
the
approach,
we recorded
the
approacher
(bear
or
wolf);
name;
sex
and
age
of first,
second,
and
third wolf and
first,
second,
and
third
bear;
distance from which approach
started;
behavior
of
approacher
(approach,
attack,
bite, etc.); whether
the
approacher
stopped
or passed;
and distance
when
the
BEARS
AND
WOLVES IN CAPTIVITY
* Koene et al. 87
approacher
stopped
or passed the approached.
Concern-
ing the short
distance interaction,
we recorded
the num-
ber of bears and wolves participating
in the interaction;
actor (bear or wolf), behavior of the actor (same as
approacher
behavior);
recipient
(bear
or wolf); behavior
of the recipient
(bear,
wolf, unknown);
distance
between
the
participants
of the interaction;
and
loser
(bear
or
wolf).
We collected some data
from interactions in the wild for
the same variables and
compared
the enclosure
data
with
data from the wild. Data were analyzed with SAS 6.01
(1990).
RESULTS
Bear-bear
Interactions
As a rule,
agonistic
interactions
between
bears
occurred
as soon as the caretaker
offered food to the bears. Some
animals gained access to the center of the feeding spot
with
the highest
food concentration.
Others
had to wait or
were even forced to stay entirely away from the place
where
all the food was thrown. As bears
gradually
left the
feeding spot, the number
of encounters
decreased
rapidly
with time after feeding. Most encounters
were without
physical
contact
and
occurred as follows: a dominant
bear
would appear
and subordinate
bears would back up or
even run or walk away, turning
the head away from the
opponent;
ears of both bears were back. If the subordi-
nate
bear
did not back
away,
the dominant
would usually
charge
it. In a charge
the dominant
bear
ran,
frontally
ori-
ented,
directly
toward
the other
with
ears
back
and
mouth
slightly open. It was possible to distinguish
the dominant
and subordinate
bears in most encounters.
We recorded
419 agonistic encounters
between the bears and deter-
mined the DS-rank
order of the 13 bears
(Table 2).
The DS-rank order yielded the following dominance
hierarchy:
(1) old males were highest in rank,
followed
by an adult
male, (2) next an old female followed by an
adult
female, the oldest blind male, and the last adult
fe-
male, and (3) finally the younger male and females, the
old blind female, and
the adult
blind male.
Although
the hierarchy
did not differ
between feeding
spots,
the
amount
of agonistic
behavior
of some individual
bears
differed
significantly
for the different
feeding spots.
For example,
Geert,
highest
in DS-rank,
was involved in
more
aggressive
interactions
with other
bears
at the place
he occupied
most
than
at
other
feeding
places. Battir,
sec-
ond in DS-rank,
was less involved in aggressive
encoun-
ters in the home range
of Geert.
Not all bears were present at all feeding places. For
example, 1 adult blind male (Koroglfi) did not walk
through
the tunnels and did not eat when food was of-
fered at the other
side of the tunnel.
The old blind
female
sometimes
continued
her
stereotypic
behavior
and
did
not
eat.
Based on the dominance
value, a somewhat different
rank
order
was found
(Table
2). Four
groups
could be dis-
tinguished
based on the dominance-value:
(1) Geert
and
Battir (DV 0.97 and 0.92, respectively), (2) Koroglu
(blind), Mascha,
Nelly, and
Mackenzie
(DV = 0.50-0.69),
(3) Niki,
Bora,
and
Fiona
(both
blind)
and
Axel (DV=0.14-
0.41) and
(4) Wolke,
Bjora, and
Tory
(DV = 0.01-0.05).
Wolf-wolf
Interactions
The behavioral
element
associated
with most observa-
tions of the total (415 of 1,491) was active submission.
Brenda
was the only female that copulated
with Cazar;
she was the alpha female. Noeska was the most active
wolf in the pack,
with
46% of the total
submissive
behav-
iors,
of which 57%
were
directed
toward
Brenda
and
39%
Table 2. Rank
ordered dominant (DOM:
rows) - subordinate (SUB: columns) matrix
of 13 brown bears based upon agonistic
Interactions
won and lost over food for a 1997 behavioral
study in a 2-ha enclosure at Ouwehand
Zoo, Rhenen, the Netherlands.
A dash indicates that no encounters were observed between this pair of animals.
Label of
dominant Total DS- DV-
animal Ge Ba Mk Ne Ma Koa Ni Ax Fia Boa To Wo Bj DOM Age rank DV rank
Ge 3 14 4 9 1 22 1 - - 3 5 7 69 old 1 0.97 1
Ba 2 19 24 30 2 25 4 3 5 11 9 3 137 old 2 0.92 2
Mk 0 2 1 0 1 0 2 - 1 10 9 11 37 adult 3 0.50 6
Ne 0 1 1 2 1 - 2 - 3 5 10 11 36 old 4 0.55 5
Ma 0 4 1 0 - 2 8 1 3 17 14 16 66 adult 5 0.61 4
Koa 0 0 1 - - 0 2 - - 4 1 3 11 old 6 0.69 3
Ni 0 2 1 0 0 0 3 2 2 16 4 5 35 adult 7 0.41 7
Ax 0 0 0 -
0 0 0 2 0 - - 2 4 young 8 0.14 10
Fia - 0 - 0 0 - 1 0 1 - - - 2 old 9 0.20 9
Boa - 0 0 0 0 - 0 2 0 2 2 2 8 adult 10 0.33 8
To 0 0 0 0 0 0 0 - - 1 1 young 11 0.01 13
Wo 0 0 0 0 0 0 1 - - 0 - 2 3 young 12 0.05 11
Bj 0 0 0 0 1 0 0 0 0 0 1 young 13 0.02 12
Total SUB 2 12 37 29 42 5 51 24 8 16 68 54 62 419
a Blind bear
88 Ursus 13:2002
toward
Cazar.
But she was also very aggressive
to Sylvia.
The majority
of all dominant
behaviors were directed
to-
ward Sylvia. The dominance-subordinate
matrix based
on active submission
(Table 3) ranked
the wolves (domi-
nant to subordinate)
as Cazar,
Brenda, Poot, Traan,
Streep,
Noeska, and Sylvia.
