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Behaviour 154 (2017) 981–996 brill.com/beh
Contrasting seasonal reactions of two sibling
woodpeckers to playback stimulation in urban areas —
implications for inventory and monitoring of the Syrian
woodpecker
Tomasz Figarski ∗
Institute of Systematics and Evolution of Animals, Polish Academy of Sciences,
17 Sławkowska, 31-016 Krakow, Poland
* Author’s e-mail address: tomasz.figarski.isez@gmail.com
Received 29 March 2017; initial decision 25 May 2017; revised 23 August 2017; accepted 5
September 2017; published online 5 October 2017
Abstract
The Syrian woodpecker (Dendrocopos syriacus) is the most synanthropic species among European
woodpeckers and can be considered to be a synurbic species benefiting from human activity and
man-made habitats. Because it is closely related to the common great spotted woodpecker (Den-
drocopos major), the interactions between the species should be investigated before conducting
any study on the Syrian woodpecker. The aim of this study was to assess, analyse and compare
both species reactions to the broadcasting of Syrian woodpecker playbacks throughout the year, in
urban areas. The study was conducted in three Polish towns. All sites hosted populations of Syrian
and great spotted woodpeckers living sympatrically. Both woodpeckers were observed throughout
each year of study and responded to playback. The Syrian woodpecker more willingly responded to
playbacks. Especially interesting was that the Syrian woodpecker female participated in territorial
activity to a greater degree than males (sex-role reversed species). This is the first woodpecker study
that shows a stronger territorial activity in females than males and the first reporting woodpecker
reactions to playback stimulation throughout the year. The Syrian woodpecker is a rare and threat-
ened species in Europe (protected under the Birds Directive of the European Union (2009/147/EC)
and is considered a keystone species for woods and associated biodiversity in urban areas. There-
fore, application of efficient methods for precise recognition of Syrian woodpecker territories (not
only breeding ones) on the basis of correct interpretation of its behaviour is crucial for its conser-
vation and urban habitat management.
© Koninklijke Brill NV, Leiden, 2017 DOI 10.1163/1568539X-00003452
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Keywords
broadcasting, urban ecology, behaviour, Dendrocopos syriacus, Dendrocopos major, urban
birds, wildlife conservation.
1. Introduction
Acoustic communication plays a major role for numerous taxa, including
birds (Bradbury & Vehrencamp, 1998). It is especially marked in singing
birds for which vocalizations (songs and calls) are of vital importance, as it
helps them to pass and acquire crucial information concerning sexual selec-
tion, defending or advertising territories and demonstrating fitness (Catch-
pole & Slater, 2008; Gil & Brumm, 2014). Woodpeckers communicate by
drumming and various calls to keep in touch with conspecifics and other
species (Gorman, 2004; Marler & Slabbekoorn, 2004). Woodpeckers play an
important ecological role in many environments, and are recognized as key-
stone and umbrella species, and indicators of local biodiversity (Mikusiński
& Angelstam, 1998; Pakkala et al., 2002; Mikusiński, 2006; Virkkala, 2006).
Most studies on woodpeckers have been conducted in forest habitats and
concerned with ecological aspects (e.g., Bütler et al., 2004; Kosiński &
Winiecki, 2005; Czeszczewik & Walankiewicz, 2006; Kajtoch et al., 2013),
and little is known about species in barely wooded areas, such as rural and
urban landscapes. Among European woodpeckers, the Syrian woodpecker
(Dendrocopos syriacus, Hemprich and Ehrenberg, 1833) (hereafter SW) is
the least known species (Pasinelli, 2006). Moreover, the SW is said to be
difficult to identify in the field and is weakly associated with forest habi-
tats (Mikusiński et al., 2001). The SW is the most synanthropic of Europe’s
woodpeckers (Gorman, 2004), and can be considered ‘synurbic’ (sensu Lu-
niak, 2004; Francis & Chadwick, 2012). In Europe it is highly dependent
on man-made habitats. Human land-use is the factor which determines its
occurrence. The species inhabits all kinds of urban and rural habitats, like or-
chards, parks, gardens, cemeteries, allotments, various clusters of trees, and
forest edges, but avoids forest interiors (Cramp, 1985; Kurek, 1984; Winkler
et al., 1995; Michalczuk & Michalczuk, 2006b, 2011, 2016a; Fröhlich &
Ciach, 2013; Figarski, 2014). These are all rather open and lightly wooded
habitats (Gorman, 2004). The primary global range of SW covers parts of
the Near East (from Iran to Israel and Turkey) and Caucasian region (Glutz
von Blotzheim & Bauer, 1980; Cramp, 1985; Michalczuk, 2014; http://maps.
