Wolf (Canis lupus) feeding habits during the first eight years of its occurrence in Germany

Abstract

Due to the fact that the feeding habits of large carnivores are the main contentious point when they start resettling regions they were absent from for several decades, the diet composition of the wolves in Germany was analysed from the beginning of this process. Wolves in Germany primarily feed on wild ungulates, which make up more than 96% of their diet. The dominating prey species is the roe deer (55.3%), followed by red deer (20.8%) and wild boar (17.7%). The second important food category are the leporids (2.9% of Biomass), whereas livestock makes up only 0.6% of all biomass consumed. Wolves clearly prefer hunting on juvenile to adult red deer; roe deer are not selected after their age. We found seasonal differences in the diet composition with a higher amount of wild boar in spring and winter, when a high amount of juveniles and weakened animals, respectively, are available. In the first years of the study the percentage of red deer was much higher, and the percentage of roe deer therefore was lower than the following years. The amount of wild boar in the wolf diet fluctuated most in the first three years. Diet composition remained constant during the last five years. Wolves needed less than two generations for adapting to the new conditions in the cultivated landscape of eastern Germany.

Full text

Please
cite
this
article
in
press
as:
Wagner,
C.,
et
al.,
Wolf
(Canis
lupus)
feeding
habits
during
the
first
eight
years
of
its
occurrence
in
Germany.
Mammal.
Biol.
(2012),
doi:10.1016/j.mambio.2011.12.004
ARTICLE IN PRESS
G Model
MAMBIO-40505;
No.
of
Pages
8
Mammalian
Biology
xxx
(2012)
xxx–xxx
Contents
lists
available
at
SciVerse
ScienceDirect
Mammalian
Biology
j
ourna
l
ho
mepage:
www.elsevier.de/mambio
Original
Investigation
Wolf
(Canis
lupus)
feeding
habits
during
the
first
eight
years
of
its
occurrence
in
Germany
Carina
Wagnera,b,∗,
Maika
Holzapfela,
Gesa
Kluthc,
Ilka
Reinhardtc,
Hermann
Ansorgea
aSenckenberg
Museum
of
Natural
History
Görlitz,
PF
300154,
D-02806
Görlitz,
Germany
bTechnische
Universität
Dresden,
Institute
of
Forest
Botany
and
Forest
Zoology,
Pienner
Straße
7,
01737
Tharandt,
Germany
cLUPUS
Wildlife
Consulting,
Dorfstrasse
16.
Spreetal/OT
Spreewitz
D-02979,
Germany
a
r
t
i
c
l
e
i
n
f
o
Article
history:
Received
29
September
2011
Accepted
23
December
2011
Keywords:
Wolf
Canis
lupus
Germany
Diet
Adaptation
a
b
s
t
r
a
c
t
Due
to
the
fact
that
the
feeding
habits
of
large
carnivores
are
the
main
contentious
point
when
they
start
resettling
regions
they
were
absent
from
for
several
decades,
the
diet
composition
of
the
wolves
in
Germany
was
analysed
from
the
beginning
of
this
process.
Wolves
in
Germany
primarily
feed
on
wild
ungulates,
which
make
up
more
than
96%
of
their
diet.
The
dominating
prey
species
is
the
roe
deer
(55.3%),
followed
by
red
deer
(20.8%)
and
wild
boar
(17.7%).
The
second
important
food
category
are
the
leporids
(2.9%
of
Biomass),
whereas
livestock
makes
up
only
0.6%
of
all
biomass
consumed.
Wolves
clearly
prefer
hunting
on
juvenile
to
adult
red
deer;
roe
deer
are
not
selected
after
their
age.
We
found
seasonal
differences
in
the
diet
composition
with
a
higher
amount
of
wild
boar
in
spring
and
winter,
when
a
high
amount
of
juveniles
and
weakened
animals,
respectively,
are
available.
In
the
first
years
of
the
study
the
percentage
of
red
deer
was
much
higher,
and
the
percentage
of
roe
deer
therefore
was
lower
than
the
following
years.
The
amount
of
wild
boar
in
the
wolf
diet
fluctuated
most
in
the
first
three
years.
Diet
composition
remained
constant
during
the
last
five
years.
Wolves
needed
less
than
two
generations
for
adapting
to
the
new
conditions
in
the
cultivated
landscape
of
eastern
Germany.
©
2012
Published
by
Elsevier
GmbH
on
behalf
of
Deutsche
Gesellschaft
für
Säugetierkunde.
Introduction
Originally
widespread
across
the
northern
hemisphere,
the
wolf
was
extirpated
in
most
of
the
west
and
middle
European
coun-
tries
till
the
19th
century.
Mostly
the
fear
of
livestock
damages
and
mythologisation
of
the
wolf
as
pest
led
to
large-scale
persecution
of
this
predator
(Fritts,
1982;
Fritts
et
al.,
2003;
Butzeck
et
al.,
1988;
Boitani,
1995,
2003;
Mech,
1995).
Additional
extinction
of
wild
ungulates
in
some
regions
accelerated
this
process
(Fernández
and
de
Azua,
2010).
During
recent
decades
and
with
the
legal
protection
of
the
wolf
in
most
European
countries,
wolves
have
started
reset-
tling
regions
they
had
been
displaced
from
(Wabakken
et
al.,
2001;
Boitani,
2003;
Valière
et
al.,
2003;
Nowak
and
Mysłajek,
2006).
