Environmental Management (2015) 56:643–652
DOI 10.1007/s00267-015-0528-7
Do Birds Avoid Railroads as Has Been Found for Roads?
Jarosław Wia˛cek1 • Marcin Polak1 • Maciej Filipiuk1 • Marek Kucharczyk1
Janusz Bohatkiewicz2
•
Received: 17 November 2014 / Accepted: 29 April 2015 / Published online: 12 May 2015
Ó Springer Science+Business Media New York 2015
Abstract The construction of railway lines usually has a
negative effect on the natural environment: habitats are
destroyed, collisions with trains cause deaths, and the noise
and vibrations associated with rail traffic disturb the lives
of animals. Cases are known, however, where the opposite
holds true: a railway line has a positive effect on the fauna
in its vicinity. In this study, we attempted to define the
influence of a busy railway line on a breeding community
of woodland birds. Birds were counted using the point
method at 45 observation points located at three different
distances (30, 280, 530 m) from the tracks. At each point,
we determined the habitat parameters and the intensity of
noise. In total, 791 individual birds of 42 species were
recorded on the study plot. Even though the noise level fell
distinctly with increasing distance from the tracks, the
abundance of birds and the number of species were the
highest near the railway line. Moreover, insectivorous
species displayed a clear preference for the vicinity of the
line. The noise from the trains did not adversely affect the
birds on the study plot. The environmental conditions
created by the edge effect meant that the birds preferred the
neighborhood of the tracks: the more diverse habitats near
the tracks supplied attractive nesting and foraging niches
for many species of birds. Trains passing at clear intervals
acted as point sources of noise and did not elicit any
& Jarosław Wia˛cek
[email protected]
1
Department of Nature Conservation, Institute of Biology and
Biochemistry, Maria Curie-Skłodowska University,
Akademicka 19, 20-033 Lublin, Poland
2
Department of Road and Bridge, Faculty of Civil Engineering
and Architecture, Lublin University of Technology,
Nadbystrzycka 40, 20-618 Lublin, Poland
negative reactions on the part of the birds; this stands in
contrast to busy roads, where the almost continuous flow of
traffic in practice constitutes a linear source of noise.
Keywords
birds
Railway Noise Edge effect Woodland
Introduction
Railway lines are an important element of any modern
transport network. Rail transport, especially the high-speed
variety, is becoming of increasing significance in both
developed and developing countries (Clewlow et al. 2014).
Along with expanding urbanization, the development of
railway networks usually takes place to the detriment of the
natural environment (Glue 1971), although one can also
give examples of its positive effects (Van Rooyen 2009;
Morelli et al. 2014). The construction of new railway lines
involves the destruction of natural habitats and leads to
collisions between animals and trains as well as considerable mortality among birds resulting from their flying into
electrical catenaries (Wells et al. 1999). Disturbances from
noise, train lights, the infrastructure’s lighting installations,
and the vibrations caused by passing trains can all disrupt
the life of animals at different stages of their life cycle
(Van Rooyen 2009). Collisions resulting from birds flying
into the catenaries and other cables supplying current to
electrified lines are of particular concern, and some groups
of birds are highly susceptible to such collisions (Van
Rooyen and Ledger 1999; Anderson 2001). Birds that fly
slowly or that are incapable of rapid flight maneuvres, as
well as young, inexperienced birds often perish in collisions with vehicles and trains (Glue 1971). The victims of
this type of collision tend to be migrants (especially at
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644
night), nomadic species and birds that move around in
flocks (Van Rooyen 2009). The noise caused by construction work and later by passing trains impairs the
breeding of birds, especially when territories are being
defended and during incubation (Morelli et al. 2014). This
can cause birds to abandon their occupied territories, nests,
and broods. The considerable mortality among some bird
species resulting from collisions with trains or the electrical
infrastructure along railway lines is due to their foraging in
the vicinity of the tracks (Barbaro et al. 2014). It is usually
raptors that hunt along lines of communication, but insectivores and granivorous forage there as well (Orłowski
2008; Van Rooyen 2009; Halfwerk et al. 2011).
On the other hand, transportation corridors, running as
they do through different habitats, can also create edge
effects, thereby increasing biodiversity (Delgado et al.
