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Published by Associazione Teriologica Italiana Online first 2019
Hystrix, the Italian Journal of Mammalogy
Available online at: doi:10.4404/hystrix–00204-2019
Research Article
Winners and losers in an urban bat community: a case study from southeastern Europe
Olga Tzortzakaki1,, Eleni Papadatou2, Vassiliki Kati3, Sinos Giokas1
1University of Patras
2Ove Arup & Partners Ltd
3University of Ioannina
land cover
Mediterranean basin
Pipistrellus kuhlii
urbanisation gradient
water bodies
Article history:
Received: 15 July 2019
Accepted: 10 October 2019
We would like to thank Prof. Athanassios Argiriou from the Laboratory
of Atmospheric Physics of the University of Patras for kindly provid-
ing the meteorological data. We are grateful to Robert Nudds for im-
proving and proofreading an earlier version of the manuscript, as well
as to Christina Kassara for statistical support. George Iliopoulos, Di-
mitris Papandropoulos and Philippos Katsigiannis supported the first
author during field work and provided important information about
the study area. Two anonymous reviewers provided constructive com-
ments, which largely contributed to improving the final version of the
Increasing urbanisation is reported to have significant effects on bat communities, due to habitat
modifications, light and noise pollution and reduced prey availability. Recent studies have indicated
that species show varying responses to urbanisation, with a few able to exploit man-made struc-
tures and adjust to the new environmental conditions. This study aimed to identify how landscape
composition influences bat diversity and community structure along the urbanisation gradient in a
coastal Mediterranean city (Patras, Greece) and whether particular species benefit from the novel
conditions. We conducted acoustic surveys along 45 transects during the post-breeding season for
two years. The effect of land cover, the number of streetlamps (a proxy of artificial illumination),
the presence of water bodies and weather conditions on bat activity, and community structure were
investigated using Generalized Linear Mixed Models, and multivariate statistics respectively. Eight
bat species and five species groups were identified. Bat communities were affected by urbanisation
in general and diversity was low in the entire study area. The community was dominated by the
synurbic species Pipistrellus kuhlii, which comprised more than 70% of the total bat activity recor-
ded. A positive relationship between built-up areas and bat activity was found, probably because P.
kuhlii usually forages around streetlamps in urban areas. In contrast, vegetation cover did not affect
bat activity, even in the less urbanised areas. The remainder of the bat species were not frequently
recorded and were mostly detected close to water bodies, highlighting their value for foraging bats
and the need for their conservation.
Urbanisation is a major cause of land use change, habitat degradation
and fragmentation, as urban areas expand rapidly at the expense of nat-
ural ecosystems. The novel ecosystems that arise are very heterogen-
eous (Pickett et al., 2011), containing a mosaic of artificial structures
such as buildings and roads, green spaces with ornamental vegetation,
bare ground, agricultural land, natural and artificial ponds, as well as
patches of remnant natural or semi-natural vegetation (Cadenasso et
al., 2007). In addition, they are characterized by new environmental
conditions, such as increased temperatures (Urban Heat Island), pollu-
tion, anthropogenic noise and artificial lighting (Grimm et al., 2008)
All these features render urban areas a challenging ecosystem for wild-
To date, numerous studies have shown the detrimental effects of urb-
anisation on biodiversity. At a global scale, urban wildlife communit-
ies have been found to become rather homogeneous along latitudinal
gradients within specific continents (Clergeau et al., 2006; McKinney,
2006) or even among different continents indicating biodiversity loss
(La Sorte et al., 2007). At a local scale, biological communities may
show varying responses to urbanisation intensity, yet, a general pat-
tern has been observed among animal communities: species richness
tends to decrease towards more urbanised areas (McKinney, 2008). A
few generalist species manage to exploit the urban habitat by adjusting
their behaviour and ecology and are usually found in high abundances
(“urban exploiters” or “synurbic”); other species can successfully occur
in both urbanised and natural areas (“urban adapters”), while moderate
Corresponding author
Email address: (Olga Tzortzakaki)
generalists and specialists tend to become scarcer (“urban avoiders”)
(McKinney, 2006; Devictor et al., 2007; McKinney, 2008; Francis and
Chadwick, 2012; Sullivan et al., 2016).
This pattern has also been observed in bat communities, as only a few
species manage to adjust to the urban environment, while other more
sensitive species avoid it (Avila-Flores and Fenton, 2005; Jung and
Kalko, 2011; Russo and Ancillotto, 2015; Jung and Threlfall, 2018).
Urban life may be harsh for many bat species due to high levels of
light and noise pollution (Schaub et al., 2008; Stone et al., 2015), in-
creased risk of collisions (Medinas et al., 2013) and spread of diseases
(Mühldorfer et al., 2011), increased exposure to predators (Ancillotto
et al., 2013; Threlfall et al., 2013), reduced prey availability and higher
inter- and intraspecific competition (Russo and Ancillotto, 2015). Fur-
thermore, artificial nighttime illumination may affect bat commuting,
breeding, roosting, foraging behaviour and hibernation (Stone et al.,
2015; Azam et al., 2016). Anthropogenic noise may also affect bat
feeding success, as noise frequencies often overlap with the frequen-
cies of sounds emitted by bat prey (Schaub et al., 2008). Similarly,
noise might affect communication with conspecifics by masking bat
social calls (Russo and Jones, 1999).
In general, urbanisation has negative effects on bat activity and
diversity, though these effects are highly species-specific (Jung and
Threlfall, 2018). Despite the generally adverse conditions in urban en-
vironments, new opportunities may arise for the more tolerant species
(Francis and Chadwick, 2012; Threlfall et al., 2012; Ancillotto et al.,
2015, 2019). Streetlamps attract large numbers of insects, offering an
easily accessible prey for some species (Rydell, 1992; Ancillotto et al.,
2015; Schoeman, 2016 but see Arlettaz et al., 2000; Stone et al., 2015).
In addition, man-made structures such as buildings, roofs and tunnels
Hystrix, the Italian Journal of Mammalogy ISSN 1825-5272 25th November 2019
©cb e2019 Associazione Teriologica Italiana
Hystrix, It. J. Mamm. (2019) — online first
may be used as roosts especially due to natural roost loss (Russo and
Ancillotto, 2015). Warmer microclimatic conditions prevailing in arti-
ficial roosts may benefit reproductive females and the growth of their
young (Lausen and Barclay, 2006; Ancillotto et al., 2015). Hence, gen-
eralists may be more widespread and abundant in the urban environ-
ment than specialists by exploiting man-made structures. However,
urban areas may also act as ecological traps for bats (Russo and An-
cillotto, 2015). Gaining an insight into bat responses to urbanisation is
important for the generation of effective management actions and their
contribution to biodiversity conservation.