In
the
late
18th
century
the
wolf
was
eliminated
from
Germany
due
to
organized
persecution
(Butzeck
et
al.,
1988;
Ansorge
and
Schellenberg,
2007).
Since
then
single
wolves
immigrated,
rarely
but
regularly,
to
eastern
Germany,
but
none
succeeded
in
establish-
ing
a
new
population
until
they
were
placed
under
legal
protection
in
the
whole
of
Germany
in
1990.
It
took
ten
more
years
until
the
first
reproduction
of
wolves
could
be
recorded
in
the
Muskau
∗Corresponding
author
at:
Senckenberg
Museum
of
Natural
History
Görlitz,
PF
300154,
D-02806
Görlitz,
Germany.
Tel.:
+49
3581
4760
5701.
E-mail
address:
carina.wagner@senckenberg.de
(C.
Wagner).
heath
in
north
eastern
Saxony.
From
that
point
the
wolves
in
Germany
reared
pups
every
year
and
the
population
started
grow-
ing
(Ansorge
and
Schellenberg,
2007).
The
feeding
habits
of
the
wolves
as
large
carnivores
and
their
influence
on
wild
ungulate
populations
and
livestock
farming
are
at
the
center
of
the
tensions
between
man
and
wolves
(Kleiven
et
al.,
2004).
Particularly
in
regions
where
the
wolf
was
absent
for
more
than
one
human
generation,
people
have
to
relearn
accept-
ing
a
large
predator
in
their
neighbourhood
(Linnell
et
al.,
2001;
Williams
et
al.,
2002;
Gärtner
and
Hauptmann,
2005).
To
prevent
speculations
and
exaggerations
it
is
of
huge
importance
to
get
pre-
cise
information
about
the
diet
composition
of
the
returned
wolves
and
its
development
during
the
adaptation
to
their
new
environ-
ment,
as
basis
of
wolf
and
game
management.
The
diet
of
the
wolf
generally
depends
on
the
availability
of
potential
prey
species,
especially
large
wild
ungulates.
Studies
in
North
America
(Rogers
et
al.,
1980;
Hughard,
1993;
Messier
1994;
Kunkel
et
al.,
1999;
Peterson,
1999;
Nelson
and
Mech,
2000;
Arjo
et
al.,
2002,
a.o.)
and
Europe
(Meriggi
et
al.,
1991;
Okarma,
1997;
J˛
edrzejewski
et
al.,
2000;
Andersone
and
Ozolins,
2004;
Fejklova
et
al.,
2004;
Gazzola
et
al.,
2005;
Nowak
et
al.,
2005,
a.o.)
show,
that
wild
ungulates
are
the
main
prey
of
wolves
living
in
game-
rich
regions.
If
there
are
not
enough
wild
ungulates
available
and
other
food
resources
like
livestock
or
waste
are
frequent,
wolves
are
able
to
change
their
feeding
habits
towards
these
categories
1616-5047/$
–
see
front
matter
©
2012
Published
by
Elsevier
GmbH
on
behalf
of
Deutsche
Gesellschaft
für
Säugetierkunde.
doi:10.1016/j.mambio.2011.12.004

Please
cite
this
article
in
press
as:
Wagner,
C.,
et
al.,
Wolf
(Canis
lupus)
feeding
habits
during
the
first
eight
years
of
its
occurrence
in
Germany.
Mammal.
Biol.
(2012),
doi:10.1016/j.mambio.2011.12.004
ARTICLE IN PRESS
G Model
MAMBIO-40505;
No.
of
Pages
8
2
C.
Wagner
et
al.
/
Mammalian
Biology
xxx
(2012)
xxx–xxx
(Boitani,
1982;
Meriggi
et
al.,
1991;
Vos,
2000;
Peterson
and
Ciucci,
2003;
Hovens
and
Tungalaktuja,
2005).
With
a
seasonal
surplus
of
other
prey
like
salmon
in
costal
British
Columbia
(North
America)
(Darimont
et
al.,
2003)
they
can
adapt
to
using
quite
nontypical
food.
The
German
wolves
originated
from
Poland,
where
the
diet
of
the
predators
has
been
intensively
studied
from
the
lowlands
of
Bialowieza
primeval
forest
to
the
mountainous
regions
in
southern
and
southeastern
Poland
(J˛
edrzejewski
et
al.,
2000,
2002;
Nowak
et
al.,
2005; ´
Smietana,
2005).
There,
red
deer
Cervus
elaphus
is
the
main
prey
and
the
only
one
positively
selected
regarding
their
share
in
ungulate
community,
whereas
roe
deer
Capreolus
capreolus
and
wild
boar
Sus
scrofa
are
used
to
a
lesser
extent.
However,
wolves
in
western
Poland,
who
belong
to
the
same
wolf
population
as
German
wolves,
seem
to
hunt
red
deer
according
to
its
relative
abundance
(Nowak
et
al.,
2011).
As
wolves
are
known
to
adopt
their
feeding
preferences
from
their
parents
(Packard,
2003),
the
implication
would
be
that
wolves
in
Germany
show
a
comparable
pattern.
Otherwise,
the
process
of
adaptation
to
new
conditions
in
the
availability
of
prey
and
environmental
circumstances
could
lead
to
specialization
on
and
preference
for
other
prey
species
and
therefore
to
a
shift
in
the
feed-
ing
habits.
This
ability
to
adapt
on
new
conditions
makes
the
wolf
one
of
the
most
widespread
mammalian
species.