2007; Morelli et al. 2014). Environmental corridors are
areas rich in food for both birds of prey (rodents and small
passerines) and insectivorous and granivorous species
(Polak et al. 2013; Morelli et al. 2014; Wia˛cek et al. 2015).
Delgado et al. (2007) found that the abundance of food
along the edge of a forest through which a road or railway
line runs is due to the special microclimate and habitat
conditions prevailing at the boundary between woodland
and a transportation corridor. Those authors point out that
the more favorable temperature and insolation along a
woodland margin, and consequently the richer species
structure there, can extend from a few to more than a dozen
meters into the woodland. This relatively narrow strip of
terrain along the edge of woodland offers far superior
living conditions for invertebrates (Vermeulen 1994; Vermeulen and Opdam 1995) and rodents, and consequently
for the birds foraging on these groups of animals (Morelli
et al. 2014). Such conditions imply that despite the risk of
colliding with vehicles/trains and of having to function in a
noise-polluted environment, birds willingly hunt along and
nest in the vicinity of transport routes so as to shorten the
distance to food-rich foraging sites (Kuitunen et al. 2003;
Palomino and Carrascal 2007). Elements of railway infrastructure like electrical catenary supports are excellent
song posts, look-out posts, and resting sites (DeGregorio
et al. 2014). Woodland margins and the vegetation along
railway lines offer good nesting sites, and the open areas by
the tracks facilitate foraging (Morelli et al. 2014). All these
factors have a positive effect on birds and may be the
reason why birds are present in larger aggregations near
railway lines than deep in a forest.
The chief objective of our study was to find out whether
the neighborhood of a railway line carrying a good deal of
traffic had any influence on the abundance and number of
species of birds nesting in the forest through which the line
passes. Another aim was to check whether the noise of
passing trains had any effect on birds belonging to the
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Environmental Management (2015) 56:643–652
various guilds associated with foraging (insectivorous or
granivorous), nest placement (on the ground or high up in
the trees), and singing frequency (low, medium, or high).
The available data on the effect of road traffic noise on
woodland birds suggest that this reduces the numbers of
birds and the numbers of species near the road (Reijnen and
Foppen 1994; Reijnen and Foppen 2006; Palomino and
Carrascal 2007). Some results indicate that birds belonging
to different guilds (those building their nests near the
ground and having a low singing frequency) occur in
smaller densities by roadsides (Dooling and Popper 2007;
Polak et al. 2013). Other figures imply that the greater
complexity of the woodland structure and the richer food
resources available by the side of a road running along a
forest margin (the edge effect) favor the presence of greater
densities of birds there, despite the nuisance of the road
noise and the risk of collisions with vehicles (Kuitunen
et al. 2003; Summers et al. 2011; Wia˛cek et al. 2014). The
basic objective of our research was to examine that birds
avoid railroad as has been found for paved roads.
Study Area and Methods
The study plot was located in the Puławy Forest District (N
51°500 0200 , E 21°910 9400 ) (belonging to the Regional Branch
of the State Forestry Administration in Lublin, eastern
Poland). Lying on slightly undulating terrain with dune
hillocks, the dominant habitat in the plot is pine forest with
predominant Scots pine (Pinus sylvestris) and a small admixture of silver birch (Betula pendula). The study was
carried out along the double-track railway line No. 7
(Warsaw–Lublin–Dorohusk) between Puławy and De˛blin,
near the village of Gołab. The intensity of traffic along this
line is 81 trains in 24 h—48 passenger trains and 33 freight
trains. The railway line and its accompanying infrastructure
run through the forest in a 50-m wide corridor. The study
plot was sited on the north side of the tracks.
The number and distribution pattern of woodland birds
were determined using the point-count method (Bibby et al.
1992). The study was done at 45 observation/listening
points in three rows of 15 running parallel to the railway
(Fig. 1). All the points were established and recorded on
GPS receivers in April 2014 before the start of the actual
counting. The first row of 15 points (henceforth called
A-points) lay 30 m from the railway, the next row of 15
points (B-points) was situated 280 m from the line, and the
last row of 15 points (C-points) ran at a distance of 530 m
from the tracks. All the points in a row were 250 m apart
from one another. Counting at each point lasted for 5 min.
