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111 2

Nikica ŠPREM, Marina PIRIA, Hrvoje NOVOSEL , Tihomir FLORIJANČIĆ,



ABSTRACT: Between 2007 and 2009, a total of 181 individual wild boar
were scored using nineteen morphological measurements from three geographical
regions to describe morphological variety of the population throughout
Croatia. In some regions we found phenotypical variability of the wild boar
population based on hybridization The results of ANOVA show that some variables
were significantly different (body weight, tail length, trunk length) but
some of them were not homogenous for all age classes (circumference of shin,
the most caudal point of scapula, circumference at chest) and were unable to
highlight differences among the areas. The redundancy analysis (RDA) showed
a connection of sampling sites with some morphological trait. Results of
cluster analysis using TREE procedure indicate separation on the two subpopulations
and suggesting the existence of morphological differences. Overall
the results confirmed that different morphotypes of wild boar are detectable in
some different areas of Croatia, and in some counties the wild boar population
has been hybridized with domestic pigs, which result in phenotypical variability
where the wild characteristics predominate. These results confirmed
the need for population genetic studies to identify the different subpopulations
of wild boar presently found in Croatia

Key words:wild boar, Croatia, morphological variability

The Eurasian wild boar (Sus scrofa L.) is one of the mountains and Mediterranean area.Therefore these re-
most widely distributed terrestrial mammals, a native gions are suitable and have high potential for the study
game of Croatia and economically very important of zoogeographic population characteristics and dynamspecies
which significantly increased in numbers during ics. One of the biggest problems of wild boar population
the last decades.Geographically, Croatia has a specific in Croatia is hybridization with domestic pigs which
position in Europe where in a narrow zone of 150 km most often occurred after the war in the 1990’s (Šprem
three different types of geographical elements are pre-2009).The consequence of this unwanted hybridization
sent, continental-Pannonian basin, continental-Dinaric can be seen in a completely different animal phenotype.
Morphological and quantitative data concerning wild
animals is still scarce and more information is needed. It


Dr. sc. Nikica Šprem, doc. dr. sc. Marina Piria,

is one of the most important phenotypic characteristics

Hrvoje Novosel mag. ing., prof. dr. sc. Tomislav Treer,
University of Zagreb, Faculty ofAgriculture, Department of Fis-of an animal, influencing fitness, life history and popuheries,
Beekeping, Game Management and Special Zoology,

lation ecology (Tymchuk etal. 2006).These data will

Svetošimunska cesta 25, 10000 Zagreb, Croatia

be interesting especially in species with some potential, tel.+385 1 2393 860, fax. +385 1 2315 300


Prof. dr. sc. Tihomir Florijančić, prof. dr. sc. Boris Antunović,
for intensive exploration such as a biological model for
University of J. J. Strossmayer in Osijek, Faculty ofAgriculture,

wild boar (Câmara Filho etal. 2003), and they pro-

Chair forWildlife, Fishery and Beekeeping, Trg Sv.Trojstva 3,

vide information on the growth and development of wild

31000 Osijek, Croatia

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N. Šprem, M. Piria, H. Novosel, T. Florijančić, B. Antunović, T. Treer: MORPHOLOGICALVARIABILITY... Šumarski list br. 11–12, CXXXV (2011), 575-583

boar, as well as on quality of the habitat. Gross external

morphological parameters have been the most common

area used by researchers to attempt to define identifying

characteristics.This is at least in part due to the relative
ease of data collection using these structures compared
to features requiring the use of more complex laboratory
methods. Morphological criteria have included a variety
of structures and parameters including external body
measurements, and coloration. Morphology is an area of
research where the shape and size of a morphological individual
or characteristic is described with quantitative


analysis, and because of very low heritability (h), morphology
characteristics are very dependent on external
conditions (Oxnard 1978). Multivariate statistical
methods were applied to classify morphology differences
and using these methods it was possible to evaluate
the most important characteristics separating the
subpopulations defined by wild boar. Relatively little
data are available on the relationships among morphological
parameters of wild boar.This data mostly dealing
with growing patterns of different parameters were
analyzed and some results were presented on the correlations
and allometry of these characteristics. Better
management of wild boar populations requires more
morphological data. Therefore, the aim of this study
was 1) to determine morphological differences among
wild boars populations located in three Croatian geographical
regions using morphological measurement;
and 2) determine the existence of hybridization.

