POLISH JOURNAL OF ECOLOGY

(Pol. J. Ecol.)

60

629634

2012

Short research contribution

Michal BAL1*, Mria BALOV1

1

Department of Biology and Ecology, Pedagogical faculty, Catholic University, Hrabovsk cesta 1,
03401 Ruomberok, Slovakia, *e-mail: michal.balaz@ku.sk (corresponding author)

DIVERSITY AND ABUNDANCE OF BIRD COMMUNITIES

IN THREE MOUNTAIN FOREST STANDS:
EFFECT OF THE HABITAT HETEROGENEITY

ABSTRACT: The structural composition of

the forest habitats is considered to be one of the
most important factors affecting the breeding
bird community composition. Structurally more
homogenous forests are usually characterized by
lower number of bird species and their densities.
This study presents results of a comparison of
the breeding bird communities in three Carpathian mountain forests, and the effect of the forest
composition on breeding bird species richness
and density. There are the bird communities occurring in semi-natural forests: natural mixed
forest (NMF) spruce-beech forest situated ca.
1000 m a.s.l. and UFZ-spruce forest in the upper
forest zone (UFZ) up to 1510 m a.s.l., and the
managed spruce forest (MSF) at altitude which is
similar to NMF. The territory mapping method
was used to estimate the number of breeding
species and their densities during years 2004
2006. The species richness as well as the density
of breeding bird communities were found to be
higher in the semi-natural habitats (33 breeding
species; 64.6 breeding pairs 101 ha in NMF and
28 breeding species; 57.6 breeding pairs 101 ha
in UFZ) than in the managed habitat (21 breeding species; 53.6 breeding pairs 101 ha). This
was true even for the high-elevated study plot
despite the fact that the altitude is usually negatively correlated with both the species richness
and density.

journal 31 v02.indb 629

KEY WORDS: forest structure, bird species

richness, Carpathians, habitat quality
The structure of a bird community is
highly influenced by composition and configuration of its habitat. The major impact of
vegetation pattern on breeding bird assemblages is considered to be one of the most important issues of the avian ecology (Ma c Art hu r and Ma c Ar t hu r 1961, C o dy 1985).
In woodlands, which are among the best
studied habitats in respect to the bird richness
(e.g. Hanss on 1994, E denius and Sj b erg
1997), the differences are often found between the plots of different size or age of tree
species as well as between the interior and exterior parts of the forest fragments. The higher values of bird diversity and density are usually found in natural forests in comparison
with the managed ones (e.g. S an i ga 1995,
B asht a 2007). Generally the natural primeval forests are considered as habitats with the
highest species richness and guild diversity
(e.g. Gowaciski and Prof us 1992, S an i ga 1995, B onci na 2000). In this study the
breeding bird communities in two Carpathian primeval forest habitats in Western Tatras
Mountains were analyzed and compared with
breeding community in a managed spruce

2012-10-12 08:43:36

630

Michal Bal, Mria Balov

forest of the same area to test the effect of the

forest composition on species richness and
density of the bird breeding communities.
The research was carried on in the Carpathian forest region (N Slovakia). The studied localities were characterized by different
forest structure. Two of them represented
semi-natural plots (natural mixed forest
NMF and upper forest zone UFZ), while
the third one represented a managed forest
(managed spruce forest MSF). Although
the distance between NMF and MSF was only
about one km, their floristic and structural
composition of the vegetation was largely different (Table 1). NMF could be characterized
as a primeval spruce-beech forest with relatively high tree species and structural diversity situated at the altitude of about 1000 m a.s.l.
UFZ study plot represents natural spruce
community situated on the border of mountain and subalpine zone at the altitude up to
1510 m a.s.l. On the other hand, MSF represents a typical even-aged managed forest with
very low tree species diversity situated at the
altitude very similar to NMF (Table 1).
The breeding bird communities of the
selected plots were studied during the years
20042006. The combined version of the
mapping method (Tomi a loj 1980) was
used to estimate population densities. Species
diversity (H) and evenness (J) were measured by Shannon-Wiener index of diversity:

where pi is proportion of i-species and S is

number of species, and t-test was used to test
the differences.