Following
this
approach
adaptation
to
the
new
environment
should
take
approx-
imately
one
wolf-generation,
two
years.
The
development
of
diet
composition
since
wolves
resettled
regions
they
have
been
displaced
from
before,
has
not
been
contin-
uously
studied
yet.
This
study
gives
new
insights
into
the
recovery
and
adaptation
strategies
of
wolves
and
would
be
very
helpful
in
regard
to
game
and
wolf
management
and
public
relation.
Study
area
The
study
area
of
about
2500
km2is
located
in
the
Lusatian
heath
in
north
eastern
Saxony
and
parts
of
southern
Brandenburg
in
Germany
and
covers
the
entire
area
occupied
by
wolves
in
Germany
during
the
examination
period
(Fig.
1).
Characterised
by
large
former
and
still
operating
opencast
coal
mines,
an
intensively
used
military
training
area
(145
km2)
and
pine
forest
monocultures,
the
region
is
under
strong
anthropogenic
influence.
But
compared
to
whole
Saxony
the
area
has
a
much
higher
amount
of
forest
cover
(52%;
Saxony:
26.8%)
and
open
areas
(6%;
Saxony:
0.6%)
than
average
and
a
lower
amount
of
settlement
and
traffic
area
(3%;
Saxony:
10.3%).
The
region
is
flat
(elevation:
120–170
m
asl.)
with
dry,
sandy
grounds
covered
by
pine
forests,
mixed
pine-oak
forest
and
open
or
scattered
heathland
including
larger
parts
of
the
biosphere
reserve
Upper
Lusatian
Heath
and
Pond
Landscape
in
the
south.
The
area
is
located
in
the
temperate
zone
with
a
semi-
continental
climate.
During
the
study
period
from
spring
2001
to
spring
2009
the
mean
annual
temperature
was
9.3 ◦C
and
the
mean
annual
precipitation
was
631.5
mm.
The
duration
of
snow
cover
differed
from
11
to
68
days
(mean
35.1
days)
per
winter.
Wolves
in
Germany
coexist
with
5
wild
ungulate
species;
two
of
them
(moufflon
Ovis
ammon
musimon
and
fallow
deer
Cervus
dama)
were
introduced
by
humans
as
game
species.
Their
share
of
the
ungulate
community
is
very
low
in
the
study
area;
moufflon
disappeared
from
the
main
areas
with
permanent
wolf
occurrence
until
2003.
With
a
mean
hunting
bag
of
1.0
animals
per
km2the
wild
boar
makes
up
the
largest
part
of
the
general
hunting
bag
in
the
area,
together
with
roe
deer
with
0.97,
followed
by
red
deer
with
a
mean
of
0.78
animals
per
km2.
The
hunting
bag
is
used
as
indication
for
the
development
of
ungulate
density,
because
no
useful
data
on
the
population
densities
of
these
ungulates
are
available.
By
establishing
in
this
area
the
wolves
recolonised
exactly
the
region
where
the
last
eastern
German
wolves
were
extirpated
in
the
18th
century.
After
the
first
reproduction
in
the
year
2000
in
the
Muskau
heath,
a
second
pack
established
in
2005,
hencefor-
ward
every
year
at
least
one
more
new
pack
could
be
confirmed.
In
the
year
2009
six
packs
and
one
territorial
pair
of
wolves
without
offspring
occupied
about
2500
km2.
Methods
Scat
collection
and
analysis
The
diet
analysis
was
conducted
using
wolf
scats,
which
were
collected
during
all
seasons
from
April
2001
till
March
2009,
by
walking
or
driving
transects
on
forest
roads
and
fire
belts.
General
characteristics
of
collectable
wolf
scats
are
a
high
amount
of
good
visible
hairs
and
bone
fragments
and
a
diameter
of
at
least
25
mm
(Weaver
and
Fritts,
1979;
Ciucci
et
al.,
1996).
Additionally,
there
is
no
sign
for
feral
dogs
in
the
study
area,
which
would
regularly
feed
on
game.
In
total
1890
scats
were
evaluated.
After
collection,
the
scats
were
frozen
until
further
analysis,
then
heated
to
free
them
from
pathogenic
organisms
like
parasites,
washed
through
a
sieve
with
1
mm
meshes
and
oven
dried
at
46 ◦C.
The
nondigested
parts
of
the
prey
items
like
bone
fragments
and
hairs
were
separated.
Hairs
were
identified
using
keys
of
Teerink
(1991)
and
Meyer
et
al.
(2002)
as
well
as
our
own
determination
key
and
reference
collections.
Cri-
teria
for
the
identification
of
hair
were
macroscopic
characteristics
like
hair
length,
colour
and
structure,
and
microscopic
features
like
the
structure
of
the
hair
medulla
and
cuticular
patterns.
Bone
frag-
ments,
teeth
and
claws
or
hooves
were
also
used
for
determining
scat
content.
For
the
differentiation
of
cervid
species
we
used
our
large
reference
collection
of
hairs
from
different
parts
of
the
animal
body
of
different
age,
sex
and
season.
Digestable
plant
material,
like
berries
and
other
fruits
were
regarded
as
food,
whereas
nondigested
plant
material
like
grass
or
pine-needles
were
not
regarded
as
food
components.
Neither
were
insects,
which
were
either
dung
or
carrion
beetles
or
parasites
of
the
prey
and
therefore
ingested
by
chance.