MATERIALAND METHODS– Material i metode)

From October 2007 to January 2009, sampling of
wild boar populations was conducted in three Croatian
geographical regions, East (Đakovo, Baranja, Bilogora)
Central (Lpolje, Banija) and West (Plominska,
Oprtalj, Grožnjan, SjVeleb, VelKapel, LicSredo) (Fig
1), where we covered three characteristic climate areas
under Köppen’s climatic classification. The Eastern
and Central region enjoys the Cfb climate, but on the
other hand theWestern region is under the influence of
the Cfa and Df climate (Šegota &Filipčić 2003).

Figure 1 Three different geographical regions and sampling sites in the study

Slika 1. Tri različite geografske regije i lokacije uzorkovanja u istraživanju

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Figure 2 RDA ordination of 11 sampling sites and 16 morphological traits a) all age classes, b) juvenile c) yearling d) adults

Slika 2. RDA koordinate 11 lokacija i 16 morfoloških svojstava a) sve dobne skupine, b) mladi c) jednogodišnjaci d) odrasli

and adults showed similar results at the most locations
(Fig 2b, 2c, 2d). Results of cluster analysis usingTREE
procedure indicate separation on the two subpopulations
and suggesting the existence of morphological differences
(Fig 3).The first canonical variable accounts for
84.8, 76.5 and 84.2 percent of the total variance for the
three ages classes respectively, but is unevenly correlated
with the original variables. At first and second
canonical variables for the young shows the highest correlation
with CS, for sub-adults HH and for adults EW,
TLand CC.The discriminant analysis results in the correct
classification of the data in the groups shows that
Central and East populations are correctly classified and
a higher percentage of misclassification can be observed
forWest population. This result is not easily explainable
but the effect of hybridization can be supposed.

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Generally, a climatic, agricultural, and forestry vertical
gradient characterizes the Croatian territory, with a flat
Eastern region of the PannonianValley strongly influenced
by the Drava and Danube Rivers. This terrain
also includes large agricultural fields and predo minantly
oak and beech forests (N 45°43’28,2’’
EO 18°50’24,7’’). A Central hilly region with small
agriculture fields and mainly beech and chestnut
fo rests (N45°15’23,4’’ EO 16°15’26,7’’), the Western
region includes big mountains (N 44°46’28,4’’
EO 15°01’53,9’’)and the typical Mediterranean region
of the Adriatic Sea with small agriculture fields
(N 45°12’59,4’’ EO 14°13’21,5’’). These locations
were used because data of genetic analysis showed similarity
among same sampling sites (Šprem 2009). All
animals presented phenotypic characteristics of the
species. The hunted wild boars were sexed, weighed
and measured. All the animals included in the study
were hunted during the legal drive hunting season.The
age of the animals were estimated using patterns of
tooth eruption and replacement (Boitani &Mattei
1992). The animals were classified into three age
classes (Pedone etal. 1991): juvenile (less than one
year of age), yearling (between 1 and 2 years of age),
and adult (older than two years of age).Wild boars were
measured from the left side, and measurements were
carried out using a Lydthin stick, tape measure and
scale.A total of 181 individuals (97 males and 84 fema les)
from six month to seven years of age were scored
for the nineteen morphological measurements: height at
withers HW; height at sacrum HS; height of back HB;
the most caudal point of scapula mCPS; height at hip
HH; depth of chest DC; circumference at chest CC;
head length HL; ear length EL; ear width EW; tail
length TaL; trunk length TrL; bristles length (at wiethers)
BL; circumference of shin (tibia) CS; circumference
of testis (left) CTL; circumference of testis (right)

The average age of studied animals was under 24
months old, which indicated a relatively young population.
The sex ratio is slightly unbalanced in favor of
males, a similar unbalanced fetal sex ratio was also
found by (Massei etal. 1996). However, the sex ratio
did not attribute to the age or, weight. Morphological
data support the hypothesis that the wild boar populations
in some regions have hybridized with domestic
pigs. Some individuals in the data set displayed white
hair on the feet, stomach, tarsus and carpus; large and
fast growth; great intrapopulation morphological variability;
and, great accumulation of subcutaneous fat.
Mean values with standard deviation of the seventeen
analyzed morphological measurements under three different
geographical zones and three different age groups
of the animals are presented inTable 1.Statistical differ-