All the birds seen and heard within a radius of 100 m were
recorded, but birds flying over the study plot were disregarded. Three counts were done at each point—on 23 April
Environmental Management (2015) 56:643–652
645
Fig. 1 The study plot with the point-count locations (45 black dots) and noise level isolines during the breeding period near a railway in Puławy
Forest (eastern Poland)
2014, 20 May 2014, and 17 June 2014. All the counts were
done from 05:30 to 08:30 h. The limiting factor in this type
of study was the fact that the observers themselves sometimes had difficulty in hearing the birds above the noise of
passing trains, so some may have gone unrecorded
(Rheindt 2003; Summers et al. 2011). Nevertheless, we
were aware of these limitations during our point counts and
tried to minimize them. Our task was made a little easier
because the railway traffic was not continuous, and we
were able to record the birds’ vocal activity in the intervals
between trains.
The counts at all the points on 1 day were performed by
three experienced observers (JW, MF, and MP), who
walked parallel to one another, each along a different row
of points (A–C). During each successive count, the observers walked along a different row of points. Before the
counts began, the study plot had been selected very carefully in order to reduce to an absolute minimum the effect
of environmental parameters on bird clustering. To minimize the edge effect, the plot was located in the depths of a
large, dense forest complex (Fig. 1). The division into
feeding, nesting, and bioacoustic guilds was based on the
publications by Tomiałojć et al. (1984), Cramp and Perrins
(1994), and Polak et al. (2013). The noise of trains and the
vegetation structure at every listening point were assessed
with the aid of environmental parameters (see Table 1).
Two different approaches were applied to determine the
model of the propagation of noise across the study plot: (1)
direct measurements of sound levels during counts and (2)
acoustic map. The level of train noise was measured at all
the points during every count in April, May, and June using
CHY 650 digital sound level meters (IEC 651-1979 type 2,
ANSI S1.4-1983 type 2, JIS C 1502). The noise level at the
center of each point was measured for 5 min and the
highest level was recorded. Train noise was measured on
weekdays in comparable and stable weather (no rain and/or
high winds). In addition, the propagation of train noise in
the study area was modeled to produce an acoustic map
(Fig. 1) using the following measurement instruments:
sound level meters (Svantek type SVAN 945A, type SVAN
955, and type Brüel and Kjaer 2238), microphones (type
40AN, type ACO 7052A, and Brüel and Kjaer type 4188)
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Table 1 The habitat variables obtained at the observation/listening points
Variables
Meaning
Tree age (years)
Age of tree stand at the point-count location (years)
Canopy cover (%)
% Canopy cover in 11 categories: 0, 0 %; 10, 1–10 %; 20, 11–20 %; 30, 21–30 %; 40, 31–40 %; 50, 51–60 %; 60,
61–70 %; 70, 71–80 %; 80, 81–90 %; 90, 91–99 %; 100, 100 % within circle (radius 30 m)
Tree height (m)
Mean height of five trees growing within circle (radius 30 m) and measured by the altimeter
Number of tree
species
Number of species of the nearest 30 trees that were [20 cm diameter at breast height
Number of deciduous
trees
Number of deciduous trees of the nearest 30 trees that were [20 cm diameter at breast height
Number of dead trees
Number of dead trees that were [20 cm diameter at breast height within circle (radius 50 m)
DBH (cm)
Diameter at breast height
Number of shrub
species
Number of shrub species and/or young trees (\20 cm diameter at breast height) growing within circle (radius 30 m)
Shrub cover (%)
% Shrub cover in 11 categories: 0, 0 %; 10, 1–10 %; 20, 11–20 %; 30, 21–30 %; 40, 31–40 %; 50, 51–60 %; 60,
61–70 %; 70, 71–80 %; 80, 81–90 %; 90, 91–99 %; 100, 100 % within circle (radius 30 m)
Herb cover (%)
% Herb cover in 11 categories: 0, 0 %; 10, 1–10 %; 20, 11–20 %; 30, 21–30 %; 40, 31–40 %; 50, 51–60 %; 60,
61–70 %; 70, 71–80 %; 80, 81–90 %; 90, 91–99 %; 100, 100 % within circle (radius 30 m)
Herb height (cm)
Mean height of herb vegetation at five places chosen randomly within circle (radius 30 m)
and an acoustic calibrator (RION type NC-74). Measurements were made at six points, two each at each distance
class from the railway, from which the noise level at each
observation point was calculated. The field studies were
done using the sampling method: three 5-min measurements were made at each point, which were then averaged
logarithmically. The measurements were made in good
weather without rainfall or wind speeds in excess of
3.0 m/s. For calculating the noise of railway traffic, we
used the French National Method of Calculation (1995).