Age
determination
For
the
determination
of
the
age
of
the
wolf
prey
we
used
the
scat
analysis
and,
additionally,
the
analysis
of
wolf
kills
found
during
field
work.
If
suitable
bone
fragments,
teeth
and
hairs
of
the
prey
in
the
wolf
scats
were
used
to
determine
the
age
of
the
prey.
Regarding
the
analysis
of
the
scats
it
was
possible
to
distinguish
young
ungulates
to
the
age
of
three
months
from
adults.
Furthermore,
the
age
of
the
prey
animals
found
as
wolf
kills
during
field
work
(roe
deer
n
=
34,
red
deer
n
=
55)
was
determined
through
stage
of
dentition
and
classified
as
young
(up
to
one
year
old)
and
adult
(more
than
one
year
old).
Due
to
the
fact,
that
very
young
ungulates
are
consumed
completely
and
no
remains
can
be
found,
we
combined
data
from
scat
analysis
and
wolf
kills
to
estimate
the
percentage
of
young
animals
in
wolf
diet
(Pj)
for
the
main
prey
species
roe
deer
and
red
deer.
For
that
we
used
formula
(1):
Pj=
Pjp ×
Bas +
Bjs
Bt(1)
where
Pjp is
the
percentage
of
biomass
of
juveniles
older
than
three
months,
from
prey
remains,
Bas the
biomass
of
non
juveniles
cal-
culated
from
scat
analysis
[kg]
and
Bjs is
the
biomass
of
very
young
juveniles
from
scat
analysis
[kg]
and
Btis
the
total
biomass
of
these
species.

Please
cite
this
article
in
press
as:
Wagner,
C.,
et
al.,
Wolf
(Canis
lupus)
feeding
habits
during
the
first
eight
years
of
its
occurrence
in
Germany.
Mammal.
Biol.
(2012),
doi:10.1016/j.mambio.2011.12.004
ARTICLE IN PRESS
G Model
MAMBIO-40505;
No.
of
Pages
8
C.
Wagner
et
al.
/
Mammalian
Biology
xxx
(2012)
xxx–xxx
3
Fig.
1.
Location
of
the
study
area
in
Central
Europe.
Table
1
Average
usable
net
weight
of
the
main
prey
species,
weight
of
small
juveniles
(age
less
than
three
months)
in
brackets.
Net
weight
[kg]
Juveniles Adults
Roe
deer
(4)8
14
Red
deer (15)30
50
Wild
boar
(5)10
40
We
used
average
usable
prey
mass
as
specified
in
Table
1
for
translating
biomass
into
numbers
of
animals
killed.
Statistics
We
calculated
the
frequency
of
occurrence
as
well
as
the
per-
centage
of
biomass
consumed
referring
to
the
general,
seasonal
and
the
annual
diet
composition.
For
the
latter
we
used
the
hunting
year
running
from
first
of
April
to
the
end
of
March.
The
percentage
of
biomass
consumed
was
calculated
using
the
method
of
Goszczy ´
nski
(1974),
where
dry
mass
of
washed
scats
is
multiplied
by
coefficients
of
digestibility
(Table
2).
Table
2
Coefficients
of
digestibility
according
to
L,
Lockie
(1961);
G,
Goszczy ´
nski
(1974);
F,
Fairley
et
al.
(1987)
(cited
in
J˛
edrzejewska
and
Jedrzejewski,
1998);
A,
Ansorge
et
al.
(2006);
juv.
Juvenile.
Prey
category
Coefficient
of
digestibility
Adult
ungulates
118G
Capreolus
capreolus
juv.
50G,A
Sus
scrofa
juv.
50G,A
Livestock
118G
Medium
sized
mammals
50G
Small
mammals
23G
Birds 35G
Fish
25F
Fruits 14L
Furthermore
we
calculated
niche
breadth
B
(Levins,
1968)
and
standardized
niche
breadth
Ba(Hurlbert
1978,
cited
in
Hofmann,
1999)
B
=1
(p2
j)(2)
where
pjis
the
percentage
of
biomass
of
prey
taxa.
Ba=B
−
1
n
−
1(3)
where
n
is
the
number
of
prey
categories.
Furthermore
we
used
the
selectivity
index
D
of
Jacobs
(1974)
(formula
(4))
to
quantify
the
different
pattern
of
utilization
of
the
game
species
by
hunters
and
wolf
and
the
selection
of
juveniles
referring
to
the
age
structure
of
an
average
cervid
population:
D
=r
−
p
r
+
p
−
2rp (4)
where
r
means
the
fraction
of
a
prey
species
in
the
total
number
of
ungulates
killed
by
the
wolf,
and
p
is
the
contribution
of
this
species
in
the
hunting
bag/of
this
age
class
in
ungulate
community.
For
evaluating
the
difference
between
the
diet
composition
(Fre-
quency)
of
different
years,
packs
or
seasons
we
used
the
Chi
square
test.
Results
Diet
composition
In
total,
33
different
food
objects,
combined
to
8
food
categories
were
detected
in
the
scats
(Table
3).
The
most
dominant
category,
concerning
both,
frequency
(F
=
97.0%)
and
percentage
of
biomass
(B
=
96.2%),
are
wild
ungulates.
With
a
frequency
of
occurrence
of
56.2%
and
a
percentage
of
biomass
of
55.3%,
roe
deer
are
the
main
prey
of
the
wolves
in
Germany,
followed
by
red
deer
and
wild
boar,
with
a
biomass
percentage
of
20.8%
and
17.7%,
respectively.