CTR; length of trunk with head TrHL; body weight
BW; color of bristles CB. Based on the age class data
where preliminary analyzed withANOVA(using GLM
procedure) for fixed effect of region, gender and their
interaction on each applicably measurement. For measurements
of CTLand CTRANOVAwas tested for effects
of region as fixed factor only on male part of
dataset. Significant effects of region were additionally
tested using Tukey HSD test. After determining variability
with ANOVA, data where merged and cluster
analysis (using CLUSTER procedure byWard method
and Euclidian distances) where preformed for determining
location groupings on averages of 14 relative morphological
measurements. Results of cluster analysis
are shown graphical (usingTREE procedure). Canonical
discriminant analysis (using DISCRIM procedure)
where preformed on region and relative morphological
measurements as given classification.ANOVA, cluster
analysis and canonical discriminate analysis where preformed
in SAS package (SAS Institute, 2007). For distinguishing
which morphological measurement allowed
different morphotypes constrained redundancy analysis
(RDA) by the CANOCO program (Braak & Smilauer
2002) which is used for sound statistical modeling
of ecological data. The Monte Carlo unrestricted
permutation test was performed to determine the significance
of the regression. For RDAanalysis, the measurement
was expressed in percentage of height at
withers.This is a very important parameter in morphological
studies of animals (Melaku 2003). Species
data (response variables) represents morphological me a
surements and environmental data (explanatory variables)
represents dummy variables.

– Rezultati
ences were observed between age classes, results put in
evidence that some variables were significantly different
(BW, TaL, TrL), but some of them were not homogenous
for all age classes (CS, mCPS, CC) and were unable
to highlight differences among the regions.
Additional effects of gender on differences between regions
were recorded on traits (BW, EL, TaL, TrL) only
in sub-adult age class.The correlation analysis shows a
significant link between measurements and age classes.
Results of RDA analysis throughout sampling sites
based on 16 morphological traits are showed in Fig 2.
There is a strong correlation between CB in Central and
West populations with the CS and TrL. West and East
populations are connected with HLand BL, but Central

and East populations are strongly correlated with HH,
EWand mCPS (Fig 2a).Analysis of young, adolescents

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Mean +SD of morphological measurements of wild boar from 3 regions in Croatia and F-values with probability for effect of region, gender and region-gender
interaction on morphometric traits

Tablica 1.Srednja vrijednost+SDmorfološkihmjerenjadivljihsvinjaiz3 regijeuHrvatskoji F-vrijednosti s vjerojatnošću utjecaja regije, spola i interakcije regija-spol
na vrijednost morfološke osobina