We found a DV-rank order
(Table
3) different from the
DS-matrix
for the
males Poot and Traan. We
distinguished
3 groups based on the dominance-value:
(1) Cazar
and
Brenda
(DV = 1.00 and
0.96, respectively),
(2) Traan, Poot,
and
Streep
(DV= 0.19-0.43), (3) Noeska and
Sylvia (DV
0.07 and
0.00, respectively).
Bear-wolf
Interactions
We recorded 19.5 hours of video including 79 bear-
wolf interactions.
The context of these interactions
in the
enclosure were as follows: interactions
over food 19%,
near wolf den 8%,
near
bear den 5%,
and unknown 68%.
In 70%
of the interactions,
only 1 bear
was involved; 8%
involved 2 bears,
3% involved 3 bears,
and 1%,
4 bears.
In 57% of the interactions
only 1 wolf was involved; in
14%,
2 wolves; in 4%, 3 wolves; in 8%,
4 wolves; and in
18%
>4 wolves. In 28%,
the bears
were the actors
(initi-
ating the interaction),
and in 72% the wolves were the
actors.
Bears
retreated
in 23% of the encounters,
wolves
in 77% (Table
4).
When approaching
another
animal, bears and wolves
stopped
or passed
in the expected frequency
(%2
= 0.01, P
= 0.92). The animal that
approached
was usually
also the
actor in the interaction
(X2
= 63.94, P = 0.001). Wolves
initiated more interactions than bears (X2 = 15.51, P =
0.001), and wolves retreated more often than bears, as
expected (X2
= 0.41, P = 0.52).
Bear and wolf behavior
during
interactions
was rather
comparable
(Table 5). Bears and wolves did not differ
in
behavior elements as approacher
(X2
= 4.83, 9 df, P =
0.78) or as an actor
(X2
= 9.82, 9 df, P = 0.28). As recipi-
ent, however,
wolves avoided bears more often than
ex-
pected (X2
= 21.52, 9 df, P = 0.011).
Most bears that interacted
with wolves were
young
bears
(Table 6); Bjorna,
a 3-year
old female, accounted
for
47%
of all bear-wolf interactions. There
was a significant
re-
lationship
between DV-rank order and number of inter-
actions
(r = 0.60, P = 0.039, n = 13);
the relationship
was
stronger
in sighted
bears
(rs
= 0.81, P = 0.016, n = 10). In
most cases the wolf or wolves involved in the interac-
tions could not be identified
(87%).
Traan
(9%)
and Poot
(3%),
the subordinate
males, were involved in most iden-
tified (recognized
individuals)
interactions,
and Brenda,
the alpha
female, only once (1%).
Table
3. Rank
ordered
dominant
(DOM:
rows)
- subordinate
(SUB:
columns)
matrix
of 7 adult wolves
based
on the behavioral
element
active
submission
for a 1997
behavioral
study
in a 2-ha enclosure
at Ouwehand
Zoo,
Rhenen,
the Netherlands.
Label of dominant
animal Ca Br Po Tr St No Sy Total DOM Age DS-rank DV DV-rank
Ca 4 17 5 25 100 9 160 adult 1 1.00 1
Br 0 2 2 4 133 73 214 adult 2 0.96 2
Po 0 2 1 1 5 0 9 adult 3 0.32 4
Tr 0 2 0 0 2 2 6 adult 4 0.43 3
St 0 1 0 0 3 3 7 adult 5 0.19 5
No 0 0 0 0 0 19 19 adult 6 0.07 6
Sy 0 0 0 0 0 0 0 adult 7 0.00 7
Total
SUB 0 9 19 8 30 243 106 415
Table
4. Pathway
representation
of bear-wolf
interactions'
for a 1997
behavioral
study
of 13 brown
bears
and
7 wolves
in a 2-ha enclosure at Ouwehand
Zoo, Rhenen, the
Netherlands.
Approacher Pass or stop Actor Bear retreat Wolf retreat
Bear pass bear 2 5
Bear stop bear 2 10
Bear pass wolf 0 0
Bear stop wolf 0 0
Wolf pass bear 0 1
Wolf stop bear 0 2
Wolf pass wolf 9 11
Wolf stop wolf 5 32
Total 18 61
a Protocol
was started
when a subject
approached
a subject
of the other
species after
which the action
of the approacher
could be either
stops or
passes.
The actor
in the interaction
could be either
the approacher
or the
approached
animal. Frequency of withdrawing
by brown bears and
wolves is presented
dependent
on the type of approacher,
the action of
the approacher
near
the approached
(pass or stop), and the type of actor
in the interaction.
Table
5. Behavioral
elements shown by brown
bears and
wolves
as approacher
(app),
actor
(act),
and recipient
(rec),
expressed as percentage
of total number
of interactions.
Data
from
a 1997
behavioral
study
of 13 brown bears
and
7
wolves in a 2-ha
enclosure
at Ouwehand
Zoo, Rhenen,
the
Netherlands. Approacher Actor Recipient
Behavior bear wolf bear wolf bear wolf
Approach 63 70 41 39 0 5
Attack 16 7 14 16 4 5
Bite 5 7 5 11 0 5
Chase 0 2 14 2 0 0
Defend 0 0 0 0 33 23
Look at 11 5 9 14 23 9
Avoid 0 0 0 0 2 27
Flee 0 0 0 0 21 18
Rob food 0 2 0 4 0 0
Sniff 0 2 0 2 2 0
Threat 0 5 5 11 9 0
Threat
bite 5 2 14 4 7 9
Total number 19 60 22 57 57 22
of interactions
BEARS
AND WOLVES
IN CAPTIVITY
* Koene et al. 89
DISCUSSION
Bear-bear
Interactions
All bear species are primarily
solitary
in the wild, and
individuals avoid each other if possible (Brown 1993).
Bears
that
do associate
are
typically
a mother
and her
cubs,
siblings that recently left their mother,
or male and fe-
male during
the breeding
season (Brown 1993). The best
known example of a social grouping
is when bears
con-
gregate
temporarily
on places where good food is abun-
dant,
such as garbage
dumps
and salmon
(Oncorhynchus
spp.) streams (Stonorov and Stokes 1972, Egbert and
Stokes 1976). In this situation,
bears
develop a stable
so-
cial structure
to use resources as efficiently as possible
without
spending
too much
time
fighting.