Similarity of species composition of bird

communities was measured by qualitative
(QS) and quantitative (CN ) Srensen index:

where A and B are the numbers of species

in samples A and B, respectively and C is the
number of species shared by the two samples,
jN is sum of the lower of the two abundances for
species found in both communities, na is total
number of individuals in community A and nb
is total number of individuals in community B.
Rarefaction method was used for better
display of species richness in all localities using the calculator program (Brzustowsk i
2005) to perform calculations. Structural
characteristics of the studied plots were measured within a 30 m radius from 12 points
evenly distributed throughout all study plots.
The cover of vegetation layers is a mean of five
measurements taken from five different points
within a 30 m radius. Amount of fallen and
standing dead wood was subjectively estimated on a nine-degree scale. Vegetation characteristics of the study plots were compared using ANOVA with Tukeys post-hoc tests.
Structural composition of vegetation differed among all three study plots, however,
MSF showed the lowest similarity in structural characteristics (Table 1). This study plot
was characterized by the lowest tree species
diversity as well as by the lowest vegetation
cover in lower vertical layers.
In total, 48 bird species were found
within the three study plots during the years

Table 1. Structural characteristics of three forest stands under study.

Co_decid cover of deciduous trees, Co_con cover of coniferous trees, Co_00.3 cover of the vegetation layer 00.3 m above the ground, Co_0.31 cover of the vegetation layer 0.31 m above the
ground, Co_13 cover of the vegetation layer 13 m above the ground, amount of fallen (Dw_fall) and
standing death wood (Dw_st). NMF nature mixed forest, MSF managed spruce forest, UFZ upper
forest zone.
Forest
Altitude
NMF
1020-1090
MSF
920-1030
UFZ
1330-1510
P
NMF vs MSF
NMF vs UFZ
MSF vs UFZ

journal 31 v02.indb 630

Co_decid
44.9
1.4
2.1
<0.001
<0.001
<0.001
0.989

Co_con
26.1
91.4
50
<0.001
<0.001
0.119
0.005

Co_00.3
0.8
0.2
0.6
0.002
0.002
0.502
0.019

Co_0.31
0.5
0.1
0.5
0.011
0.023
0.998
0.021

Co_13
0.3
0.1
0.5
0.019
0.387
0.198
0.015

Dw_st
2.3
0.6
1.6
0.011
0.009
0.041
0.121

Dw_fall
4.3
1.1
3
0.007
0.005
0.309
0.112

2012-10-12 08:43:36

Impact of habitat structure on breeding bird communities

631

Table 2. Characteristics of breeding bird communities in three study forest stands in Carpathians. Density average number of pairs per 10 ha., dom. % dominance. S(50) and S(90) number of breeding species
in equal abundance 50 and 90 individuals; H index of diversity and J index of evenness. NMF
nature mixed forest, MSF managed spruce forest, UFZ upper forest zone; + pair breeding on the
edge of study plot, its density is not included in the overall estimate; non-breeding species.
Species
Fringilla coelebs L.
Periparus ater (L.)
Erithacus rubecula (L.)
Regulus regulus (L.)
Sylvia atricapilla (L.)
Phylloscopus trochilus (L.)
Prunella modularis (L.)
Troglodytes troglodytes (L.)
Turdus torquatus L.
Certhia familiaris L.
Phylloscopus collybita (Viellot)
Lophophanes cristatus (L.)
Regulus ignicapillus (Temminck)
Turdus philomelos C. L. Brehm
Phylloscopus sibilatrix (Bechstein)
Pyrrhula pyrrhula (L.)
Sitta europaea L.
Ficedula parva (Bechstein)
Turdus merula L.
Columba oenas L.
Anthus trivialis (L.)
Picoides tridactylus (L.)
Poecile montanus Baldenstein
Dendrocopos major (L.)
Columba palumbus L.
Dendrocopos leucotos (Bechstein)
Dryocopus martius (L.)
Picus canus Gmellin
Accipiter gentilis (L.)
Strix uralensis Pallas
Lyrurus tetrix (L.)
Carduelis spinus (L.)
Garrulus glandarius (L.)
Phoenicurus phoenicurus (L.)
Cuculus canorus L.
Aquilla chrysaetos (L.)
Tetrastes bonasia (L.)
Buteo buteo (L.)
Corvus corax L.
Falco tinnunculus L.
Glaucidium passerinum (L.)
Scolopax rusticola L.
Loxia curvirostra L.
Ficedula albicollis (Temminck)
Muscicapa striata (Pallas)
Aegithalos caudatus (L.)
Coccothraustes coccothraustes (L.)
Turdus viscivorus L.
Total
S(50)
S(90)
H
J