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iucnredlist.org/map.html?id=22681127). Furthermore, since the end of the
XIXth century the SW has been expanding, and has settled in the Balkans
and central Europe, reaching northern Poland, the Czech Republic, Austria
and Slovenia. In the east there are populations in Ukraine, Belarus and Rus-
sia (Gorman, 2004; Michalczuk, 2014).
SWs produce the range of calls and sounds typical for Dendrocopos wood-
pecker species. There are a variety of calls, such as contact, alarm, courtship
or calls performed when disturbed (Cramp, 1985; Winkler et al., 1995; Gor-
man, 2004). Calls of the SW are a good identification factor for this species,
especially to distinguish it from the great spotted woodpecker (Dendrocopos
major, Linnaeus, 1758) (hereafter GW). SW and GW differ also with respect
to drumming activities, which can be observed in both sexes (Winkler et al.,
1995; Gorman, 2004).
The SW is very territorial (Winkler, 1972; Gorman, 2004) and readily
answers to calls of conspecifics and other woodpeckers. The species ac-
tively defends its territories from intruders especially in the pre-breeding
season. This is why the inclusion of broadcasting of calls/drums (playback)
can improve a population inventory method for this species (Michalczuk &
Michalczuk, 2006a, b). On the other hand, its behaviour during the breed-
ing season is secretive (Michalczuk & Michalczuk, 2006a, b; Michalczuk
et al., 2011). There is no agreement about which species — the SW or GW
— is more aggressive and dominant over the other when living sympatrically
(Gorman, 2004; Michalczuk & Michalczuk, 2011, 2016b). Also, the involve-
ment of both sexes in disputes is not clear, but males have been observed to
participate in fierce disputes and answer to playbacks in the breeding sea-
son (Gorman, 2004). Interactions between the species may also be disrupted
by hybridization, which is no rare phenomenon, especially in urban areas
(Dudzik & Polakowski, 2011; Michalczuk et al., 2014, Figarski & Kajtoch,
in press) and can lead to the presence of viable and fertile hybrids, whose
behaviours are unknown.
Some aspects of the biology and ecology of the SW have been studied in
rural environments (e.g., Michalczuk & Michalczuk, 2006a). But such stud-
ies in urban habitats are very scarce and are concerned almost exclusively
with the occurrence, and rarely with the habitat preferences of this species
(Mošanský & Mošanský, 1999; Ciach & Fröhlich, 2013; Fröhlich & Ciach,
2013; Figarski, 2014). Despite the fact that urban populations of SW may
make up a sizeable proportion of regional or national populations (Fröhlich
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& Ciach, 2013; Kajtoch & Figarski, 2017), there are still knowledge gaps
with respect to this interesting phenomenon.
Taking the above background into account, the aim of this study was to
assess, analyse and compare SW and GW reactions to the broadcasting of
SW calls and drums throughout the year, in urban areas.