Their
repertoire
of threatening
behaviors enables them to keep distance
without
showing overt aggression.
The Bear
Forest,
with
its high
food availability, compares
well to the brown
bears
in Alaska studied
by Stonorov
and Stokes (1972). Weber
(1988) reported
that 252 of 478 (52%)
bear-bear
encoun-
ters at natural
feeding places in Romania were aggres-
sive. Unfortunately,
no rank order
was presented.
Weber
(1988) found
the same
repertoire
in his bears as Stonorov
and Stokes (1972) found, except for the behavior
"jaw-
ing", which was not recorded
in the Romanian
bears.
We provoked
agonistic
interactions
among
the bears in
the Bear Forest
by offering all the food for a meal in one
place instead of scattered
around.
The dominance
hierar-
chy we found was based on the dominance-subordinate
matrix and on the dominance value of each individual.
Blind bears
were ranked
lower than
expected
on the basis
of their
age and
sex. However,
based on dominance-value,
the blind female Fiona
was the only one with a lower
rank
than expected based on her age and sex. Despite their
handicap,
the blind bears still dominated
younger bears
in the Bear Forest.
Literature
on social structure
in brown bears
revealed
the existence of temporary
hierarchies
in groups
of bears
on rich
feeding places (Stonorov
and Stokes 1972, Weber
1988). Large
adult
males were highest in rank,
followed
by females with cubs, adult
males, and
other
females. Fi-
nally young bears
deferred
to all the other bears
(Brown
1993). The hierarchy
found in the Bear Forest was very
similar to that in the wild under favorable food condi-
tions. Furthermore,
as in nature, agonistic interactions
without
physical contact
were used to maintain
this hier-
archy,
with strong
bear-bear
tolerance
(Colmenares
and
Rivero 1983).
Wolf-wolf
Interactions
Whereas in bears social grouping
is an exception, in
wolves it is expected.
Therefore,
dominance
relationships
are well established. It is nearly impossible to observe
social relationships
of a wolf pack in the wilderness,
be-
cause wolves have home ranges
between 50-1,000 km2.
As a result, nearly all studies of social relationships
of
wolves have been conducted on captive packs in large
enclosures.
The enclosure
in Zimen's
study
(1982) of wolf
pack sociogram
was 6 ha. Mech (1970) used Isle Royale
as a natural
enclosure with an area of 54,400 ha. Van
Hooff
and
Wensing
(1987) described
pack structure under
cap-
tive conditions
in a small
enclosure
(0.3 ha)
and concluded
that: (1) there are different
rank orders for females and
males, especially for the higher
ranking
members,
(2) the
sexual rank order
is mainly structured
according
to age,
with older animals
generally
dominant over younger
ani-
mals;
(3) the rank differences are
larger
in the
higher
ranks
and lower between the lower
ranking
wolves;
and
(4) there
is no cross-sex dominance relationship,
as long as the
wolves have the same rank
level in their sexual rank
or-
der;
if the rank
levels are different and there are signifi-
cant age differences, there are cross-sex dominance
Table 6. Brown bear-wolf interactions in relation to bear dominance in descending order, for a 1997 behavioral study of 13
brown bears and 7 wolves in a 2-ha enclosure at Ouwehand Zoo, Rhenen, the Netherlands.
Bear Sex Age DS-rank DV-rank IA (No.) IA (%)
Bj female young 13 12 39 49.4
Wo female young 12 11 4 5.0
To female young 11 13 4 5.0
Mk male adult 3 6 12 15.2
Ax male young 8 10 9 11.4
Ge male old 1 1 8 10.1
Ni female adult 7 7 1 1.3
Ma female adult 5 4 1 1.3
Ne female old 4 5 1 1.3
Ba male old 2 2 0 0
Koa male old 6 3 0 0
Boa male adult 10 8 0 0
Fia female old 9 9 0 0
79 100%
a Blind bear
90 Ursus 13:2002
relationships.
The wolf dominance
hierarchy
is thus
best
described
as a pyramid
with the alpha
male and
female at
the highest
position,
followed by the beta
male and
some-
times the beta
female, then
adult subordinate
females and
males.
In the Rhenen
wolf pack,
there were 4 especially active
wolves:
Cazar,
Brenda, Noeska,
and
Sylvia.
Cazar showed
no submissive
behavior,
much dominant
behavior,
and
was
apparently
the alpha male of the pack. He was also the
only intact male in the pack (with testicles and thus hor-
monal
activity).
Cazar was the only wolf observed
copu-
lating with a female (Brenda,
the alpha female). There
was a clear linear
hierarchy
within
the females
of the
pack:
Brenda,
Noeska, and
Sylvia. The relationships among
the
wolves were sometimes
very aggressive and led to inju-
ries, especially for Sylvia, the lowest in rank.
Only 6% of all interactions
occurred
in the non-alpha
males
Traan,
Streep,
and
Poot. It was difficult to rank these
wolves based
on a small number of interactions.
Poot and
Streep showed more submissive behaviors than Traan;
Traan
could be the beta male and the others the adult sub-
dominants. This agrees
with the dominance
rank from the
dominance-value
(DV). On the other
hand,
Landau's
in-
dex of linearity
(Lehner
1996) for the pack was 0.77, in-
dicating
no clear
linear
hierarchy
(a linearity
index of 0.9
is considered
as indicative
of a linear
hierarchy).
Comparison
of our
results with the hierarchy
of a free-
ranging
wolf pack is difficult. In the Rhenen wolf pack,
the social interactions
seemed to be more
aggressive
and
more concentrated
on only some individuals
than in a free-
ranging wolf pack. Mech (1999) emphasized
that free-
living wolf packs are
family groups
in which dominance
displays are uncommon except during competition for
food. He suggested
that
active submission
is primarily
a
food-begging
gesture
and not primarily
an indicator
of a
dominance
relationship.
At the beginning
of each breed-
ing season, Brenda, the alpha female, severely injured
Sylvia by biting
her in the back
near
the tail, causing deep
open wounds. In the wild, biting will not occur because
low ranked
females leave the group
before the breeding
season
(Mech 1999). In captivity
there
is no possibility
to
leave the area and to avoid the alpha
female. We specu-
late that
in the wild, Sylvia probably
would have left the
pack
to avoid
being
attacked
by Brenda
and Noeska.
