Adaptive Diet Strategy of the Wolf (Canis lupus L.) in Europe: a Review

Abstract

The diet strategy of the wolf in Europe is reviewed on the basis of 74 basic and 14 additional literature
sources. The comparative analysis reveals clear dependence on the latitude (and, therefore, on the changing
environmental conditions) correlated with the wild ungulate abundance and diversity. Following a
geographic pattern, the wolf is specialised on different species of ungulates: moose and reindeer in Scandinavia,
red deer in Central and Eastern Europe and wild boar in Southern Europe. Where this large prey
is taken, the roe deer is hunted with almost the same frequency in every region. The wolf diet in Europe
shows two ecological adaptations formed by a complex of variables: 1. Wolves living in natural habitats
with abundance of wild ungulates feed mainly on wild prey. 2. In highly anthropogenic habitats, with low
abundance of wild prey, wolves feed on livestock (where husbandry of domestic animals is available) and
take also a lot of plant food, smaller prey (hares and rodents) and garbage food. The frequency of occurrence
of wild ungulates in the diet of wolves in North Europe varies from 54.0% in Belarus to 132.7% in
Poland, while that of livestock is in the range from 0.4% in Norway to 74.9% in Belarus. In South Europe,
the frequency of occurrence of wild prey varies from 0% in Italy and Spain to 136.0% in Italy, while of domestic
ungulates ranges between 0% and 100% in Spain. The low density or lack of wild prey triggers the
switch of the wolf diet to livestock, plant food (32.2-85% in Italy) or even garbage (up to 41.5% in Italy).

Full text

439
ACTA ZOOLOGICA BULGARICA
Acta zool. bulg., 66 (4), 2014: 439-452
Invited Review
Introduction
The wolf (Canis lupus L.) has always been a com-
petitor with humans for the wild prey as well as a
conict species, which attacks domestic animals. It
is also one of the wild species, which spreads dis-
eases and parasites, such as rabies, tapeworms and
others. Therefore, the strong opposition and ght
against this species through the centuries with all
available means was inevitable. As a result of this,
the wolf was exterminated in Great Britain in the 14th
century and in many countries in Western Europe as
early as the 18th century (Ay b e s , yA l d e n 1995). This
could happen also in the Balkans in the 20th cen-
tury but due to the relatively less disturbed nature
and the high adaptability of the species to the con-
stantly changing environmental conditions, the wolf
managed to survive till the time when its important
role in nature was properly assessed. The rst as-
sessments of the wolf’s role in the ecosystems were
published in the 1960s by Pi m l o t (1967) and me c h
(1970). These assessements were further developed
by Fi l o n o v , KA l e t s K A y A (1985), Ru K o v s K y (1985),
several Russian authors in An o n y m o u s (1986),
bo i t A n i (1996), me c h , bo i t A n i (2003), Fe R R A R i
(2012) and others. Many studies on the wolf diet
have been conducted since the 1950s, aiming to as-
sess the wolf impact on nature, especially in coun-
tries as Italy, where the wolf numbers were reduced
to about 100 in the 1970s and afterwards increased.
Adaptive Diet Strategy of the Wolf (Canis lupus L.)
in Europe: a Review
Diana Zlatanova1, Atidje Ahmed2, Albena Valasseva2, Peter Genov2
1Department of Zoology and Anthropology, Faculty of Biology, Soa University ”St. Kliment Ohridski”, 1164 Soa, Bulgaria;
E-mail: zlite2@mail.bg
2 Institute of Biodiversity and Ecosystem Research, Bulgarian Academy of Sciences, 2 Gagarin Street, 1113 Soa;
E-mail: genov_bg@yahoo.it
Abstract: The diet strategy of the wolf in Europe is reviewed on the basis of 74 basic and 14 additional literature
sources. The comparative analysis reveals clear dependence on the latitude (and, therefore, on the chang-
ing environmental conditions) correlated with the wild ungulate abundance and diversity. Following a
geographic pattern, the wolf is specialised on different species of ungulates: moose and reindeer in Scan-
dinavia, red deer in Central and Eastern Europe and wild boar in Southern Europe. Where this large prey
is taken, the roe deer is hunted with almost the same frequency in every region. The wolf diet in Europe
shows two ecological adaptations formed by a complex of variables: 1. Wolves living in natural habitats
with abundance of wild ungulates feed mainly on wild prey. 2. In highly anthropogenic habitats, with low
abundance of wild prey, wolves feed on livestock (where husbandry of domestic animals is available) and
take also a lot of plant food, smaller prey (hares and rodents) and garbage food. The frequency of occur-
rence of wild ungulates in the diet of wolves in North Europe varies from 54.0% in Belarus to 132.7% in
Poland, while that of livestock is in the range from 0.4% in Norway to 74.9% in Belarus. In South Europe,
the frequency of occurrence of wild prey varies from 0% in Italy and Spain to 136.0% in Italy, while of do-
mestic ungulates ranges between 0% and 100% in Spain. The low density or lack of wild prey triggers the
switch of the wolf diet to livestock, plant food (32.2-85% in Italy) or even garbage (up to 41.5% in Italy).
Keywords: Wolf, Canis lupus, prey, adaptive strategy

440
Zlatanova D., A. Ahmed, A. Valasseva, P. Genov
Several papers reviewed the wolf preybase prefer-
ences in Europe (oK A R m A 1995, me R i g g i , lo v A R i
1996, mA R s i l i 2007, me R i g g i et al. 2011) but these
articles usually omitted data and analyses published
in Cyrillic alphabet and in grey literature (papers,
study reports, theses).
In the present paper, we assess the diet strategy
of the wolf in Europe based on a review of the latitu-
dinal and londitudinal gradients of the prey and the
real wolf preferences in relation to multiple factors.
We also try to summarise the wolf hunting strategies
by using commonly accessible published sources
and by including some grey literature, which is gen-
erally inaccessible to the West-European authors.
Material and Methods
We reviewed 74 scientic papers (Appendix 1) on
the wolf preybase in Europe, published in the pe-
riod 1953-2010 and originating from various coun-
tries (Fig. 1). These papers are written in 7 languag-
es: 32 in English, 13 in Italian, 17 in Russian, 8 in
Bulgarian, one in German, one in Portuguese and
one in Ukrainian. These are 44 papers published in
scientic journals, 8 in books, 7 reported at confer-
ences or symposia, 3 PhD theses, 6 MSc theses and 5
project reports. In addition, 19 further sources were
used to clarify the wolf adaptability.
The reviewed papers include diverse methods
of wolf diet assessment, the scat analyses being most
frequently used, followed by the analyses of the
stomach content and prey remains. The papers also
report different indices which describe the utilisation
and selectivity of the food components.
Two main methods used for the scat analyses are:
1) Diet diversity following the method of ci u c c i et al.
(1996); 2) Diet volume percentage over dry weight.
The stomach content analyses are rarely used,
especially if the species is not hunted because dead
specimens are difcult to retrieve. Several papers
deal with data on prey remains, as a result of snow
tracking during winter or tracking based on telem-
etry study. The prey animals are sexed, and the age
and physical condition of the prey are identied.
The main statistical analyses in the reviewed
papers follow the standard procedures of lo c K i e
(1959), namely: frequency of occurrence (Fi%) of
the different types of food (1); relative frequency of
occurrence (rFi%) (2); mean volume (Vі) of the food
remains in %. (3)
Fig. 1. Map of the reviewed literature sources study areas