2. Material and methods
2.1. Study areas
The study was conducted in the years 2013-2015 in three study areas: the
town of Radom (27.1 km2 ; approx. midpoint 51.400°N 21.150°E), the towns
of Grodzisk Mazowiecki and Milanówek (25.4 km2 ; 52.111°N 20.655°E)
(both Mazovia Province) and the town of Wieliczka (12.5 km2 ; 49.987°N
20.066°E) (Małopolska Province). Observations in each study area were
performed in subsequent years (Radom, 2013, Grodzisk Maz.-Milanówek,
2014, Wieliczka, 2015). These are typical Polish towns with a variety of
building types: from compact downtown buildings and tenements, single-
and multi-family houses (old as well as modern ones), to industrial buildings,
etc. Buildings are surrounded by various vegetation types, such as parks, or-
chards, allotments, various woodlots, dispersed urban greenery, grasslands,
and undeveloped lands, etc. All sites hosted populations of SW and GW liv-
ing sympatrically (Figarski, 2014; Kajtoch & Figarski, 2017).
2.2. Collection of woodpecker responses
Woodpecker data was collected with the help of a combined cartographic
method, with observation points located systematically in a 500 × 500 m
grid, and call stimulation (playback; see Michalczuk & Michalczuk, 2006a,
b; Michalczuk et al., 2011). There were 122 observation points in Radom,
110 in Grodzisk Maz.-Milanówek and 55 in Wieliczka. This method was
used due to the diverse nature of these urban habitats and because territories
of the SW in these habitats are often fragmented, where favourable habitat
patches are divided by other areas (sensu Gorman, 2004). The inventories
were conducted over the entire study areas throughout the year. Playback
was used to detect birds. High quality records of SW male contact calls and
drumming were collected prior to the study from local individuals during the
pre-breeding season (March-April). Calls were typical for SW, a single call
repeated every second (9 calls/10 s) with an average frequency of 3.25 kHz,
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and drumming, which was a repeated 1.2-s series of 25–27 strikes with a fre-
quency of 1.02 kHz (analysed in BatSound 3.31 Pettersson-Elektronik AB).
This method is also sufficient for the GW, which responds equally well to
sounds (calls and drums) produced by SW as to the sounds produced by
its own species (Michalczuk & Michalczuk, 2016a), which was additionally
verified in an investigation of local populations prior to the study. There-
fore, for inventories and monitoring involving simultaneously SW and GW
it is common to use records of SW sounds. Stimulation was carried out
according to the following protocol (modified from Michalczuk & Michal-
czuk, 2006a, b): 2 min of playback (drumming), 3 min of listening, 2 min
of playback (calls), 3 min of listening. Stimulation was stopped immediately
after a woodpecker responded. The equipment used for stimulation was a
RadioShack amplifier/speaker and a MuVo v100 mp3 player. At least one
inspection per month covering the entire study area (lasting 4–7 days), was
undertaken in favourable weather conditions. Each day of observation started
in the morning (between 5 and 8 a.m., depending on season) and lasted 5–6 h.
This resulted in 4–5 inspections during the pre-breeding period (March–mid-
May), 4–5 inspections during the post-breeding period (July–October) and 4
inspections during the winter period (November–February) in each area. In
order to not disturb the birds during breeding, playback was not used be-
tween mid-May and the end of June each year; this period was excluded
from the analysis. Birds of apparent hybrid morphology were also omitted
from the analysis.
SW and GW responses to playback were noted and analysed at two basic
levels. Firstly, four time categories of reaction were considered: (1) during
first playback, (2) during first listening, (3) during second playback, (4) dur-
ing second listening. Secondly, seven main type categories of reaction were
considered: I, response (vocalization) from afar; II, flying to loudspeaker,
vocalization, observation; III, flying to loudspeaker, without vocalization,
observation; IV, flying to loudspeaker, vocalization, foraging; V, flying to
loudspeaker, without vocalization, foraging; VI, flying to loudspeaker, drum-
ming; VII, without a visible marked reaction. Woodpeckers were observed
by one observer with the use of Vortex Crossfire 10 × 42 binoculars.
During the study period, in total 189 observations of SW and 278 obser-
vations of GW were collected (Table 1). Both woodpeckers were observed
throughout each year of study and responded to playback. Because inspec-
tions were done systematically and each point (territory) was checked the
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Table 1.