Morphological measurementMorfološke mjere East/Istok Central/Središnji West/Zapad Region/RegijaF-value (P>F)
Gender/SpolF-value (P>F)
Region*GenderRegija*SpolF-value (P>F)
Young/MladiNumber/Broj 47 32 15
Body weight/Tjelesna težina 31.032±7.955A 37.325±10.826 B 39.1±6.574 B 7.06 (0.0014) 0.32 (0.5742) 2.13 (0.1249)
Height at withers/Visina u grebenu 61.2±6.883A 64.656±7.196AB 67.54±7.271 B 5.25 (0.007) 0.28 (0.5967) 0.29 (0.747)
Height of back/Visina leđa 60.534±6.598 63.631±8.576 65.473±7.899 3.2 (0.05) 1.42 (0.23) 0.26 (0.773)
Height at sacrum/Visina križa 59.63±6.626 61.609±8.898 62.933±6.781 1.32 (0.2682) 0.41 (0.526) 0.19 (0.83)
The most caudal point of scapula/
Visina prednje noge od vrha plećke do tla57.053±6.435 58.541±6.924 59.6±9.216Height at hip/Visina kuka 57.272±6.447 56.847±7.478 56.8±8.809 0.04 (0.9647) 0.45 (0.5029) 0.22 (0.8068)
Depth of chest/Dubina prsiju 33.134±3.916A 33.856±6.608A 38.013±2.995 B 5.04 (0.0085) 0.08 (0.7771) 1.19 (0.3105)
Circumference at chest/Opseg prsiju 73.726±7.562 79.356±10.918 81.42±5.841Head length/Dužina glave 51.461±3.468 50.169±6.555 53.236±7.422 2.12 (0.1257) 5.32 (0.0234) 3.05 (0.05)
Ear length/Dužina uške 11.066±1.492 11.288±1.732 11.48±1.629 0.47 (0.6259) 0.12 (0.7319) 2.18 (0.1196)
Ear width/Širina uške 10.191±1.253 9.838±1.2 9.427±1.125 2.23 (0.1135) 0.47 (0.4943) 0.11 (0.8945)
Tail length/Dužina repa 17.326±3.784A 19.191±3.609AB 16.247±3.352 B 4.27 (0.0169) 0.06 (0.7994) 0.82 (0.4419)
Trunk length/Dužina trupa 68.287±7.83A 75.434±7.63 B 80.94±4.791 C 19.48 (<0.0001) 0.31 (0.5763) 0.51 (0.604)
Bristles length (at wiethers)/
Dužina čekinja na grebenu10.128±1.65A 11.141±2.329 AB 12.38±1.35 B 9.12 (0.0003) 0.22 (0.6373) 3.08 (0.05)
Circumference of shin (tibia)/Opseg cjevanice 11.496±1.103 13.122±1.469 13.32±1.348
Circumference of testis (left)/
Opseg lijevog testisa6.06±1.74 7.30±5.64 5.8±0.64 1.45 (0.2543)
Circumference of testis (right)/
Opseg desnog testisa6.23±1.91 7.43±5.71 8.22±0.98 1.84 (0.1802)
Sub-adults/ Srednjedobni jednogodišnjaciNumber/Broj 10 12 10
Body weight/Tjelesna težina 75.9±5.77A 86.13±10.613 B 66.625±8.878 C 15.36 (<0.0001) 0.88 (0.3568) 5.25 (0.0095)
Height at withers/Visina u grebenu 79.358±3.628 77.49±3.899 74.317±6.577 2.83 (0.0761) 0.28 (0.5979) 1.32 (0.2823)
Height of back/Visina leđa 78.017±4.317A 74.8±3.747AB 71.075±4.874 B 6.9 (0.0037) 0.15 (0.6974) 1.34 (0.2784)
Height at sacrum/Visina križa 74.383±4.873 71.97±3.549 69.65±5.364 2.62 (0.0903) 0.01 (0.9086) 0.22 (0.8059)
The most caudal point of scapula/
Visina prednje noge od vrha plećke do tla72.533±5.852 67.88±6.238 64.725±5.96Height at hip/Visina kuka 75.3±4.416A 65.45±4.351 B 61.942±6.043 B 22.23 (<0.0001) 1.87 (0.1829) 0.37 (0.6925)
Depth of chest/Dubina prsiju 46.417±1.676A 42.52±4.879AB 41.717±6.446 B 4.08 (0.0279) 0.59 (0.4496) 3.16 (0.0577)