In recent years, there has been a growing interest in the study of
bat foraging and roosting ecology and behaviour in urban areas, par-
ticularly in Europe (e.g. Lintott et al., 2015; Border et al., 2017;
Suarez-Rubio et al., 2018; Ancillotto et al., 2019), Australia (Threlfall
et al., 2012; Caryl et al., 2016), North America (Dixon, 2012; Krauel
and LeBuhn, 2016; Schimpp and Kalcounis-Rueppell, 2018), Cent-
ral America (Jung and Kalko, 2011; Rodríguez-Aguilar et al., 2017),
South America (Oprea et al., 2009) and South Africa (Schoeman,
2016). However, studies on bat responses to urbanisation in south-
eastern Europe are lacking, despite its high bat diversity (Dietz et al.,
2009). In this study, we aimed to fill this knowledge gap and identify
(a) how bat activity varies along the urbanisation gradient, (b) the en-
vironmental factors that affect bat activity and (c) whether urbanisation
favours particular species in a densely-built Mediterranean coastal city
(Patras, Greece).
Materials and methods
Study area and site selection
The study area covered the city of Patras and its surroundings (total
area 110 km2), located in SW Greece (38°140N, 21°440E). Patras is the
third largest city in Greece with approximately 200000 inhabitants (EL-
STAT, 2011). The city has an elongated expansion along the coast of
Patraikos Gulf and is delimited by Mount Panachaikon (1926 m a.s.l.,
Natura 2000 site GR2320007) to the east-southeast (Fig. 1). Due to
various historical reasons and inadequate spatial planning, the town has
a very dense urban core with severely reduced green space (Papadatou-
Giannopoulou, 1991; Tzortzakaki et al., 2019). The surrounding peri-
urban and rural areas consist of agricultural land, orchards (mostly olive
groves), Mediterranean shrublands, and a few scattered small remnant
patches of coniferous or deciduous forests and riparian vegetation. Due
to complex geological processes (i.e. the tectonic uplift caused by the
rifting of the Corinth and Patraikos Gulfs), a number of gorges and
gullies containing streams were formed on Mount Panachaikon, which
flow through the study area into the Patraikos Gulf (Fig. 1). The cli-
mate is typical Mediterranean with a long dry period and mild winter
(average annual temperature 17.9 C), and relatively high precipitation
levels (average annual rainfall: 607 mm; HNMS, 2017).
The study area was stratified into three zones of decreasing built
cover (b.c.): the "urban zone" (b.c.>50%), the "suburban zone"
(30%<b.c.<50%), and the “peri-urban zone” (b.c.<30%), after overlay-
ing a grid of 500×500 m cells and calculating the proportion of built
cover in each cell using the Urban Atlas (EEA, 2010; Tzortzakaki et
al., 2019). In each zone, 15 grid cells were randomly selected, on the
basis that they were situated at least 500m from their nearest neigh-
bouring cell (Tzortzakaki et al., 2019). Each gird cell encompassed
one bat sampling site (45 sites in total; Fig. 1).
Bat surveys and sound analysis
Bat surveys were conducted during the post-breeding period from the
end of August to the beginning of October in two consecutive years
(2015–2016). In each of the 45 grid cells, a 300 m line transect was
established, which was walked in one direction once in each year (90
transects in total). Surveys started half an hour after sunset (Vaughan
et al., 1997) and were completed within three hours. On average, three
sites were sampled on each survey night, in random order (Dixon, 2012)
and regardless of their location along the urbanisation gradient to re-
duce possible temporal or spatial sampling bias. Sampling was con-
Figure 1 Map of the study area, showing the distribution of the sampling sites in the
urban (squares), suburban (circles) and peri-urban zone (triangles). Urbanisation zones
were delineated based on land use data from the Urban Atlas (EEA, 2010). Grey diagonal
lines represent the Natura 2000 site of Mount Panachaikon (GR2320007).
ducted by the same individual and under good weather conditions, i.e.,
without rain or strong winds and minimum nightly temperature >19 C.
Bat vocalizations were recorded using a Pettersson M500 USB ultra-
sound microphone (Pettersson Electronik AB, Uppsala, Sweden), set at
approximately 1.8m above the ground and connected to a tablet com-
puter. Recordings were performed with the Pettersson Bat Sound Touch
Lite software at a 500 kHz sampling rate, set in the automatic mode
with a sound power level of -10 dB and a bandwidth ranging from 19
to 120 kHz, which was defined after testing microphone sensitivity in
the field to avoid recording low-frequency noise. The bat detector was
automatically triggered by sounds above the lower threshold and was
set to record for 6 seconds each time (5 seconds from the trigger point +
1 second before). Social calls and echolocation calls of Tadarida teni-
otis were recorded while recording other species’ echolocation calls.
Bat ultrasound analysis was performed using SonoBat 3.0 (SonoBat
Bat Call Analysis Software); echolocation call parameters were auto-
matically measured, after applying manual filters to minimize masking
by noise where necessary. Batsound 4.2 (Pettersson Electronik AB,
Uppsala, Sweden) was additionally used to explore the possible pres-
ence of more than one individual or species or social calls within sound
files. Spectrograms were visualized in Batsound with a 512 samples
Hanning FFT window.
Bat species identification was conducted with the open source auto-
mated software for the identification of European bats “iBatsID” (Wal-
ters et al., 2012). iBatsID uses the parameters measured by Sonobat and
assigns each call sequence to one or more species based on a classific-
ation probability. We considered identification at the species level as
trustworthy, when classification probability was 90%. When prob-
ability was <90%, call sequences were initially assigned to the spe-
cies group with the highest classification probability. They were sub-
sequently evaluated manually and identified to species level where pos-
sible using BatSound. Where frequency of echolocation call and time
parameters fell into the overlap zone of at least two species (Dietz and
Kiefer, 2014), identification remained at the species group level, unless
social calls were present (Russo and Jones, 1999; Pfalzer et al., 2003;
Russo and Papadatou, 2014; Nardone et al., 2017). Call sequences that
Bat community of an Eastern Mediterranean city
could not be assigned to species or species groups were recorded as
unknown and were only included in total bat activity estimates.
Calls of Myotis species were grouped, as their identification is of-
ten ambiguous (Lintott et al., 2015). Call sequences classified as P.
nathusii by iBatsID were considered as P. kuhlii in the analysis, be-
cause P. nathusii is rare in southern Greece (Georgiakakis and Papad-
atou, unpubl. data) and none of the social calls recorded belonged to
the latter species.
The number of bat passes of each species or species group within
each recording, i.e., within 6 seconds, was used as an index of bat activ-
ity. A bat pass is defined as a sequence of two or more echolocation
pulses emitted by a bat (Thomas, 1988).
Environmental variables
In each sampling site, mean hourly temperature (MHT) and relative
humidity (MHRH) were extracted from the meteorological data that
were provided by the Laboratory of Atmospheric Physics of the Uni-
versity of Patras, given the known influence of weather conditions on
bat activity (O’Donnell and Sedgeley, 2001). Second, the percent cover
of the predominant land-cover types (buildings, impervious surfaces,
woody vegetation, open green spaces and water bodies) was assessed
within a 200 m radius circular buffer zone around each transect centroid
(Tzortzakaki et al., 2019), so that the buffer zones fit within the re-
spective grid cells (500×500 m) and do not overlap with nearest neigh-
bouring buffers. Proximity of sampling sites to the closest natural (i.e.,
streams and marshes) or artificial water body (hereafter all referred as
water bodies), and to motorway (Fig. 1) was measured using ArcGIS
10.1 (ESRI). Finally, the number of streetlamps along each transect
was recorded as a proxy of the intensity of illumination in the sampling
Data analysis
First, pairwise comparisons of mean daily temperature (°C), mean
daily relative humidity (%) and total daily rainfall (mm) were per-
formed for each month between the two sampling years with Analysis
of Variance (ANOVA), in order to test for possible differences in the
meteorological conditions between the two years.