Traan,
Streep,
and
Poot would have been more socially active if
they had retained
their
natural
hormonal
level. No sexu-
ally related behavior was observed in these 3 animals.
Although
the hierarchy
appeared
to be somewhat
normal
for wolves, it seemed to be restricted
by the non-intact
males and the restrictions
of the enclosure.
Bear-wolf
interactions
Magoun
(1976) observed
bears
and wolves near
a car-
cass simultaneously
in the wild, and during 173 5-min
periods she recorded 39 aggressive acts by the bears to-
ward
wolves and
only 1 aggressive
act
by a wolf toward
a
bear.
Unfortunately,
the behavior is not described
in de-
tail;
most aggressive
acts of bears involved a short
lunge
or swipe, although
sometimes a bear
chased a wolf. Murie
(1981) also described
wolves as the losers in bear-wolf
interactions,
but others observed
the opposite (see Mech
1970, Brown 1993). In Nelchina
Basin, brown bears con-
tested
wolves in 13.1% of 130 observations of wolves on
kills (Ballard
1982; Table
7).
A more recent
description
of interactions showed that
wolves attack bears fiercely, especially when bears are
near the wolves' den (Kehoe 1995). Wolves
mainly
initi-
ated the bear-wolf interactions in the Bear Forest.
Wolves
also retreated
the most from interactions and were there-
fore
the
probable
losers. The interactions were sometimes
related to fights concerning the bear den, wolf den, or
food. However,
most interactions
(68%) were still of an
unknown
origin.
Bears and
wolves showed approach
be-
havior as actors and
defensive or flee behavior as recipi-
ents.
The wolves did not often harass the blind bears.
Only
one video showed interactions
between Bora and wolves
and
Koroglu
and
wolves. The first
confrontation
between
the blind
bears
and
the wolves was described
earlier,
"all
blind bears were at least once attacked
by the group of
wolves, but the
bears did not react and
the attack
stopped"
(Koene 1998: 582). A possible explanation
is that
the in-
teractions
were mostly play motivated
and that the blind
bears
reacted
inadequately
to the wolf behavioral
initia-
tive.
Comparison
between the bear-wolf interactions
in the
enclosure and interactions
in the wild is rather
compli-
cated. Bears
do not
normally
conflict
with
wolves. Wolves
cannot win an aggressive encounter
with an adult bear.
Still wolves often kill bears, but only when the bear is
rather
young, ill, or trapped
in the den. In North
America,
cases of bears as victims of wolves have been reported,
but also the reverse
is found.
We made
an
attempt
to evalu-
ate bear-wolf interactions
as reported
in the available
lit-
erature
(Table
7).
Based on the data
collected in the Bear Forest
and the
data
extracted
from
the literature
(Table
8), we made
some
general
comparisons.
Contrary
to expectation,
interactions
in the Bear Forest
were more of unknown
origin than
in
the wild (x2=37.14, 4 df, P=0.001). Wolves and
bears
in
the Bear
Forest
began
interactions
in the same
proportion
as in the wild (X2= 1.31, P=0.25). Losers of the interac-
tions
in the wild were
wolves in fewer
cases than
expected
(x2=47.24, 3 df, P =0.001), but wolves were more
often
the losers in the Bear
Forest.
Conditions
of the Bear Forest
may explain these dis-
BEARS AND WOLVES IN
CAPTIVITY
* Koene et al. 91
Table
7. Literature
review
of bear-wolf
interactions
recorded
in the wild.
Behavior Loser
chase bear
kill wolf
attack unknown
attack unknown
attack bear
approach unknown
eat unknown
eat unknown
eat bear
kill wolf
attack unknown
kill bear
kill bear
kill wolf
attack wolf
attack wolf
attack wolf
attack none
attack none
attack wolf
attack bear
chase wolf
chase wolf
chase wolf
eat bear
chase wolf
chase bear
displace? wolf
chase? bear
displace unknown
kill bear
displace unknown
displace unknown
displace unknown
kill bear
eat bear
eat wolf
kill bear
kill bear
kill bear
approach unknown
kill wolf
attack unknown
attack unknown
attack unknown
approach bear+cubs
chase bear
attack wolf
stand unknown
chase unknown
chase unknown
attack bear
Bears
(No.) Wolves (No.)
1 5
1 5
4 5
4 5
1 5
3 2
1 1
1 1
1 5
~~1 4
1 5
1 5
1 5
? 1
1 5
4 5
4 5
1 1
1 1
3 1
1 12
1 1
>3 1
1 1
>3 9
4 2
4 2
1 1
2 3
1 4
1 2
1 1
1 3
1 4
? 1
3 2
1 4
1 10?
1 2
1 5
3 1
1 3
1 3
1 1
2 5
3 1
1 2
4 1
1 1
1 1
1 1
1 1
Bear
species
brown
brown
brown
brown
brown
brown
brown
brown
brown
black
black
black
brown
brown
brown
brown
brown
brown
brown
brown
brown
black
black
black
black
brown
brown
brown
brown
brown
brown
brown
brown
brown
black
polar
brown
black
black
black
brown
brown
black
black
brown
brown
brown
brown
brown
brown
brown
brown
Source
Mills 1919
Mills 1919
Murie 1944
Murie
1944
Murie 1944
Murie 1944
Lent 1964
Lent 1964
Pulliainen
1965
Joslin 1966
Rutter
and
Pimlot 1968
Rutter and
Pimlot 1968
trapper
(in Mech) 1970
Ballard
1980
Herning
(in Murie)
1981
Murie
1981
Murie 1981
Murie
1981
Murie 1981
Murie 1981
Murie
1981
Rogers
and
Mech 1981
Rogers
and
Mech 1981
Rogers
and Mech 1981
Rogers
and Mech 1981
Ballard
1982
Ballard
1982
Ballard 1982
Ballard
1982
Ballard
1982
Ballard 1982
Ballard
1982
Ballard 1982
Ballard 1982
Horejsi
et al. 1984
Ramsay
and
Stirling
1984
Hornbeck
and
Horejsi
1986
Paquet
and
Carbyn
1986
Paquet
and
Carbyn
1986
Paquet
and
Carbyn
1986
Hayes and
Mossop 1987
Hayes and Baer 1992
Gehring
1993
Veitch et al. 1993
Kehoe 1995
Koenea 1995
Follmannb 1997
Jamesb 1997
Magob
1997
Reynoldsb
1997
Reynoldsb
1997
Stephensonb
1997
a Koene, personal
observation.