Number of collected observations by species/sex group (SWf, Syrian woodpecker females;
SWm, Syrian woodpecker males; GWf, great spotted woodpecker females; GWm, great
spotted woodpecker males), taking into account (i) if the bird was single or in the company of
a partner, (ii) or the period of the year: pre-breeding (March–mid-May, III–V), post-breeding
period (July–October, VII–X) and winter (November–February, XI–II).
Species and sex
SWf SWm GWf GWm
Birds
Single 60 53 90 108
With partner 38 38 40 40
Total 98 91 130 148
Period (months)
III–V 54 42 70 72
VII–X 22 14 25 37
XI–II 22 35 35 39
Total 98 91 130 148
same numbers of times, the data was not affected by overrepresentation of
some individuals.
2.3. Data analysis
The above-mentioned aspects were analysed for the whole set of observa-
tions and also separately for birds observed individually (single birds) and
for those in the company of a partner. Spontaneous reactions (those not fol-
lowing stimulation) were excluded. A Mann–Whitney U -test was used to
compare the times and the types of reaction showed by each species/sex
group (SW: males, SWm; females, SWf; GW: males, GWm; females, GWf)
and also by birds observed singly and with a partner.
Seasonal differences in responses by each species/sex group (SWm, SWf,
GWm, GWf) among the three periods defined previously (pre-breeding,
post-breeding and winter) were assessed, using a Kruskal–Wallis test, and
(as a post-hoc test) a Mann–Whitney U -test with Bonferroni multiple testing
correction.
In all the statistics, the significance level adopted was 0.05. All calcula-
tions were performed using Statistica 7.0 software.
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3. Results
3.1. Time of reaction
The comparisons between pairs of species/sex groups for the time of reaction
gave significant differences between females and males of SW, and between
females of SW and GW (Table 2; Figure 1A). In both cases, SWf reacted dis-
tinctly faster, mostly during the first sequence of stimulation. Most reactions
were recorded during stimulation sequences rather than listening sequences.
However, it should be noted that, when analysed separately, the differences
were significant only among single birds and not among birds observed with
a partner (Table 2, Figure 1A). Within each species/sex group, a significant
difference in time of reaction between single birds and birds with a partner
was observed only in SWf (Table 3, Figure 1A): single females reacted faster.
Figure 1. Distribution of woodpeckers’ reactions according to time of reaction (A), time
of reaction in each season (B), type of reaction (C) and type of reaction in each season
(D). SWf, Syrian woodpecker females; SWm, Syrian woodpecker males; GWf, great spot-
ted woodpecker females; GWm, great spotted woodpecker males. S, single birds; P, birds
with a partner; A, all birds. Time of reaction: (1) during first playback; (2) during first lis-
tening; (3) during second playback; (4) during second listening. Type of reaction: I, response
(vocalization) from afar; II, flying to loudspeaker, vocalization, observation; III, flying to
loudspeaker, without vocalization, observation; IV, flying to loudspeaker, vocalization, for-
aging; V, flying to loudspeaker, without vocalization, foraging; VI, flying to loudspeaker,
drumming; VII, without a visible marked reaction.
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Table 2.
Differences in times and types of reaction between woodpecker species/sex groups, for single birds, birds observed with a partner and all birds,
and (separately) in each season.