Two
more
species
of
ungulates,
fallow
deer
and
moufflon,
are
rarely
found
in
the
wolf
scats,
as
they
are
in
ungulate
community
in
the
study
area.
The
majority
of
all
scats
contained
remains
of
only
one
food
object
(64%),
in
28%
of
all
faeces
two
different
food
objects
were

Please
cite
this
article
in
press
as:
Wagner,
C.,
et
al.,
Wolf
(Canis
lupus)
feeding
habits
during
the
first
eight
years
of
its
occurrence
in
Germany.
Mammal.
Biol.
(2012),
doi:10.1016/j.mambio.2011.12.004
ARTICLE IN PRESS
G Model
MAMBIO-40505;
No.
of
Pages
8
4
C.
Wagner
et
al.
/
Mammalian
Biology
xxx
(2012)
xxx–xxx
Table
3
Food
categories
and
diet
composition
of
wolves
in
a
eight
year
development
and
in
total
(calculated
after
Goszczy ´
nski,
1974);
+,
less
than
0.05%.
Percentage
of
biomass
01/02
02/03
03/04
04/05
05/06
06/07
07/08
08/09
Total
Capreolus
capreolus
36.0
49.9
40.2
48.7
63.8
53.7
53.0
50.8
55.3
Cervus
elaphus
34.9
39.3
19.6
28.0
19.4
25.1
23.2
26.4
20.8
Sus
scrofa 19.2
8.9
36.1
19.4
11.1
12.6
17.1
15.2
17.7
Ovis
ammon
musimon 8.6
0.3
0.7
1.4
0.9
Cervus
dama 0.3
1.1
2.3
3.5
1.5
Artiodactyla
98.7
98.1
95.9
96.6
94.4
93.1
97.0
95.9
96.2
Leporidae
1.3
1.7
3.8
2.9
4.1
4.9
2.5
3.9
2.9
Nyctereutes
procyonoides
0.1
0.1
+
Vulpes
vulpes 0.1
+
Mustela
erminea ++
Ondatra
zibethicus 0.4
0.1
Medium
sized
mammals
0.1
0.4
0.1
0.1
Apodemus
sylvaticus +
+
Apodemus
spec.
+
+
Arvicola
terrestris
+
+
+
+
Clethrionomys
glareolus
+
+
+
Microtus
agrestris
+
+
+
Microtus
arvalis
+
+
+
Microtus
spec. + 0.1
+ 0.1
0.2
0.1
+
0.1
Rattus
norvegicus
+
+
Erinaceus
europaeus 0.1
+
Small
mammals
indet
+
+
+
Small
mammals
+
0.2
+
0.3
0.2
0.1
+
0.1
Felis
sylvestris
f.
catus
0.2
+
Gallus
gallus
f.
domestica
+
+
+
+
Ovis
ammon
f.
aries 0.2
0.5
1.1
0.2
0.1
0.4
Oryctolagus
cuniculus
f.
domestica
0.5
0.2
0.1
0.1
Domesticated
animals 0.2
+
0.5
0.7
1.3
0.2
0.2
0.6
Aves
+
+
+
+
+
+
+
Rubus
fruticosus
0.1
+
+
+
Malus
domestica 0.2
+++
Zea
mays
+
+
+
+
Prunus
cerasus +
+
Pyrus
communis
+
+
+
Fruits
0.1
0.3
0.1
+
0.1
Pisces
+
+
+
detected
and
only
8%
of
all
samples
consisted
of
more
than
two
different
items
(up
to
four).
With
a
percentage
of
biomass
of
less
than
5%,
leporids
are
by
far
the
second
most
important
food
category.
Remains
of
domes-
ticated
animals
were
found
in
1.4%
of
all
scats,
making
up
0.6%
of
the
biomass
consumed.
Among
the
domesticated
animals,
the
domestic
sheep
dominated
with
a
proportion
of
74%
of
this
cate-
gory,
followed
by
rabbit
(17%)
and
one
type
of
domestic
cat
(8%).
Barn
fowl
appears
occasionally,
but
makes
only
1%
of
the
biomass
in
this
food
category.
Two
percent
of
all
faeces
contained
fruit,
such
as
apple
(Malus
domestica)
and
pear
(Pyrus
sp.)
which
appeared
mainly
in
autumn
and
winter
when
they
are
used
for
attracting
game
to
feeding
sites.
In
summer,
blackberry
(Rubus
fruticosus)
and
cherry
(Prunus
sp.)
could
be
determined
in
some
wolf
scats.
Other
items
like
small
mammals
(several
species
of
Muridae
and
Arvicolidae),
medium
sized
mammals,
birds
and
fish
were
found
in
the
wolves’
diet
too,
but
with
a
percentage
of
biomass
less
than
0.2%,
so
their
proportion
was
very
low.
Anthropogenic
waste
did
not
play
any
role
in
the
diet
of
wolves
in
Saxony.
Juvenile
ungulates
in
wolf
diet
Regarding
the
percentage
of
biomass,
calves
make
up
to
49.5%
of
all
red
deer
consumed,
while
just
15.4%
of
roe
deer
biomass
is
made
up
by
fawns.