During the excursion of the Tenth
IBA (International
Bear Association) conference
in Fairbanks,
Alaska, 1995, a
wolf coming from a long distance
encountered a grizzly bear
mother
with 2 cubs. The cubs were directed
higher up hill, while the mother stood on
her hind legs. The wolf stopped
and made a half circle around the bear and her cubs and continued
on its way. Immediately
thereafter,
the cubs ran
toward the mother
(as if they had been called) and the mother bear suckled them for a short while (as if to calm them). They then continued
their
foraging
behavior.
b Data from northwestern
Brooks Range near the Kokolik River, Alaska (H. Reynolds, Alaska Department
of Fish and Game, Fairbanks, Alaska,
USA, personal
communication,
1998).
crepancies. First, the food was provided and, although ity and
solution of the bear-wolf conflict. Second,
wolves
there was competition, food was abundant. The size of normally
outnumber bears
in interactions
in the wild. In
the food (chicken carcasses in contrast with large prey the Bear Forest
bear
density was exceptionally
high, re-
carcasses
in the wild) probably played a role in the sever- sulting
in a higher
potential
danger
for the wolves. Third,
Actor
wolf
bear
bear
wolf
wolf
bear
bear
wolf
wolf
bear
bear
wolf
wolf
bear
bear
bear
bear
bear
bear
bear
wolf
bear
bear
bear
wolf
bear
wolf
bear
wolf
wolf
wolf
wolf
wolf
wolf
wolf
wolf
bear
wolf
wolf
wolf
bear
bear
wolf
wolf
wolf
wolf
wolf
wolf
wolf
bear
wolf
wolf
92 Ursus 13:2002
Table
8. Comparison
of bear-wolf
interactions in the wild
(Table
7)
and
in
Bear Forest
for a 1997
behavioral
study
of 13
brown
bears
and 7 wolves in
a 2-ha
enclosure at Ouwehand
Zoo,
Rhenen,
the Netherlands.
Interaction Item Bear
Forest Wild
Context bear den 4 3
wolf
den 6 9
food 15 20
cubs 0 3
unknown 54 10
Actor bear 22 17
wolf 57 28
Loser bear 18 16
wolf 61 10
none 0 3
unknown 0 16
neither bears
nor wolves could leave the area.
Fourth,
no
reproducing
bears or wolves were kept so no interactions
concerning
cubs
could
be recorded.
Fifth,
bears and
wolves
in the enclosure were familiar with each other,
a situation
that
hardly
will occur under wild conditions.
This famil-
iarity implies that
social contacts
between individuals in-
clude more learned
(predictable
and
controllable)
aspects,
and hence implies less exploration
and emotional reac-
tions. Bears and wolves seemed to tolerate
each other
in
the Bear
Forest,
as sometimes
also occurs
in natural
situ-
ations (Lent 1964).
MANAGEMENT
IMPLICATIONS
During
the
research
no severely
injurious
bear-bear
and
bear-wolf interactions
were recorded.
However,
wolves
sometimes bit and injured
other wolves in attacks that
could be fatal. Recently,
Mech (1999) pointed to differ-
ences in social organization
of captive wolf packs and
natural
wolf packs.
The typical
natural
wolf pack
is a fam-
ily, a breeding pair
with 1-3 generations
of offspring.
This
may imply that a stable organized
wolf pack should be
setup
from the start
from a pair
that
develops stable
rela-
tions by breeding.
As long as there are no reproducing
animals
in the en-
closure, no severe problems are expected from having
bears
and wolves together.
However,
we had 2 concerns.
The distances
between bear
and wolf dens were unnatu-
rally small, and because young bears, wolves, and cubs
seem to induce bear-wolf interactions
in the wild, wolf
and bear cubs in enclosures may cause severe manage-
ment
problems.
Also, the feeding schedule should
be var-
ied to keep the animals
more active, so that
they do not
expect food every day at the same time.
To keep a wolf pack as natural
as possible, the males
should not be castrated
- sterilized if necessary - to
maintain
a natural
hormonal level within the pack.
Thus,
the pack will be more stable and activities will be more
equally
distributed
among
the wolves. Whether
the same
is true for bears is not clear.
If a group
of intact
bears and
intact wolves are
kept together, many
interactions should
be expected. It is probably
difficult - maybe even im-
possible - to keep intact animals in such an enclosure.
CONCLUSIONS
Brown bears in the Bear
Forest
developed
a social hier-
archy
based
upon
sex and
age. The blind
bears
occupied
a
lower rank than
expected based on their
age and sex. As
in a free-ranging pack, the rank
order
of the wolves was
sexually
related.
The linear
hierarchy
in the females was:
(1) alpha
female Brenda,
(2) beta female Noeska and (3)
lowest ranking
female Sylvia. The interactions
among
these females were very aggressive.
Bear-wolf interactions occurred
more
often
in the Bear
Forest than at first thought.
Interactions of an unknown
origin
were
more common
than
in the wild, and
may have
been play interactions between
bears and
wolves familiar
with each other.
However,
more
research
concerning spe-
cific behavior
sequences
is needed to elucidate
this ques-
tion.
Comparing
our results of interactions in the enclosure
to interactions
in the wild, we found that (1) the domi-
nance hierarchy
in bears is similar to that found in the
wild during
salmon
fishing, that
(2) the wolf hierarchy
is
most probably
similar
to that
found in the wild, and that
(3) bear-wolf interactions are
less severe in the enclosure
than
in the wild. In the enclosure wolves killed no bears
and bears killed no wolves. There seemed to be a high
tolerance
level, as was sometimes described for natural
interactions
between bears and wolves.
ACKNOWLEDGMENTS
Special
thanks are due to Ouwehand
Zoo, Rhenen,
and
the Netherlands.