Adaptive Diet Strategy of the Wolf (Canis lupus l.) in Europe: a Review
441
(1) Fi%=ni/N.100
(2) rFi%= Fi/Fn
(3) Vі%=∑vі/N
where: Fі% – frequency of occurrence of one
type of food; nі – number of samples containing the
particular type of food; i – type of food; N – number
of all samples; rFi% – relative frequency of occur-
rence; Fn – overall frequency of occurrence of the
particular type of food; Vі% – volume of the particu-
lar food; Vі – volume of the different types of food
In the cases when more than one type of food
is found in the samples, the percentage of food is
dened according to the 7-scale method of KR u u K
(1989).
This standardised approach allows for a com-
parison of the data between the different studies,
published in the papers. Some of the papers also deal
with the seasonal importance of the different prey
base. However, these analyses are not consistent
though all the studies and are not taken into consider-
ation in the current paper. For the aims of the current
review we consider mainly the frequency of occur-
rence (Fi%) of the different types of food as a basic
comparison tool for the wolf preferences in the dif-
ferent parts of Europe. The diverse methods of wolf
diet assessment do not allow for a sound statistical
cross country/ wolf study approach. Therefore, we
mainly consider the type of preybase and frequency
of occurrence ranges as a basis for comparison.
Results
The rst systematic analyses of the current wolf
distribution and basic wild preybase (ungulates) in
Europe were made by Pe t e R s (1993) and oK A R m A
(1995). Both authors state that the main wolf preybase
in Europe consists of 8 species: reindeer (Rangifer
tarandus), moose (Alces alces), European bison
(Bison bonasus), red deer (Cervus elaphus), roe deer
(Capreolus capreolus), wild boar (Sus scrofa), fal-
low deer (Dama dama) and saiga (Saiga tatarica).
Additionally, several other species play an impor-
tant role for the wolf in some regions, owing to their
isolated distribution but high abundance locally: the
chamois (Rupicapra rupicapra), the Pyrenean cham-
ois (Rupicapra pyrenaica), the Alpine ibex (Capra
ibex), the Caucasian tur (Capra caucasica), the
Iberian ibex (Capra pyrenaica), the Common ibex
(Capra aegagrus), and some introduced species such
as the mouon (Ovis orientalis musimon) (ge n o v ,
pers. comm). The mentioned species do not play
a vital role for the wolf but they may constitute an
important food source in case of deciency of other
preys in certain periods. The most widely distributed
and abundant prey species, which are the main prey
for the wolf in many places, are the following three
species: the red deer, the roe deer and the wild boar.
According to its distribution from the Polar cir-
cle to 40˚ N latitude, the wolf feeds mainly on the
ungulates, which are most abundant in the area. In
the most northern parts near the Polar circle up to
50˚ N latitude, the main prey is the reindeer, which
in many areas is semi-domesticated (Ko j o l A et. al.
2004), followed by the moose. In South Sweden, the
preybase is enriched by added values of the roe deer
and wild boar. Between 60˚ and 50˚ N, the reindeer
is replaced by the red deer, which is the most abun-
dant and signicant food source. Although the food
base in Bialowieza Primary Forest is very rich (there
are 5 prey species, i.e. red deer, moose, European
bison, wild boar and roe deer), the red deer has been
selected by the wolf (je d R z e j e j e w s K i et al. 2000).
The main prey of the wolf from the Polar circle
to 50˚ N latitude is the wild ungulates, Fi% of which
ranges from 40.3% to 100% of all the food sources
and constitute from 78.8 to 99.9% of the volume of
the food taken (Vi%) (Table 1).
The low frequency of occurrence of the wild
ungulates (Fi%=40.3%) and the high frequency
of domestic ungulates (Fi%=31.7%) presented by
gA v R i n , do n A u R o v (1954) for the Belorussian part
of Bialowieza at that time results from the high wolf
density (due to less cull), on the one hand, and the
low wild ungulate density due to poaching and mis-
management, on the other hand.
The wolf diet between 50˚ and 40˚ N latitude
is more complicated. There are three wolf prey spe-
cies, which represent different shares: the red deer,
roe deer and wild boar. In the semi-desert regions of
the lower Volga River, the main food source for the
wolf is the saiga antelope. Further southwards, the
more important is the share of domestic ungulates
and other animals. The main wolf prey species there
are the wild boar and the roe deer, with a lesser share
of the red deer. Some other species appear as an al-
ternative prey, such as the fallow deer, chamois and
the mouon (Table 2).
There are several areas, in which the wild boar
is a prevailing prey. In Italy, in the Casentino for-
est, the species constitutes 52.5% of the samples and
45.6% of the volume (mA t t i o l i et al. 1995). In the
Western Rhodope Mountains, Bulgaria, the wild boar
occurs in 38.5 % (Fi) of the wolf diet (se R A F i m o v et.
al. 2009), being prefered to the roe deer.
In this part of Europe the frequency of occur-
rence and share of the domestic ungulates is rising.
In some region as the Northern Caucasus the live-
stock Fi is 94.5% of all samples (bi b i K o v et al. 1985),

442
Zlatanova D., A. Ahmed, A. Valasseva, P. Genov
while in the Northeastern Portugal 76.8% of Fi in the
scats (Vi = 84.7%) is due to domestic ungulates.
The synanthropisation of the wolf in that lati-
tude is not only based on the take of livestock but
also on the usage of garbage dumps. According to
bo i t A n i (1996), the analysis of 220 scat samples in
Italy reveal that the Fi in the food of garbage dumps
is 33.4% (Vi= 44.1%). The consumption of fruits is
increased as in the Northern Italy the dog rose (Rosa
canina) constitutes 31.5% of all the food (me R i g g i
et al. 1991), while Pe z z o et al. (2003) report that the
fruits of hawthorn (Crataegus monogyna) are the
most frequent plant food in the wolf diet. In Bulgaria,
in the lowland, the wolf is also feeding on grapes and
sweet corn (ge n o v , pers. comm).
The wolf in Italy is more anthropogenically in-
uenced in the lowlands. The results of mA c d o n A l d
et al. (1980) in the Majella National Park show a
higher take of plant food (Fi = 64.7%), domestic un-
gulates (Fi = 41.0%), garbage dumps (Fi = 14.2%),
and others (Fi = 37.8%). Almost the same results
are obtained by RA g n i et al. (1985) during a study
in Umbria, where the Fi of domestic ungulates is
extremely high (71.0%) owing to the lack of wild
ungulates. In Abruzzo NP, the Fi of wild ungulates
(38.0%) is almost equal to the Fi of domestic ungu-
lates (34.5%), with a high share of plant food (32.8%)
and garbage (12.0%) (PA t A l A n o , lo v A R i 1993). A
recent review of the wolf diet in Italy (me R i g g i et al.
2011) reported signicant variable trends in the fre-
quency of occurrence of the wild boars, roe deer, red
deer, and the chamois in the wolf diet over time. The
authors discovered signicant and positive relation-
ships between the ungulate abundance and the ungu-
late presence in the wolf diet only for wild boars and
roe deer. These two species are pointed out as the
most important prey for the wolf in Italy.
The frequency of occurrence of domestic ungu-
lates, plant food and garbage is high in other countries
as well. In Spain, Fi of wild ungulates is 2/3 less than
that of the livestock (52.3%); the plant food and gar-
bage are also frequent food (8.5% to 41.5%, respec-
tively) (sA l v A d o R , Ab A d 1987). In Greece, the ratio
wild / domestic ungulates is 1:8 (Fi domestic ungu-
lates = 64.3%) and the plant food consumption is ex-
tremely high (Fi = 57.1%) (PA P A g e o R g i u et al. 1994).
Between the 1980s and 1990s, there was a clear
dependence of the wolf in the Southern Europe on
Table 1. Frequency of occurrence (%) of different types of food in the wolf diet in Northern Europe above 50˚ N lati-
tude: 1. Wild ungulates; 2. Domestic ungulates; 3. Plant food; 4. Others
Country Region 1 2 3 4 Source
Scandinavia South-central 102.0* 0.4 -40.0 ol s s o n et al. (1997)
Sweden 91.1 - - 48.9 mü l l e R (2006)
Finland East-central Karelia 72.0 3.0 2.0 24.5 gA d e -jø R g e n s e n , st A t e g A A R d (2000)
Kainuu 75.0 - - 35.8 hu i t u (2000)
German Sassonia 110.5 - - 9.8 An s o R g e et al. (2006)
Poland
Bieszczady 79.9 - - 29.1 su m i n s K i , Fi l i P i A K (1977)
Bialowieza 132.7 0.6 30.5 26.6 je d R z e j e j e w s K i et al.(2000)
Western Beskids 99.4 4.4 57.2 24.5 no w A K et al. (2005)
Estonia Alam-Pedja Reserve 79.6 5.0 3.0 23.1 Kü b A R s e P P , vA l d m A n n (2003)
Latvia Latvia 75.0 13.0 6.5 33.3 An d e R s o n e , oz o l i n s (2004)
Belarus
Polesija 46.9 25.2agA t A h (1979)
Bialowieża 40.3 31.7 1.1 27.9 gA v R i n , do n A u R o v (1954)
90.2 7.9 2.4 10.7 Bu n e v i č (1988)
North-eastern part 94.7 10.5 3.4 27.2 si d o R o v i c h et al. (2003)
Russia
Voronezh state reserve 18.6 10.1 -71.3bme R t z (1953)
89.5 2.9 3.3 16.9cli K h A t c K y et al. (1995)
Arkhangelsk, Onega
peninsula 24.4 - 5.5 78.7dRu K o v s K i , Ku P R i y A n o v et. al. (1972)
Verhnevolzie 68.8 11.3 5.2 28.9 Ko č e t K o v, so K o l o v (1979),
Pskovski region 60.5 10.4 -36.2 Ru s s A K o v (1979)
62.1 12.1 7.7 33.5 Ru s s A K o v , ti m o F e e v A (1984)
*The values above 100% are due to the fact that in a single sample (scat or stomach) there is more than one type of food
a dog; b hare (Lepus sp.) 20.4%, dog 19.1% and beaver 14.0%; c beaver 3.0%; d mountain hare (Lepus timidus) and
muskrat (Ondatra zibethicus).