Compared groups of Birds Period
woodpeckers (species/sex)
Single With a partner All III–V VII–X XI–II
Z p Z p Z p Z p Z p Z p
Time of reaction
SWf–SWm −2.22 0.026∗ −0.90 0.366 −2.30 0.021∗ −3.43 0.000∗ 0.054 0.957 0.30 0.762
GWf–GWm 1.41 0.159 0.22 0.826 1.32 0.185 1.43 0.152 −0.89 0.373 1.88 0.060
SWf–GWf −3.74 0.000∗ −0.13 0.899 −3.10 0.002∗ −3.40 0.000∗ −0.01 0.991 −0.84 0.403
SWm–GWm −0.10 0.916 1.17 0.242 0.67 0.501 1.70 0.089 −0.91 0.363 0.07 0.945
Type of reaction
SWf–SWm 1.02 0.309 −2.10 0.036∗ −0.77 0.438 −1.08 0.282 −0.58 0.558 1.29 0.198
GWf–GWm −1.17 0.240 0.43 0.663 −0.86 0.390 −0.64 0.522 −0.96 0.335 −0.11 0.908
SWf–GWf −0.48 0.631 −1.34 0.179 −1.28 0.199 −0.75 0.450 −0.50 0.614 −0.63 0.531
SWm–GWm −2.57 0.010∗ 0.98 0.327 −1.30 0.194 −0.04 0.966 −1.21 0.224 −1.83 0.067
Reactions of woodpeckers to playback in urban areas
Species/sex groups: SWf, Syrian woodpecker females; SWm, Syrian woodpecker males; GWf, great spotted woodpecker females; GWm,
great spotted woodpecker males. Period (season, in months): pre-breeding (March–mid-May, III–V), post-breeding period (July–October, VII–
X) and winter (November–February, XI–II). For sample size see Table 1.
∗ Significant difference, P < 0.05.
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Table 3.
Differences in time and type of reaction between single birds and birds observed with a
partner in each group of woodpecker species/sex.
Woodpecker Time of reaction Type of reaction
species/sex Z p Z p
SWf −2.14 0.033∗ 1.96 0.049∗
SWm −0.94 0.347 −1.44 0.150
GWf 1.20 0.230 0.33 0.737
GWm 0.28 0.777 1.48 0.138
SWf, Syrian woodpecker females; SWm, Syrian woodpecker males; GWf, great spotted
woodpecker females; GWm, great spotted woodpecker males. For sample size see Table 1.
∗ Significant difference, P < 0.05.
Considering each season, time of reaction differed significantly in the pre-
breeding season (III–V) between females and males of SW, and between
females of SW and GWs; males of the SW and GWs also tended to be
different (Table 2, Figure 1B). In other seasons there were no significant dif-
ferences; only females and males of GW presented a tendency to be different
in the winter period (Table 2, Figure 1B).
A comparison, for each species/sexes group, of the time of reaction be-
tween each pair of periods/seasons (III–V vs. VII–X, III–V vs. XI–II, VII–X
vs. XI–II) showed differences for SWf and GWm (Table 4; Figure 1B). For
SWf differences were between the pre-breeding period (III–V) and the two
other periods; in the pre-breeding period reactions were clearly faster. For
GWm there was a difference between the pre-breeding period and the post-
breeding period (VII–X); in the former period birds also reacted faster.
3.2. Type of reaction
Considering all the birds, there was no significant difference in the type of
reaction between the pairs of species/sex groups (Table 2, Figure 1C). How-
ever, considering only single birds, males of SW and GW differed signifi-
cantly. Most reactions of SWm relied on observation (reaction type numbers
II and III), while in GW the type of reaction varied to a large degree. In turn,
for the birds observed with a partner, a difference was observed between fe-
males and males of SW: females called and observed more often (number
II), whereas males drummed more often (number VI) (Table 2, Figure 1C).
Within each season no significant difference was observed between the
species/sex groups (Table 2, Figure 1D). Only males of SW and GW showed
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Table 4.
Differences in time and type of reaction between each pair of seasons in each group of
woodpecker species/sex.
Woodpecker Kruskal–Wallis Mann Whitney
species/sex test U -test
H p III–V vs. VII–X III–V vs. XI–II VII–X vs. XI–II
Z p Z p Z p
Times of reaction
SWf 8.72 0.013∗ −2.64 0.008∗ −2.21 0.027∗ −0.11 0.909
SWm 0.48 0.786 – – – – – –
GWf 2.03 0.362 – – −1.38 0.167 – –
GWm 6.08 0.048∗ −2.40 0.016∗ −0.84 0.403 1.61 0.107
Types of reaction
SWf 7.04 0.030∗ 0.73 0.466 −2.22 0.026∗ −2.43 0.015∗
SWm 0.77 0.681 – – – – – –
GWf 5.55 0.062 0.36 0.717 −2.23 0.025∗ −1.75 0.079
GWm 2.15 0.342 – – – – – –
SWf, Syrian woodpecker females; SWm, Syrian woodpecker males; GWf, great spotted
woodpecker females; GWm, great spotted woodpecker males. Period (season, in months):
pre-breeding (March–mid-May, III–V), post-breeding period (July–October, VII–X) and win-
ter (November–February, XI–II). For sample size see Table 1.