Funding
for the project
and
travel
to the
Eleventh
International Conference
on Bear Research
and
Management
in Graz
in 1997 came
from the International
Bear Foundation,
The Netherlands.
Also thanks are due
to 2 referees
and
H. Reynolds who provided
information
about bear-wolf interactions
in the wild and stimulated
us to explore the comparison
of interactions
in the 2-ha
enclosure with interactions
between bears
and wolves in
the wild.
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Received: 4 September 1997.
Accepted: 7 June 2002.
Associate Editor: Kate Kendall.
... where y =individual bear, W =number of interactions where the individual displaced another bear (wins), T = number of interactions where neither bear was supplanted by the other (ties), and N = total number of interactions the individual was involved in (Lehner 1996, Chi 1999, Koene et al. 2002. I selected daily bear use, expressed as the number of minutes that any bear was present during a sampling session relative to the length of the ...
... where y =individual bear, W =number of interactions where the individual displaced another bear (wins), T =number of interactions where neither bear was supplanted by the other (ties), and N = total number of interactions involving the individual (Lehner 1996, Chi 1999, Koene et al. 2002. I ranked these OS giving OSR, where one is the most dominant individual corresponding to the highest OS. ...
Thesis
Full-text available
Wildlife-based ecotourism is rapidly increasing in popularity, especially when featuring large mammals in their natural environment. Researchers have questioned the sustainability of wildlife-based ecotourism because it may compromise the survival and reproduction of focal animals. I investigated the potential spatio-temporal effects of bear viewers on grizzly bears at a proposed bear viewing site along the Fishing Branch River, Yukon. Spatial river use of grizzly bears was largely explained by habituation status. Bears consumed 24% less salmon when viewers were present, posing serious energetic consequences if spatio-temporal compensation does not occur. Dominance status had no measurable effect on bears' fishing behaviour presumably because abundant salmon and few conspecifics minimized resource-driven competition. However, dominance status could influence feeding behaviour in years with reduced salmon abundance, which would compound viewer-induced reductions in fish consumption. I recommend further investigation into spatio-temporal compensatory behaviours of grizzly bears along the Fishing Branch River.
... Resource competition (i.e., resource guarding (RG)) is suggested to be part of the normal behavioural repertoire of animals, observed across a wide range of species such as lizards (Stamps, 1977), mice (Wolff and Cicirello, 1989), birds (Gowaty, 1993), and baboons (Henzi et al., 2003). Related behaviours have also been described in a number of canid species including wolves kept in captivity (Keone et al., 2002), free-ranging domestic dogs (Cafazzo et al., 2010;Mangalam and Singh, 2013), and domestic dogs kept as companion animals (described between dogs and humans only) item and consume the meal together; this cooperative hunting and feeding is observed to be relatively harmonious among group members (Mech, 1970). It has been suggested that members of the pack use body postures and expressions to confirm social bonds and cooperation, which aids in maintaining low levels of aggression between members of the group (Bradshaw and Nott, 1995). ...
... It has been suggested that members of the pack use body postures and expressions to confirm social bonds and cooperation, which aids in maintaining low levels of aggression between members of the group (Bradshaw and Nott, 1995). Interestingly, when wolves are placed in captivity or a limited spatial area and composition is artificially determined or managed by people, aggression and competition for resources are reported at increased frequencies in comparison to wild populations (Keone et al., 2002;Wilmers and Stahler, 2002). Further, when the wolves are in captivity, the likelihood and type of aggressive threat displays tend to be reliant on external factors, such as hunger level (Kappe, 1997). ...
Article
Resource guarding (RG) involves the use of specific behaviour patterns to control access to an item of potential “value” (as perceived by the dog), and can be expressed in the presence of various animals, including other dogs. The current study aimed to identify factors associated with RG patterns expressed around other dogs. Dog owners (n = 3068) were recruited through social media to answer questions regarding dog- and household-related factors, as well as their dog's current and past behaviour around resources in the presence of other dogs. Participants were screened for their ability to identify different forms of resource guarding from video, and were removed from the study if they incorrectly identified any of the videos provided. This resulted in a final sample of 2,207 participants (n = 3,589 dogs). Multiple multi-level logistic regression models were developed to determine the association between independent variables of interest and RG patterns (i.e., RG aggression, avoidance, and rapid ingestion) when in the presence of other dogs. Namely, dogs living in multi-dog households were more likely to display RG aggression, avoidance, and rapid ingestion (p < 0.01) compared to dogs that live without other dogs. Dogs with higher levels of impulsivity and fear were more likely to display RG aggression (p < 0.001). Neutered males (p < 0.01) were more likely to be RG aggressive compared to dogs of other sexes and neuter statuses. Teaching dogs to reliably “drop” items when requested was associated with a reduced likelihood of biting RG aggression (p <0.05). Distinct associative relationships between the patterns of RG in the presence of other dogs were identified. Dogs that express RG aggression were less likely to express RG avoidance or RG rapid ingestion; however, the latter two types were likely to co-occur, perhaps dependent on the type of resource involved. This suggests that dogs may be relatively more fixed in their action around items in the presence of dogs. However, a longitudinal study is important to determine the exact nature of relationships between RG patterns. The results identified in this study can be used as a basis for further investigation on factors influencing RG behaviour in the presence of other dogs.
... Recently, large bear enclosures (LBE) have become popular, demanding more from zookeepers and management, i.e., electric fencing, feeding, environmental and veterinary management. Some scientific research has been done in LBEs and is helpful in feeding and social management of bears [51][52][53]. The social interactions between brown bears have been studied [52] and may play an important role in the positive social network [53]. ...
... Some scientific research has been done in LBEs and is helpful in feeding and social management of bears [51][52][53]. The social interactions between brown bears have been studied [52] and may play an important role in the positive social network [53]. In large bear enclosures (LBE) many brown bears can be housed, and these supposedly solitary animals are forced to develop social contacts, especially in feeding situations. ...
