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Prey choice and diet of wolves related to ungulate communities and wolf
subpopulations in Poland
Author(s): Wlodzimierz Jdrzejewski , Magdalena Niedzialkowska , Matthew W . Hayward , Jacek
Goszczyski , Bogumila Jdrzejewska , Tomasz Borowik , Kamil A . Barto , Sabina Nowak , Joanna
Harmuszkiewicz , Andrzej Juszczyk , Tomasz Kalamarz , Agnieszka Kloch , Joanna Koniuch , Katarzyna
Kotiuk , Robert W . Myslajek , Monika Ndzyska , Anna Olczyk , Marta Teleon , and Mariusz
Wojtulewicz
Source: Journal of Mammalogy, 93(6):1480-1492. 2012.
Published By: American Society of Mammalogists
DOI: http://dx.doi.org/10.1644/10-MAMM-A-132.1
URL: http://www.bioone.org/doi/full/10.1644/10-MAMM-A-132.1
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Journal of Mammalogy, 93(6):14801492, 2012
Prey choice and diet of wolves related to ungulate communities and
wolf subpopulations in Poland
WODZIMIERZ JEDRZEJEWSKI, MAGDALENA NIEDZIAKOWSKA, MATTHEW W. HAYWARD, JACEK GOSZCZY

NSKI,
BOGUMIA JEDRZEJEWSKA,* TOMASZ BOROWIK, KAMIL A. BARTO

N, SABINA NOWAK, JOANNA HARMUSZKIEWICZ,
ANDRZEJ JUSZCZYK, TOMASZ KAAMARZ, AGNIESZKA KLOCH, JOANNA KONIUCH, KATARZYNA KOTIUK,
ROBERT W. MYSAJEK, MONIKA NEDZY

NSKA, ANNA OLCZYK, MARTA TELEON, AND MARIUSZ WOJTULEWICZ

Mammal Research Institute, Polish Academy of Sciences, 17-230 Biaowie za, Poland (WJ, MN, MWH, BJ, TB, KAB)
Department of Forest Protection and Ecology, Faculty of Forestry, Warsaw Agricultural University, ul. Nowoursynowska
159, 02-776 Warsaw, Poland (JG, JH, AJ, TK, MW)
Association for Nature Wolf, Twardorzeczka 229, 34-324 Lipowa, Poland (SN, RWM)
Department of Ecology, Faculty of Biology, Warsaw University, ul. Banacha 2, 02571 Warsaw, Poland (AK)
Department of Zoology, Faculty of Biology and Chemistry, University of Biaystok, ul. S

wierkowa 20 B, 15-950 Biaystok,

Poland (JK, KK)
Collegium Biologicum, Department of Systematic Zoology, Adam Mickiewicz University, ul. Umultowska 89, 61-614
Pozna n, Poland (MN)
Department of Genetics and Animal Husbandry, Faculty of Animal Sciences, Warsaw Agricultural University, ul.
Nowoursynowska 159, 02-776 Warsaw, Poland (AO)
Department of Ecology and Environmental Protection, Institute of Biology, Academy of Podlasie, ul. Prusa 10, 08-110
Siedlce, Poland (MT)
* Correspondent: bjedrzej@ibs.bialowieza.pl
Wolves (Canis lupus) belong to 3 genetically distinct subpopulations despite the absence of topographic barriers
limiting dispersal. Based on data on wolf diets from 13 localities and wolf kill remains from a national-scale
census, we investigated regional variation in wolf diet in relation to species structure of ungulate communities
and spatial genetic differentiation of wolf populations. We also tested if various sources of data on wolf prey
(scats and kills) and availability of ungulates (game inventory and harvest) yielded comparable results on prey
selection. The main prey of wolves was red deer (Cervus elaphus), roe deer (Capreolus capreolus), and wild
boar (Sus scrofa). The proportion of main prey in wolf diets increased with prey availability in the community,
yet wolves selected red deer, preyed on roe deer proportionally to their relative abundance, and avoided wild
boar. Large prey was recorded among kills more often than small prey. Despite similar species structure of
ungulate communities throughout Poland, there were signicant regional differences in wolf diet, which
corresponded to the genetic structure of populations. In northeastern Poland, wolves frequently hunted red deer,
roe deer, wild boar, beavers (Castor ber), and moose (Alces alces). In eastern Poland, roe deer dominated kills.
In southeastern Poland, wolves were strongly specialized on red deer. We propose that prey and habitat
specialization of wolves, rather than geographic distance or topographic barriers to dispersal, are responsible for
the observed ecological divergence of wolf populations, as reected in their diet composition.
Key words: Canis lupus, deer, diet composition, genetic structuring, Poland, wild boar
2012 American Society of Mammalogists
DOI: 10.1644/10-MAMM-A-132.1
Poland supports some of the least disturbed landscapes, and
the most diverse communities of native ungulates, on the
European continent (Okarma 1995). The wolf (Canis lupus) is
a dominant large carnivore in Poland, reaching abundances far
higher than in most other countries in western Europe (Boitani
2000; Jedrzejewski et al. 2008). This high abundance provides
excellent conditions for studying the role of wolves in
regulating the abundance of ungulates and structure of ungulate
w w w . m a m m a l o g y . o r g
1480
communities. Interactions between large predators and their
prey in Poland have been studied in the Carpathian Mountains
(Lesniewicz and Perzanowski 1989; Nowak et al. 2005;
S