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Morphological measurementMorfološke mjere East/Istok Central/Središnji West/Zapad Region/RegijaF-value (P>F)
Gender/SpolF-value (P>F)
Region*GenderRegija*SpolF-value (P>F)
Circumference at chest/Opseg prsiju 106.483±6.989 106.77±8.133 91.2±11.193Head length/Dužina glave 52.22±5.209 49.89±3.525 50.173±3.522 1.14 (0.333) 0.17 (0.6858) 0.67 (0.5193)
Ear length/Dužina uške 12.2±1.134A 14.72±2.68 B 12.367±0.657A 11.9 (0.0003) 4.26 (0.0485) 20.94 (<0.0001)
Ear width/Širina uške 11.833±1.096 A 11.77±1.523 A 9.85±0.703 B 12.09 (0.0002) 1.18 (0.2872) 0.73 (0.4907)
Tail length/Dužina repa 22.492±6.106A 23.95±3.287A 17.55±1.973 B 10.44 (0.0004) 0.3 (0.5862) 8.41 (0.0014)
Trunk length/Dužina trupa 91.167±4.747 92.57±7.6 92.025±7.079 0.08 (0.9224) 2.76 (0.1076) 7.73 (0.0021)
Bristles length (at wiethers)/
Dužina čekinja na grebenu10.692±1.151 11.77±0.7803 10.467±2.955 1.27 (0.2956) 1.12 (0.2995) 0.24 (0.7908)
Circumference of shin (tibia)/Opseg cjevanice 10.692±1.151 11.77±0.78 10.467±2.955Circumference of testis (left)/
Opseg lijevog testisa6.06±1.74 7.30±5.64 5 8.0±0.64 1.45 (0.2543)
Circumference of testis (right)/
Opseg desnog testisa6.23±1.91 7.43±5.71 8.22±0.98 1.84 (0.1802)
Adults/OdrasliNumber/Broj 14 20 21
Body weight/Tjelesna težina 89.607±8.242A 111.445±13.685 B 110.524±18.612 B 12.49 (<0.0001) 0.08 (0.7768) 1.56 (0.2194)
Height at withers/Visina u grebenu 84.593±3.689A 86.155±5.133A 90.986±4.406 B 9.24 (0.0004) 0.10 (0.7521) 0.88 (0.4205)
Height of back/Visina leđa 81.586±4.757A 84.27±9.463AB 87.41±4.45 B 3.55 (0.0365) 0.16 (0.6877) 0.63 (0.5346)
Height at sacrum/Visina križa 78.907±5.44A 80.425±5.337A 84.562±6.496 B 6.05 (0.0045) 1.19 (0.2806) 2.32 (0.1093)
The most caudal point of scapula/
Visina prednje noge od vrha plećke do tla78.45±5.403 78.655±5.564 79.21±6.546Height at hip/Visina kuka 78.007±4.296 75.585±6.402 76.257±6.449 0.4 (0.6712) 0.5 (0.4813) 2.29 (0.1117)
Depth of chest/Dubina prsiju 49.807±2.724 48.445±9.481 50.2±4.341 0.72 (0.4918) 0.69 (0.4099) 1.21 (0.3068)
Circumference at chest/Opseg prsiju 112.086±6.715 118.145±10.787 115.648±14.287Head length/Dužina glave 51.714±3.799 53.05±4.686 51.774±3.156 0.83 (0.4404) 0.15 (0.6956) 1.96 (0.152)
Ear length/Dužina uške 13.407±3.14 14.025±2.081 14.714±2.11 0.56 (0.5758) 0 (0.99) 0.46 (0.6316)
Ear width/Širina uške 12.143±2.982A 13.3±2.056A 10.081±0.743 B 10.68 (0.0001) 2.18 (0.1463) 0.35 (0.7092)
Tail length/Dužina repa 23.457±4.085A 26.19±4.635AB 21.652±4.206 B 6.15 (0.0041) 3.33 (0.0739) 1.02 (0.3693)
Trunk length/Dužina trupa 100.457±8.814A 102.105±7.58AB 109.676±13.645 B 3.78 (0.0298) 1.08 (0.304) 1.43 (0.2484)
Bristles length (at wiethers)/
Dužina čekinja na grebenu11.014±0.687 A 10.67±2.253A 14.495±2.082 B 18.54 (<0.0001) 0.63 (0.432) 0.08 (0.9211)
Circumference of shin (tibia)/Opseg cjevanice 14.821±1.126 16.205±1.15 17.833±2.455Circumference of testis (left)/
Opseg lijevog testisa16.27±3.36 14.22±5.38 16.97±5.24 0.51 (0.6076)
Circumference of testis (right)/
Opseg desnog testisa16.52±3.78 14.84±5.60 16.90 ±5.12 0.09 (0.9139)

Regions with same letter are not significantly different – Regije sa istim slovom nisu značajno različite