To examine bat activity patterns along the urbanisation gradient, bat
activity was compared among the urbanisation zones for each year sep-
arately, using Kruskal-Wallis tests, because the data were not normally
distributed. A Wilcoxon signed-rank test was used to compare bat
activity at each sampling site between the two years.
To investigate the effect of the environmental parameters on bat
activity, Generalized Linear Mixed Models (GLMMs) were used. Prior
to model building, the relationships among the response and explan-
atory variables were investigated (Zuur et al., 2010). Land-cover
types were highly correlated, therefore Principal Components Analysis
(PCA) was used to reduce their number and only the first two axes were
retained: PC1 represented a gradient of decreasing vegetation cover to
increasing built cover, while PC2 characterised a gradient of decreas-
ing open spaces to increasing water cover (Tzortzakaki et al., 2019).
The distance to motorway and the number of streetlamps were excluded
from the analyses, because they were both collinear with PC1 (Pearson
r=0.63 and r=0.78, respectively), as well as MHRH, because it was
correlated with MHT (r=-0.66). No evidence for spatial and temporal
autocorrelation was found after plotting bat activity vs. spatial coordin-
ates and Julian date, respectively (Zuur et al., 2010).
GLMMs were carried out using negative binomial error distribu-
tion with a log link function. Since two visits per site were conduc-
ted, sampling site was considered as a random factor, while PC1, PC2,
MHT, distance to water bodies (DistW) and year of study (Year) were
included as fixed terms. Continuous explanatory variables were stand-
ardised (Schielzeth, 2010).
Model selection was performed with backward stepwise deletion:
the global model was constructed, and then non-significant covari-
ates were sequentially removed until the best model with the smallest
Akaike’s Information Criterion (AIC) was achieved. Tests of homogen-
eity of variance, normality of residuals, independence of observations
Figure 2 Total bat activity (number of bat passes) per urbanisation zone and year of
study. Bat activity was significantly higher in 2016, but no statistically significant dierences
were found among the urbanisation zones in either year.
and model overdispersion were carried out for model validation (Zuur
et al., 2010). Analyses were carried out with lme4 package (Bates et
al., 2015) in R v.3.3.1 (R Core Team, 2016).
To examine the relationship between community composition and
environmental variables, Redundancy Discriminant Analysis (RDA)
was performed separately for each year using the vegan package (Ok-
sanen et al., 2016) in R. Permutation tests with 999 permutations were
used to assess the significance of the relationships between species
composition and the explanatory variables. For this analysis, calls at
the frequency overlap zone that could not be identified to species level
were excluded, as well as species with less than two bat passes. Prior
to the analysis, bat activity data were Hellinger-transformed (Oksanen
et al., 2016), while environmental data were standardised (Leps and
Smilauer, 2003).
To assess the relationship between species composition and the urb-
anisation gradient, Permutational Multivariate Analysis of Variance
(PERMANOVA) was carried out separately for each year (Anderson,
2001). PERMANOVA allowed for pairwise comparisons of the com-
munity composition among the urbanisation zones. The analysis was
performed on the community data matrix (same as RDA) using the
Bray-Curtis dissimilarity index with 999 permutations with the “ad-
onis” function of the vegan package (Oksanen et al., 2016), after ap-
plying a Hellinger transformation. Pairwise comparisons among the
urbanisation zones were conducted with the pairwise.Adonis pack-
age (Martinez Arbizu, 2017). Multivariate homogeneity of group dis-
persions (PERMDISP), performed with the “betadisper” function (Ok-
sanen et al., 2016), did not indicate violation of the homogeneity of
variances assumption (p>0.05 for both years).
A total of 653 bat passes were analysed, 67% of which were identified to
species level. Eight bat species and five species groups were identified
(Tab. 1). P. kuhlii was the most common species in the study area,
comprising more than 70% of the bat passes in both years and occurring
at approximately 80% of the sites (Tab. 1). Excepting P. pygmaeus and
T. teniotis, other species were recorded very infrequently (<10% of the
sites in both years). The occurrence of H. savii could not be assessed
precisely, as respective calls were mostly in the overlap zone of H. savii
and P. kuhlii (Tab. 1).
Bat activity did not vary considerably along the urbanisation gradi-
ent in 2015. In contrast, it was remarkably higher in the urban and sub-
urban zones than in the peri-urban zone in 2016 (Fig. 2). However, stat-
istical comparisons of bat activity among the urbanisation zones did not
show any significant differences in either year (2015: Kruskal-Wallis
H=1.281, p=0.527; 2016: H=2.893, p=0.235). Bat activity was signi-
ficantly higher in the second year (Wilcoxon test: V=229, p=0.015),
noting that significantly higher temperatures occurred in the study area
during the winter and spring of 2016 (Tab. S1). However, summer and
autumn temperatures were similar across the two years and no differ-
ences were found in most cases for relative humidity (Tab. S1).
Hystrix, It. J. Mamm. (2019) — online first
Table 1 Proportion of bat passes assigned to bat species or species group in each urbanisation zone per year of study (calculated as the proportion of the total annual bat passes).
Occupancy indicates the proportion of sampling sites (out of 45) where a given species or species group was recorded.
2015 2016
Bat passes (%) Occ. (%) Bat passes (%) Occ. (%)
Species / Group Urban Suburban Peri-urban Total Urban Suburban Peri-urban Total
Eptesicus serotinus / Nyctalus
0 0 2.14 2.14 2.22 0 0 0 0 0
Nyctalus noctula 0 0 0 0 0 0 0 0.24 0.24 2.22
Hypsugo savii 0.43 0 0 0.43 2.22 0.48 0.72 0 1.2 4.44
Pipistrellus kuhlii 24.79 22.65 24.36 71.79 80 33.17 23.56 14.9 71.64 82.22
P. kuhlii / H. savii 1.71 3.42 2.99 8.12 31.11 4.09 1.68 1.2 6.97 37.78
Pipistrellus pipistrellus 0.43 0.85 0.85 2.14 8.89 0 3.85 0 3.85 4.44
P. pipistrellus / P. kuhlii 0.43 0 0.43 0.85 4.44 0.48 0.72 0 1.2 8.89
Pipistrellus pygmaeus 0.43 3.85 2.14 6.41 13.33 0 6.97 0.48 7.45 13.33
Miniopterus schreibersii 0 0 0 0 0 0 0.48 0 0.48 2.22
M. schreibersii / P. pygmaeus 0 0 0 0 0 0 1.44 0 1.44 2.22
Myotis spp. 0.43 0.43 2.99 3.85 6.67 0.24 0 0 0.24 2.22
Tadarida teniotis 3.85 0.43 0 4.27 8.89 4.57 0.24 0.48 5.29 13.33
Total 32.48 31.62 35.9 100 43.03 39.66 17.31 100
The GLMMs indicated a positive effect of built cover (PC1), water
cover (PC2) and the year of study on bat activity (Tab. 2). Temperature
and distance to water bodies were non-significant and not included in
the final statistical model. Bat activity increased with increasing build-
ing and water cover, but it was strongly correlated with P. kuhlii activity
(r = 0.92). Therefore, a positive relationship between these variables
and P. kuhlii activity can be inferred.