Adaptive Diet Strategy of the Wolf (Canis lupus l.) in Europe: a Review
443
Table 2. Frequency of occurrence (%) of the different types of food in the region between 40˚ and 50˚ N latitude (South
Europe): 1. Wild ungulates; 2. Domestic ungulates; 3. Plant food; 4. Garbage food; 5. Others
Country Region 1 2 3 4 5 Source
Portugal North-East 8.6 76.8 - - 14.8 Ro q u e et al. (2001)
Spain
León 35.3 52.3 85.0 41.5 14.3 sA l v A d o R , Ab A d (1987)
Galicia - 80.0 10.0 - 38.3 cu e s t A et al. (1991)
Cantabria 82.0 10.0 1.3 - 18.0 cu e s t A et al. (1991)
Douro Meseta 107.7 3.8 6.3 2.5 106.0 cu e s t A et al. (1991)
Demanda mountains 62.0 57.0 - - 5.0 cu e s t A et al. (1991)
Sierra Morena 100.0 - - - - cu e s t A et al. (1991)
France Alpi Marittime 80.0 18.0 - - 3.0 Po u l l e et al. (1997)
Italy
Abruzo NP 38.0 34.5 32.8 12.0 52.3 PA t A l A n o , lo v A R i (1993)
Majella NP -41.0 64.7 14.2 37.8 mA c d o n A l d et al. (1980)
Umbria -71.0 - - 29.0 RA g n i et al. (1985)
Forli 107.7 3.8 6.3 2.5 6.3 mA t t i o l i et al. (1995)
Arezzo – Foreste Casen-
tinesi- FC 105.0 5.0 - - 5.0 mA t t i o l i et al. (1995),
(2004), gA z z o l A (2000);
Av A n z i n e l l i (2001)
Arezo – lto Mugello
(SAF) 110.0 1.0 - - 5.0
Arezzo – Vallesanta (VS) 107.0 7.0 - - 5.0 gA z z o l A (2000); Av A n z i n e l l i
(2001); gi u s t i n i (2002)
Arezo – Pratomagno (PM) 103.0 1.0 - - 6.0 cA P i t A n i et al. (2004);
mA t t i o l i et al. (2004)
Arezo – Alpe della Luna
-Valtiberina 102.0 6.0 - - 5.0 mA t t i o l i et al. (2004)
Arezo – Alpe di Catenaia 122.0 1.0 - - 6.0 Al b o n i 2004, lA m b e R t i
2004; co l o m b o 2005
Genova 17.2 22.9 64.9 9.5 76.3 me R i g g i et al. (1996)
La Spezia 36.0 56.3 42.2 -21.9
Val di Susa- Alpi Cozie 86.4 6.7 1.7 - 4.5 gA z z o l A et al. (2005)
Val di Cecina 110.9 9.3 - - 9.3 cA P i t A n i et al. (2004)
Val di Susa 91.4 5.7 - - 3.0 cA P i t A n i et al. (2004)
PN Orechella 136.0 32.0 11.0 -73.0 ci u c c i et al. (1996)
Central Italy* 55.3 28.6 38.2 - 33.1 Pe z z o et al. (2003)
Greece North Greece** 7.8 64.3 57.1 - 53.4 PA P A g e o R g i u et al. (1994)a
Central Greece** 22.2 154.6 8.3 - 2.8 mi g l i et al. (2005)b
Bulgariа
Central Balkan, Rositsa 65.0 22.0 - - 13.0 st e P A n o v (2009)
West Rhodopi, Shiroka
poljana 80.3 11.9 - - 2.4 ge n o v et al. (2008)
West Rhodopi, Beglika 74.2 20.9 2.0 - 5.5 se R A F i m o v et al. (2008)
West Rhodopi, Chepino 93.0 5.0 - - 2.0 ge o R g i e v et al. (2008)
West Rhodopi, Laki 88.1 9.6 2.7 - 1.8 ge n o v et al. 2010
Ukraine East Carpathians 32.6 48.9 48.1 66.2 Ko R n e e v , 1950
Azerbajdzan Caucas 37.0 35.0 8.0 20.0 Gidаyatov, 1970
Russia
West Caucas 1.0 95.2 2.9 -2.9 bi b i K o v et. al. (1985)
Voronz Region* 2.0 99.5 81.0c- 16.3
Caucas reserve 78.7 - 5.4 - 15.9 Ku d A K t i n , 1978
Caucas reserve 85.9 6.0 8.1 Ku d A K t i n , 1986
Sayano-Shushenski
reserve 98.7 1.1 0.2 Zavazkiy, 1981
a Autumn and winter; b winter; c 48.4% pears; * stomachs/ guts; ** stomachs.

444
Zlatanova D., A. Ahmed, A. Valasseva, P. Genov
the livestock due to the rapid decline of wild prey.
This is one of the reasons for the elevated conict.
This is the time of appearance of the so called ‘synan-
thropic’ wolves and the differentiation between the
‘wild’ and ‘synanthropic’ packs (bi b i K o v et. al. 1985,
oK A R m A 1995). The reversed situation is observed in
the Eastern Europe, and in particular in Bialowieza
(je d R z e j e w s K i et. al. 2000), where the wolf diet con-
sists mainly of wild prey (Fi=132.7%) with insigni-
cant consumption of livestock (0.6%) but still with
high consumption of plant food (30.5%).
The clear adaptable nature of the wolf to the
availability of prey is conrmed also by mA t t i o l i et
al. (1995) in a study in Casentino forest in Italy, where
the high wild prey consumption (Fi = 107.7%) cor-
relates with the wild prey abundance, resulting also
in very low livestock losses (Fi = 3.8%) and garbage
usage (2.5%). Another study of me R i g g i et al. (1996)
in three different regions in Italy with different wild
ungulate abundance also shows clear correlation be-
tween the low wild prey densities due to hunting and
Fi of wild ungulates = 17.2% to Fi domestic = 22.9%,
garbage = 9.5 and plant food = 64.9% (Genova re-
gion). With clear wild prey abundance (Casentino
forest, Flori) the situation changes to Fi of wild prey
= 107.7% and Fi of domestic animals = 3.8%
meRiggi, lovARi (1996) have found a signi-
cant inverse correlation between the Fi% of wild
and domestic ungulates in the diet. This was later
conrmed by meRiggi et al. (2011) and shows that
when wolves can choose between the two prey
categories, they may prefer wild prey. In Italy, the
consumption of rubbish / fruit and that of ungulates
are signicantly negatively correlated (meRiggi et
al. 2011). When the wild herbivores are scarce, the
wolves are forced to use alternative food sources
(e.g. small mammals, lagomorphs, fruits and gar-
bage). The same results are obtained by cuestA et
al. (1991), who compared 5 regions with different
wild prey abundance in Spain: the Fi of wild prey
varies from 0% (Galicia) to 100% (Sierra Morena)
according to its abundance.
There are also some studies of the conict with
farmers in regions where the wolf appears for the rst
time since its local extinction – Po u l l e et al. (1997) in
Mercantour, the French Alps, with Fi of domestic un-
gulates of 17.0%; gA z z o l A et al. (2005) in Val di Susa,
with Fi of domestic ungulates of 6.7%; and cA P i t A n i
et al. (2004) in Val di Cecina, with Fi of domestic un-
gulates of 9.3%. In all of these studies, there is also a
high Fi for the wild prey (above 80%), which is a clear
sign of wild prey preference. si d o R o v i c h et al. (2003)
in Belarus also studied two regions, with high wild
prey abundance (Fi for wild prey = 94.7%; domestic
= 10.5 %) and with low wild prey abundance (Fi for
wild prey = 54.0%; domestic = 74.9 %).
In all cases of low wild prey density, the share
of other prey (smaller alternative prey) is becoming
high and (or) the wolf feeds more on livestock (Table
3 and 4). Regarding the volume (Vi) of the different
food, the domestic ungulates (usually sheep) are tak-
ing around 4.5%, while the rest is taken by hares/
rabbits, rodents, fruits, grass and garbage food.
According to the reviewed published sources,
the wolves found between 40˚ and 50˚ N latitude
show a clear preference for the wild boar, with the
roe deer as a secondary prey in cases the red deer
is scarce or absent. Fi of the wild boar compared to
that of the other species shows an increment corre-
lated with the decrease in the latitude. In some plac-
es, the two prey species may have the same share or
there may be a slight prevalence of the roe deer (in
Arezzo – Vallesanta, gAzzolA et. al. 2000, or Val
Tiberina, mAttioli et al. 2004) but, in general, the
prevalence in most of the regions in Italy is for the
wild boar (Fi > 60%, for example in Pratomagno
– cAPitAni et al. 2004, mAttioli et al. 2004, or in
Alpe di Catenaia – Alboni 2004, lAmbeRti 2004,
colombo 2006). This is probably due to the wider
distribution, higher productivity and abundance of
the wild boar, followed by the roe deer, the red deer
and the mouon. Yet, there are some local devia-
tions where the wild boar is not always preferred by
the wolf and this is most probably a consequence of
other variables (local persecution, low habitat qual-
ity, etc.), which are not studied.
The longitudinal review in the east of 17˚ E
(Central and East Europe) shows a different situa-
tion. The main prey there is the deer species, roe deer
as the most important in the south and red deer and
moose in the north.
Discussion
The studies on the wolf diet are numerous but they
are usually using a similar approach. They reveal
that the wolf preybase strategy in Europe differs
signicantly from that of their counterparts in North
America and Asia, mainly due to the highly frag-
mented habitats and the lack of enough wild ungu-
lates to feed upon. In many places, the original un-
gulate diversity of 5-6 species is decreased to 2 or 3
species (oK A R m A 1995).
The uctuation in the abundance of the wild
undulates due to anthropogenic pressure led to the
complexity of the wolf feeding ecology (sP A s s o v
2007). The wolf uses all available sources and shows
exibility in its attempts to survive.