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SAŽETAK: Između 2007. i 2009. godine, ukupno 181 jedinki divljih svinja,
koristeći devetnaest morfoloških mjera sa tri geografske regije, korišteno je za
opis morfološke raznolikosti populacije diljem Hrvatske. U nekim regijama
pronašli smo fenotipsku varijabilnost populacije divljih svinja temeljenu na
hibridizaciji. Rezultati ANOVA-e pokazuju da su neke varijable bile značajno
različite (masa tijela, dužina repa, dužina rila), ali neke od njih nisu homo-
gene za sve dobne skupine (opseg cjevanice, visina prednje noge od vrha plećke
do tla, opseg prstiju). Multivarijatna statistička metoda (RDA) pokazala
je povezanost lokaliteta s nekim morfološkim osobinama. Rezultati klaster
analize pomoću TREE postupka ukazuje na razdvajanje dviju podpopulacija i
sugerira postojanje morfoloških razlika. Ukupni rezultati potvrđuju da su različiti
morfotipovi divljih svinja detektirani u različitim područjima Hrvatske, a
u nekim regijama populacija divljih svinja je hibridizirala s domaćim svinjama,
što ima za posljedicu fenotipsku varijabilnost gdje ipak karakteristike
divljih svinja prevladavaju. Ovi rezultati potvrđuju potrebu za populacijsko
genetskim istraživanja kako bi identificirali različite podpopulacije divlje svinje
koje trenutno obitavaju u Hrvatskoj.

Ključne riječi:divlja svinja, Hrvatska, morfološka varijabilnost

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Figure 3 Results of cluster analysis (usingTREE procedure)

Slika 3. Rezultat klaster analize (koristeći TREE postupak)

Population structure is similar to the cited in studies
in Europe,America and Oceania (Herrero &Fernandez
deLuco 2003) approximately 70% of individuals
were under 24 months old and 6 animals were over
72 months. Morphology data in adult animals can enable
one to determine if a specimen in question resembles a
pure wild boar or a hybridized animal with domestic
pigs, but data presented in this study supported the hypothesis
that the wild boar populations have hybridized
with domestic pigs. In some regions of Croatia we found
phenotypical variability of the wild boar population
based on hybridization, but results suggested that the
wild characters predominate. Results for some morphological
measurement were different than in others studies
(Martinoli et al. 1997;Herrero&Fernandezde
Luco 2003; Mayer & Lehr Brisbin 2006). For
example, TaL and HW were smaller, but EL was the
same compared to data presented byMayer &Lehr
Brisbin (2006).The data forTrHLwere smaller than
was shown by Martinoli et al. (1997). Data from this
study revealed that the CC, HL and BW was much
higher than fromHerrero &Fernandez deLuco
(2003). Some of these morphological measurements
supported the hypothesis that the wild boar population
has hybridized with domestic pig, and the same hypothesis
was given in similar studies (Martinoli et al.
1997; Herrero & Fernandez de Luco 2003). In
generally standard morphological analyzes did not show
clear area of separation in any morphological trait, but
on the other hand, RDAanalysis showed the connection
of sampling sites with some morphological trait.The re

– Diskusija
sults of cluster analysis using TREE procedure show
that East and Central populations are morphologically
very close and belong to one subpopulation, but the
West population belongs to another subpopulation. This
argument can explain on the basis of different habitat
and climate conditions.Western population is under the
influence of Mediterranean climate and mountain region
constitute a natural barrier between other populations.In
some respects this result also confirm hypothesis of the
existence of two European subpopulations, western and
central populations(Larson etal. 2005).Eigen values
were performed and we can be confirmed that body
shape analysis using multivariate statistical methods
may be useful in the evaluation of conformation and
other applications.When all parameters are analyzed, it
can be concluded that wild boar from Croatia present a
certain degree of variability. These results confirmed
that different morphotypes of wild boar are detectable
in some different regions of Croatia. These morpho-
types are differentiated on the basis of height and
length measurements and can be relevant only for older
animals. Presently the subpopulation is the accepted
way of giving formal recognition to these differences,
the origin of which may have been an adaptation to different
geographical situations.The average data values
presented here are slightly higher than those estimated
for other European wild boar populations (Pedone et
al. 1991; Ernhaft &Csányi 1995; Amici et al.
2010). However, it should be noted that a large number

of the studies reported by various authors from other
countries (Randi et al. 1987; Apollonio et al.

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N. Šprem, M. Piria, H. Novosel, T. Florijančić, B. Antunović, T. Treer: MORPHOLOGICALVARIABILITY... Šumarski list br. 11–12, CXXXV (2011), 575-583

1988;Genovetal. 1995; Tinellietal. 1999), refer cupied by wild boar we can concluded that the popula

to variables not directly comparable with those presented
in this study. On the basis of morphological
characteristics, the western Croatian wild boar population
fits the description of the Italian wild boar (Amici
et al. 2010). Because of the large geographic range oc-

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