Assessing
a
healthy
and
average
cervid
pop-
ulation
with
a
growth
rate
of
25%
in
red
deer
and
30%
in
roe
deer
(Niethammer
and
Krapp,
1986),
red
deer
calves
are
clearly
positively
selected
(D
=
0.75),
whereas
roe
deer
fawns
are
chosen
according
to
their
share
in
ungulate
community
(D
=
0.0).
Every
third
roe
deer
killed
is
juvenile,
whereas
about
70%
of
all
red
deer
killed
by
the
wolves
are
less
than
one
year
old
(Table
4,
Fig.
2).
The
percentage
of
very
young
wild
boar
in
the
wolf
diet
is
even
higher
than
in
the
red
deer
(Table
4),
suggesting
that
the
per-
centage
of
all
juveniles
is
even
higher
for
wild
boar,
too.
But
due
to
the
fact
that
we
do
not
have
enough
data
on
wild
boar
wolf-
kills,
the
real
percentage
of
juvenile
boar
in
the
wolf
diet
remains
unknown.
Fig.
2.
Percentage
of
juveniles
in
the
two
main
prey
species
red
deer
and
roe
deer
calculated
as
percentage
of
animals
killed.

Please
cite
this
article
in
press
as:
Wagner,
C.,
et
al.,
Wolf
(Canis
lupus)
feeding
habits
during
the
first
eight
years
of
its
occurrence
in
Germany.
Mammal.
Biol.
(2012),
doi:10.1016/j.mambio.2011.12.004
ARTICLE IN PRESS
G Model
MAMBIO-40505;
No.
of
Pages
8
C.
Wagner
et
al.
/
Mammalian
Biology
xxx
(2012)
xxx–xxx
5
Table
4
Percentages
of
biomass
and
number
of
individuals
from
scat
analysis
(n
=
1384)
and
prey
remains
found
during
wolf
monitoring
(Roe
deer:
n
=
42,
Red
deer:
n
=
47);
n.d.,
no
data.
Biomass
[%] Number
[%]
Scat
analysis
Prey
remains
Combined
Scat
analysis
Prey
remains
Combined
Capreolus
capreolus Juvenile
5.7
5.6
15.4
16.9
9.3
30.4
Adult 94.3
94.4 84.6
83.1
90.6
69.6
Cervus
elaphus Juvenile
21.9
39.4
49.5
32.4
47.0
68.2
Adult
78.1
60.6
50.5
67.6
53.0
31.8
Sus
scrofa Juvenile
15.3
n.d.
49.6
n.d.
Adult 84.7
n.d. 50.4
n.d.
Development
of
diet
composition
Wild
ungulates
are
the
main
prey
of
the
wolves
during
the
whole
time
period
2001–2009,
amounting
to
at
least
93.1%
of
the
diet.
The
percentage
of
roe
deer
in
the
wolf
diet
is
increasing
in
the
first
years
of
the
examination
period
and
then
it
remains
constant
with
a
peak
in
the
hunting
year
05/06.
The
role
of
red
deer
however
is
decreasing
after
two
years
of
high
proportion,
and
the
percentage
of
wild
boar
is
fluctuating
throughout
the
years
without
any
obvi-
ous
trend.
Other
food
categories
did
not
show
any
trend
during
the
eight
year
development
(Fig.
3,
Table
3).
Medium-sized
mammals,
small
mammals,
fish
and
birds
as
well
as
fruits
are
supposed
to
be
fed
on
by
occasion
and
are
not
actively
searched
for
by
the
wolves.
Only
the
frequency
of
prey
species
in
the
first
year
01/02
shows
significant
differences
to
the
others
(p
=
0.031).
The
frequency
of
roe
deer
in
the
diet
was
much
less
than
in
the
following
years
and
the
amount
of
wild
boar
and
red
deer
was
comparably
high.
Further-
more
mouflon
was
quite
an
important
prey,
which
became
much
less
important
in
the
following
years.
The
niche
breadth
was
the
highest
in
the
first
year
of
the
study
(2001/02),
with
B
=
3.3
(Ba=
0.6)
and
decreases
to
an
index
level
between
B
=
2.2
(Ba=
0.1)
and
B
=
2.8
(Ba=
0.2)
(Table
5).
Livestock
in
the
diet
of
the
wolves
did
occur
in
seven
out
of
eight
years
with
a
peak
in
the
year
06/07
(1.3%
of
biomass
consumed)
but
there
no
trend
could
be
confirmed.
Sheep
as
the
main
domestic
prey
species
peaked
in
the
year
06/07
too,
with
1.1%
of
the
biomass
consumed.
Utilization
of
the
game
species
by
hunters
and
wolf
The
composition
of
the
wolf
diet
in
relation
to
the
percentages
of
the
same
species
in
the
hunting
bag
shows
the
differences
in
the
utilization
of
the
ungulate
game
species.
Whereas
hunters
shot
nearly
the
same
amount
of
roe
deer
and
wild
boar
and
only
a
few
less
red
deer,
wolf
diet
is
more
based
on
roe
deer,
being
the
main
Fig.
3.
Eight
year
development
of
diet
composition.
Table
5
Niche
breadth
(Levins,
1968)
and
standardized
niche
breadth
(Hofmann,
1999)
over
an
eight
year
development.
Hunting
year
B
Ba
01/02
3.3
0.6
02/03 2.4
0.3
03/04
3
0.3
04/05 2.8
0.2
05/06
2.2
0.1
06/07
2.7
0.2
07/08
2.7
0.2
08/09
2.8
0.2
Total
2.6
0.1
prey.