This is the standard way of measuring traffic noise in
Poland and other EU countries. However, it takes no account of propagated sound being absorbed by vegetation. In
view of this, sound propagation was calculated according to
the Polish standard PN-ISO 9613-2:2002 Acoustics. For the
model calculations, we used SoundPLAN, version 7.2. In the
present analyses, parametric tests were applied after
checking whether the distribution was consistent with the
normal distribution (Kolmogorov–Smirnov test). A bilateral
critical region was assumed in the tests, and results were
deemed significant if the probability of committing an error
of the first kind was equal to or less than 0.05. Redundancy
analysis (RDA) was used to analyze the relationship between numbers of particular bird species and distance from
the railway—this parameter was used as the environmental
variable. The Monte Carlo test with 500 permutations was
used to determine the significance of canonical axes. Only
species with abundance C10 individuals were included in
RDA. Analysis of variance (ANOVA) was used to assess
noise propagation over the study plot and to determine the
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differences in species richness and numbers in specified
categories of observation points. The measure of species
richness was taken to be the sum of all species recorded
during the three counts, and the number of birds was the sum
total of all individuals discovered during all three counts.
Multiple regression analysis was performed to test the impact of the seven environmental parameters on the abundance and number of species. Therefore, multivariate
ANOVA (MANOVA) was conducted to test for variation in
avian ecological groups at points located at different distances from the railway. The means are given together with
their standard deviations ± SD. The computations were
performed using STATISTICA 6.0 (Statsoft, Inc., 2001) and
Canoco 4.0 (ter Braak and Smilauer 1998).
Results
Environmental Parameters
The mean noise intensity during the counts was
73.2 ± 20.1 dB (range 36.1–96.8 dB; n = 45) at the
A-points, 57.3 ± 8.5 dB (42.8–77.6 dB; n = 45) at the
B-points, and 51.1 ± 8.5 dB (31.0–67.5; n = 45) at the
C-points. There were significant differences in noise
propagation between the point categories during the counts
in May (Fig. 2; ANOVA; F2,42 = 26.33; P \ 0.0001) and
June (F2,42 = 15.79; P \ 0.0001), but not in April
(F2,42 = 1.09; P = 0.346). Figure 1 shows the acoustic
map and the model of noise propagation across our study
Environmental Management (2015) 56:643–652
647
species was Chaffinch (Fringilla coelebs), 37 % of the
birds counted. The dominants (C5 %) also included Great
Tit (Parus major) and European Robin (Erithacus rubecula). The numbers of the most common birds ([10 individuals) differed widely in relation to distance from the
railway (Fig. 3; Monte Carlo test of the significance of the
first axis; F ratio = 3.137; P \ 0.05; Monte Carlo test of
the significance of all axes; F ratio = 2.13; P \ 0.05).
There were no significant differences between the mean
abundance of birds (all individuals/point) in April:
6.4 ± 2.2 (range 2–10; n = 45), May: 5.7 ± 2.1 (range
1–10; n = 45), and June: 5.5 ± 2.8 (range 0–13; n = 45;
ANOVA; F2,132 = 1.89; P = 0.16). The mean number of
birds at the A-points was 18.7 ± 4.4 (range 13–29;
n = 15) and was similar to the numbers at the B-points
(16.3 ± 5.5; 9–30; n = 15) and C-points (17.9 ± 3.9;
16–24; n = 15). The number of individual birds/point of
all the species recorded was unrelated to distance from the
railway line (ANOVA; F2,42 = 1.11; P = 0.34). Multiple
regression analysis indicated that the only one of the seven
environmental parameters (in all cases P [ 0.128) influencing abundance (all individuals/point) was herb cover
(B = 1.415; SE = 0.255; P \ 0.001).
plot. Meticulous analysis of the environmental parameters
showed that the area on which the birds were counted was
homogeneous with respect to habitat (Table 2). Four of the
11 habitat parameters differentiated the study area. The
number of tree species, number of deciduous trees, and
herb cover decreased with distance from the railway line
and only canopy cover increased with distance from the
line.