mietana and Klimek 1993) and the lowland forest of

Biaowie za (Jedrzejewska and Jedrzejewski 1998; Jedrzejewski
et al. 1992, 2000, 2002). These studies have shown that wolves
preferentially prey on red deer (Cervus elaphus) and avoid wild
boar (Sus scrofa), while taking roe deer (Capreolus capreolus)
in proportion to its abundance (Jedrzejewska and Jedrzejewski
1998; Nowak et al. 2005).
These local case studies have so far covered a small part of
the variation in prey resources observed throughout Poland. In
Poland, ungulate communities consist of 2 (roe deer and wild
boar) to 5 (European bison [Bison bonasus], moose [Alces
alces], red deer, roe deer, and wild boar) native species. In the
Tatra Mountains of southern Poland, a small population of
chamois (Rupicapra rupicapra) persists, and in several other
localities introduced nonnative species (fallow deer [Dama
dama], sika deer [Cervus nippon], and mouon [Ovis aries])
occur in small numbers (Wawrzyniak et al. 2010). Moreover,
the local communities of ungulates strongly vary in relative
abundance of species, in terms of both numbers and crude
biomass. This variation provides an opportunity to study the
dietary plasticity of wolves in the same biogeographic region
and similar environmental conditions of temperate forests.
Based on a review of continental-scale (European forest
zone) data on wolf prey choice in relation to available ungulate
species, Okarma (1995) suggested that the food niche of
wolves widens as the ungulate community becomes more
diverse, while at the same time regularly showing a preference
for red deer. The degree to which wolves select a given species
may depend not only on the relative abundance of that species
but also on available alternative prey. For example, in central
and eastern Europe where large cervids (moose or red deer, or
both) predominate ungulate communities, wolves notably
avoid preying on wild boar (Filonov 1989; Jedrzejewska et
al. 1994). Conversely, in southern Europe where large cervids
are absent or rare, wolves often rely on wild boar and prefer it
to the more numerous roe deer (Mattioli et al. 2004).
Varying availability of prey species may not be the only
factor shaping variation in wolf diet. Recently, Pilot et al.
(2006) studied genetic variability of central and eastern
European wolves and found that a nonrandom spatial genetic
structure of the population in the absence of topographic
barriers limited dispersal. They demonstrated a signicant
correlation between genetic structuring and ecological factors,
namely climate, habitat type, and composition of wolf diets.
Polish wolves appear to belong to 3 or 4 genetically distinct
subpopulations delimited based on frequencies of mitochon-
drial DNA (mtDNA) haplotypes and to 2 subpopulations based
on microsatellite loci. Both mtDNA and nuclear DNA markers
suggested restricted gene ow among wolf populations
inhabiting northeastern, mideastern, and southeastern Poland
(Pilot et al. 2006; W. Jedrzejewski and coworkers, pers. obs.).
Moreover, the grouping of Polish wolves into mtDNA
subpopulations results from a steep gradient in frequencies of
haplotypes belonging to 2 main haplogroups, which represent a
major subdivision within the worldwide wolf population (Pilot
et al. 2010).
In this paper, based on data on wolf diet collected from the
species range in Poland, we evaluated relationships in wolf
choice of ungulate prey in relation to their relative abundances
in local communities and wolf genetic differentiation among
subpopulations. We tested if differences in diet, if any,
reected species structure of local ungulate communities or
resulted from different hunting specialization; if regional
variation in wolf diet composition conformed to the described
spatial pattern of their neutral genetic variation; and whether
analysis of various sources of information on wolf prey (scats
versus kill remains) and ungulate abundance (ofcial data on
game inventories versus hunting harvest) yielded comparable
results.
MATERIALS AND METHODS
Study areas.Data on wolf prey were collected at 2 spatial
scales: the whole range of Polish wolves during the National
Census of Wolves (Jedrzejewski et al. 2004, 2005), and in 11
sites where detailed local studies were performed (Fig. 1;
Appendix I). Published data from 2 other sites (sites 8 and 13;
see Appendix I) were included in analyses. Sites of local
studies were situated in the northeastern, eastern, southeastern,
and southern parts of Poland (Fig. 1). Areas of studies ranged
from 170 to 8,590 km
2
(Appendix I) and in total covered
22,640 km
2
(7.3% of the country).
Polands area (about 311,900 km
2
; 49800
0
54850
0
N, 14808
0