Article
Full-text available
It may become advantageous to keep human-managed animals in the social network groups to which they have adapted. Data concerning the social networks of farm animal species and their ancestors are scarce but essential to establishing the importance of a natural social network for farmed animal species. Social Network Analysis (SNA) facilitates the characterization of social networking at group, subgroup and individual levels. SNA is currently used for modeling the social behavior and management of wild animals and social welfare of zoo animals. It has been recognized for use with farm animals but has yet to be applied for management purposes. Currently, the main focus is on cattle, because in large groups (poultry), recording of individuals is expensive and the existence of social networks is uncertain due to on-farm restrictions. However, in many cases, a stable social network might be important to individual animal fitness, survival and welfare. For instance, when laying hens are not too densely housed, simple networks may be established. We describe here small social networks in horses, brown bears, laying hens and veal calves to illustrate the importance of measuring social networks among animals managed by humans. Emphasis is placed on the automatic measurement of identity, location, nearest neighbors and nearest neighbor distance for management purposes. It is concluded that social networks are important to the welfare of human-managed animal species and that welfare management based on automatic recordings will become available in the near future.
... To our knowledge, this study is the first to document brown bears and wolves feeding simultaneously on a whale carcass. Previous studies have shown that brown bear-wolf interactions often are antagonistic or aggressive, particularly at kill sites (Koene et al. 2002, Ballard et al. 2003, Gunther and Smith 2004. Ballard et al. (2003) summarized 108 brown bear-wolf interactions outside of Yellowstone National Park and found that only 6% involved brown bears and wolves feeding at the same carcass, 7% involved brown bears and wolves in the same vicinity, 24% involved brown bears and wolves fighting and chasing each other, and 21% involved brown bears displacing wolves from kill sites. ...
... Milleret (2011) found that wolves stayed closer to their prey carcasses after brown bear den emergence than before brown bear emergence, possibly to defend their prey from the larger scavenger. Koene et al. (2002) hypothesized that carcass size has a role in bear-wolf conflict severity with conflict over larger carcasses being less severe. Our results from a large carcass support this assertion because we saw little evidence of aggression between species-no instances of wolves attempting to displace brown bears and few instances of brown bears displacing wolves. ...
Article
Full-text available
Brown bears (Ursus arctos) are generally solitary animals, although they are known to aggregate at concentrated food resources. Using remote cameras, we documented brown bears aggregating while scavenging a whale carcass from 19 May to 17 September 2010 in Glacier Bay National Park, Alaska, USA. Such aggregations have not been reported in Glacier Bay, a recently deglaciated fjord where bear food resources are dispersed and less diverse than in other regions of coastal Alaska. We documented multiple brown bears and wolves (Canis lupus) scavenging the carcass—at times, simultaneously. This study provided a rare opportunity to document brown bear– wolf interactions over several months associated with a large-magnitude resource event with little evidence of aggression between species. Our results suggest that the humpback whale (Megaptera novaeangliae) carcass provided a substantial food resource for brown bears and wolves in Glacier Bay, potentially influencing space use and interspecific interactions.
... Although brown bears have been recorded to kill and displace wolves at ungulate carcasses, wolves have also been documented displacing female brown bears and killing bear cubs (Gunther & Smith 2004;Jimenez et al. 2008). Abundant food sources appear to increase tolerance and reduce interspecific aggression (Koene et al. 2002;Lewis & Lafferty 2014; see also Chapter 3). ...
Chapter
Bears have fascinated people since ancient times. The relationship between bears and humans dates back thousands of years, during which time we have also competed with bears for shelter and food. In modern times, bears have come under pressure through encroachment on their habitats, climate change, and illegal trade in their body parts, including the Asian bear bile market. The IUCN lists six bears as vulnerable or endangered, and even the least concern species, such as the brown bear, are at risk of extirpation in certain countries. The poaching and international trade of these most threatened populations are prohibited, but still ongoing. Covering all bears species worldwide, this beautifully illustrated volume brings together the contributions of 200 international bear experts on the ecology, conservation status, and management of the Ursidae family. It reveals the fascinating long history of interactions between humans and bears and the threats affecting these charismatic species.
... Wolves and brown bears also exhibit interference competition with one another, although outcomes appear to depend on strength of participants (e.g., pack size, bear body mass) rather than fall out consistently between species (Koene et al., 2002;Gunther and Smith, 2004;Tallian et al., 2017). Although direct predation of wolf pups by brown bears has been observed (Hayes and Baer, 1992), the absence of wolves on islands with high densities of salmon-supported brown bears plausibly results from kleptoparasitism of wolf kills by brown bears in much the same way that kleptoparasitism by lions (Panthera leo) and hyenas (Crocuta crocuta) seems to negatively affect African wild dogs (Lycaon pictus) (Carbone et al., 2005;van der Meer et al., 2011). ...
Article
Full-text available
Apex predators play keystone roles in ecosystems through top-down control, but the effects of apex omnivores on ecosystems could be more varied because changes in the resource base alter their densities and reverberate through ecosystems in complex ways. In coastal temperate ecosystems throughout much of the Northern Hemisphere, anadromous salmon once supported abundant bear populations, but both taxa have declined or been extirpated from large parts of their former ranges with limited research on the consequences of diminished or absent interactions among species. Here we review the biogeography of bear-salmon interactions and the role of salmon-subsidized bears in (1) resource provisioning to plants and scavengers through the distribution of salmon carcasses, (2) competition among bears and other large carnivores, (3) predation of ungulate neonates, (4) seed dispersal, and (5) resource subsidies to rodents with seed-filled scats. In addition to our review of the literature, we present original data to demonstrate two community-level patterns that are currently unexplained. First, deer densities appear to be consistently higher on islands with abundant brown bears than adjacent islands with black bears and wolves, and moose calf survival is higher at low bear densities (<∼25 bears per 100 km²) but is constant across the vast majority of bear densities found in the wild (i.e., ∼>25 bears per 100 km²). Our review and empirical data highlight key knowledge gaps and research opportunities to understand the complex ecosystem effects related to bear-salmon interactions.
... Home ranges of females and their offspring can shift as a result of roadkill. It is has been documented that adult males will dominate over females and younger age classes (Miller et al. 2003, Koene et al. 2002. Due to the dominant male behavior, younger bears may be forced to move out of resident bear home ranges and therefore may be more susceptible to impact by roadkill. ...