RDA performed on the 2015 data did not show any significant re-
lationships between the environmental variables and bat community
composition (total variance explained=15.8%; model: F=1.888,
p=0.064; Fig. 3 a). In contrast, in 2016 a significant relationship
was found between PC2 (presence of water) and community com-
position (PC2: F=3.539, p=0.034; Fig. 3 ). The environmental vari-
ables accounted for only 14.4% of the total variance (model: F=1.675,
PERMANOVA showed a significant relationship between com-
munity composition and the urbanisation gradient, but only for 2016
(2015: F=0.359, p=0.912; 2016: F=2.329, p=0.037). Pairwise com-
parisons indicated a statistically significant difference between the sub-
urban and the urban zone in 2016 (F=3.319, p=0.026), while no differ-
ences were found between the urban and the peri-urban zone (F=0.345,
p=0.721) and between the suburban and the peri-urban zone (F=2.266,
This current study provides the first insights into bat activity and
diversity patterns in an urban area in southeastern Europe (Patras,
Greece), demonstrating the sensitivity of bat communities to urban-
isation and the differential responses among species. Bat diversity was
generally low in the city of Patras and its surroundings, although the
area lies within a biodiversity hotspot containing a high number of spe-
cies (Dietz et al., 2009, Papadatou and Georgiakakis, unpubl. data).
The bat community was dominated by P. kuhlii, which together with
the group P. kuhlii /H. savii comprised approximately 80% of the total
bat activity in the study area in both years. The rest of the species were
Table 2 Parameter estimates (±standard error) of fixed eects of the best GLMM.
Fixed effects Estimate (±S.E.) Z p
Intercept 1.257 (±0.191) 6.604 <0.001
PC1 0.323 (±0.151) 2.142 0.032
PC2 0.384 (±0.155) 2.479 0.013
Year 0.477 (±0.202) 2.365 0.018
much less abundant, likely implying decreased tolerance to the urban
environment in the study area.
Bat activity and species richness did not differ along the urbanisa-
tion gradient, contradicting the generally observed pattern of bat activ-
ity declining with increasing urbanisation intensity (Walsh and Harris,
1996; Hale et al., 2012; Russo and Ancillotto, 2015; Jung and Threlfall,
2016) or studies showing higher diversity in suburban areas (Hourigan
et al., 2010; Threlfall et al., 2011). Contrary to our expectations, the
peri-urban area did not hold greater bat activity or more species than
the urban and suburban areas, despite the fact that it includes less dis-
turbed landscapes with high vegetation cover (Tzortzakaki et al., 2019).
Bat activity was highest in the urban zone in the second sampling year.
As indicated by several studies, species richness may be generally low
in urban areas, but total abundance and biomass is often increased due
to the occurrence of a few synurbic species (Avila-Flores and Fenton,
2005; Francis and Chadwick, 2012; Krauel and LeBuhn, 2016).
A positive relationship between built cover and bat activity was in-
dicated by the GLMMs. These results may reflect the ability of P. kuh-
lii to exploit the urban landscape and forage around sources of artificial
illumination (Tomassini et al., 2014; Russo and Ancillotto, 2015; An-
cillotto et al., 2019), as the number of streetlamps was highly correlated
with built cover in the study area. Streetlamps seem to play an import-
ant role in P. kuhlii foraging, as they prey upon insects that are attracted
by the light (Rydell, 1992; Tomassini et al., 2014).
Surprisingly, vegetation cover did not have any positive effect on bat
activity, although it is considered a key landscape element for foraging
bats in urban environments, as it is associated with increased insect prey
abundances (Avila-Flores and Fenton, 2005 and references therein).
Furthermore, urban green spaces such as parks (Glendell and Vaughan,
2002; Gehrt and Chelsvig, 2003; Russo and Ancillotto, 2015), gardens
(Hale et al., 2012; Lintott et al., 2016) or even small green areas (Avila-
Flores and Fenton, 2005) can enhance insect abundances, while tree
networks in green spaces can facilitate bat commuting and foraging
(Dixon, 2012; Hale et al., 2012).
The lack of a positive relationship between vegetation cover and
urban green spaces, and bat activity in this study may in part be ex-
plained by the dominance of P. kuhlii. However, green spaces in the
study area may be too small or generally inadequate (e.g. due to in-
adequate vegetation structure) for most bat species, even in the peri-
urban zone. The latter is an agricultural area covered mainly by agri-
cultural mosaics and olive groves, which have been found to support
relatively high bat diversity in other Mediterranean areas (Russo and
Jones, 2003). A negative relationship between rural areas and bat activ-
ity was reported in Illinois, USA, where, unlike Patras, intensive crop
agriculture prevails (Gehrt and Chelsvig, 2003). More natural areas in
Bat community of an Eastern Mediterranean city
Figure 3 RDA ordination plots showing the relationship between the environmental variables (PC1: built cover, PC2: water cover, Temp: mean hourly temperature, DistW: distance to
water) and bat community composition in (a) 2015 and (b) 2016. No significant relationships were found, except for a significant relationship between water cover (PC2) and community
composition in 2016.
the surroundings of the study area (e.g. within the adjacent Natura 2000
site) may provide higher prey availability and, thus, be more attractive
to foraging bats.
Similar to other studies (e.g. Li and Wilkins, 2014), the highest bat
activity was in most cases recorded in specific sites with presence of
water, i.e. along the banks of water bodies. However, these sites varied
between the study years. In general, the highest total bat activity was
recorded at the main wastewater treatment plant of the city, while the
upstream section of Glafkos river (Fig. 1) and a remnant natural mire
also provide important foraging grounds. In the rest of the sites where
water is present such as streams, bat activity was lower than expected.
In contrast, one of the few remaining seaside open green spaces was
also important for foraging bats. Despite the positive effect of water
cover for foraging bats, site proximity to water did not have any ef-
fect on bat activity contrary to the findings of previous studies (Dixon,
2012; Ancillotto et al., 2015; Krauel and LeBuhn, 2016). These find-
ings imply that the main factors influencing bat activity are possibly
associated with water cover and/or particular characteristics of the wa-
ter bodies (e.g. shape or structure) and not with the site proximity to
water per se.
Water bodies in Patras have generally undergone degradation due
to modification of riverbeds, residential and industrial development,
draining and dumping of refuse, and, consequently, degradation of their
riparian character. Lintott et al. (2015) showed that even if waterways
are accessible to bats, their use may largely depend on the riparian ve-
getation cover, while it may also be negatively affected by the propor-
tion of the adjacent sealed surfaces.