Adaptive Diet Strategy of the Wolf (Canis lupus l.) in Europe: a Review
445
Wild or domestic animals?
The studies on the wolf diet in Europe from the
1980s till today showed two ecological tendencies:
1. Wolves living in natural habitats with a high abun-
dance of wild ungulates feed mainly on wild prey.
2. If the habitats are highly anthropogenic, with low
abundance of wild prey, the wolves feed on live-
stock, also taking a lot of plant food, smaller prey
(hare and rodents) and garbage food.
On the European scale, there is a clear geo-
graphic pattern showing a different diet strategy be-
tween North (above 50˚ N latitude), where the wolf
hunts on reindeer, moose and red deer (bjARvAll et
al. 1982, Filonov 1989, jedRzejewsKi et al. 2000,
KočetKov 1988, Okarma 1995, ozolinš, A ndeRsone
2003, smietAnA, K limeK 1993), and South (below 50˚
N), where the wolf preys upon a variety of wild un-
gulates, anthropogenic food and fruits (mAcdonAld
et al. 1980, RAgni et al. 1985, sAlvAtoR, AbAd
1987, cuestA et al. 1991, PAtAlAno, lovARi 1993,
PAPAgeoRgiu et al. 1994). Yet, over time there is a
clear tendency to an increased use of wild ungulates
in Southern Europe (meRiggi et al. 2011).
There is another split between Western and
Eastern Europe (around 17oE longitude) affecting the
choice of prey, especially in Ukraine and Moldova,
where large forests have been converted into agri-
cultural lands and open elds, forcing the wolf to
adapt to new spatial, diet and reproductive condi-
tions (including hybridisation with dogs) (bi b i K o v et
al. 1985, Ry A b o v 1993)
In Southern Europe, where all wolf habitats
Table 3. Wolf diet (Fi%) in the region above 50˚ N latitude
1. N – number of samples; 2. Reindeer; 3. Moose; 4. Red deer; 5. Roe deer; 6. Wild boar; 7. Others; 8. Wild ungulates,
total; 9. other food
Source 123456789
mü l l e R (2006) 1594 92.3 -----92.3 7.7
gA d e -yo R g e n s e n , st A g e g A A R d (2000) 370 92.0 ----92.0 8.0
hu i t u (2000) 120 34.5 44.3 - - - - 78.8 21.2
An s o R g e et al. (2006) 152 - - 43.0 34.0 16.0 3.2a92.2 3.8
Kübarsepp, Valdmann (2003) 119 - 77.5 2.0 20.4 - - 99.9 0.1
Andersone, Ozolinš (2004) ? - - 59.0 -26.0 -85.0 15.0*
Ję d r z e J e w s K i et al. (2000) 411 - - 60.9 2.9 15.4 18.2b97.4 2.6**
dA n A i l o v et al. 1979 978 -47.0 -0.7 - 15.4c47.7 36.9
a mouon; b European bison; c mountain hare; * 2.3% domestic ungulates; ** 1.2% domestic ungulates.
Table 4. Wolf diet (Fi%) between 40˚ and 50˚ N latitude (Southern Europe)
1. N – number of samples; 2. Red deer; 3. Roe deer; 4. Wild boar; 5. Others; 6. Wild ungulates; 7. Domestic ungulates;
8. Plant food; 9. Other food
Source 123456789
no R e s et al. (2008) 1456 16.6 16.7 12.9 11.0 57.2 33.2 - 9.6
me R g g i et al. 1996. 519 0.8 5.0 27.3 12.5 45.6 24.1 11.5 17.9
mA t t i o l i et al. (1995) 240 14.6 32.9 44.8 3.9 92.6 6.1 0.3 1.0
Pe z z o et al. (2003) 38/45* 14.7 - 35.6 - 46.0 23.5 7.0 b 23.5-
mi g l i et al. 2005a36** -2.45 10.42 -12.87 74.3 12.14 0.69
ce l l i n A (2001) 58 - 45.0 28.7 -73.7 9.7 7.6 9.0
PA l u m b o (2003) 50*** -28. 0 62.0 8.0 98.0 - - 2.0
mA R u c c o et al. (2010) 2806 ----82.1 15.2 -2.7
st e P A n o v (2009) 84 27.0 13.5 24.5 -65.0 22.2 -12.8c
se R A F i m o v et al. (2008) 197 9.7 26.4 36.4 - 72.5 20.6 - 6.9
ge n o v et al. (2008) 84 - 35.7 50.0 -85.7 11.9 - 2.4
ge o R g i e v et al. (2008) 80 22.5 46.3 25.0 - 93.8 5.0 -1.2
ge n o v et al. (2010) 109 13.8 24.8 40.3 8.2 87.1 9.2 1.9 1.8
a winter; b fruits (Crataegus monogyna); c food remains bear/ wolf;
* stomachs / guts; ** stomachs; *** prey remains.

446
Zlatanova D., A. Ahmed, A. Valasseva, P. Genov
have been destroyed, the species uses both types
of the feeding strategies mentioned above, and the
exibility depends on the level of anthropogenic in-
uence. In Spain in the mountain regions, the wolf
is specialised on wild ungulates, while in the low-
lands it feeds more on domestic ungulates (cu e s t A
et al. 1991). In Italy the rst studies (around 1960s
and 1970s) showed a large share of livestock in the
wolf diet, but with an increase in the wild prey abun-
dance, the wolf switches to the wild prey as more
risk-free source of food (mA t t i o l i et al. 1995, ci u c c i
et al. 1996, me R i g g i et al. 1996, cA P i t A n i et al. 2004,
mA t t i o l i et al. 2004, gA z z o l A et al. 2005, mA R s i l i
2007). This is conrmed by me R i g g i , lo v A R i (1996),
me R i g g i et al. 2011 and si d o R o v i c h et al. (2003), who
state that the selection of wild and domestic ungulate
prey is inuenced mainly by their local abundance,
but also by their accessibility. Therefore, the diet
breadth increases with the decrease in the presence
of large prey in the diet. Furhermore, in areas (for
example in Bulgaria and France), where the wolf had
a local extinction and reappeared again, with the in-
crease in the wolf numbers, the livestock damages
also increase (sP A s s o v 2007, Po u l l e et al. 1997).
This is mainly due to the loss of livestock protec-
tion habits and skills in shepherds. The restocking of
wild ungulates in many regions led to a signicant
decrease in the conict wolf-man (me R i g g i , lo v A R i
1996, Po u l l e et al. 1997, vo s 2000). Since the 1990s,
there has been a constant and persistent increase in
the share of the wild ungulates in the wolf diet in
Europe. On the European scale, this phenomenon is
linked to the increase in the distribution and abun-
dance of wild ungulates since the 1970s and the total
protection of the wolf in many European countries.
This is followed by the appearance of the wolf in
new regions, but also by the formation of negative
attitudes in farmers and other local people in these
regions, and therefore, to illegal persecution (Po u l l e
et al. 1997, gA z z o l A et al. 2005).
A critical element of the quick wolf recovery in
many areas is the ability of performing long distance
travels compared to that of the prey. As a result, the
wolf may recolonise areas with low densities of wild
prey and later to switch to livestock as alternative
prey. Noting this remarkable adaptability in the wolf
expansion, li n n e l l et al. (2008) point out that in the
future the conict wolf-man will be increased by in-
cluding newly recolonised areas with higher human
population density.
Specicity of diet in the different regions
The review of the literature shows a clear dif-
ference in the diet strategy corresponding to the eco-
logical conditions. On latitudinal gradient, the wolf
is mainly taking a larger prey: moose and reindeer in
Scandinavia (An s o R g e et al. 2006, gA d e -yo R g e n s e n ,
st A g e g A A R d 2000, hu i t u 2000, mü l l e R 2006), red
deer in Central and Eastern Europe (oK A R m A et.
al. 1995, iv A n o v 1988; je d R z e j e w s K i et. al.2000;
sP A s s o v et al. 2000, sP A s s o v 2007) and wild boar in
Southern Europe. The review shows that although
this is generally true, the wolf may show a local ad-
aptation to another type of prey and even carrion in
relation to miltiple variables. Thus, the wolf offen
plays a vital sanitary role in the prevention of spread
of deseases (iv A n o v 1988), especially during the
outbreak of zoonoses, such as the swine plague.
Where this large prey is taken, the roe deer is
hunted with almost a similar frequency in every re-
gion. The other ungulate species are with local im-
portance only.
The wolf is often referred to as an opportunis-
tic species, which takes the most abundant and easy
to acquire prey species (oK A R m A 1995, me R i g g i ,
lo v A R i 1996). This is generally argued by bA R j A
(2009) who states that the trophic position of the
wolf in Galicia, Spain, is closer to a facultative spe-
cialist (feeding mainly on roe deer in the presence
of other alternatives – red deer and wild boar), than
to an opportunistic species. This is supported by
data from Osogovo Mountain, Bulgaria, where the
roe deer is mostly preferred (more than 70% of the
prey) in the presence of the extremely abundant wild
boar (Al e x A n d e R du t s o v , unpublished). This may
be due to the inability or unwillingness of the wolf
pack to undertake the risk attacking large, potential-
ly dangerous prey, when smaller but enough prot-
able prey is available. Yet, bA R j A (2009) states that
the wolf may change the key food item when other
protable prey, not so dangerous as the wild boar,
is available. The red deer preference in Poland also
conrms that hypothesis (je d R z e j e w s K i et al. 2000,
no w A K et al. 2005),
There is also a difference in ‘where’ the prey is
taken. In Scandinavia, the moose is preyed upon in
the forests (Fi l o n o v 1989), while the roe deer and the
wild reindeer – in the open areas and elds (bj ä R v A l l ,
is A c K s o n 1981, ol s s o n et al. 1997). The red deer is
mainly taken by the wolf in coniferous forests with
patches of mixed forests and also in the lowlands
(oK A R m A 1995). The wild boar and the roe deer in
Southern Europe are hunted mainly in mountainous
areas. So the local specicity may be also a result of
the types of habitat available and their coverage.
One important question is the risk taking by
the wolf while hunting the prey. The study in Arezzo
Province in Italy (cA P i t A n i et al. 2004) reveals a