journal 31 v02.indb 631

NMF
density
dom.
17.8
28
10.3
16
5.6
9
4.4
7
3.8
6
+
0.6
<1
2.8
4
0.7
1
2.2
3
1.8
3
1.7
3
1.7
3
2.4
4
1.7
3
0.3
<1
1.7
3
2.1
3
0.3
<1
1.4
2

0.8
+
0.3
0.3
0.3
(1)+
(1)+

MSF
density
15.5
9.1
6.2
5.7
1.4
+
0.5
1.9
1.4
3.3

1.4
1.9
1.4
1
1
+

3
4
3
2
2

dom.
14
8
12
5
9
14
10
3
7
0.7
5
4
<1

0.8
0.4

1
<1

<1

0.8

2
<1

1.2
+
0.4

1
0.5

<1

0.4

<1

1
<1
<1
<1

+
+

+
+

64.6
22.1
28.7
2.61
0.81

<1
4
3
6

UFZ
density
8
4.7
6.9
2.8
5.1
8.2
5.9
2
3.9
0.4
3.1
2.4
0.4
+

0.5

+
+

dom.
29
17
12
11
3

(1)+
+

+
+

+
+
+

53.6
16.2
20.1
2.17
0.81

57.6
19.3
24.7
2.72
0.87

2012-10-12 08:43:36

Michal Bal, Mria Balov

Number of species

632
35
30
25
20
15
10
5
0

NMF
UFZ
MSF

50
100
Number of individuals

Fig. 1. Rarefaction (species-individuals) curves

for three study forest stands in Carpathian Mts:
NMF nature mixed forest, UFZ upper forest
zone, MSF managed spruce forest.

20042006. The highest number of the species

was observed in UFZ, while the lowest species richness was recorded in MSF (Table 2).
Altogether, 39 bird species were found to
breed directly in one of the study habitats
at least and 8 species breed constantly in all
of them. The highest number of breeding
species and the highest density of breeding
community were found in NMF. The lowest
number of the breeding species and the lowest density was observed in MSF (Table 2).
MSF was also characterized as a habitat with
the lowest species richness by the rarefaction
species-individual curves (Fig. 1). There was
found no difference in biodiversity between
NMF and UFZ (P> 0.05, t = 0.57, df = 126).
On the other hand, MSF differed significantly
from both NMF (P <0.05, t = 2.19, df = 108)
and UFZ (P <0.01, t = 2.72, df = 105). The
similarity of communities structure reached
values in the range from 0.7 to 0.78 (qualitative index) and from 0.51 to 0.74 (quantitative index). The lowest similarity was found
between MSF and UFZ and between MSF
and NMF in qualitative and quantitative index, respectively. There were nine species detected as breeding species only in NMF. Six of
them: Strix uralensis Pallas, Tetrastes bonasia
(L.), Picus canus Gmelin, Dryocopus martius
(L.), Dendrocopos leucotos (Bechstein) and
Ficedula parva (Bechstein) are species of European importance. (according to NATURA
2000 criteria). Four species were registered
only in UFZ and three of them Glaucidium
passerinum (L.), Lyrurus tetrix (L.) and Phoenicurus phoenicurus (L.) are species of Euro-