Nevertheless, the value of the remaining water bodies for the local
bat community should be emphasised, as the highest bat activity was
recorded at some of these sites. In particular, P. pygmaeus was found
to have a positive (although not statistically significant) relationship
with water cover (Fig. 3b), which supports the findings of Nicholls
and Racey (2006) and Lintott et al. (2016) who found a strong asso-
ciation with freshwater (but see Hale et al., 2012). Water bodies such
as rivers and lakes are highly important for insectivorous bats, because
they provide high insect prey availability and drinking water (Vaughan
et al., 1997; Russo and Jones, 2003; Li and Wilkins, 2014). Linear
water bodies (waterways) may also function as corridors facilitating
bat movements between the urban core and peri-urban natural habitats
(Rouquette et al., 2013; Lintott et al., 2016). Hence, their maintenance
and management are important for bat conservation in large urban set-
tlements (Russo and Ancillotto, 2015).
The year of the study affected bat activity and this was more evid-
ent in sites located along water bodies (Tab. S2). In the second year,
bat activity was significantly higher, underlying that changes in spatio-
temporal patterns of bat activity may be due to either stochastic or spe-
cific factors related to bat biology. Differences between years may re-
flect changes in environmental conditions or bat behaviour (O’Donnell
and Sedgeley, 2001). In this study, differences in bat activity observed
between the two years may be attributed to the prevailing weather con-
ditions preceding the sampling, as the winter and spring of the second
year were significantly warmer. Higher temperatures may increase in-
sect abundance (Scanlon and Petit, 2008), but may also influence bat
reproductive success (Ancillotto et al., 2015). They may lead to ad-
vanced parturition, which in turn may result in increased juvenile sur-
vival (Frick et al., 2010) or more females reaching sexual maturity dur-
ing the first autumn (Dietz et al., 2009). The scenario of increased prey
availability is considered more plausible, since winters in the study area
are generally mild and do not show strong fluctuations among years,
while the influence of other stochastic undetermined factors cannot be
Our findings are in agreement with the conclusion of other studies
that the response of bats to urbanisation is highly species-specific (e.g.
Avila-Flores and Fenton, 2005; Threlfall et al., 2012; Russo and Ancil-
lotto, 2015). The bat community was mainly represented by P. kuhlii,
particularly in the urban zone in the second year of the study. Pipistrelle
bats such as P. kuhlii,P. pipistrellus and H. savii have been reported to
successfully inhabit urban areas and expand their range across Europe
(Russo and Ancillotto, 2015; Ancillotto et al., 2019). Specifically in
southern Europe, P. kuhlii has shown notable behavioural and ecolo-
gical flexibility and has adjusted to artificial structures such as build-
ings and streetlamps for roosting and foraging, respectively (Russo and
Jones, 2003; Ancillotto et al., 2015), while P. pipistrellus occupies a
similar niche in Northern Europe (Vaughan et al., 1997).
Similarly, T. teniotis was mainly recorded in the urban zone, demon-
strating its ability to exploit human-modified landscapes and man-made
structures (Avila-Flores and Fenton, 2005; Threlfall et al., 2012; Krauel
and LeBuhn, 2016). Tadarida species have been documented flying
over cluttered environments and large areas such as the dense city core
(Russo and Jones, 2003; Avila-Flores and Fenton, 2005; Krauel and Le-
Buhn, 2016), foraging on passing insect swarms (Marques et al., 2004).
It should be noted that in this study, its occurrence may have been un-
derestimated, as it usually flies high above the ground and produces low
frequency sounds (Zbinden and Zingg, 1986).
Hystrix, It. J. Mamm. (2019) — online first
Pipistrellus pygmaeus and P. pipistrellus were rarely detected in the
urban zone but were in general not common in the study area (<10%
of the total bat activity). These two pipistrelle species together with
M. schreibersii were principally recorded in the suburban zone and
they showed higher activity in sites near water bodies especially in the
second year (Tab. S2). As mentioned above, P. pygmaeus is associated
with freshwater and avoids densely built areas (Nicholls and Racey,
2006; Lintott et al., 2016), whereas P. pipistrellus is considered a gen-
eralist and more tolerant of intermediate urbanisation levels (“urban
adapter”, Hale et al., 2012), and thus, occurs in a broader spectrum of
habitats (Lintott et al., 2016). The latter species appears to avoid hab-
itats with high freshwater coverage, in order to avoid competition with
P. pygmaeus (Nicholls and Racey, 2006; Lintott et al., 2016). In this
study, P. pipistrellus was principally found in the wastewater treatment
plant foraging together with P. pygmaeus and other species, but it was
rare in any other habitats. Miniopterus schreibersii was only found at
the wastewater treatment plant.
Larger species such as Nyctalus sp./E. serotinus were only detected
in one site in the peri-urban zone. Nyctalus species have been reported
to roost in buildings and forage in parks and city edges, if high veget-
ation and insect densities are provided (Dietz and Kiefer, 2014), and
in urban waterways (Lintott et al., 2015), while E. serotinus often for-
ages in settlements and big cities (Dietz and Kiefer, 2014). Both N.
leisleri and E. serotinus have been recorded to forage at streetlights
(Azam et al., 2015). However, in this study these species were rarely
recorded, thus, no conclusions on their occurrence can be drawn. Sim-
ilarly, Myotis spp. were mostly found in sites out of the urban zone and
close to water bodies, in accordance with other studies (Vaughan et al.,
1997; Russo and Jones, 2003; Avila-Flores and Fenton, 2005; Dixon,
2012; Lintott et al., 2015).
The negative impact of urbanisation on many bat species due to hab-
itat fragmentation and patch isolation (Hale et al., 2012; Threlfall et
al., 2012), reduced vegetation cover (Lintott et al., 2016), and artificial
lighting (Azam et al., 2016) possibly explains the low number of spe-
cies recorded in the urban and the suburban zones. However, it is un-
clear why bat diversity was low in the peri-urban zone, despite retaining
significant natural and semi-natural characteristics (Tzortzakaki et al.,
2019). The effects of some landscape aspects such as land-cover were
investigated at a relatively small scale (200 m), considering that bats are
highly mobile organisms and that species responses may vary depend-
ing on their different foraging and roosting requirements (Dixon, 2012;
Hale et al., 2012; Lintott et al., 2016). Previous studies have underlined
the importance of local habitat features such as vegetation structure,
water bodies and streetlamps on urban bat communities, but also of the
surrounding landscape (Walsh and Harris, 1996; Gehrt and Chelsvig,
2003; Threlfall et al., 2012; Lintott et al., 2015). Hence, further in-
vestigation of the local habitat characteristics (e.g. vegetation structure
and the existence of buildings providing bat roosts) and the landscape
structure (e.g. habitat heterogeneity and connectivity) at a larger scale
may provide insight into the factors affecting bat community response
to urbanization, especially in the peri-urban zone. Bat surveys in the
adjacent natural areas would be useful as it would allow the magnitude
of the effect of urbanisation on bat communities to be assessed.
Finally, we need to acknowledge that the low bat diversity and the
rare occurrence of most species may have resulted from the sampling
methodology (i.e., short acoustic transect surveys), which was designed
to be consistent with two companion studies of birds (Tzortzakaki et al.,
2018) and butterflies (Tzortzakaki et al., 2019). Additionally, surveys
were not conducted during the breeding season (late spring – mid sum-
mer) and this would likely provide a better assessment of bat activity
patterns in the urban environment.