Adaptive Diet Strategy of the Wolf (Canis lupus l.) in Europe: a Review
447
great uctuation in the wolf prey’s preference under
different circumstances in six areas with different
prey availability (as species and as abundance). The
observed uctuations are not only due to the differ-
ent temporary prey densities but also to the tness
of the pack. The wolf packs might be specialised on
certain prey species according to the body size of the
individuals in the pack and the numbers of the pack
members. This is closely connected with the size
of the prey (body mass) and tness, so the energy
expenditure during hunting is balanced on income-
outcome level. The predator assesses every victim
prior to the attack for the probability of killing it
with minimum efforts. There is also a risk injuries
or even death involved when attacking a larger prey.
According to me c h , Pe t e R s o n (2003), a great role in
the selection of the prey is played by a combination
of efciency of the attack and amount of food to be
obtained and the risk of injury involved, when the
prey is actively resisting.
This gives the roe deer a second place in many
areas as it is a difcult prey to get – it leads more or
less solitary life (gathering in small herds only in win-
ter or in open elds) and represents a smaller amount
of food for one take (ge R A R d et al. 1995). Although
the moose is much more risky to get, it is preferred
by the wolf in the north (ol l s s o n et al. 1997) by pre-
senting a large portion of food for one take.
More complex is the question why the wild
boar is not preferred since the abundance in Central
and Eastern Europe (Poland in particular) is simi-
lar as in Southern Europe. The wolves prefer to take
down wild boars with a weight between 10 to 35 kg.
(mA t t i o l i et al. 1995 me R i g g i et al. 1996, mA t t i o l i
et al. 2011). The juveniles with this weight are poor-
ly defended by the mothers, live in a big herd, which
is easily seen by the predators, or can be separateed
from the mothers before the end of their rst year.
The juveniles with smaller weight (below 10 kg) are
better defended by the mother and do not worth the
risk as supply small amount of food (me R i g g i et al.
1991, 1996, 2011).
The preferred prey in Central and Eastern
Europe – the red deer is also carefully chosen by the
wolf. These are usually subadults (1-2 years old) with
a body weight around or a bit higher than that of the
wild boar (je d R z e j e w s K i et al. 2000, no w A K et al.
2005). The hypothesis is that the red deer is preferred
because of the bigger biomass at one take and the
greater pack size of the northern wolves compared to
their southern counterparts. These bigger packs can
be supported by the extensive forests of Bialowieza,
where most of the studies on the wolf prey were done,
while the fragmented forests of Southern Europe
(Italy in particular) can sustain only smaller packs.
Additionally, the red deer populations are not as de-
pendent to the forest mast productivity as the wild
boar whose number may greatly vary after years with
low mast production and this could provide more sta-
ble income for the bigger packs.
Conclusions
The wolf is one of the most exible species of large
mammals which is able to survive in diverse habitats
and food conditions. In the northern part of its dis-
tribution it feeds mainly on wild ungulates (reindeer,
moose and red deer) and livestock is not of signi-
cance to its diet. In Southern Europe the wolf diet is
more diverse, as wild ungulates (mostly wild boar
and the share of roe and red deer) are still predomi-
nantly taken. In some regions the wolf also feeds on
livestock, which may constitute a greater share of its
diet. It is especially true for regions where the spe-
cies recolonises back after extinction and important
reason for higher livestock losses is the loss of pre-
venting habits in livestock husbandry.
The species may adapt its hunting/ food aquisi-
tion strategy in every region according to the status
of the prey, the habitat allowance and anthropogenic
factors. In regions, in which the wild ungulates are
abundant the wolf chooses its strategy in a balance
of the prey biomass, energy spent in catching the
prey, and the defence capabilities of the prey itself.
The wolf may quickly switch its main prey from one
type to another according to the shifts of prey density
and the anthropogenic factors. Nowadays, because
of that adaptability, it is able to expand, reestablish
and gain some of the lost territories of his historical
distribution in Europe.
References
An o n y m o u s 1986. The role of the large carnivores and ungulates
in the reserves biocenoses. Sbornik Nauchnih trudov. Rus-
sian Federation Council of Ministries Head Management
of Hunting Enterprises and Reserves. Мoskow (Central
Scientic Laboratory for hunting enterprises and reserves).
152 p. (In Russian).
Al b o n i m. 2004 . Biologia del lupo (Canis lupus) nell’Oasi di
protezione dell’Alpe di Catenaia in Provincia di Arezzo.
– Tesi di Laurea, Università degli Studi di Sassari. 57 p.
(In Italian).
An d e r s o n e z., J. oz o l i n љ . 2004. Food habits of wolves Canis
lupus in Latvia. – Acta Theriologica, 49 (3): 357-367.