journal 31 v02.indb 632

pean importance. The spruce plantation in

MSF hosted no exclusive breeding species in
respect to NMF and UFZ. The comparison of
foraging guilds among localities revealed that
NMF is the most valuable habitat for bird diversity. Although canopy and ground feeders
were the most abundant groups in all studied
forest habitats, NMF was characterized by the
highest number of foraging specialist groups.
Breeding species which belonged to the airspace foragers and predators were detected
only in NMF. On the other hand, MSF as the
habitat with the lowest structural heterogeneity, and UFZ as the plot situated in higher
altitude, showed lower diversity of foraging
specialists (Fig. 2).
As expected, what is already known from
literature (e.g. B onci na 2000, Te ws et al.
2004), these results show strong effect of the
forest composition on the breeding bird
communities. The habitats more resembling
the unaffected natural conditions host bird
communities with higher diversity and density than do managed and changed habitats.
Species composition and density of breeding
pairs in NMF corresponded to what had already been known about breeding bird communities from other parts of the Carpathian
mountain forests which are usually inhabited by 3050 species with the density 6070
breeding pairs 101 ha (e.g. Ko c i an 1981,
Ki e 1991, Kropi l 1996, Kor an 2004).
Bird community in another study plot with
the semi-natural habitat, UFZ, is affected by
the occurrence of the plot in the upper forest zone which itself could have had some effect on the community composition. Usually
only low numbers of bird species could be
found to breed in the upper mountain zone
of the central European mountains, with
the density varying from 23 to 60 breeding
pairs 101 ha (Koz l owsk i 1974, Top e rce r 1989, G owaciski and Prof us 1992,
li z ows ki 1991, Wi nd i ng et al. 1993). The
density of the bird community in the plot
with the managed forest habitat, MSF, was
found to correspond to the most Carpathian spruce forest communities (ca. 30 to 50
breeding pairs 101 ha) (e.g. Kr it n 1990,
B asht a 2007), and also to other Central European mountain communities (e.g. D y rc z
1973, F l ous ek 1989, Wi nd i ng et al. 1993).
On the other hand, the species richness was

2012-10-12 08:43:36

Impact of habitat structure on breeding bird communities

25
Number of species

NMF
20

UFZ
MSF

15
10
5
0
BF

CF

AF

GF

Fig. 2. Bird foraging guilds (BF bark feeders,

CF canopy feeders, AF airspace foragers, GF
ground feeders, P predators) in three study
forest stands in Carpathian Mts: NMF nature
mixed forest, UFZ upper forest zone, MSF
managed spruce forest.

lower in comparison with most of recorded

data for such a type of habitat (21 in comparison to 1350; D y rcz 1973, Kr it n 1990,
Winding et al. 1993, B asht a 2007). This
could presumably be attributed to the effect
of the forest structure. This forest was characterized by underdeveloped shrub level and
young even aged trees, attributes preventing
the understory and interior species (especially the hole-nesters) from occurrence in
higher species number.
Taking all this together, species richness,
number of breeding species, density of breeding pairs and the diversity of foraging guilds
were highest in the semi-natural study plot
NMF. Forests which are heterogeneous in
their structure, are characterized by higher
number of ecological niches and consequently by higher bird species diversity and density (B onc ina 2000, Te w s et al. 2004). In
summary, the presented results show strong
influence of habitat structural heterogeneity
on bird species composition. The managed
spruce monoculture characterized by low values of most of the important parameters important for breeding birds offers less suitable
conditions than the upper forest zone and the
primeval spruce-beech forest. As diversity
and density of bird communities are generally
highly dependent on vertical and horizontal
diversity of the forest offering high variety of
suitable microhabitats, both these character-

journal 31 v02.indb 633

633

istics usually decrease with increasing habitat

uniformity (B onc i na 2000, B asht a 2007).
The fact that the recorded diversity and
abundance was significantly higher in the
study plots with semi-natural habitat than in
the managed forest despite that one of the former ones was situated at considerably higher
altitude (300600 m), suggests that the habitat composition and structure resembling the
natural conditions could compensate the negative effect of altitude on breeding bird communities. This builds a framework hypothesis
which could be tested on a broader altitudinal range of habitats with different levels of
change in their natural composition on the
parameters of breeding bird communities.
ACKNOWLEDGEMENTS: We would like
to thank David Jandzik, Martina Juroviov and
two anonymous reviewers for their helpful comments. This study was partially supported by the
grant VEGA 2/0068/10.
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Received after revision April 2012

journal 31 v02.indb 634

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