Numbers of Individuals
During the three counts, a total of 791 birds belonging to
42 species (Table 3) were recorded. The most numerous
Numbers of Species
The number of species/point was the highest near the
railway line (ANOVA; F2,42 = 3.89; P \ 0.05; Fig. 4).
The mean number of species at the A-points, lying closest
to the tracks, was 10.2 ± 3.2 species (range 6–15; n = 15)
and differed significantly from the number at the B-points
(7.5 ± 2.3; range 5–14; n = 15). In the row of C-points,
the number of species was 8.5 ± 2.3 (range 5–13; n = 15).
Herb cover was the only environmental factor (multiple
regression; in all cases P [ 0.169) influencing species diversity (B = 0.700; SE = 0.171; P \ 0.001). There were
Fig. 2 Relationship between A-weighted railway traffic noise (dB) at
point-count locations and distance from the railway in April, May,
and June. The arrow shows significant differences between points
(Tukey’s test, *P \ 0.001)
Table 2 Vegetation at the
point-count locations in relation
to distance from the railway (Apoints: 30 m, B-points: 280 m,
C-points: 530 m)
Variables
A-points
B-points
C-points
Tree age (years)
55
55
55
0.090
Canopy cover
5
7
7
12.550
P
H2,45
ns
\0.005
Tree height (m)
13
14.5
13.9
3.133
ns
Number of tree species
2
1
1
8.624
\0.05
Number of deciduous trees
2
0
0
10.189
\0.01
Number of dead trees
4
3
2
1.875
ns
DBH (cm)
19.9
19.0
19.6
0.239
ns
Shrub cover
3
3
3
2.726
ns
Density of shrubs
115
40
120
2.726
ns
Herb cover
Herb height (cm)
3
32.6
1
23.2
1
20
7.130
4.879
\0.05
ns
Data presented as median values. Differences between points were tested by Kruskal–Wallis test
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Table 3 Forest bird community composition in relation to distance from railway (A-points: 30 m, B-points: 280 m, C-points: 530 m) in eastern
Poland
Species
Guilds
Total number
Number of individuals (%)
A-points (n = 15)
B-points (n = 15)
C-points (n = 15)
105
Fringilla coelebs
Hn, G, Hf
295 (32, 3)
88
102
Parus major
H, I, Hf
70 (8, 8)
33
29
8
Erithacus rubecula
Sylvia atricapilla
Ln, I, Hf
Ln, I, Mf
51 (6, 4)
34 (4, 3)
14
14
11
6
26
14
Anthus trivialis
Ln, I, Hf
34 (4, 3)
10
15
9
Oriolus oriolus
Hn, I, Mf
32 (4, 2)
8
6
18
Garrulus glandarius
Hn, G, Mf
31 (3, 9)
7
7
17
Lophophanes cristatus
H, I, Hf
25 (3, 1)
7
6
12
Lullula arborea
Ln, I, Mf
23 (2, 9)
15
1
7
Periparus ater
H, I, Hf
22 (2, 8)
7
7
8
Turdus merula
Hn, I, Hf
21 (2, 6)
9
7
5
Phoenicurus phoenicurus
H, I, Mf
17 (2, 3)
8
5
4
Cuculus canorus
Ln, I, Lf
17 (2, 3)
7
7
3
Dendrocopos major
H, G, Mf
16 (2, 0)
7
6
3
Phylloscopus collybita
Ln, I, Mf
11 (1, 4)
4
4
3
Carduelis spinus
Hn, G, Mf
11 (1, 4)
2
4
5
Phylloscopus trochilus
Ln, I, Hf
9 (1, 1)
7
1
1
Turdus philomelos
Turdus viscivorus
Hn, I, Mf
Hn, G, Mf
8 (1, 0)
8 (1, 0)
3
4
2
1
3
3
Columba palumbus
Hn, G, Lf
8 (1, 0)
5
3
0
C. coccothraustes
Hn, G, Mf
7 (0, 9)
4
2
1
Poecile montanus
H, I, Hf
6 (0, 7)
3
3
0
Phylloscopus sibilatrix
Ln, I, Hf
4 (0, 5)
2
0
2
Serinus serinus