24809
0
E) is mainly covered by plains, with 91% of the territory
below 300 m above mean sea level. The lowland landscape has
been shaped by glaciations (mainly the Riss, 310,000130,000
years ago, and the W urm, 70,00010,000 years ago).
Mountains, which extend along the southeastern and southern
border of Poland, are part of the Carpathian Mountain range.
Poland is situated in the temperate climate zone of the
Palearctic region, and its climate is characterized by mixed
Atlantic and continental inuences. About 60% of the
countrys area is covered by agricultural land, with a
dominance of arable elds, and smaller amounts of meadows,
pastures, and orchards. Forests cover 29% of the country (Fig.
1). Mean human population density is 124 individuals/km
2
(Statistical Yearbook of the Republic of Poland [Central
Statistical Ofce 2007]).
Most of the local study sites were located in the lowlands
(sites 19), 1 was in the uplands (site 10), and 3 were in the
mountains (sites 1113). Elevation of study sites ranged from
75 to 1,557 m above mean sea level. Sites were characterized
by mixed temperate forest, largely in the form of commercial
stands managed by the State Forestry but also included forests
protected as national parks (Biaowie za, Biebrza, Roztocze,
and Wigry National Parks) and small nature reserves.
Dominant tree species were Scots pine (Pinus sylvestris),
Norway spruce (Picea abies), beech (Fagus sylvatica), and r
(Abies alba), the last 2 species occurring mainly in the
December 2012 1481 JEDRZEJEWSKI ET AL.CHOICE OF UNGULATE PREY BY POLISH WOLVES
mountains. Other trees include oak (Quercus robur), ash
(Fraxinus excelsior), birch (Betula pubescens and B. verruco-
sa), hornbeam (Carpinus betulus), maple (Acer platanoides),
and black alder (Alnus glutinosa).
Mean temperature in July (summer) was from 15.68C to
18.88C, whereas in January (winter) temperature varied from
38C to 5.78C (Appendix I). Annual average temperature was
from 5.58C to 7.38C. Snow cover remained on the ground for
60100 days. Total annual rainfall was 510898 mm. The
ungulate community consisted of 35 species (red deer, roe
deer, wild boar, moose, and European bison). Population
density of wolves in the studied sites ranged from 0.9 to 4.7 (on
average 2.0) individuals/100 km
2
(Appendix I). Data on body
mass of wolves in Poland are available from the Carpathian
Mountains (southeastern Poland). Adult males weighed, on
average, 46.4 kg (SE 0.86 kg, minimummaximum3567
kg) and adult females weighed 39.3 kg (SE 0.70 kg,
minimummaximum2750 kgOkarma 1989). In addition
to wolves, other large predators present in the study areas were
lynx (Lynx lynx) and brown bear (Ursus arctos; Appendix I).
All these predators are protected by law in Poland.
Data collection and analyses.Data on wolf prey remains
(1,913 ungulates and 6 other animals, wild-living species only)
representing the entire range of wolves in Poland were
collected during the National Wolf Census in 20002007.
The census was conducted by the services of State Forests and
National Parks under the supervision of the Mammal Research
Institute, Polish Academy of Sciences, in Biaowie za
(Jedrzejewski et al. 2004, 2005). Year-round, forestry and
national parks personnel recorded wolf tracks, sightings, kills,
and howls. Most records, however, come from winter (with
snow cover). Data were sent to the Mammal Research Institute,
Polish Academy of Sciences, and analyzed using the program
MapInfo 6.5 (MapInfo Corporation 2001). Species structure of
wolf prey in Poland was compared with estimates of game
populations and harvest data, averaged for 2000 and 2005
(after Wawrzyniak et al. 2010) and with data on game
inventories in those state forest districts and national parks
where wolf kills were reported.
In local study sites, wolf diet was investigated on the basis of
feces (scats) and kill remains of wild prey collected between
1998 (in one study1985) and 2006 (Appendix I). Scats and
kills of wolves were detected by snow tracking and walking
along forest roads and paths. Data on wolf kills collected by the
authors were supplemented by information obtained from local
forestry and National Park Services. Most kill remains were
found in winter and were differentiated from carcasses of
animals dead from other causes by presence of wolf tracks and
sign in snow at carcasses. Seasonal variation in diets of wolves
in Poland is small (cf. Jedrzejewski et al. 2000; Nowak et al.
2005), so the dominance of winter-collected scats and kills in
our sample was not likely to affect interpretation of data.
Throughout eastern and southern Poland, 963 scats and 2,335
kills were recorded from the 11 study sites (Appendix I).
Number of kills and scats recorded depended upon sampling
effort, accessibility of the terrain, and length of study period
(Appendix I). Data collected in the 11 sites come from a series
of Master of Science degree projects supervised by WJ or JG,
or both (sites 1 and 2AO, 3 and 4AK, 5aJH, 5bKK,
6MW, 7JK, 9MT, 10MN, 11TK, and 12AJ).
Data on the research projects published by Nowak et al. (2005)
and Jedrzejewski et al. (1992, 2000, 2002), which were
conducted using the same methodology (see sites 8 and 13 in
Appendix I), also were used for the analysis of variation in
wolf diets in Poland.
Analysis of scats was performed according to the standard
method of drying and washing feces through a 0.5-mm mesh
sieve (Goszczy nski 1974; Jedrzejewska and Jedrzejewski
1998; Lockie 1959). Prey were identied by bone remains,
hooves, claws, feathers, and hair (microscopic identication in
the last case) according to taxonomic keys (Debrot 1982;
Dziurdzik 1973; Pucek 1984; Teerink 1991) as well as by
comparison to reference material of Mammal Research
Institute, Polish Academy of Sciences, in Biaowie za. Com-
position of food was expressed as the percentage of fresh
biomass ingested of a particular food component relative to
total biomass consumed by wolves. Biomass of food
FIG. 1.Location of the 3 regions (IIII) inhabited by genetically
distinct subpopulations of wolves delimited based on Pilot et al.
(2006) and W. Jedrzejewski and coworkers (Mammal Research
Institute, Polish Academy of Sciences, pers. comm.) and the 13 sites
of detailed local studies on diet composition and prey preferences of
wolves (Canis lupus). Description of sites is given in Appendix I.
1482 Vol. 93, No. 6 JOURNAL OF MAMMALOGY
components was assessed by multiplying weight of prey
remains found in scats by coefcients of digestibility (obtained
from studies by Goszczy nski [1974], Jedrzejewska and
Jedrzejewski [1998], Jedrzejewski et al. [1992], Lockie
[1961], Lode [1990], and Roger et al. [1991]). The following
coefcients of digestibility were used: ungulates118,
medium-sized mammals50, small mammals23, birds
35, and plant material4. Breadth of the food niche was
calculated using Levins (1968) formula:
B
1
X
p
2
i
; 1
where p
i
is the contribution of group i of wolf prey to the total
biomass of food consumed by wolves. We pooled all prey
species identied into 4 groups: wild ungulates, domestic
ungulates, medium-sized mammals, and others (e.g., small
mammals and plants). Consequently, Levins index (B) could
vary from 1 (strong specialization in 1 group of prey) to 4
(opportunistic preying on all available groups of prey).
Species structure of wild ungulates killed by wolves was
based on kill remains found and the proportion of occurrence
of ungulates alone in scats. Occurrence of a given species in 1
scat was treated as 1 specimen. In cases where deer could not
accurately be classied (Cervidae undetermined), proportions
of red and roe deer in the whole sample were determined based
on proportions of these species among accurately identied
samples. Structure of ungulate communities in all study sites
was estimated using ofcial data on game inventories and
hunter harvest reports, both provided by the local state forest
districts. Unit areas, for which both data had been collected,
were hunting districts of an average size of 59 km
2
. During the
study, no single census method was used consistently in all
areas, and the ofcial data on ungulate numbers were derived
from a combination of various methods of censuses done in
forest districts and national parks (see Wawrzyniak et al. 2010).
The traditional method of censusing ungulates was based on
snow tracking after new snowfall. Observers walked the
sample grid, counted all tracks of ungulates, and mapped their
direction. Difference between number of tracks entering and
leaving a grid cell (usually a forest compartment) gave the
number of animals present in that cell (Jedrzejewska et al.
1997; Pucek et al. 1975). In site 8, drive censuses were done,
that is, counts of animals driven out of a grid cell by a dense
battue line of people. Each censused cell was surrounded by
stationed observers who counted escaping animals (Jedrze-
jewska et al. 1994). In some regions, a method based on snow
tracking and counts of animals observed on censused plots was
applied (Bobek et al. 2001); however, most often inventory
was based on year-round observations gathered by forest
personnel and hunters. Consequently, ofcial statistics are most
probably reliable as far as they document the presenceabsence
and relative abundance of various ungulate species but do not
precisely show actual numbers of species (Wawrzyniak et al.
2010).
The diversity index of the ungulate community and the
ungulates killed by wolves were calculated according to
Levins (1968) formula (see equation 1), where p
i
was the
contribution of each species of ungulate. To assess selection by
wolves of particular species of ungulates, the selectivity index
D (after Jacobs 1974) was calculated as:
D r p=r p 2rp; 2
where r was the fraction of a species among all wild species
killed by wolves and p was the fraction of that species in the
ungulate community. D ranged from 1 (total avoidance of a
species) to 0 (selection proportional to occurrence) to 1
(maximum positive selection). According to Hayward et al.
(2006), sample sizes of .20 scats or kill records do not bias the
values of Jacobs selectivity index. In our study, the smallest
sample was 40 scats from site 1 and 18 kills plus 72 scats from
site 3.
Pearsons correlation coefcient and MannWhitney U-tests
were used to check if the 2 estimates of species structure of the
ungulate communities (ofcial inventories and harvest) yielded
similar results. We also performed pairwise correlations and
MannWhitney U-tests to compare the 2 methods of assessing
the proportion of red deer, roe deer, and wild boar occurrence
in wolf diet (kills versus scats). Linear regression analysis was
applied to see how the proportions of the 3 ungulate species
occurring in wolf diets using the 2 methods (kills and scats)
depended on their occurrence in the community and in hunter
harvests. We 1st checked for outliers (Cooks distance . 1),
which, as highly inuential points, may force regression lines
close to outliers. Only site 12 appeared to be a consistent
outlier in most of the models; we therefore removed it from
further analyses. A G-test for homogeneity of percentages was
used to test for differences in percent species structure of
ungulate communities and wolf kills among the 3 regions.
Overlap of wolf feeding niches (wild ungulate prey only)
was calculated according to Pianka (1973) as:
a
ij