As
the
hunting
bag
depends
on
the
different
reproduction
rates
and
lots
of
other
parameters,
it
is
just
a
weak
indication
of
the
real
ungulate
density.
But
this
high
index
value
(Fig.
4)
indicates,
that
roe
deer
might
be
positively
selected
by
the
wolves,
whereas
red
deer
and
wild
boar
are
not.
During
the
eight
year
development
this
pattern
remains
stable,
with
a
positive
trend
of
the
roe
deer
index
value.
Seasonal
differences
in
the
wolf
diet
For
eliminating
errors
based
on
differences
between
the
years
we
used
the
data
of
only
one
year
(08/09)
who
are
corroborating
the
data
of
the
whole
study
period:
We
found
significant
differ-
ences
between
the
average
and
the
diet
composition
of
spring
(p
=
0.027)
and
winter
(p
=
0.045).
The
main
difference
between
these
seasons
is
that
the
amount
of
wild
boar
is
higher
and
the
percentage
of
deer
is
lower
in
spring
and
winter
(Fig.
5).
Further-
more
the
niche
breadth
is
the
highest
in
spring
(B
=
3.1;
Ba=
0.35)
and
winter
(B
=
3.2;
Ba=
0.31)
too,
meaning
that
the
wolf
diet
was
Fig.
4.
Comparison
between
the
utilization
of
the
three
main
prey
species
by
hunters
and
wolf;
positive
values
mean
a
higher
percentage
of
the
prey
species
in
the
wolf
diet,
negative
values
a
higher
percentage
in
the
hunting
bag.

Please
cite
this
article
in
press
as:
Wagner,
C.,
et
al.,
Wolf
(Canis
lupus)
feeding
habits
during
the
first
eight
years
of
its
occurrence
in
Germany.
Mammal.
Biol.
(2012),
doi:10.1016/j.mambio.2011.12.004
ARTICLE IN PRESS
G Model
MAMBIO-40505;
No.
of
Pages
8
6
C.
Wagner
et
al.
/
Mammalian
Biology
xxx
(2012)
xxx–xxx
Fig.
5.
Diet
composition
in
different
seasons
in
the
year
08/09.
more
diverse
in
these
seasons
than
in
summer
(B
=
2.6;
Ba=
0.23)
and
autumn
(B
=
2.5;
Ba=
0.19).
Discussion
Diet
composition
The
diet
of
the
wolves
in
Germany
is
dominated
by
wild
ungu-
lates,
as
it
is
characteristic
for
wolves
living
in
game
rich
regions.
Studies
in
eastern
and
central
Europe
agree
that
wolves
hunt
for
wild
or
domestic
ungulates
(i.e.
J˛
edrzejewski
et
al.,
2000;
Kübarsepp
and
Valdmann,
2003;
Sidorovich
et
al.,
2003;
Andersone
and
Ozolins,
2004;
Gazzola
et
al.,
2005;
Nowak
et
al.,
2005;
Hovens
and
Tungalaktuja,
2005)
depending
on
game
abundance
(Meriggi
et
al.,
1991;
Mattioli
et
al.,
1995;
Sidorovich
et
al.,
2003).
But
unlike
our
results
from
German
wolves,
who
primarily
hunt
on
roe
deer,
most
of
the
other
studies
confirmed
the
wolf
hunt
preferably
on
the
largest
ungulate
species
in
high
abundance,
available
in
the
region.
J˛
edrzejewski
et
al.
(2000)
and
Nowak
et
al.
(2005)
report
that
wolves
in
Bialowieza
primeval
forest
and
the
Beskid
moun-
tains
in
Poland,
where
the
composition
of
the
ungulate
community
is
comparable
to
that
in
our
study
area,
clearly
prefer
hunting
on
red
deer.
On
the
other
hand
in
western
Poland
roe
deer
is
the
most
consumed
prey
and
red
deer
is
obviously
not
preferred
by
the
wolf
(Nowak
et
al.,
2011).
The
percentage
of
red
deer
in
the
ungulate
community
is
given
with
38.5%
in
Bialowieza
(J˛
edrzejewski
et
al.,
2000),
21%
in
the
Beskid
mountains
(Nowak
et
al.,
2005)
and
in
western
Poland
22.2%
(Nowak
et
al.,
2011)
respectively.
As
we
do
not
have
comparable
data
about
the
real
density
of
red
deer
in
Lusa-
tia,
we
can
only
use
the
hunting
bag,
where
21%
are
red
deer,
and
41%
are
roe
deer.
According
to
Okarma
(1995)
roe
deer
is
the
main
prey
of
wolves
in
Europe
when
it
is
very
frequent,
and
larger
cervids
like
red
deer
or
reindeer
are
rare.
Nonetheless,
Bunewich
(1988)
found
wolves
in
Belarus
preying
preferentially
on
roe
deer.
He
refers
to
the
smaller
pack
sizes
due
to
legal
hunting
of
the
wolf
in
Belarus
to
explain
the
preference
of
roe
deer
in
presence
of
high
numbers
of
red
deer.
As
wolves
in
Germany
are
a
strictly
protected
species,
packs
are
normal
in
size
(about
eight
in
annual
mean),
so
this
should
not
be
the
reason
for
the
preference
of
roe
deer
in
this
case.
The
roe
deer
is
one
of
the
two
most
common
cervid
game
species
in
the
study
area
and
occurs
all
throughout
the
country
extensively.