... Weaker, smaller bodied or younger individuals might avoid on purpose the most suitable habitats in an attempt to reduce the possibility of meeting other individuals and especially dominant males during mating season (Steyaert et al., 2013), a behavior that could further be justified by the dominant hierarchical social system of the species (Clapham et al., 2012). This structure is highly reflected at places where food is abundant: in these sites bears develop a stable social structure to use resources as efficiently as possible, with dominance hierarchies being related to size and large adult males being highest in rank (Koene et al., 2002). ...
... Social behavior in which grizzly bears occurred in dense numbers in open habitat have been documented (Hornocker 1962, Craighead et al. 1969, and these studies revealed that sometimes a large percent of time was spent interacting with conspecifics. Other work that addressed grizzly bear associations includes observations of captive grizzly bears (Koene et al. 2002), incidental mating observations (Mundy and Flook 1964, Herrero and Hamer 1977, Clevenger et al. 1992, and inferred interactions based on VHF (very high frequency) telemetry locations (Wielgus andBunnell 1994, 1995;Mace and Waller 1997). When association is inferred from the closeness of telemetry points, this approach assumes that bears close together interact in some way and are not in the same area due to chance overlap of utilized areas. ...
Article
Full-text available
 We used Doncaster's test to differentiate home range overlap in range use from mutual attraction in grizzly bears (Ursus arctos) based on global positioning system (GPS) telemetry data. From a sample of 61 collared bears, 404 pairs of GPS locations placed 2 or more bears ≤500 m from each other at about the same time (within 3 hr). From these 404 pairs, 68 were significantly positive associations (mutual attraction) in which 65% were male–female (MF) and 35% were the same sex. Most MF associations involved adults. Male and female bears had associations with 1.8 and 1.2 partners/year, respectively. Associations between males occurred twice as often in the pre-berry season than in the berry season, whereas female–female (FF) associations occurred more frequently in the berry season. The length of same-sex associations was significantly shorter than MF associations. Fifty-one percent of MF pairs associated more than once within a single year. For MF associations, the mean distance between individuals was 152 m. Our findings suggest that grizzly bears can spend a considerable amount of time interacting with conspecifics and that behavioral interactions between grizzly bears are more complicated than we understand. Human activity that affects grizzly bear associations could disrupt social behavior and ultimately reproduction.
Article
We report the first photographic and genetic evidence of a brown bear (Ursus arctos) on Wrangel Island, Russia, located north of 71 in the Arctic Ocean. The sequenced control region (D-loop) of mitochondrial DNA obtained from hair of a sighted bear was indistinguishable from one of the most widespread haplotypes of the Eurasian brown bear. Molecular genetic analysis indicated that the bear was male. We photographed what may have been a second brown bear on a remote camera trap. It is unknown whether the bear(s) were transients or indicative of a range expansion associated with warming temperatures. Wrangel Island currently supports muskoxen (Ovibos moschatus), reindeer (Rangifer tarandus), and several top predators including a high density of polar bears (Ursus maritimus) during the summer and autumn. Thus, the presence of brown bears could lead to novel interspecies interactions with potentially cascading ecological effects.
Article
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Fourteen European brown bears (Ursus arctoś) were confiscated in Turkey in October 1993. Three bears-1 female and 2 males-were blind and showed mainly stereotypies and behavior toward others that was passive, aggressive, or both. These blind bears were transported to the Zoo in the Netherlands and placed in an enclosure with 8 resident wolves and 7 resident bears, all with normal vision. Problems were expected concerning the adaptation to the new environment and the interactions with other animals. Three ethological studies were done in 3 phases: (1) experimental behavioral study during release, (2) behavioral description in the stable phase following release, and (3) detailed description of play behavior of blind and other bears. The blind bears had very few contacts with the electric fencing. After such a contact the bears immediately returned to the quarantine facility and stayed there for a long period. The female bear sometimes showed stereotypies during the first 6 stages of the release, but they were not observed after stage 6. Bears played much of the time in phases 2 and 3. The decrease in stereotypies and the increase in playing behavior may reflect improved welfare of the blind bears. However, comparison between individual reactions of the bears suggested different coping styles, as is found in other species. The 2 male bears developed an active coping style with many playful interactions with other bears, whereas the female showed a more passive coping style.
Article
Full-text available
The Observer is a general-purpose software package for event recording and data analysis in behavioral research. It allows any IBM-type personal computer to serve as an event recorder. In addition, The Observer can generate dedicated event-recording programs for several types of non-IBM-compatible portable and hand-held computers and transfer files between the PC and such computers. The user specifies options through menus. The configuration can be either used directly for event recording on the PC or passed on to a program generator that creates a program to collect data on a hand-held computer. Observational data from either type of computer can be analyzed by the program. Event-recording configurations can be tailored to many different experimental designs. Keys can be designated as events, and modifiers can be used to indicate the limits of an event. The program allows grouping of events in classes and distinction between mutually exclusive versus nonexclusive events and duration events versus frequency events. Timing of events is accurate to 0.1 sec. An on-line electronic notepad permits notes to be made during an observation session. The program also includes on-line error correction. User comments as well as independent variables can be stored together with the observational data. During data analysis, the user can select the level of analysis and the type of output file. The Observer calculates frequency of occurrence and duration for classes of events, individual events, or combinations of events. For analysis of concurrence, one can select the number of nesting levels and the order of nesting. Output can be generated in the form of sorted event sequence files, text report files, and tabular ASCII files. The results can be exported to spreadsheet and statistical programs.
Article
The prevailing view of a wolf (Canis lupus) pack is that of a group of individuals ever vying for dominance but held in check by the 'alpha' pair, the alpha male and alpha female. Most research on the social dynamics of wolf packs, however, has been conducted on non-natural assortments of captive wolves. Here I describe the wolf-pack social order as it occurs in nature, discuss the alpha concept and social dominance and submission, and present data on the precise relationships among members in free-living packs, based on a literature review and 13 summers of observations of wolves on Ellesmere Island, Northwest Territories, Canada. I conclude that the typical wolf pack is a family, with the adult parents guiding the activities of the group in a division-of-labor system in which the female predominates primarily in such activities as pup care and defense and the male primarily during foraging and food-provisioning and the travels associated with them.
Article
Thesis (M.A.)--University of Toronto, 1966. Includes bibliographical references (leaves 96-98). Photocopy. n