X
p
ia
p
ja

X
p
2
ia

X
p
2
ja
h i
1=2
; 3
where a
ij
was the overlap between region i and j, p
ia
was the
fraction of ath ungulate species in the summed number of
ungulate prey killed by wolves in the ith region, and p
ja
is the
fraction of the ath ungulate species in the summed number of
ungulates killed by wolves in jth region. Values of a range
from 0 (exclusive niches) to 1 (complete overlap). Piankas
index a also was used to calculate similarity of species
structure of ungulate communities among regions. Regional
differences between Piankas indexes were compared with
MannWhitney U-tests. Results were regarded statistically
signicant at P , 0.05.
RESULTS
Ungulate prey of wolves in Poland.The community of
wild-living ungulates was dominated by roe deer (68.9% of
ofcial numbers of ungulates and 49.4% of hunting harvest),
wild boar (15.6% and 36.8%), and red deer (13.8% and 12.9%;
Table 1). These 3 species made up 98.3% of estimated numbers
December 2012 1483 JEDRZEJEWSKI ET AL.CHOICE OF UNGULATE PREY BY POLISH WOLVES
of ungulates and 99.1% of ungulates harvested in Poland.
Among wolf kill remains recorded in the country, 7 species of
wild ungulates (as well as 2 species of nonungulate prey, the
red fox [Vulpes vulpes4 specimens] and the brown hare
[Lepus europaeus2 specimens]) were recorded (Fig. 2; Table
1). Red deer (nearly 60% of ungulates killed) and roe deer
(32.6%) dominated wolf diets, followed by wild boar and
moose. Comparison of wolf prey with species structure of the
ungulate community in Poland showed that red deer was a
preferred species of prey, whereas wild boar was signicantly
avoided (Table 2). Roe deer seemed to be taken less than
expected from their relative abundance, but only 1 of the
comparisons (wolf kills versus estimated population size) was
signicant (Table 1).
Wolf diet composition and choice of ungulate species in 13
local study sites.Analysis of scats showed that wild
ungulates averaged 86.6% (range 68.396.4%) of the total
food biomass consumed by wolves in Poland (Table 2).
Secondary food components, according to their biomass, were
domestic ungulates (mainly cattle and sheep, in total from 0%
to 15.3%,

X 5.2%) and beaver (024.6%,

X 5.2%). The
latter species played an important role in northeastern Poland
(especially in sites 1, 2, and 7; Table 2). Niche breadth B of the
wolf ranged from 1.08 to 1.85 (

X 1.24); thus, wolves in

Poland appeared to specialize on wild ungulates.
Throughout the local study sites, the most numerous species
in the ungulate communities were roe deer (

X 48.3% based
on game inventories and 38.0% based on harvest), red deer (

X
30.5% and 24.7%) and wild boar (

X 18.4% and 37.2%;

Table 3). The 2 estimates of species structure of ungulate
communities yielded highly correlated results throughout all
study sites. For 3 dominant species, the number of harvested
individuals increased with their abundance according to game
inventory data (red deer: R
2
0.98, n 10, P , 0.0001; roe
deer: R
2
0.96, n 10, P , 0.0001; wild boar: R
2
0.97, n
10, P , 0.0001, based on data in Appendix II). For the relative
abundance of ungulates, harvest data yielded a mean
percentage contribution of wild boar twice as high as that
obtained from game inventories (Table 3). For red deer and roe
deer, estimates based on harvest were lower than those based
on game inventories, but the difference was signicant only for
red deer (Table 3).
FIG. 2.Distribution of wolf kill remains in Poland recorded during the National Census of Wolves in 20002007, shown on the background
of the 3 genetically distinct subpopulations of wolves (IIII). Records of kill remains by wolves colonizing central and western Poland also are
shown (scattered points in region IV).
1484 Vol. 93, No. 6 JOURNAL OF MAMMALOGY
Among wild ungulate kills, the species that occurred in wolf
diets most often were red deer (

X 45.6%, range 2181%)

and roe deer (

X 43.9%, range 471%; Table 3; Appendix

II). Based on proportion of ungulate occurrences in scats, roe
deer (

X 49.9%, range 1583%) and red deer (

X 27.1%,
range 458%) were most frequent prey. Wild boar were
found in 9.4% (range 022%) of kill remains and 22.7%
(range 051%) of ungulates recovered from scats. Moose
only occur in northeastern Poland, and were substantially
preyed upon by wolves in 1 site (extensive marshes of Biebrza
National Park, site 6), where they constituted 2426% of all
ungulates killed (Appendix II).
Proportions of red deer, roe deer, and wild boar in wolf diets
assessed by the 2 methods (kills versus scats) were positively
correlated for all species, but the relationships were statistically
signicant in roe deer (r 0.923, n12, P ,0.0005, based on
data in Appendix II) and wild boar (r 0.828, n 12, P
0.001), but not red deer (r 0.448, n 12, P 0.14). If the 2
methods (analysis of kill remains and analysis of scats) would
yield similar results, the relative proportions of each species
from a multispecies community of prey recovered from wolf
scats and found among kills would approach 1:1; however,
searches for kill remains detected large prey (moose and red
deer) more often than small prey. The difference was
signicant in red deer (MannWhitney U-test, n
1
10, n
2