In
the
whole
of
Saxony,
the
yearly
hunting
bag
of
the
roe
deer
doubled
since
1990.
Typical
habitats
of
roe
deer
are
widely
distributed
in
the
wolf
region:
edges
of
woods
with
dense
undergrowth
and
access
to
field,
grassland
or
scrub.
Thus
roe
deer
is
the
prey
species
which
wolves
encounter
most
frequently
during
their
ramble.
Furthermore,
the
smaller
deer
species
is
of
a
suitable
prey
size
with
low
risk
for
the
wolf.
Nevertheless,
the
anti-predator
behavior
of
roe
deer
such
as
vigilance
and
barking
(Reby
et
al.,
1999)
impedes
the
wolf
hunting
success.
Quoting
a
recent
study
from
Scandinavia,
wolves
run
only
47%
successful
attacks
on
roe
deer,
but
none
of
the
roe
deer
escaped
after
being
injured
by
a
wolf
(Wikenros
et
al.,
2009).
The
seasonal
differences
in
the
diet
result
from
a
high
avail-
ability
of
young
wild
boar
in
spring
and
more
weakened
boar
in
winter.
Particularly
a
higher
amount
of
wild
boar
in
the
wolf
diet
in
spring
was
reported
from
other
regions
(Meriggi
et
al.,
1991;
Okarma,
1995;
J˛
edrzejewski
et
al.,
2000)
and
results
from
a
posi-
tive
selection
of
juveniles
due
to
the
potential
risk
in
killing
a
well
fortified
adult
wild
boar.
Potential
for
conflict
Livestock
makes
up
only
a
very
small
part
of
the
diet
of
wolves
in
eastern
Germany.
This
fact
is
based
on
very
efficient
livestock
pro-
tection
methods
like
fencing
and
livestock
guarding
dogs,
which
are
financially
supported.
In
the
flat
regions,
flocks
are
fenced
behind
90
cm
high
electrical
mash
for
keeping
the
livestock
and
protection
against
wild
boar
and
dogs,
so
a
basic
wolf
protection
is
quite
common.
Several
authors
(Meriggi
et
al.,
1991;
Mattioli
et
al.,
1995;
Sidorovich
et
al.,
2003;
Nowak
et
al.,
2005
and
oth-
ers)
prove
that
damage
to
livestock
by
wolves
mainly
depends
on
the
quality
of
livestock
protection
methods
(Okarma,
1995;
Nowak
and
Mysłajek,
2004)
and
the
availability
of
wild
ungulates
(Okarma,
1995;
Capitani
et
al.,
2004;
Nowak
et
al.,
2005).
The
wildlife
stock
in
the
study
area
is
high,
so
that
the
wolves
do
not
need
to
prey
on
live-
stock
and
thereby
take
the
risk
of
a
confrontation
with
shepherds,
guarding
dogs
or
fences.
Even
during
the
eight
year
study
period,
where
the
wolf
pop-
ulation
was
growing
from
one
to
seven
packs
within
Lusatia,
wolf
attacks
on
domestic
animals
never
exceeded
22
per
year
and
the
last
two
years
of
the
examination
period,
damage
declined.
According
to
J˛
edrzejewski
et
al.
(2000)
and
Nowak
et
al.
(2005)
the
potential
for
conflicts
in
Germany
is
comparably
low.
Juvenile
ungulates
in
the
wolf
diet
Lots
of
studies
prove
that
juveniles,
females,
old
animals
and
those
with
bad
condition,
especially
of
the
larger
prey
species,
are
used
by
wolves
above
average
(Mattioli
et
al.,
1995;
Okarma,
1995;
J˛
edrzejewski
et
al.,
1992,
2000,
2002;
Gula,
2004;
Gazzola
et
al.,
2005;
Nowak
et
al.,
2005,
and
others).
This
corresponds
well
to
our
first
results
combining
the
analysis
of
the
wolfkills
and
scat
analy-
sis,
where
juvenile
red
deer
are
clearly
preferred,
whereas
neither
the
female
roe
deer
nor
the
fawns
are
preferred
by
the
wolf.
In
the
opposite
a
study
from
Italy
shows
preference
of
young
individuals
within
the
roe
deer
prey
(Mattioli
et
al.,
2004).
The
percentage
of
very
young
juvenile
wild
boar
in
the
wolf
scats
indicates
that
juve-
nile
wild
boars
are
even
more
positively
selected
by
the
wolves
than
young
red
deer,
as
observed
in
other
studies
(J˛
edrzejewski
et
al.,
2000,
2002;
Capitani
et
al.,
2004;
Nowak
et
al.,
2005).
But
without
data
from
wild
boar
prey
remains
we
cannot
give
an
imperative
statement
on
the
percentage
of
juvenile
wild
boar
in
the
wolf
diet.
Development
of
diet
composition
In
the
eight
year
development
of
the
food
composition
the
per-
centage
of
roe
deer
shows
an
upward
trend
during
the
first
years
without
any
indication
of
growing
roe
deer
density
in
the
region.
On
the
other
hand
the
proportion
of
red
deer
in
the
wolf
diet
declines
to
a
lower
level
after
two
years,
while
the
amount
of
wild
boar
is
fluctuating.
Especially
the
percentage
of
wild
boar
in
the
eight
year
development
can
be
explained
by
the
different
availability
of
this
prey
species
because
of
changing
density
and
availability
of
juveniles
due
to
weather
conditions
and
acorn
crop.