∗ Significant difference, P < 0.05.
a tendency to be different from each other in the winter season (XI–II).
Reactions of the former relied mainly on observations (numbers II and III)
and the latter more often foraged (number V) or showed no reaction (number
VII).
An important difference between single birds and birds observed with a
partner was found in SWf (Table 3, Figure 1C): females observed in the
company of a partner reacted mostly with calls and observations (number
II), while single females were more often quiet (numbers III and V).
In turn, a comparison of the types of reaction between pairs of seasons
(III–V vs. VII–X, III–V vs. XI–II, VII–X vs. XI–II) for each species/sex
group showed differences in a few cases (Table 4, Figure 1D). Reactions
of SWf differed between winter and the two other periods. GWf showed
the same pattern of differences (although weaker between winter and post-
breeding period). In both females, in winter more reactions were based on
foraging than in the other periods (numbers IV and V) (Table 4, Figure 1D).
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4. Discussion
4.1. General findings
This study has confirmed that playbacks can be successfully used for SW
inventories in urban populations, as is the case in rural populations (Michal-
czuk & Michalczuk, 2006a, b; Michalczuk et al., 2011). Both SWs and GWs
reacted to playback stimulation using the SW’s calls and drums, and through-
out the year.
4.2. Syrian woodpecker’s reactions to playback
In woodpeckers that communicate by drumming and calls, males are gener-
ally regarded to participate in defending territories more often than females
(Gorman, 2004). However, the present study questions this hypothesis for
the case of the SW. The reactions of SWf to playback were generally faster
than those of SWm. However, this was only evident in single birds. Without
a partner, SW individuals of both sexes mainly responded during stimu-
lation sequences, whereas in pairs responded more often during listening
sequences. There were also differences between the sexes in the types of
reaction while in pairs: SWf reactions relied mainly on alarm calls and ob-
servation, while SWm, besides vocal responses, often drummed (females
never drummed when accompanied by males). With SW, it seems that in
pairs the responsibilities for defending territories are shared equally between
both partners (co-operative territory defence, e.g., Amundsen, 2000; Hall,
2000; Hall et al., 2015). Woodpeckers that faced an intrusion in its territories
with a partner were also calmer, with more delayed responses (more frequent
during listening sequences). Generally, it appears that the SW is a sex-role
reversed species with respect to territorial activity (see Amundsen, 2000;
Illes & Yunes-Jimenez, 2009). In agreement with a previous study, calling
and drumming are the most frequent reactions in this species (Michalczuk &
Michalczuk, 2006a).
SW responses to playback were observed throughout the year. Clear dif-
ferences were found in SWf in the times of reaction between the pre-breeding
period and the two other periods. Responses in spring were much faster
than those performed later in the year. There was also a significant differ-
ence between females and males in spring. This result seems to confirm that
SWf were clearly (and to a greater extent than SWm) involved in defend-
ing territories. Woodpeckers displayed highly territorial activity, which was
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especially marked in the pre-breeding season (Michalczuk & Michalczuk,
2006a, b; Michalczuk et al., 2011). There is agreement in the literature (e.g.,
Kurek, 1984; Gorman, 2004) that SWf drum during the breeding season, but
no detailed information has been available for other periods till now. SWm
drummed in the pre-breeding season and, less often, in winter, while SWf
drummed only in pre-breeding season. Reaction types in SWf differed signif-
icantly between winter and the two other seasons. In winter SWf responded
with foraging activity much more often. There was no such difference be-
tween seasons in SWm. This can be explained by the higher demand for
food in females before the breeding season. Females may try to take every
opportunity to forage, even in the presence of another bird (mimicked by the
playback).