10, P0.019; Table 3). In the case of moose, sample size was
too small for statistical analyses, but in 4 of 6 sites moose were
recorded as kills but not in scats (Appendix II). For roe deer,
scats and kills yielded similar results (Table 3). Wolves
predominantly hunted young wild boar and often consumed
them completely (Jedrzejewski et al. 1992, 2002). Not
surprisingly, wild boar were found in signicantly lower
proportions among kill remains than among ungulates
recovered from scats (Table 3). Average bias from a 1:1 ratio
TABLE 2.Diet composition (percentage of consumed biomass) of wolves (Canis lupus) in eastern and southern Poland based on analysis of
scats. See Fig. 1 for location of regions and study sites and Appendix I for description of sites and sample sizes. Scats were not collected in site 12.
A plus symbol () denotes shares of prey below 0.05%. Food niche breadth B is after Levins (1968).
Prey item
Study site

X 6 SE (n 13)
Region I Region II Region II
1 2 3 4 5a 5b 6 7 8 9 10 11 13
European bison 1.7 0.1 6 0.13
Moose 14.8 0.3 1.1 6 1.06
Red deer 8.6 6.2 1.3 30.5 29.9 10.2 15.9 22.7 35.8 19.5 44.2 42.2 19.1 6 4.13
Roe deer 42.9 27.5 60.4 38.0 23.1 24.4 41.9 30.6 1.7 57.2 57.7 26.6 32.5 33.2 6 4.95
Undetermined Cervidae 8.3 24.5 20.4 38.5 14.9 8.3 19.5 6.2 49.2 13.4 9.5 16.4 17.6 6 3.75
Wild boar 31.2 10.5 6.8 9.4 9.9 18.5 15.6 20.0 1.6 5.8 15.1 4.2 10.6 6 2.35
Total wild ungulates 82.4 71.1 93.8 87.2 78.4 81.1 86.4 68.3 95.6 94.6 96.4 95.4 95.3 86.6 6 2.70
Livestock 0.8 14.0 3.5 3.5 15.3 13.7 4.9 3.1 4.0 0.6 0.8 2.7 5.2 6 1.51
Domestic dog 1.2 1.8 0.3 0.1 1.1 2.1 3.6 0.1 0.8 6 0.31
Wild Carnivora 0.2 0.1 0.1 0.1 0.2 0.2 0.1 6 0.02
Beaver 15.4 13.4 3.8 5.6 4.5 24.6 0.3 5.2 6 2.17
Brown hare 0.2 0.1 0.8 5.3 0.5 5.9 1.8 0.4 0.2 0.8 1.6 1.3 6 0.54
Small mammal 0.1 0.4 0.1 0.2 0.1 6 0.03
Undetermined mammal 7.3 0.6 6 0.56
Birds
Plant material 1.2 0.1 0.1 6 0.09
TABLE 1.Number and percentages of various species of ungulates in the community, hunting harvest, and wolf (Canis lupus) kills in Poland.
A plus symbol () denotes proportions , 0.05%. DJacobs selectivity index. Positive selection or avoidance are statistically signicant at * P
, 0.05, *** P , 0.001 (G-test for homogeneity of percentage). Diversity index B is after Levins (1968).
Species
Wolf kills Estimated population size
D
Annual hunting harvest
D n % n, thousands % n, thousands %
Roe deer 623 32.6 644.5 68.9 0.64*** 153 49.4 0.34
Wild boar 96 5.0 145.5 15.6 0.56* 114 36.8 0.83***
Red deer 1,146 59.9 129 13.8 0.81*** 40 12.9 0.82***
Fallow deer 5 0.3 11.1 1.2 2.4 0.8
Moose 40 2.1 3 0.3
Mouflon 1 0.1 0.2 0.1
European bison 2 0.1 0.6 0.1
Chamois 1 0.1
Sika deer 0.1 0.02
Diversity index B 2.14 1.93 2.52
December 2012 1485 JEDRZEJEWSKI ET AL.CHOICE OF UNGULATE PREY BY POLISH WOLVES
(proportion of the same species among all ungulates found in
scats and as wolf kills) was 1:0.4 in wild boar (median body
mass of boar falling as prey to wolves was 12.5 kg
Jedrzejewski et al. 2002), 1:0.8 in roe deer (20 kg), 1:1.4 in red
deer (62.5 kg), and 1:1.6 in moose (.80 kg).
Proportions of red deer and wild boar in wolf diet increased
with their proportion in the community, regardless of the
method used, although the relationship was statistically
signicant in 8 of 12 pairs of variables: 4 in wild boar (kills
versus community: r
2
0.72, F
1,11
31.64, P , 0.0005; kills
versus harvest: r
2
0.71, F
1,9
25.09, P 0.001; scats versus
community: r
2
0.47, F
1,11
11.79, P 0.006; scats versus
harvest: r
2
0.39, F
1,8
6.77, P 0.03), 2 in roe deer (kills
versus community: r
2
0.48, F
1,11
11.96, P 0.005; kills
versus harvest: r
2
0.34, F
1,9
6.14, P 0.03), and 2 in red
deer (kills versus harvest: r
2
0.45, F
1,9
9.32, P0.01; kills
versus community: r
2
0.27, F
1,11
5.41, P 0.04; Fig. 3).
Jacobs selectivity indexes (D) revealed that wolves
generally positively selected red deer, preyed on roe deer
proportionally to its relative abundance, and avoided wild boar
(Table 3). We found that wolf niche breadth B (both for kills
and scats) grew with increasing diversity of the wild ungulate
community, but the relationship was not statistically signicant
(B for kills versus community: r
2
0.202, F
1,11
2.79, P
0.12; B for scats versus community: r
2
0.141, F
1,11
1.81, P
0.2; data points in Appendix II).
Variation in wolf diet and choice of ungulate prey in relation
to genetic differentiation of wolf population.Data on wolf
kills and ungulate community structure originating from the 2
sources (National Census of Wolves and 13 detailed local
studies) were grouped into 3 regional groups to conform to the
genetic structuring of the Polish wolf population. Records of
kill remains by wolves colonizing central and western Poland
were not used in this analysis because of the small sample size
(n 32). Interestingly, data from National Census of Wolves
and those from 13 local studies yielded a similar picture of
species structure of ungulate communities (G-test for
homogeneity of percentages; G
6
1.092.91, P . 0.1; 3
regions compared) as well as wolf prey composition in the
same regions (G
6
1.776.76, P . 0.05; Fig. 4).
Percentage species structure of ungulate communities
differed signicantly among the 3 regions (G
6
17.98, P ,
0.01 for National Census data and G
6
29.04, P , 0.001 for
values averaged from local studies) as did species composition
of wolf prey (G
6
54.65, P , 0.001 and G
6
29.33, P ,
0.001, respectively). Species structure of wolf kills was more
dissimilar among regions (mean Piankas a 0.87, SE 0.04)
than were ungulate communities (mean a 0.96, SE 0.01)
and the difference between niche overlap (as) for communities
and wolf kills was signicant (MannWhitney U-test, U
31.50, n
1
6, n
2
6, P0.03). In northeastern Poland (region
I), wolves had the widest feeding niche (B 2.682.88
compared to B 1.772.64 in other regions; calculated from
data shown in Fig. 4) and they frequently hunted 2 large