Despite its limitations, our study provides an insight into the effects
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Associate Editor: L. Ancillotto
Supplemental information
Additional Supplemental Information may be found in the online version of this arti-
Table S1 Monthly mean temperature (T), relative humidity (RH) and total monthly
rainfall recorded for each sampling year.
Table S2 Total number of sampling sites with species occurrences, number of sites
near water bodies (<100 m) with species records, and proportion of bat passes
of each species recorded in sites near water bodies for each survey year.
... Secondly, animals are limited by different morphological, physiological and behavioural constraints that make adjusting to urban ecosystems difficult ensuring that only a few species dominate this ecosystem. Urban ecosystems have often shown reduced diversity and dominance by a handful of species that have adapted well in human-modified landscapes (Tzortzakaki, et al., 2019). Urban landscapes often have a limited area available for foraging. ...
... The results show that the influenced by vegetation parameters such as vegetation species richness and canopy height on bat activity was inconclusive at the scale of sampling that is, 15 m radial plots ( However, another study reported that vegetation cover did not influence the bat activity in areas of varying degrees of urbanisation (Tzortzakaki et al., 2019). In this study, exploratory analysis using bar graphs and boxplot were made to observe patterns of bat activity and individual vegetation parameters. ...
... P. ceylonicus/ Scotophilus species group was the most commonly recorded in two out of three sites. Urban ecosystems often tend to support a few species which dominate the ecosystems (Tzortzakaki et al., 2019). This species group showed hourly variation in activity peaks and low activity throughout the night. ...
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The increase in urbanisation over the years has affected wildlife and negatively impacted their habitats. The impact of urbanisation can be at a global-scale such as carbon emissions from cities and at a regional-scale like the effects of urban sprawl on neighbouring habitats and species. The effect of urbanisation is often documented as negatively influencing wildlife. Although urbanisation does have a negative impact on different species, some species have learnt to tolerate and even thrive in human habitations. Such species-specific responses to different urban areas dictate which species dominate the urban ecosystem. The aim of this study was to provide information about bat activity in urban green spaces, to understand what factors influence bat activity, to know the species-specific responses to urbanisation parameters and test the effectiveness of active and passive acoustic survey methods in an urban landscape. This study was conducted between January 2020 and February 2020. To understand the use of these green spaces by different species, variables such as green space area, edge perimeter of green space, distance to main road and distance to floodlights and vegetation parameters including canopy height, canopy cover, species richness, tree density, floristic composition of trees and shrubs, phenological state of trees and shrubs were obtained during the sampling period. I found 3 species Pipistrellus tenuis, Hipposideros speoris, Tadarida aegyptiaca, and a species group Pipistrellus ceylonicus or Scotophilus species during the period of this study. There was a difference in the number of detections of bats between the study sites. From personal observations, bats foraging around floodlights in one of the sites evidently increased the bat activity in the area. The influence of vegetation parameters on bat activity was not conclusive. Size of a green patch might have an influence on the activity of P.tenuis. This study highlights the first information on ‘light-opportunistic' bats in India. Pipistrellus tenuis, Tadarida aegyptiaca, and the species group Pipistrellus ceylonicus /Scotophilus species seem to be well-adjusted to the urban environment. It is necessary to note that although artificial lighting at night seems to positively influence the activity of these species, it could deter other ‘light-shy’ or ‘light-averse’ species which were not detected during this study. Activity patterns vary between species and the overlap is higher between the P. ceylonicus /Scotophilus species x group and P. tenuis than with T. aegyptiaca. Active monitoring (transects) detects P. tenuis well but misses out on rare species and underestimates T. aegyptiaca and P. ceylonicus / Scotophilus. The findings of this study are preliminary further studies are required to understand species-specific responses to urbanisation.
... The urban environment is perhaps the most human-modified type of landscape. Nevertheless, it also has its own fauna; ecologically flexible animal species (including bats) adapt to such uncharacteristic living conditions, populating not only park zones but also buildings and other urban developments [2,3]. At the same time, the attention of researchers, especially in the tropics, has been mainly focused on the inhabitants of primary natural habitats, while urban fauna remain "behind the scenes". ...
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Bats are the second largest order of mammals, with about 1400 known species [...]
... Bats use a variety of urban habitats throughout the year. Research indicates that different bat species vary in their response to urbanization [5]. Urbanization provides roosting opportunities and facilitates range expansion of particular Central European bat taxa [6,7]. ...
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Introduction: Bats are considered natural reservoirs for lyssaviruses. A total of 17 out of 19 known lyssaviruses circulate in bat populations. Lyssaviruses cause rabies in animals and humans. The transmission of lyssaviruses from European bats to terrestrial animals and humans is rare, but the risk of infection still exists even in developed countries. Slovakia is currently a rabies-free country. Objective: The aim of the study was to assess the potential circulation of EBLV-1 in synanthropic bats present in human inhabited buildings, and to give an overview of human exposure to bats. Material and methods: A passive serological survey targeted the prevalence of antibodies to bat lyssaviruses in synanthropic bats between 2009 - 2019. A total of 598 bats of the species Pipistrellus pipistrellus, Pipistrellus pygmaeus, Eptesicus serotinus, Nyctalus noctula and Vespertilio murinus were captured in buildings mainly in Eastern Slovakia, and examined by the rapid fluorescent focus inhibition test (RFFIT). Results: Lyssavirus-specific antibodies were detected in 2 (0.3%) of the 598 examined bats. Additionally, brain tissues of bats found dead were examined using the standard fluorescent antibody test (FAT) with negative results. An overview of available data on human exposure to bats recorded in Slovakia from 2007 - 2019 is also included. Conclusions: The study confirmed the presence of lyssavirus antibodies in synanthropic bats in Slovakia, suggesting the active circulation of bat lyssaviruses in bat populations exploiting human buildings. Although the seroprevalence was found to be extremely low, the results show that any case of human exposure to bats must be treated with caution in order to protect public health.
... Bats inhabit urban areas worldwide. Certain bat species are better at adapting to the urban environment than others [33][34][35]. Nocturnal insectivorous bat activity peaks within the first few hours following sunset [36,37], which is a pattern that overlays temporally with human leisure nightlife. In the current literature, there is no evidence suggesting that urban bats avoid flying over human crowds or areas of high human activity. ...
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In the urban environment, wildlife faces novel human disturbances in unique temporal patterns. The weekend effect describes that human activities on weekends trigger changes in the environment and impact wildlife negatively. Reduced occurrence, altered behaviors, and/or reduced fitness have been found in birds, ungulates, and meso-carnivores due to the weekend effect. We aimed to investigate if urban bat activity would differ on weekends from weekdays. We analyzed year-round bat acoustic monitoring data collected from two sites near the city center and two sites in the residential area/park complex in the city periphery. We constructed generalized linear models and found that bat activity was significantly lower on weekends as compared to weekdays during spring and summer at the site in the open space near the city center. In contrast, during the same seasons, the sites in the city periphery showed increased bat activity on weekends. Hourly bat activity overnight suggested that bats might move from the city center to the periphery on weekends. We demonstrated the behavioral adaptability in urban wildlife for co-existing with human. We recommend that urban planning should implement practices such as adding new greenspaces and/or preserving old-growth vegetation to form continuous greenways from the city center to the city periphery as corridors to facilitate bat movements and reduce possible human-wildlife conflict.