Ln, G, Hf
3 (0, 4)
0
1
2
Cyanistes caeruleus
H, I, Hf
3 (0, 4)
2
0
1
Ficedula hypoleuca
H, I, Hf
3 (0, 4)
0
2
1
Certhia familiaris
H, I, Hf
2 (0, 25)
0
1
1
Sylvia communis
Ln, I, Hf
2 (0, 25)
2
0
0
Emberiza citrinella
Ln, I, Hf
2 (0, 25)
2
0
0
Upupa epops
H, I, Lf
2 (0, 25)
1
0
1
Loxia curvirostra
Hn, G, Mf
2 (0, 25)
2
0
0
Corvus corax
Hn, R, Mf
2 (0, 25)
2
0
0
Buteo buteo
Hn, R, Hf
1 (0, 12)
1
0
0
Regulus regulus
Luscinia luscinia
Hn, I, Hf
Ln, I, Mf
1 (0, 12)
1 (0, 12)
0
1
0
0
1
0
Dryocopus martius
H, I, Mf
1 (0, 12)
0
0
1
Hippolais icterina
Hn, I, Mf
1 (0, 12)
0
1
0
Dendrocopos minor
H, I, Mf
1 (0, 12)
0
0
1
Accipiter nisus
Hn, R, Mf
1 (0, 12)
0
1
0
Sylvia nisoria
Ln, I, Mf
1 (0, 12)
1
0
0
Lanius collurio
Ln, I, Mf
1 (0, 12)
1
0
0
Gallinago gallinago
Ln, I, Lf
1 (0, 12)
0
0
1
283
241
267
Total
791
The explanation of birds classification according to nesting, foraging, and bioacoustics guilds: H, hole-nesters; Hn, high nesters; Ln, low nesters;
R, raptorial; G, granivorous–insectivorous; I, insectivorous; Hf, high-frequency singers; Mf, medium frequency singers; Lf, low-frequency
singers. The percentages shown in parentheses
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649
Fig. 3 Ordination diagram of redundancy analysis (RDA) with the
most common bird species recorded in Puławy Forest (eastern
Poland) explained by the point-count locations in relation to distance
from the railway. Abbreviations of species names include the first
three letters of the genus and specific names
Fig. 4 Relationship between number of species at point-count
locations and distance from the railway. The arrow shows significant
differences between points (Tukey’s test, *P \ 0.001)
no significant differences between the mean number of
species in April: 4.3 ± 1.6 (range 2–9; n = 45), May:
4.3 ± 1.7 (range 1–8; n = 45), and June: 3.7 ± 2.3 (range
0–11; n = 45; ANOVA; F2,132 = 1.40; P = 0.25).
Ecological Guilds
We thus used MANOVA to test for the effects of the
railway line on the proportion of individuals in different
ecological groups. Species foraging on invertebrates preferred the proximity of the railway (MANOVA;
F4,126 = 4.54; P \ 0.005; Fig. 5a). There was an increase
with distance from the railway in the number of highnesting species (F4,126 = 3.33; P \ 0.05; Fig. 5b) and
Fig. 5 a Foraging (G granivorous–insectivorous, I insectivorous,
R raptorial) guilds in relation to the distance from railway. Vertical
lines indicate 95 % confidence intervals. b Nesting (Hn high nesters,
Ln low nesters, H hole-nesters) guilds in relation to the distance from
railway. Vertical lines indicate 95 % confidence intervals. c Bioacoustic (Hf high-frequency singers, Mf medium frequency singers, Lf
low-frequency singers) guilds in relation to the distance from railway.
Vertical lines indicate 95 % confidence intervals
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650
birds using high-frequency calls for communication
(F4,126 = 5.78; P \ 0.001; Fig. 5c).