ungulates, red deer and moose. Moreover, wolves had the
largest share of medium-sized mammals (especially beavers;
see Table 2) in diets. In eastern Poland (region II), roe deer
made up the majority of wolf kills (.65%), yet red deer,
despite their small share in ungulate community, were
positively selected (D 0.30 and 0.37). In that region, mean
body mass of wolf prey (46 kg) was markedly smaller than in
the 2 other regions (74.5 kg in northeastern Poland and 79 kg
in southeastern Poland). In southeastern Poland (region III),
wolves were strongly specialized on red deer (more than 50%
of kills, D 0.39 and 0.74). Thus, the regional feeding habits
of wolves diverged more than expected from variation in
available prey.
TABLE 3.Mean (6 SE) percentages of the 3 most common species in communities of ungulates based on ofcial game inventories (Census)
and hunting harvest (Harvest), and among ungulates killed by wolves (Canis lupus) based on analysis of scats and kill remains. Mean values are
calculated for results of local studies presented in Appendix II (only studies with all 4 data sources included). Mean (6 SE) values of food niche
breadth (B) and Jacobs selectivity index D are calculated based on the 4 data sources. B index also includes moose and European bison wherever
present or killed by wolves. D-values are signicantly different from 0 at ** P , 0.01, *** P , 0.001 (MannWhitney U-test performed on
respective percentage shares of prey species in wolf diet and ungulate community, n
1
10, n
2
10; NS not signicant).
Parameter Red deer Roe deer Wild boar Ungulate diversity B
Species structure of the living community of ungulates
Census 30.5 6 3.4 48.3 6 4.5 18.4 6 2.9 2.58 6 0.14
Harvest 24.7 6 4.7 38.0 6 5.2 37.2 6 6.2 2.43 6 0.13
MannWhitney U-test P 0.041 NS P 0.013 NS
Species structure of ungulates killed by wolves
Wolf kills 45.6 6 5.9 43.9 6 6.7 9.4 6 2.4 2.14 6 0.14
Wolf scats 27.1 6 5.0 49.9 6 6.6 22.7 6 4.0 2.33 6 0.16
MannWhitney U-test P 0.019 NS P 0.010 NS
Selectivity indexes D
Censuskills 0.31 6 0.08** 0.13 6 0.11 0.46 6 0.08***
Censusscats 0.11 6 0.14 0.02 6 0.11 0.12 6 0.08
Harvestkills 0.45 6 0.06*** 0.08 6 0.12 0.73 6 0.04***
Harvestscats 0.02 6 0.14 0.23 6 0.11 0.29 6 0.12***
D averaged 0.17 0.05 0.34
1486 Vol. 93, No. 6 JOURNAL OF MAMMALOGY
DISCUSSION
Wolves in Poland, living with multispecies communities of
ungulates, preyed on all native species present and the 4 most
common and widespread ungulates formed a bulk of their diet.
Proportion of these 4 species (red deer, roe deer, wild boar, and
moose) in wolf diets increased with their proportions in the
ungulate communities; however, only 1 prey species, the red
deer, was regularly positively selected.
Our study also demonstrated some methodological problems
that should be considered in studies on the feeding habits of
large carnivores. First, we showed that, if ungulate community
structure is estimated based on hunter harvest, the relative share
of a productive species, such as the wild boar, is higher
compared to low-productive species. This problem was earlier
identied by Wawrzyniak et al. (2010), who reported that wild
boar hunting harvest throughout Poland often exceeds ofcial
estimates of population numbers with no long-term conse-
quences for population size. Yet another source of error
originates from size-related detectability of wolf prey based on
kill remains. The larger the prey, the more likely it is to be
found. Small prey often is eaten completely. Moreover, the
remains of large prey persist in the forest longer. For instance,
Selva et al. (2003) reported that European bison carcasses were
still well visible and utilized by scavengers (including wolves)
FIG. 3.Relationships between percentage of a given ungulate species among all ungulates killed by wolves and the proportion of that species
in the local ungulate community (based on game inventory, denoted as percentage in community; and on hunting harvest, denoted as percentage in
harvest) in the 13 local study sites. Black dots and lines indicate percentage of a given species among wolf kills, and open points and broken lines
indicate percentage of a given species among ungulate remains recovered from wolf scats. Thin dotted line indicates proportion of ungulates in
wolf diet equal to that in a community. See text for correlation coefcients and signicance. Data points can be found in Appendix II.
FIG. 4.Species structure of ungulate communities and wolf kill
remains in 3 regions of Poland (see Fig. 1 for delimitation of regions).
In each pair of bars the left bar shows data averaged from local study
sites, and the right bar presents data collected during the National
Census of Wolves.
December 2012 1487 JEDRZEJEWSKI ET AL.CHOICE OF UNGULATE PREY BY POLISH WOLVES
4 months after a bisons death. This methodological bias
affects not only the results of studies based on traditional
methods (e.g., snow tracking) but also those applying telemetry
techniques (Jedrzejewski et al. 2002). Furthermore, Marucco et
al. (2008), who studied wolf diet in the Western Alps (Italy and
France), found that common species of ungulates were
overrepresented among wolf kills.
To minimize error of estimating wolf diet composition and
prey selection in a multispecies community, we recommend
that both methods of data collecting (search for kill remains
and analysis of scat) are applied together, as was done in this
study. These results can even be combined as proposed by
Jedrzejewski et al. (2002) and Marucco et al. (2008) to obtain a
comprehensive picture of wolf feeding habits. If only 1 source
of data is available and the aim of the research is to assess wolf
prey selection from a multispecies community of ungulates, we
suggest that in addition to being aware of the weaknesses of
each method, data on kill remains should be compared to
community structure assessed by game inventories (both yield
higher estimates of the relative shares of larger species