448
Zlatanova D., A. Ahmed, A. Valasseva, P. Genov
An s o R g e h., g. Kl u t h , s. hAhne. 2006. Feeding ecology of
wolves Canis lupus returning to Germany. – Acta Therio-
logica, 51 (1): 99-106.
AvAnzinelli e. 2001. Consistenza, dinamica di popolazione e
dieta del lupo (Canis lupus) nel versante toscano del Parco
Nazionale delle Foreste Casentinesi, Monte Falterone e
Campigna. Tesi di Laura, Università degli studi di Pisa.
77 p. (In Italian).
Ay b e s c., d. yA l d e n . 1995. Place – name evidence for the former
distribution and status of wolves and beavers in Britain. –
Mammal Review, 25 (4): 201-227.
bA R j A I. 2009. Prey and Prey-Age Preference by the Iberian
Wolf Canis lupus signatus in a Multiple-Prey Ecosystem.
– Wildlife Biology, 15 (2):147-154.
Bi B i K o v d., А. Ku d A K t i n , l. ry A B o v . 1985. Synanthropic wolves:
distribution and diet. – Zoologicheskiy zhurnal, 64: 429-
441. (In Russian; English and Russian summaries).
bj A R v A l l A., e. is A K s o n . 1982. Winter ecology of a pack of three
wolves in north western Sweden. – In: F. H. hA R R i n g t o n
and P. C. PA q u e t . Parkridge (eEds.): Wolves of the World
-Perspective of Behaviour, Ecology, and Conservation.
New Jersey (Noyes Publications), 146-157.
bo i t A n i l. 1996. Dalla parte del lupo. Milano, I libri di Airone,
G. Mondadori e Ass. 215 p. (In Italian).
Bu n e v i c h А. 1988. Dynamics of wolf numbers and diet in Bialow-
ieza Forest. In: Zapovedniki Belorusii. Minsk. – Urozhay
12: 108-113 (In Russian).
cA P i t A n i c., i. be R t e l l i , P. vA R u z z A , m. sc A n d u R A , m. AP o l l o n i o .
2004. A comparative analysis of wolf (Canis lupus) diet in
three different Italian ecosystems. – Mammalian Biology
69: 1-10.
cellinA s. 2001. La dieta del lupo (Canis lupus, Linnaeus,
1758) nel Parco Regionale dei Cento Laghi nell’Alta Val
Parma e Cedra (PR). – Tesi di laurea effettuata e discussa,
l’Università degli Studi di Parma, 72 p. (In Italian).
ci u c c i P., l. bo i t A n i , e. Pe l l i c c i o n i , m. Ro c c o , i. gu y . 1996. A
comparison of scat analysis methods to assess the diet of
the wolf. – Wildlife Biology, 2: 37-48.
co l o m b o A. 2005. Ecologia del lupo (Canis lupus) nell’Oasi di
protezione dell’Alpe di Catenaia in Provincia di Arezzo.
– Tesi di Laurea, Università degli Studi di Milano, 69 p.
(In Italian).
cu e s t A l., F. bA R c e n A , F. PA l c i o s , s. Re i g . 1991. The trophic
ecology of the Iberian Wolf (Canis lupus signatus Cabrera,
1907) – A new analysis of stomach’s data. – Mammalia 55
(2): 239-255.
dAnilov i., s. Ru s A K o v, l. tumAnov. 1979. The carnivorous
mammals of North-West USSR. Leningrad (Nauka), 161
p. (In Russian).
Fe R R A R i m. 2012. La via del lupo. Bari, Sedit, 197 p. (In Italian).
Fi l o n o v K. 1989. Ungulate and large predators in wildlife re-
serves. Moscow, Nauka, 251 p. (In Russian).
Fi l o n o v K., m. KA l e t s K A y A . 1985. Inuence of wolf predation on
wild ungulates. – In: bi b i K o v D. I. (ed.): The Wolf – History,
Systematics, Morphology, Ecology. Moscow (Nauka), 14
(3): 336-354. (In Russian).
gA d e -j o R g e n s e n i., R. st A g e g A A R d . 2000. Wolf Canis lupus diet
composition of in East-Central Finland. – Acta Theriologica
45 (4): 537-547.
GA v r i n В., s. do n A u r o v . 1954. The wolf in in Bialowieza Forest.
– Zoologicheskiy zhurnal, 33: 904-924. (In Russian).
gA t A h V. 1979. Role of the wolf in the biogeocoenoses in the
forests of Belarus. – In: Ecological brackground of the
conservation and sustainable use of carnivorous mammals.
Moscow (Nauka), 94-95. (In Russian).
gA z z o l A A. 2000. Distribuzione ed ecologia alimentare del lupo
nel Parco Nazionale delle Foreste Casentinesi, Monte
Falterone e Campigna. – Tesi di Laura, Università degli
studi di Pisa, 83 p. (In Italian).
gA z z o l A A., i. be R t e l l i , e. Av A n z i n e l l i , A. to l o s A n o , P. be R -
t o t t o , m. AP o l l o n i o . 2005. Predation by wolves (Canis
lupus) on wild and domestic ungulates of western Alps,
Italy. – Journal of Zooogyl London, 266: 205-213.
ge n o v P., d. di m i t R o v A ., t. ge o R i e v, v. dR A g A n o v , P. bA n c h e v ,
d. AR A b A d z i e v , R. mi R c h e v . 2008. A study on the bear and
wolf in State Hunting Station „Shiroka Poyana“, for the
aim of their management. – Godishnik na Shumenskiya
Universitet „Episkop Konstantin Preslavski”, XVIII B6,
Prirodni nauki, Biologia, 173-189. (In Bulgarian).
Ge n o v P., А. dz i n d z i e v A , G. be d R o v . 2010. The diet of the wolf
(Canis lupus L.) in the area of State Hunting Station „Ko-
rmisosh”, Western Rhodopi. Anniversary Scientic Confer-
ence „Bulgaria and the Bulgarians in Europe”, 17.10.2009.
(In Bulgarian).
Ge o r i e v v., G. Ge o r i e v , n. ni n o v , А. dz i n d z i e v A P. Ge n o v . 2008.
A study on the wolf (Canis lupus L.) diet in the area of
State Hunting Station „Chepino“, Western Rhodopi. – In:
Anniversary Scientic Conference of Ecology, dedicated
to the 20th anniversary of the Department of “Ecology”
at the Faculty of Biology of the University of Plovdiv
“Paysii Hilendarski”, 01.11.2008, Plovdiv, 216-224. (In
Bulgarian).
ge R A R d j., y. le Pe n d u , m. mA u b l A n c , j. vi n c e n t , m. Po u l l e ,
c. ci b i e n 1995. Large group formation in European roe
deer: an adaptive feature? – Rev. Ecol. (Terre Vie), 50:
391-401.
GidАyAtov Y. 1970. About wolf (Canis lupus L.) ecology in
Azerbajdzan. Izvestiya AN АZSSR. Ser. Biol. Nauk., 1:
50-56 (In Russian).
giustini d. 2002. Indagine sul comportamento alimentare e
spaziale di tre branchi di lupo (Canis lupus) nelle Foreste
Casentinesi, Monte Falterona e Valle Santa. – Tesi di Lau-
rea, Università degli studi di Pisa, 78 p. (In Italian).
huitu O. 2000. Wolf (Canis lupus L.) diet and prey species
selectivity in Kainuu, Finland. – Master of science thesis.
University of Javaskyla, 30 p.
iv A n o v , v. 1988. Observations on the behaviour and biology of
the wolf (Canis lupus L., 1758) in Ihtimanska Sredna Gora
mountain. – Ecology. Soa, 21: 25-33
je d R z e j e w s K i w., b. je d R z e j e w s K A , h. oK A R m A , K. sc h m i d t , c.
zu b , m. mu s i A n i . 2000. Prey selection and predation by
wolves in Bialowieza Primary Forest, Poland. – Journal
of Mammalogy, 81: 197-212.
Ko č e t K o v v. 1988. Wolf biology in Verhnelolzie, in the region of
Central-Forestry State Reserve. Summary of PhD thesis.
Moskva, 20 p. (In Russian).
Ko č e t K o v V., A. so K o l o v . 1979. Wolf diet in the Central-Forestry
State Reserve. – In: Ecological brackground of the con-
servation and sustainable use of carnivorous mammals.
Moscow, Nauka, 112-114. (In Russin).
KojolA i., s. heiKKinen, o. huitu, K. toPPinen, K. heiKuRA,
s. Ro n K A i n e n . 2004. Predation on European wild forest
reindeer (Rangifer tarandus) by wolves (Canis lupus) in