Discussion
In general, most of papers demonstrate the decline in
numbers and species in the vicinity of roads carrying heavy
traffic (Reijnen and Foppen 1994; Reijnen and Foppen
2006; Polak et al. 2013). However, the results of this study
indicate that plot railway traffic did not lower the abundance
of birds or the number of species using the area near the
tracks. Most papers assessing the effect of tarred roads on
birds, i.e., analyzing mainly the effect of road noise, but also
disturbances due to vehicle lights, environmental pollution,
or the presence of humans, demonstrate that there is a distinct drop in both the abundance of birds and the numbers of
bird species in the vicinity of a busy road (Reijnen and
Foppen 1994; Rheindt 2003; Reijnen and Foppen 2006;
Palomino and Carrascal 2007; Benitez-López et al. 2010;
Polak et al. 2013; Wia˛cek et al. 2015). One of the principal
reasons for the decrease in the numbers of birds and species
is road noise (McClure et al. 2013). The mean levels of
noise emitted by passing trains measured at the observation/
listening points showed that they are similar to the average
levels of road noise; moreover, these levels decrease distinctly with increasing distance from the road or railway
line (Polak et al. 2013). But the effects of such noise on
birds are quite different in these two instances. The abundance of birds at the observation/listening points closest to
the railway tracks was higher than that along the other two
rows of points deeper in the forest, although these differences were not statistically significant. In contrast, the
abundance of birds close to a busy road usually rises with
increasing distance from the road, which would suggest that
a road adversely affects the birds living near it (Reijnen and
Foppen 1994; Reijnen and Foppen 2006; Polak et al. 2013).
Likewise, the number of bird species detected by the railway line was significantly larger than at the points within
the forest, which suggests that rail traffic did not have a
negative effect on birds (Fig. 4). In their study of bird
densities along railway lines in Tibet, Li et al. (2010) obtained results similar to ours—they found more birds on the
transects situated closer to the tracks. The larger numbers of
birds near the tracks is due, among other things, to the
railway infrastructure. Electrical catenary supports and the
catenaries themselves make excellent song or look-out
posts, or resting posts safe from the attentions of predators
(Morelli et al. 2014). The open terrain along the tracks also
provides good foraging for various guilds of birds hunting
along transportation routes (granivorous, insectivores, raptors) (Barbaro et al. 2014; Morelli et al. 2014). During our
study, we found species preferring ecotone environments:
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Environmental Management (2015) 56:643–652
Raven (Corvus corax), Buzzard (Buteo buteo), Red-backed
Shrike (Lanius collurio), Barred Warbler (Sylvia nisoria),
Whitethroat (Sylvia communis), and Yellowhammer (Emberiza citrinella) were sighted only along the tracks. The
same applied to granivorous birds like Jay (Garrulus
glandarius), Hawfinch (Coccothraustes coccothraustes),
and Wood Pigeon (Columba palumbus), and insectivores
such as Great Tit (P. major), Blackbird (Turdus merula),
Willow Warbler (Phylloscopus trochilus), Great-spotted
Woodpecker (Dendrocopos major), and Woodlark (Lullula
arborea), most of which also displayed a distinct preference
for the vicinity of the railway (Table 3). The abundance of
food available along woodland margins by the side of
railway lines or roads may also be due to the special habitat
conditions and microclimate prevailing there (Delgado
et al. 2007; Barbaro et al. 2012). Many papers dealing with
the edge effect highlight the different environmental conditions obtaining along forest margins (Barbaro et al. 2014;
Batary et al. 2014). These differences are due primarily to
the superior insolation and the consequent slightly higher
temperatures there (Delgado et al. 2007). Such conditions
promote the growth of invertebrates and rodents, and this
attracts birds that feed on these groups of animals (Vermeulen 1994; Vermeulen and Opdam 1995; Morelli et al.
2014). The greater amount of light at the edge of a forest
means better growth of the ground and shrub layers, as well
as a generally higher number of species of flora compared
with the forest interior: more suitable habitat conditions are
thus created for birds nesting along a woodland margin
(Šálek et al. 2010; Halfwerk et al. 2011). Our results appear
to confirm this, as the only environmental factor significantly affecting the abundance of birds and the numbers
of species was the herb cover. Moreover, the larger numbers
of tree species in general and of deciduous species in particular, together with the lower canopy density along the
first line of points (A-points) provided a superior habitat for
birds (Table 2). The greater amount of light reaching the
floor of the forest along its edge promoted the growth of a
denser ground layer, which in turn supplied richer food
resources for the birds, especially the insectivorous species.