compared to smaller ones), whereas prey composition derived
from scat analysis can be related to hunting harvest by humans
(both overestimate the shares of small species and highly
productive species).
Wolf diet composition is known to vary over time in
response to changes in ungulate abundance (for the magnitude
of that variation in central and eastern Europe [see Jedrze-
jewska and Jedrzejewski 1998; Sidorovich et al. 2003]). Thus,
for spatial comparison, it is important to consider samples
collected in similar periods. In our study, the biggest temporal
differences occurred between sites 8 (19852000) and 5
(20002006). We believe that such a time span did not affect
the interpretation of data, because, despite some changes in
ungulate numbers, the relative proportion of various species in
ungulate communities remained remarkably stable at both the
local scale (Jedrzejewski et al. 2000) and in the entire country
(Wawrzyniak et al. 2010) during recent decades.
European wolves inhabiting forest ecosystems are known for
their notable plasticity in diet. Okarma (1995), who reviewed
more than 50 studies on trophic ecology of wolves in Europe,
found a signicant positive relationship between species
richness of the local ungulate community and feeding niche
breadth of wolves. Yet, in most studies, red deer (as well as the
introduced sika deer) were preferred prey of wolves. Because
wolves balance difculty in killing prey with the reward in
food biomass obtained, red deer was shown to be the optimal-
size prey for typical central European packs of 46 wolves
(Jedrzejewski et al. 2002). In the whole of Europe, selection of
moose and roe deer varied greatly, but the mean values of the
selectivity index of wolves for these 2 species did not differ
from 0 (Okarma 1995). Wild boar was usually (with few
exceptions) a markedly avoided prey species. Our results are
consistent with that general picture. The novel nding of our
study was that variation in relative abundance of ungulate
species in a community is not the sole factor determining wolf
diet composition. We propose that another factor contributing
to the observed pattern of variation in wolf feeding habits
among regions may result from distinct ecotypes of wolves,
the occurrence of which is reected in the genetic structuring of
their population (see Pilot et al. 2006).
Similar diet-related genetic structuring of animal populations
was reported in North American wolves (Carmichael et al.
2001), killer whales (Orcinus orcaHoelzel et al. 1998), and
arctic foxes (Alopex lagopusDal en et al. 2005). In the last
study, 2 distinct feeding ecotypes of foxes (lemming and
coastal), often reported by ecological studies, appeared
concordant with the genetic structuring of fox populations in
their wide circumboreal range (Dal en et al. 2005). Evidence
that adaptations to distinct ecological niches (food, habitat, and
migration routes) can result in genetic and phenotypic
divergence also has been provided by studies on European
blackcap warblers (Sylvia atricapillaRolshausen et al. 2009)
and wolves in Canada (Musiani et al. 2007). The latter study
showed that barren-ground migratory wolves hunting for
migratory caribou (Rangifer tarandus) and those resident in
boreal forest and hunting for resident prey differ not only in
genetic markers but also in phenotypic features (coat color).
Musiani et al. (2007) concluded that prey-habitat specializa-
tion, rather than distance or topographic barriers to dispersal,
were responsible for the observed divergence of wolf
populations.
In that respect, the situation of Polish wolves described in
our study should be viewed in a wider geographic context. The
3 genetic subpopulations of wolves in Poland are the
westernmost (or northernmost in the case of Carpathian
Mountains) tip of the vast array of subpopulations identied
in central and eastern Europe (see map in Pilot et al. [2006]).
This large-scale genetic differentiation among subpopulations
was correlated with climate, habitat type, and diet composition.
There was substantial gene ow within each subpopulation but
it was restricted among them (W. Jedrzejewski and coworkers,
pers. obs.). Therefore, the specic predatory adaptations of
wolves from different genetic subpopulations may explain why
in our national-scale study, dietary composition of wolves in
regions IIII differed among each other more than expected
when compared to the regional variation in ungulate
communities.
However, it should be kept in mind that all the above cited
papers on diet-related genetic structuring of animal populations
reported on the neutral genetic variation, studied by analyses of
noncoding microsatellite markers or noncoding fragments of
control region of mtDNA, or both. It remains to be studied
whether and how ecological factors also shape the adaptive
genetic variation of the species (see Joost et al. 2007).
Moreover, the spatial pattern of genetic structuring of a
population may vary depending on markers analyzed. Based on
microsatellite markers, Carmichael et al. (2007) detected 8
subpopulations of North American gray wolves, whereas
Knowles (2010), through the use of a large set of single
nucleotide polymorphism markers, found evidence for only 5
subpopulations. Nonetheless, 4 of them corresponded to those
reported by Carmichael et al. (2007) and supported the
1488 Vol. 93, No. 6 JOURNAL OF MAMMALOGY
ecological separation of subpopulations. Thus, the hypothesis
of relationship between feeding habits and adaptive genetic
variation in European wolves should be a subject of future
research.
Finally, our results suggest a need for further study on
preference of wolves for red deer. Based on mtDNA
phylogeography, Polziehn and Strobeck (2002) and Ludt et
al. (2004) suggested that the modern species of Cervus started
their speciation .0.52 million years ago, assisted by recurring