A atividade humana é um elemento chave na transformação do ambiente natural.O avanço da urbanização, o aumento de poluentes e a degradação de áreasambientais são apenas alguns pontos presentes no processo transformador queo homem exerce sobre a natureza. Não obstante, os impactos antrópicos afetamdiretamente o comportamento e ecologia dos animais que habitam em seuentorno, podendo alterar seus hábitos. Dessa maneira, o objetivo deste trabalhofoi avaliar as alterações comportamentais oriundas do contato com a poluiçãosensorial nos mamíferos não humanos presentes nas cidades. Para tal, foirealizada uma revisão bibliográfica, buscando elencar os principais dados sobrea temática. Foram utilizados os portais de busca Google Acadêmico e Web of Science, com combinações de palavras em português e inglês. Os trabalhosselecionados passaram pelo critério de inclusão de apresentar informaçõesrelevantes sobre poluentes sensoriais e sua relação com a mastofauna. Após aanálise do conteúdo, obtiveram-se 36 publicações que abordavam o tema deforma relevante. Através delas ficou evidente que a poluição sensorial e suarelação com os mamíferos ainda é um campo a ser estudado. Os organismosapresentam reações específicas aos diferentes tipos de poluentes,demonstrando que a atividade humana pode afetar os indivíduos de diversasmaneiras, afetando suas relações sociais e sua busca por recursos. Constatase que maiores estudos são necessários para a compreensão dos reais impactosda presença antrópica na história de vida desses animais, também buscandoangariar informações para a construção de planos de manejo e urbanos queconsigam mitigar os efeitos adversos da atividade humana no ambiente.
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In ever larger and denser cities, spaces with a nature character provide ecosystem services, enable biodiversity conservation and are meeting places for humans and non-humans. It is therefore key for decision-makers, city designers and city managers to be able to rapidly characterise the biodiversity housed in these spaces in order to reconcile the population’s expectations and environmental issues in development projects and management methods. Here, we aim to identify the factors determining the species assemblages observed in the 10 parks of the city of Aix-en-Provence (characteristics of the parks, management methods, types of urban context), and this for two taxa with different ecological requirements: avifauna and chirofauna. We contacted 39 species of birds. Bird communities become more homogenous from the periphery to the city centre, and the richest communities are found in the larger parks, with a greater diversity of habitats and surrounded by more greenery. We also recorded more than 53,000 passings of, bats, indicating a rather high level of activity for urban Mediterranean parks. There is a very strong heterogeneity of activity between parks, which is not explained by the characteristics of the parks. The park with the highest activity rate is a medium-sized park embedded in the dense city. Our results confirm that the question of biodiversity in the city must be asked by considering several spatial scales and several taxa. The habitat potential and vegetation cover in and around the park is a critical dimension for the planning and management of the city.
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Urbanization is a severe threat to global biodiversity, often leading to taxonomic and functional homogenization. However, current urban ecology research has focused mostly on urban birds and plants, limiting our ability to make generalizations about the drivers of urban biodiversity globally. To address this gap, we conducted a global meta-analysis of 87 studies, including 180 bat species (Chiroptera) from urban areas in Asia, Australia, Europe, North and South America. We aimed to (i) understand the importance of functional traits and phylogeny in driving changes in urban bat assemblages, and (ii) assess the capacity of traits for predicting which types of species are most sensitive to urbanization. Our results indicate that species-specific functional traits explain differences in the intensity of urban habitat use. Urban tolerance mainly occurred within the open and edge space foraging and trawling species as well as in bats with flexible roosting strategies. In addition, across bioregions and independent of phylogeny, urban tolerance correlated with higher aspect ratio, a trait enabling fast flight but less agile manoeuvres during aerial food acquisition. Predictive success varied between bioregions, between 43 and 83%. Our analysis demonstrates that the local extinction of bat species in urban areas is non-random, trait-based and predictable, allowing urban landscape managers to tailor local conservation actions to particular types of species.
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Structural complexity is known to determine habitat quality for insectivorous bats, but how bats respond to habitat complexity in highly modified areas such as urban green spaces has been little explored. Furthermore, it is uncertain whether a recently developed measure of structural complexity is as effective as field-based surveys when applied to urban environments. We assessed whether image-derived structural complexity (MIG) was as/more effective than field-based descriptors in this environment and evaluated the response of insectivorous bats to structural complexity in urban green spaces. Bat activity and species richness were assessed with ultrasonic devices at 180 locations within green spaces in Vienna, Austria. Vegetation complexity was assessed using 17 field-based descriptors and by calculating the mean information gain (MIG) using digital images. Total bat activity and species richness decreased with increasing structural complexity of canopy cover, suggesting maneuverability and echolocation (sensorial) challenges for bat species using the canopy for flight and foraging. The negative response of functional groups to increased complexity was stronger for open-space foragers than for edge-space foragers. Nyctalus noctula, a species foraging in open space, showed a negative response to structural complexity, whereas Pipistrellus pygmaeus, an edge-space forager, was positively influenced by the number of trees. Our results show that MIG is a useful, time- and cost-effective tool to measure habitat complexity that complemented field-based descriptors. Response of insectivorous bats to structural complexity was group- and species-specific, which highlights the need for manifold management strategies (e.g., increasing or reinstating the extent of ground vegetation cover) to fulfill different species’ requirements and to conserve insectivorous bats in urban green spaces.
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Time of peak bat activity during the night differs among bat species due to temperature, prey availability, habitat availability, and/or interactions between species. Habitat availability is altered in urban areas, which may affect insect prey availability and interspecies interactions. Our objectives were to use mobile acoustic monitoring to determine when bat species were active in a single night in urban and nonurban sites and if nightly bat activity patterns differed in urban versus nonurban sites. Bat echolocation call sequences were recorded using Anabat acoustic detectors while driving transects through the night at five sites (three “urban” and two “nonurban”) located in the Piedmont region of north-central North Carolina from May through August 2016. Transects were driven three times per night starting 45 min, 180 min, and 300 min after sunset. Recorded echolocation call sequences were analyzed manually using AnalookW and automatically using Bat Call Identification and Echoclass software. Total bat activity was not different between urban and nonurban sites. However, total bat activity was lower later in the night in urban sites, but stayed the same in nonurban sites. Species specifically, there were more Eptesicus fuscus, Lasionycteris noctivagans, and Tadarida brasiliensis call sequences and fewer Lasiurus borealis, Nycticeius humeralis, and Perimyotis subflavus call sequences in urban sites than nonurban sites. There were also fewer E. fuscus, L. noctivagans, and N. humeralis call sequences later in the night in both urban and nonurban sites. Only Lasiurus borealis activity in urban sites later in the night reduced and L. borealis activity in nonurban sites remained at the same. These results suggest that bats in urban areas partition time differently, which is important to consider for urban conservation efforts and planning.