Adaptive Diet Strategy of the Wolf (Canis lupus l.) in Europe: a Review
449
Finland. – Journal of Zooogy London, 263: 229-235.
KoRneev А. 1950 Wolf and its importance. Kiev, 104 p. (In
Ukrainian).
KR u u K , H. 1989. The social badger. Ecology and behaviour of
a group-living carnivore (Meles meles). Oxford, Oxford
University Press, 253 p.
Kü b A R s e P P m., h. vA l d m A n n . 2003. Winter diet and movements of
wolf (Canis lupus) in Aalampedja Nature Reserve, Estonia.
– Acta Zoologica Lituanica, 13 (1): 28-33.
Ku d A K t i n А. 1978. About wolf hunting selectivity on ungulates
in Caucas reserve . – Bulletine М. Obshestva Ispitanie
Priroduy, Otd. Biol., 83 (3): 19-28. (In Russian).
Ku d A K t i n А. 1986. The inuence of wolf on the ungulates in
Влияние волка на копытных в Caucas reserve, Caucas
biosphere reserve. – In: The role of large carnivores and
ungulates in the biocoenoses of reserves. Sbornik Nauchnih
trudov, Moscow, 21-35 (In Russian).
lAmbeRti P. 2004. L’analisi di un sistema coevoluto: il caso
lupo-ungulati selvaticinell’Oasi di protezione dell’Alpe
di Catenaia. – Tesi di Dottorato, Università degli studi di
Pisa, 153 p. (In Italian).
li K h A t c K y u., d. bi b i K o v , l. Ry A b o v . 1995. Wolf (Canis lupus)
and red deeer (Cervus elaphus) in Voronezh state reserve. –
Zoologicheskiy zhurnal, 74 (11): 110-121, (In Russian).
li n n e l l j., v. sA l v A t o R i , l. bo i t A n i . 2008. Guidelines for popula-
tion level management plans for large carnivores in Europe.
A Large Carnivore Initiative for Europe report prepared for
the European Commission (contract 070501/2005/424162/
MAR/B2). 83 p.
lo c K i e j. 1959. The estimation of the food of foxes. – The Journal
of Wildlife Management, 23: 224-227.
mA c do n A l d d., l. bo i t A n i , P. bA R A s s o . 1980. Foxes, wolves and
conservation in the Abruzzo Mountains. – Biogeographica
18: 223-235.
mA R s i l i s. 2007. Ecologia troca del lupo Canis lupus L., revisione
degli studi effettuati in Europa nell’ultiemo trentennio, con
particolare riferemento a un’area dell’Appennino cenrto-
orientale. Italy. – Tesi di laurea, Universita’ degli studi di
Pisa, 118 pp. (In Italian).
mA R u c c o F., l. bo i t A n i , e. Av A n z i n e l l i , s. dA l m A s s o , l. oR-
l A n d o . 2010. Progetto lupo regione Piemonte. Torino, –
Rapporto 1999-2010, 138 p. (In Italian).
mA t t i o l i l, m. AP o l l o n i o , v. mA z z A R o n e , e. ce n t o F A n t i . 1995.
Wolf food habits and wild ungulate availability in the For-
este Casentinesi National Park; Italy. – Acta Theriologica,
40: 387-402.
mA t t i o l i l., c. cA P i t A n i , e. Av A n z i n e l l i , A. gA z z o l A , m. AP o l -
l o n i o . 2004. Predation by wolves (Canis lupus) on roe deer
(Capreolus capreolus) in north-eastern Apennine, Italy. –
Journal of Zooogyl London, 264: 249-258.
mA t t i o l i l., c. cA P i t A n i , A. gA z z o l A , m. sc A n d u R A , m. AP o l -
lonio m. 2011. Prey selection and dietary response by
wolves in a high-density multi-species ungulate community.
– Eur J Wildl Res., 57:909-922
mech l. 1970. The Wolf: The Ecology and Behavior of an
Endangered Species. Natural History Press (Doubleday
Publishing Co., N.Y.) 389 p.
me c h l., l. bo i t A n i . 2003. Wolves: behavior, ecology, and con-
servation. USA, University of Chicago Press. 448 p.
me c h l., R. Pe t e R s o n . 2003. Wolf–prey relations. In me c h l., l.
bo i t A n i (eds): Wolves: behavior, ecology, and conservation.
University of Chicago Press, Chicago, Illinois: 131–157
me R i g g i A., s. lo v A R i . 1996. A review of wolf predation in south-
ern Europe: does the wolf prefer wild prey to livestock?
– Journal of Applied Ecology, 33: 1561-1571.
me R i g g i A., A. bR A n g i , c. mA t t e u c c i , o. sA c c h i . 1996. The feed-
ing habits of wolves in relation to large prey availability in
northern Italy. – Ecography, 19: 287-295.
me R i g g i A., A. bR A n g i , l. sc h e n o n e , d. si g n o R e l l i , P. mi l A n e s i .
2011. Changes of wolf (Canis lupus) diet in Italy in relation
to the increase of wild ungulate abundance. – Ethology,
Ecology & Evolution, 23:3, 195-210
me R t z P. 1953. The wolf in the Voronez region. – In: Reconstruc-
tion of the vertebrate fauna in our country. Moscow, Mos-
cow Society for Nature Study, 117-135. (In Russian).
mi g l i d., d. yo u l A t o s , y. il i P o u l o s . 2005. Winter food habitats
of wolves in central Greece. – Journal of Biological Re-
search, 4: 217-219.
mü l l e R s. 2006. Diet composition of wolves (Canis lupus) on
the Scandinavian peninsula determined by scat analysis.
Sweden. PhD thesis, Swedish University of Agricultural
Sciences, Uppsala, 238 p.
no R e s c., l. ll A n e z A , m. Ál v A R e z . 2008. Wild boar (Sus scrofa)
mortality by hunting and wolf (Canis lupus) predation: an
example northern Spain. – Wildlife Biology, 14: 44-51.
no w A K s., R. my s l A j e K , b. je d R z e j e w s K A . 2005. Pattern of wolf
Canis lupus predation on wild and domestic ungulates in
the western Carpathian Mountains. – Acta Theriologica,
50 (2): 263-276.
oK A R m A h. 1995. The trophic ecology of wolves and their preda-
tory role in ungulate communities of forest ecosystems in
Europe. – Acta Theriologica, 40 (4): 335-38.
oKARmA h., b. jedRzejewsKA, w. j edRzejewsKi, z. KRAsinsKi,
l. mi l K o w s K i . 1995. The roles of predation, snow cover,
acorn crop, and man-related factors on ungulate mortality
in Białowieża Primeval Forest, Poland. – Acta Theriologica
40 (2): 197-217.
ol s s o n o., j. wi R t b e R g , m. An d e R s s o n , i. wi R t b e R g . 1997. Wolf
(Canis lupus) predation on moose (Alces alces) and roe
deer (Capreolus capreolus) in south central Scandinavia.
– Wildlife Biology, 3: 13-23.
ozolinš J., Ž. Andersone. 2003. Management Plan for Wolf
(Canis lupus) in Latvia. Latvian State Forestry Research
Silava. State Forest Service of Ministry of Agriculture.
36 p.
PA l u m b o d. 2003. I lupi del parco del Corno alle Scale. Bo-
lognia, Riccrche e monotoraggi sulla presenza dell lupo
nell’Appennino Bolognese, 52 p. (In Italian).
PA P A g e o R g i o u n., c. vl A c h o s , A. sF o u g A R i s , e. ts A c h A l i d i s . 1994.
Status and diet of wolves in Greece. – Acta Theriologica
39 (4): 411-416.
PA t A l A n o m., s. lo v A R i . 1993. Food habits and trophic niche
overlap of the wolf Canis lupus, L. 1758 and the red fox
Vulpes vulpes (L. 1758) in a mediterranean mountain area.
– Revue d’écologie, 48: 279-294.
PeteRs, g. (1993): Canis lupus Linnaeus, 1758 – Wolf. – In:
st u b b e m., F. KR A P P (eds.): Handbuch der Säugetiere Eu-
ropas, Band 5: Raubsäuger – Carnivora (Fissipedia), Teil I:
Canidae, Ursidae, Procyonidae, Mustelidae 1. – Wiesbaden
(Aula-Verlag), 47-106. (In German).
Pezzo F., l. PARigi, R. Fico. 2003. Food habitat of wolves in
central Italy based stomach and intestine analyses. – Acta
Theriologica, 48 (2): 265-270.
Pimlot d. 1967. Wolf predation and ungulate populations. –

450
Zlatanova D., A. Ahmed, A. Valasseva, P. Genov
American Zoologist, 7 (2): 267-278
Po u l l e m., l. cA R l e s , b. le q u e t t e . 1997. Signicance of ungu-
lates in the diet of recently settled wolves in the Mercantour
Mountains (Southern France). – Revue d’écologie, 52:
357-368.
RA g n i , b., A. mA R i A n i , F. in v e R n i , m. mA g R i n i . (1985) Il lupo
in Umbria. Atti del Convegno Nazionale Gruppo Lupo
Italia (ed. G. Boscagli), Gruppo Lupo Italia, Pescasseroli,
22-36. (In Italian).
Ro q u e s., F. Ál v A R e s , F. Pe t R u c c i -Fo n s e c A . 2001. Utilización
espacio-temporal y hábitos alimentarios de un grupo re-
productor del lobos en el noroeste de Portugal. – Galemys,
13: 1-20. (In Portuguese).
RuKovsKy n. 1985. Feeding and biocoenotic relationship. . –
In: bi b i K o v d. i. (ed.): The Wolf – History, Systematics,
Morphology, Ecology. Moscow (Nauka), 14 (3): 336-354.
(In Russian).
RuKovsKy m., A. KuPRiyAnov. 1972. Some peculiarities of
distribution and ecology of Canis lupus in the Onega
peninsula. – Zoologicheskiy zhurnal, 31 (10):1593-1596
(In Russin).
ru s A K o v О. 1979. Feeding habits of the wolf in the northern part
of the Pskov region. – In: Ekologicheskiye osnovy okhrany
i ratsionalnogo ispol’zovaniya khishchnykh mlekopitay-
ushchikh. – Nauka, 132-134 (In Russian).
ru s A K o v О., e. ti m o t e e v A . 1984. Wild boar. Leningrad, Leningrad
University, 205 p. (In Russian).
RyAbov L. 1993. Wolves of the Black Earth Belt. Izdatelstvo
Voronezhskogo Universiteta, Voronezh, 1-167. (In Rus-
sian).
sA l v A d o R A., P. Ab A d . 1987. Food habits of a wolf population
(Canis lupus) in León province, Spain. – Mammalia, 51
(1): 45-52
se r A f i m o v G., B. so f u , А. dz i n d z i e v A , P. Ge n o v . 2008. Place and
role of the wolf (Canis lupus L.) in the State Hunting Station
„Beglika”, Western Rhodopi. – In: Anniversary Scientic
Conference of Ecology, dedicated to the 20th anniversary of
the Department of “Ecology” at the Faculty of Biology of
the University of Plovdiv “Paysii Hilendarski”, 01.11.2008,
Plovdiv, 225-235. (In Bulgarian).
sidoRovich v., l. tiKhomiRovA, b. jedRzejewsKA. 2003. Wolf
Canis lupus numbers, diet and damage to livestock in
relation to hunting and ungulate abundance in northeast-
ern Belarus during 1990-2000. – Wildlife Biology 9 (2):
103-111.
sm i e t A n A w., A. Kl i m e K . 1993. Diet of wolves in Bieszczady
Mountains, Poland. – Acta Theriologica, 38: 245-251.
sP A s s o v n. 2007. Wolf Canis lupus (Linnaeus, 1758). – In: mi t e v A
s., B. mi h o v A , К. Ge o r G i e v , B. Pe t r o v , d. v A n si n K (eds.):
Mammals important to preserve in Bulgaria. Dutch Mam-
mal Society VZZ, Silistra (Neo Art), 222-233.
sPAssov, n., n. ninov, K. geoRgiev, R. gunchev, v. ivAnov.
2000. Status of the large mammals (Macromammalia). –
In: Biodiversity of the Central Balkan National Park. Soa
(Pensoft), 425-490.
st e P A n o v i. 2009. Study on food of the wolf (Canis lupus L.) on
the territory of State Hunting Station „Rositsa“, Central
Stara Planina. Proceedings of the Anniversary Scientic
Conference of 80 years Innstitute of Forest BAS. (In Bul-
garian).
su m i n s K i P., w. Fi l i P i A K. 1977. Beitrag zur Nahrungsuntersuchung
des Wolfes (Canis lupus L.). – Zeitschrift für Jagdwissen-
schaft, 23 (1): 1-5 (In German).
vo s j. 2000. Food habits and livestock depredation of two Iberian
wolf packs (Canis lupus signatus) in the north of Portugal.
– J. Zool. (Lond.), 251: 457-462.
zA v A z K i y Б. 1981. The role of wolf in the biocoenoses of Sayano-
Shushenski reserve. In: The role of large carnivores and
ungulates in the biocoenoses of reserves. Sbornik Nauchnih
trudov, Moscow, 35-54. (In Russian).
Received: 08.05.2014
Accepted: 03.11.2014