glaciations. Having originated from the area between Kyrgyz-
stan and northern India, red deer colonized the western
Palearctic (western red deer) as well as eastern Palearctic and
North America (eastern red deerLudt et al. 2004). Although
no such comprehensive studies have thus far been done on
wolves, the analysis of mtDNA sequences showed that the
most ancient lineages of extant wolves are found in northern
India and the Himalayas (Sharma et al. 2004), which suggests
that the same region could have been the region of both red
deer and wolf evolution. Evolutionary adaptation of the wolf to
C. elaphus would explain why the red deer is that predators
preferred prey. If so, we could expect that in the entire
Holarctic zone wolves will selectively hunt for C. elaphus,
wherever they occur sympatrically.
ACKNOWLEDGMENTS
The study was nanced by the budget of the Mammal Research
Institute, Polish Academy of Sciences, the General Directorate of State
Forests (Poland), European Natural Heritage FundEuronatur
(Germany), and the Polish Ministry of Science and Higher Education
(grant 2 P06L 00629). MWH was supported by a Marie Curie Transfer
of Knowledge project BIORESC (MTKD-CT-2005-029957) from the
European Commission. SN and RWM also were supported by the
International Fund for Animal Welfare and Wolves and Humans
Foundation. We thank the personnel of all state forest districts and
national parks who helped with eldwork. We are grateful to 4
anonymous reviewers for their comments.
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Associate Editor was Jonathan A. Jenks.
1490 Vol. 93, No. 6 JOURNAL OF MAMMALOGY
APPENDIX I
Description of the sites where wolf (Canis lupus) diet composition and prey choice were studied by the analysis of scats and kill remains.
Numbers of regions and sites refer to Fig. 1. In August ow Forest (site 5) 2 studies were done, hereafter referred to as 5a and 5b.
Site Name
Area
(km
2
)
Mean temperature (8C) Annual
rainfall
(mm)
Snow
cover
(days)
Altitude
(meters
above
sea level)
Wolf
density
(no./
100 km
2
)
Predators
(other than
wolf)
Period
of data
collecting
Sample size
(in numbers)
Annual January July Scats Kills
Region INortheastern Poland
1 Skaliskie Forest 200 6.4 4.1 18.0 648 83 75201 0.9 Lynx 19992004 40
2 Rominta Forest 170 6.4 4.1 18.0 648 83 150256 0.9 Lynx 19992004 84 49
3 Pisz Forest 1,350 5.8 3.0 16.5 627 75 100180 1.0 Lynx 19982003 72 18
4 Napiwoda-Ramuki Forest 1,940 6.2 3.1 16.8 583 94 100225 1.0 19982003 63 46
5 August ow Forest 1,350 6.8 4.2 18.1 569 90 100167 2.4 Lynx a) 20002005 193 284
b) 20042006 180 57
6 Biebrza National Park 640 6.8 4.3 18.8 609 94 100126 2.0 Lynx 20012004 48 35
7 Knyszyn Forest 1,500 6.9 4.0 18.4 592 90 100205 2.6 Lynx 20002002 53 52
8 Biaowie za Forest
a
600 7.1 4.3 18.0 588 83 135202 2.3 Lynx 19852000 1,022 269
Region IIEastern Poland
9 Sobib or Forest 1,910 6.8 4.1 18.0 510 72 125132 2.8 Lynx 20002005 59 38
10 Roztocze 8,590 7.3 4.0 18.8 574 60 125391 2.0 Lynx 20002003 84 39
Region IIISoutheastern Poland (Carpathian Mountains)
11 Bieszczady Mountains 780 7.1 4.3 17.5 847 101 4001,199 4.7 Lynx, bear 19982005 87 1,050
12 Beskidy Mountains 2,190 5.6 5.0 15.6 789 83 2501,262 1.7 Lynx, bear 19992004 667
13 Western Beskidy Mountains
b
1,410 5.5 5.7 16.4 898 79 3001,557 1.6 Lynx, bear 19972001 390 93
a
Data from Jedrzejewski et al. (1992, 2000, 2002).
b
Data from Nowak et al. (2005, 2008).
December 2012 1491 JEDRZEJEWSKI ET AL.CHOICE OF UNGULATE PREY BY POLISH WOLVES
APPENDIX II
Spatial variation in species structure of wild ungulate communities, based on data from game inventories (Census) and hunters records
(Harvest), and species composition of wolf prey, based on kill remains (Wolf kills) and analysis of scats (Wolf scats). Study sites and regions are
as in Fig. 1 and Appendix I. No data on harvest were available for sites 1, 6, and 13; no data on wolf kills were available from site 1; and wolf
scats were not analyzed in site 12. Prey diversity index B is after Levins (1968).
Site
Community or
wolf prey
Percentage of a species in the community or wolf prey
Diversity
index B Moose Red deer Roe deer Wild boar
Region INortheastern Poland
1 Census 2.0 15.0 59.0 24.0 2.33
Wolf scats 48.6 51.4 2.00
2 Census 5.0 37.0 30.0 28.0 3.25
Harvest 20.7 9.6 69.7 1.86
Wolf kills 35.0 43.0 22.0 2.81
Wolf scats 16.5 57.5 26.0 2.35
3 Census 0.3 34.8 53.2 11.7 2.39
Harvest 26.7 42.7 30.6 2.88
Wolf kills 33.0 67.0 1.79
Wolf scats 8.0 83.0 9.0 1.42
4 Census ,0.1 26.9 55.3 17.8 2.44
Harvest 22.4 46.0 31.6 2.76
Wolf kills 43.0 48.0 9.0 2.36
Wolf scats 4.0 75.0 21.0 1.64
5a Census 6.2 33.2 47.4 13.2 2.81
Harvest 19.5 46.5 34.0 2.70
Wolf kills 1.1 33.4 59.5 6.0 2.13
Wolf scats 38.0 41.0 21.0 2.80
5b Census 7.0 25.0 51.0 18.0 2.78
Harvest 19.5 46.5 34.0 2.70
Wolf kills 5.0 54.0 32.0 9.0 2.47
Wolf scats 32.4 41.5 26.1 2.90
6 Census 23.5 20.3 37.5 18.7 3.68
Wolf kills 25.7 51.3 20.1 2.9 2.70
Wolf scats 23.7 23.7 52.6 2.57
7 Census
a
1.6 21.6 53.8 22.7 2.58
Harvest 0.4 19.0 39.7 40.9 2.77
Wolf kills 1.9 42.3 42.3 13.5 2.66
Wolf scats 25.5 41.8 32.7 2.88
8 Census
b
0.4 35.4 20.8 37.8 3.21
Harvest 22.0 9.6 68.4 1.90
Wolf kills 1.1 72.7 4.1 22.1 1.73
Wolf scats 2.5 31.1 15.1 51.3 2.61
Region IIEastern Poland
9 Census 2.9 13.4 66.0 17.7 2.06
Harvest 1.3 11.1 52.5 35.1 2.43
Wolf kills 2.7 21.0 71.0 5.3 1.81
Wolf scats 38.0 53.0 9.0 2.31
10 Census 0.3 24.0 64.7 11.0 2.05
Harvest 20.0 55.6 24.4 2.45
Wolf kills 40.0 54.0 6.0 2.20
Wolf scats 20.0 64.0 16.0 2.10
Region IIISoutheastern Poland
11 Census 53.3 40.5 6.2 2.21
Harvest 65.7 30.8 3.5 1.89
Wolf kills 81.1 17.9 1.0 1.45
Wolf scats 57.6 27.2 15.2 2.33
12 Census 35.2 57.3 7.5 2.18
Harvest 26.7 67.8 5.5 1.87
Wolf kills 41.2 52.5 6.3 2.22
13 Census 21.0 74.0 5.0 1.68
Wolf kills 46.0 50.0 4.0 2.16
Wolf scats 44.0 47.3 8.7 2.35
a
European bison constituted 0.3% of the ungulate community.
b
European bison constituted 5.6% of the ungulate community.
1492 Vol. 93, No. 6 JOURNAL OF MAMMALOGY