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Urbanization causes rapid changes in the landscape and land use, exerting a significant pressure on bird communities. The effect of urbanization on bird diversity has been widely investigated in many cities worldwide; however, our knowledge on urban bird communities from the eastern Mediterranean region is very scarce. In this context, we aimed to investigate the effect of the different land-cover types on bird species richness and abundance in a densely built coastal Mediterranean city (Patras, Greece) during the breeding and wintering seasons. We sampled the bird community in 90 randomly selected sites along an urbanization gradient. Open green spaces proved to be the most significant factor favouring bird diversity in both seasons. In winter, woody vegetation and impervious surfaces had a positive effect on species richness as well. The local bird community consisted of a large number of species associated with open and semi-open unmanaged green areas, 12 of which are Species of European Conservation Concern (SPECs) showing a declining trend in Europe. On the other hand, in winter the number of forest-dwellers increased significantly. Species richness was significantly higher in winter indicating that the urban environment provides important wintering grounds. Thus, management actions in cities with similar characteristics in the Mediterranean region should focus on the maintenance of open green spaces and woody vegetation patches to enhance bird diversity.
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Bat social calls are specifically tailored for communication and play different roles according to their structure. Their structure is stereotyped and species-specific so they may aid acoustic identification of bats. To provide the first quantitative description of social call repertoire of Hypsugo savii and offer a way to identify this bat in free flight we made audio recordings in four areas of Central and Southern Italy in summers 2011–2015. We identified single and multiple-component social calls and categorized them into five structure types. Within each structure type, call frequency and / or duration differed between single and multiple-component calls, the latter being shorter and showing higher frequencies. In multiple-component calls both the number of syllables and the way they were associated were highly variable, making it difficult to recognize patterns. Some motifs, however, showed the same first component type and final sequence. We also recorded trill-like calls and two complex sequences of multiple-component social calls (songs) lacking repeated motifs. The complex association of syllables, the rarity of recurrent motifs and the significant structural flexibility suggest that social calls serve a range of scopes and that they might convey acoustic signatures and other individual-specific features.
Ponds have an important role in the ecology of urban areas, as they provide essential habitats to aquatic species, as well as fundamental resources to terrestrial wildlife. Artificial water sites such as urban ponds provide foraging and drinking resources to synurbic wildlife, among which bats stand out as an important group. Availability of water sources may thus strongly influence the persistence of animal populations in urban habitats. Pond characteristics, as well as landscape structural patterns in the surrounding area, may modulate the use of such water sites by bats. We investigated bat species richness and activity levels in a pond archipelago within the city of Rome, one of the largest urban areas in Italy. We hypothesized that the presence of woody vegetation and hedgerows affects activity rates over ponds and that bat responses to habitat and landscape structures as well as artificial illumination are species-specific. Bat species richness was mainly influenced by the availability of wooded vegetation within 1000 m around ponds, with minor effects of the amount of bank habitat, while bat activity was affected by different habitat features in a species-specific way. All species responded positively to pond proximity to linear landscape elements such as hedgerows and to the amount of bank habitat. The presence of natural banks, the amount of woodland and that of open green areas positively influenced the activity of different species at different scales, while distances between ponds and artificial lights had a species-specific effect direction. Our results highlight the importance of key factors characterizing ponds and the surrounding habitat in urban landscapes whose appropriate management may improve the viability of synurbic bat populations.
Urbanization induces rapid landscape and habitat modifications leading to alterations in species distribution patterns and biodiversity loss. As pollinating insects such as butterflies are particularly susceptible to urbanization, it is important to pinpoint the factors that could enhance their diversity in the urban areas in order to design adequate management and conservation actions. Our study aims to investigate the influence of land cover and local habitat characteristics on the butterfly diversity patterns and community structure in a densely built city in the eastern Mediterranean region. We carried out butterfly surveys (line transects) in 45 randomly selected sites, distributed along an urbanization gradient. In each site, we assessed the surrounding landscape by measuring the land cover in a 200-m buffer zone, and the local habitat by estimating the available plant resources along each transect. Overall, 1805 individuals belonging to 41 butterfly species were recorded. Land cover was found to have the strongest influence on butterfly species richness, abundance and community structure. Although plant resources were sufficiently available within the whole study area, the butterfly community was significantly poorer in the more urbanized areas, indicating the potential role of habitat fragmentation and patch isolation. In contrast, butterfly diversity was significantly higher in the peri-urban area, underlying its conservation value for butterflies in the urban landscape. We attribute these findings to the degradation of the more urbanized areas due to long-term inadequate planning and the disorganized expansion of the city.
Human-induced alterations often lead to changes in the geographical range of plants and animals. While modelling exercises may contribute to understanding such dynamics at large spatial scales, they rarely offer insights into the mechanisms that prompt the process at a local scale. Savi's pipistrelle (Hypsugo savii) is a vespertilionid bat widespread throughout the Mediterranean region. The species' recent range expansion towards northeastern Europe is thought to be induced by urbanization, yet no study actually tested this hypothesis, and climate change is a potential alternative driver. In this radio-telemetry study, set in the Vesuvius National Park (Campania region, Southern Italy) we provide insights into the species' thermal physiology and foraging ecology and investigate their relationships with potential large-scale responses to climate, and land use changes. Specifically, we test whether H. savii i) exploits urbanisation through a selection of urban areas for roosting and foraging, and ii) tolerates heatwaves (a proxy for thermophily) through a plastic use of thermoregulation. Tolerance to heatwaves would be consistent with the observation that the species' geographic range is not shifting but expanding northwards. Tracked bats roosted mainly in buildings but avoided urban habitats while foraging, actively selecting non-intensive farmland and natural wooded areas. Hypsugo savii showed tolerance to heat, reaching the highest body temperature ever recorded for a free-ranging bat (46.5 °C), and performing long periods of overheating. We conclude that H. savii is not a strictly synurbic species because it exploits urban areas mainly for roosting, and avoids them for foraging: this questions the role of synurbization as a range expansion driver. On the other hand, the species' extreme heat tolerance and plastic thermoregulatory behaviour represent winning traits to cope with heatwaves typical of climate change-related weather fluctuations.
Urban land cover is the fastest growing land-use form globally and there is concern that urbanisation will negatively impact native biodiversity. Bats are ecologically diverse predators and their responses to urban development may provide insights into wider biodiversity responses to urbanisation. Developments in bat detection methods mean it is now possible for citizen scientists to collect detailed bat distribution data. The geographical and habitat coverage of such data make them ideal for addressing urban planning issues. In this paper we quantify the impact of planned housing on bat populations and evaluate possible mitigation measures. We combined data on 12 bat species collected through a large citizen science project in Norfolk, UK, with spatially explicit housing plans for the next decade and tested the impact of mitigation planning scenarios operating at different spatial scales. The planned housing was predicted to decrease occurrence or activity for all 12 bat species. Locally, these decreases could be substantial, leading to a reduction in the likelihood of occurrence from 40% to 1%. However, at a county-scale the proposed level of housing is equivalent to less than a 2% decrease in total occurrence and abundance across all species. The negative effect of planned housing could be reduced by 46% on average by preferentially building on less preferred habitats and in areas with low populations of urban-sensitive bat species. This paper demonstrates an easily transferable method for determining rich habitats where new developments should be avoided and for investigating the potential of mitigation strategies.