Adaptive Diet Strategy of the Wolf (Canis lupus l.) in Europe: a Review
451
Appendix 1. Literature review of the main sources on the wolf diet strategy in Europe
Country Location Latitude
(oN)
Longitude
(oE or oW) Source
- Europe – overall comparison - - me R i g g i , lo v A R i (1996)
- Europe – overall comparison - - oK A R m A (1995)
Azerbajdzan Caucas Gi d А y A t o v , 1970
Belarus Bialowieza 52°2′-52o4′ 23°2′-24°2′ E gA v R i n , do n A u R o v (1954)
Belarus Bialowieza 52°2′-52°4′ 23°2′-24°2′ E bu n e v i c h (1988)
Belarus North-eastern Belarus 55°0′ 29°3′ E si d o R o v i c h et. al. (2003)
Belarus various regions 53°3′-56o0′ 23°0′ -32°5′ E gA t A h (1979)
Bulgaria Western Rhodopes 41°5′ 24°5′ E ge n o v et. al. (2010)
Bulgaria Western Rhodopes 41°5′ 24°1′ E ge n o v et. al. (2008)
Bulgaria Western Rhodopes 41°5′ 23°6′ E ge o R g i e v et. al. (2008)
Bulgaria Western Rhodopes 41°5′ 24°0′ E se R A F i m o v et. al. (2009)
Bulgaria Central Balkan region 42°5′ 25°0′ E sP A s s o v et al. 2000; st e P A n o v (2009)
Bulgaria Ihtimanska Sredna Gora
mountain 42°5′ 23°8′ E iv A n o v (1988)
Estonia Alam-Pedja Nature Reserve 58°2′ 26°1′ E Kü b A R s e P P , vA l d m A n n (2003)
Finland Central Finland 62°3′ 25°5′ E gA d e -j o R g e n s e n , st A g e g A A R d (2000)
Finland Kainuu 64°2′ 28°3′ E hu i t u (2000)
Finland East-Central Finland 62°3′ 26°3′ E Ko j o l A et. al. (2004)
France Southern France 44°1′ 7°10′ E Po u l l e et. al. (1997)
Germany all country 51°0′ 13°1′ E An s o R g e et. al. (2006).
Greece Central Greece 39°4′ 21°3′ E mi g l i et. al. (2005)
Greece North Greece 41°1′ 24°1′ E PA P A g e o R g i o u et. al. (1994)
Italy Provincia di Arezzo 43°3′ 11°5′ E Al b o n i (2004)
Italy Parco Nazionale, Monte
Falterone 43°6′ 11°4′ E Av A n z i n e l l i (2001)
Italy all country 37°-47° 7°3′ -17°3′ E bo i t A n i (1996)
Italy North-Eastern Apennines 46°1′ 12°5′ E cA P i t A n i et. al. (2003)
Italy three different Italian ecosys-
tems 43°3′ 11°5′ E cA P i t A n i et. al. (2004)
Italy Parco Regionale dei Cento
Laghi 44°3′ 10°1′ E ce l l i n A (2001)
Italy all country 37°-47° 7°3′ -17°3′ E ci u c c i et. al. (1996)
Italy Provincia di Arezzo 43°3′ 11°5′ E co l o m b o (2005)
Italy Western Alps 44°1′ 7°1′ E gA z z o l A et. al. (2005)
Italy Parco Nazionale, Monte
Falterone 43°6′ 11°4′ E gA z z o l A et. al. (2000)
Italy Foreste Casentinesi, Monte
Falterona e Valle Santa 43°6′ 11°5′ E gi u s t i n i (2002)
Italy Alpe di Catenaia 43°4′ 11°6′ E lA m b e R t i (2004)
Italy Abruzzo mountains 41°4′ 13°5′ E mA c do n A l d et. al. (1980)
Italy Central-East Apennine 43°4′ 12°4′ E mA R s i l i (2007)
Italy Piemonte region 44°1′ 7°2′ E mA R u c c o et. al. (2010)
Italy Foreste Casentinesi National
Park 43°6′ 11°4′ E mA t t i o l i et. al. (1995)
Italy North-eastern Apennine 46°1′ 12°5′ E mA t t i o l i et. al. (2004)
Italy Northern Italy 46°0′ 10°0′ E me R i g g i et. al. (1996)
Italy all country 37°-47° 7°3′ -17°3′ E me R i g g i et. al. (2011)
Italy BologneseAppenne 44°3′ 11°2′ E PA l u m b o (2003)
Italy Abruzzo NP, Central Italy 41°4′ 13°5′ E PA t A l A n o & lo v A R i (1993)
Italy Central Italy 43°3′ 11°5′ E Pe z z o et. al. (2003)
Italy Umbria, Central Italy 44°5′ 12°4′ E RA g n i et. al. (1985)
Latvia all country 55°4′- 58°0′ 20°5′- 28°1′ E An d e R s o n e , ozolinš (2004)

452
Zlatanova D., A. Ahmed, A. Valasseva, P. Genov
Country Location Latitude
(oN)
Longitude
(oE or oW) Source
Latvia all country 55°4′- 58°0′ 20°5′- 28°1′ E oz o l i n š , An d e R s o n e (2003)
Poland Bialowieza 52°3′- 55°0′ 23°3′- 23°5′ E je d R z e j e w s K i et. al. (2000)
Poland Western Carpathian Mnts 49°3′- 50°0′ 19°0′- 23°3′ E no w A K et. al. (2005)
Poland Bieszczady Mountains 49°3′ 22°5′E sm i e t A n A , Kl i m e K (1993)
Poland all Poland 49°3′-54°5′ 15°0′E-23°3′
Esu m i n s K i , Fi l i P i A K (1977)
Portugal North-east Portugal 41°4′ 7°3′W Ro q u e et. al. (2001)
Russia West Caucasus
Voronz Region 43°0′ 42°1′ E bi b i K o v et. al. (1985)
Russia Darvinskii Reserve 58°5′ 37°3′E Fi l o n o v (1989)
Russia all Russia - - Ru K o v s K y (1985); Fi l o n o v , KA l e t s K A y A
(1985)
Russia Voronezh state reserve 51°5′ 39°2′E li K h A t c K y et. al. (1995)
Russia Arkhangelsk, Onega peninsula 64°3′ 38°0′E Ru K o v s K i , Ku P R i y A n o v et. al. (1972)
Russia North-West USSR - - dA n i l o v et. al. (1979)
Russia Tversk region 56°5′ 34°5′E Ko č e t K o v , so K o l o v (1979)
Russia Caucas reserve Ku d A K t i n , 1978
Russia Caucas reserve Ku d A K t i n , 1986
Russia Voronez region 51°5′ 39°2′E me R t z (1953)
Russia North Pskov region 58°3′ 28°4′E Rusakov (1979)
Russia Pskov region 58°0′ 28°4′E RussAKov, timoFeevA (1984)
Russia Sayano-Shushenski reserve zA v A z K i y , 1981
Spain ve areas to cover all country 36°0-43°5′ 9°0′W-3°0′ E cu e s t A et. al. (1991)
Spain Northern Spain 43°0′ 4°5′W no R e s et. al. (2008)
Spain León province 42°4′ 5°5′W sA l v A d o R , Ab A d (1987)
Spain Galicia (North-western Spain) 42°3′ 8°1′W bA R j A (2009)
Sweden South central Scandinavia 60°3′ 15°0′E ol s s o n et. al. (1997)
Sweden North-Western Sweden 65°1′ 16°0′E bj A R v A l l , is A K s o n (1982)
Sweden all country 55°0′-68°3′ 11°0′-19°0′ E mü l l e R (2005)
Ukraine East Carpathians Ko R n e e v , 1950
Appendix 1. Continued