4.3. Differences in Syrian and great spotted woodpeckers’ reactions to
playback
There were clear differences in the times of reaction between females of
SW and females of GW. Responses of the former were faster (more fre-
quent during the first sequence of stimulation), especially in single birds,
whereas there was no distinct pattern in the latter. GWf reacted consistently
over the entire period of observations. There were also no significant dif-
ferences between females and males of GW. A general comparison of the
types of reactions of SW and GWs showed that the latter displayed a lower
degree of interest: reactions from afar or even no responses were more often
observed. Important differences were found between single males of both
species. In the SW, most responses were observations, whereas the GW’s
responses were more diverse, with foraging and drumming more often ob-
served. Females and males of GW did not differ in their types of reaction.
Further, the responses of GWs observed either solitarily or with a partner
also did not differ in their times or types of reaction. Basically, there was
no clear distinction in the reactions to playback between the sexes in GW. In
contrast to SW, females of GW did not show an increased commitment to de-
fending territory. It seems that in this species both sexes are equally involved
in reacting, and defending activities.
As in SW, GW responded to playback throughout the year. SW and GW
females had significantly different times of reaction in the pre-breeding sea-
son. On the other hand, in both species, the variations in the types of reactions
between the seasons were similar. Interestingly, the difference between win-
ter and two other seasons observed in SW females was also observed in GW
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females. The latter also more often responded with foraging activity in winter
than in the other seasons. And as in SWs, such differences were not observed
in GW males.
4.4. Conclusions
There are many gaps in the knowledge of SW biology, and the most pro-
nounced concerns its behaviour (Pasinelli, 2006), including its interactions
with GW. SW is an important species in urban areas as it increases bio-
diversity in cities due to its status as a keystone species for urban woods
(Figarski & Kajtoch, unpubl.) and its capacity to create nesting places for
secondary cavity-nesters (Ciach & Fröhlich, 2013; Fröhlich & Ciach, 2013).
The SW uses man-made habitats of traditional character with softwood and
fruit trees (Figarski, 2014). However, recently its habitats are in danger due
to intensive works on urban and rural greenery (Kajtoch, 2017). Therefore
application of efficient methods for its inventories and precise recognition of
its territories (not only breeding ones) on the basis of correct interpretation
of its behaviour is crucial for conservation and urban habitat management.
Furthermore, the SW is protected under the Birds Directive of the European
Union (2009/147/EC). Unfortunately, all Natura 2000 sites that have been
designed for the protection of the SW are located in rural landscapes. Nu-
merous populations in cities have been neglected, mainly because of lack of
research in such environments (Kajtoch & Figarski, 2017; Figarski & Kaj-
toch, unpubl.). A few interesting findings were obtained in the present work.
It was especially surprising that SWf participated in territorial activity to an
evidently greater degree than males (sex-role reversed species). Although the
species live sympatrically, it seems that their ability to thrive in urban ecosys-
tems depends more on the structure and composition of preferred habitat than
intra- and interspecific interactions (Figarski, data not shown). Finally, this
study confirms that any study on the SW needs to consider the GW as an
important factor influencing numerous aspects of SW occurrence, ecology
and behaviour.
Acknowledgements
I am grateful to Ł. Kajtoch for his support in the planning and conducting of
research and for valuable comments to the final version of the manuscript.
The help of A. Jakubowska in characterization of acoustics of woodpecker
Downloaded from Brill.com06/16/2022 11:53:04PM
via free access994 Reactions of woodpeckers to playback in urban areas
voice records is greatly appreciated. The paper has greatly benefited from
the constructive comments of two anonymous reviewers. This work was
partially covered by the grant for young scientists 6085/E-47/M/2015 funded
by the Institute of Systematics and Evolution of Animals, Polish Academy
of Sciences from resources provided by Polish Ministry of Higher Education
and Science.
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