Foraging Guild
According to our results, there were more insectivorous
species near the railway tracks (Fig. 5a). The more diverse
and richer vegetation structure in the ecotone along a forest
margin provides an optimal living habitat for many invertebrate species (Helle and Muona 1985; Vermeulen 1994),
which constitute the staple diet of many insectivorous
birds. Similar results were obtained during studies performed in France and New Zealand, where the rich resources of food available along woodland margins attracted
insectivorous species to that part of the forest (Barbaro
Environmental Management (2015) 56:643–652
et al. 2014). Similar results were obtained from a study in
deciduous woodland in central Germany, where the greater
specific diversity of trees and the edge effect led to larger
numbers of birds being found along the forest edge (Batary
et al. 2014).
Nesting Guild
The noise from fairly busy roads adversely influences birds
that build their nests on the ground or just a small distance
above it (Polak et al. 2013). Noise propagates along railway tracks in much the same way as it does along tarred
roads. But we did not observe any such interaction on our
study plot (Fig. 5b). The lack of a negative reaction to train
movements may be due to the nature of the noise source:
the noise from trains passing once every 5–15 min does not
have such an adverse effect as the almost continuous noise
coming from passing motor vehicles on the road. The same
lack of reaction was found in hole-nesting birds, which
occupied tree holes near the tracks or deeper in the wood
with equal frequency. Likewise, such birds did not react to
road noise, regardless of whether they were nesting in
natural tree holes or in nesting boxes (Polak et al. 2013;
Wia˛cek et al. 2014).
Masking of Signals by Noise
The noise level decreased with increasing distance from the
tracks (Fig. 2), but this in itself did not raise the numbers of
individual birds or species (Fig. 4). Some authors have
pointed out that road noise can mask signals biologically
important for birds, like song (Brumm 2004; Brumm and
Slabbekoorn 2005). This applies especially to species that
make low-frequency calls, which are thus masked by traffic
noise (Dooling and Popper 2007). It has been shown that
species like Cuckoo (Cuculus canorus), Wood Pigeon,
Turtle Dove (Streptopelia turtur), and some woodpeckers
avoid the vicinity of noisy transport routes (Polak et al.
2013). Other authors have confirmed this, expressing the
view that species emitting low-frequency calls avoid the
neighborhood of a road to prevent their biological signals
from being masked (Trombulak and Frissell 2000; Brumm
2004; Halfwerk et al. 2011). Our results regarding train
noise do not support this hypothesis. Species making lowfrequency calls were present in larger numbers near the
railway line (Wood Pigeon, Cuckoo), or else were observed
both near the tracks and within the forest (Hoopoe Upupa
epops) (see Table 3). Generally speaking, species with
low-frequency calls did not specially avoid the railway line
(Fig. 5c). The lack of a clear reaction among this group of
birds to noise can be put down to the short time during
which a single passing train can be heard and the clear
intervals of relative quiet in between trains. This is in
651
evident contrast to the almost continuous noise from vehicles on tarred roads. In the intervals between trains,
communication between individual birds was normal. This
is because a passing train is a moving point source of noise,
which does not impair communication between birds. On
the other hand, a busy road is a linear source of noise: the
noise generated by a constant stream of traffic adversely
affects some bird species (Brumm 2004; Halfwerk et al.
2011).
In summary, the noise generated by passing trains did
not have a negative impact on either the abundance of birds
or the numbers of woodland species near the village of
Goła˛b. The railway line carrying 81 trains in 24 h cuts
through a forest; the edge effect thereby created was
manifested in the larger number of bird species observed
close to the tracks. The more complex structure of a forest
along its margins provides more optimal habitats for birds,
thus attracting them to the forest edge. The railway infrastructure (catenary supports, catenaries, and other electrical cables) in itself offers good conditions for courtship
(song posts), territorial behavior (look-out and resting
posts), and foraging (waiting for a victim). Train noise does
not clearly differentiate the distribution of birds from various guilds living in the vicinity of railway lines, in contrast
to the situation by very busy roads. This is due in the first
place to the different nature of train noise (a moving point
source passing by at fairly infrequent intervals) compared
with that of noise from vehicles on a very busy road (in
practice, this is a linear source).
Acknowledgments We thank two anonymous referees for critically
reviewing previous version of the manuscript. This research was
supported by the Institute of Biology and Biochemistry UMCS. Our
special thanks go to Maciej Hałucha and Janusz Bohatkiewicz from
EKKOM Ltd. for preparing the acoustic map of the study area.
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