Landscape and Urban Planning 122 (2014) 186–195

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Landscape and Urban Planning

journal homepage: www.elsevier.com/locate/landurbplan

Research Paper

The effect of plant richness and urban garden structure on bird

species richness, diversity and community structure
Yair Paker a , Yoram Yom-Tov b , Tal Alon-Mozes c , Anat Barnea d,∗
a
Porter School of Environmental Studies, Tel-Aviv University, Israel
b
Department of Zoology, Tel-Aviv University, Israel
c
Faculty of Architecture and Town Planning, Technion – Israel Institute of Technology, Israel
d
Department of Natural and Life Sciences, The Open University of Israel, Israel

h i g h l i g h t s

• Bird species diversity is positively related to shrubs species richness.

• Richness of trees and shrubs, small lawns and tree cover invite most bird species.
• Native birds prefer to forage on native trees, while alien birds – on alien trees.
• Bird species diversity changes with migrating seasons.
• Presence of people and dogs has a negative effect on birds’ presence.

a r t i c l e i n f o a b s t r a c t

Article history: Urban green areas improve the standard of living in cities and affect people’s attitude to nature and
Received 27 September 2012 conservation. Zoological knowledge may provide data that will help designers to enhance bird diversity
Received in revised form 13 October 2013 in gardens. We studied the effect of plant species richness and structure on bird species richness, diversity
Accepted 16 October 2013
and community structure in 25 public gardens in Tel-Aviv city and, neighboring suburbs, Israel. A total
Available online 13 November 2013
of 65 bird species were observed, of which nine were urban, exploiters or alien species. These latter
species composed 54% of all individuals seen. Additional 13 bird species, mostly migrants, were observed
Keywords:
in gardens further from the observation fixed radius. We found that shrubs species richness positively
Birds
Gardens affected bird species diversity. Most bird species were found where trees and shrubs species richness
Native and aliens plants was high, and trees and lawn cover were medium or low. High trees or high lawn cover attracted only
Urban biodiversity conservation a few bird species, mostly aliens and urban exploiters. Native birds preferred to forage on native trees
Urban ecology and alien birds preferred to feed on alien trees. Bird species diversity was higher during spring and fall
Urban planning because of the presence of migrating bird species. Dogs and people had a negative effect on bird presence.
Accordingly, we recommend that when planning new gardens, designers will avoid large lawns, prefer
diverse and dense shrubberies, native trees, and will create some areas that will not be accessible to dogs
and people. Finally, we emphasize the importance of multidisciplinary studies conducted in collaboration
between landscape designers and zoologists.
© 2013 Elsevier B.V. All rights reserved.

1. Introduction
Currently, almost half the world’s human population lives in
cities and by 2030 the proportion living in cities is expected to reach
60% (United Nations, 2004). Urban population growth causes nat-
ural habitat loss by conversion of natural habitats into urbanized
areas (McKinney, 2002), resulting in biodiversity homogenization
∗ Corresponding author at: Department of Natural and Life Sciences, 1 University
Road, Raanana 43107, Israel. Tel.: +972 9 7781753.
E-mail addresses: yairpk@gmail.com (Y. Paker), yomtov@post.tau.ac.il
(Y. Yom-Tov), artal@technion.ac.il (T. Alon-Mozes), anatba@openu.ac.il (A. Barnea).
(Blair, 1996) and decreased native biodiversity (Bino et al., 2008;
Blair, 1999; Czech, Krausman, & Devers, 2000; Kendle & Forbes,
1997; Mason, Moorman, Hess, & Sinclair, 2007). Because urbaniza-
tion and its consequences occur worldwide, there is an agreement
that the ecological outcome could be staggering, and therefore it is
essential to monitor patterns and trends in urban areas. For exam-
ple, Puth and Burns (2009) used species richness, a widespread
ecological metric, to assess the status and changes in biologi-
cal diversity of flora and fauna in the New York metropolitan
area over time. They argue that using such quantitative metrics is
advantageous, as they can serve as indicators for trends in produc-
tivity, invasibility, extinction, and stability. Nevertheless, in a recent
review, Magle, Vernon, and Crooks (2012) show that although

0169-2046/$ – see front matter © 2013 Elsevier B.V. All rights reserved.
http://dx.doi.org/10.1016/j.landurbplan.2013.10.005
 Y. Paker et al. / Landscape and Urban Planning 122 (2014) 186–195
urbanization impacts on wildlife, trends in urban wildlife stud-
ies have not been evaluated systematically, and nearly all were
conducted in North America, Europe, or Australia.
Urban nature, found in parks and smaller urban green areas
(such as public and private gardens), improves the standard of
living in cities (Miller, 2005) and can affect people’s attitude to nat-
ural ecosystems and conservation (Savard, Clergeau, & Mennechez,
2000; Tilghman, 1987). Thus, the quality of urban environments,
and particularly of urban green spaces, is increasingly regarded as
an important issue (Gaston et al., 2007). These green spaces that
are set aside for recreational use include parks (which are rela-
tively large and can be in their natural or semi-natural state), or
gardens (which are smaller and planned-spaces). In assessing the
quality of urban gardens, birds became a subject for considerable
research (Sandstrom, Angelstam, & Mikusinski, 2006) for several
reasons: most birds are diurnal, conspicuous and can be identified
and observed with relative ease and hence their spatial variation
can be well recorded (Bino et al., 2008). In addition to this practical
reason, in comparison with other taxa (e.g. invertebrates, reptiles,
amphibians), birds are probably the one taxon that people are
most familiar with, and can identify to a certain extent. Moreover,
birds are the taxon that people most actively attempt to engage
with, for example by providing them with feeders or nesting boxes
(reviewed by Baker, Thomas, Newson, Thomson, & Paling, 2010).
In England, for example, as a consequence, the status of bird popu-
lations is utilized by the government as a ‘quality of life’ metric
for urban occupants (Dept. of Environment, Food and Rural Affairs
[Defra], 2003).
Bird communities can be evaluated in several ways: (1) Bird
species richness, which is the number of bird species found in
one habitat. (2) Bird species diversity, which can be calculated
by using indices such as Shannon index that takes into account
both the number of species present and their relative abundance
in the community (Magurran, 1988). (3) Bird community struc-
ture, which is typified by an assemblage of several species that
are associated with a certain habitat and tend to appear together
in this habitat. For example, a recent study in England identified
three breeding bird communities in Bristol: a rural community
(associated with woodland, managed grassland and inland water),
suburban community (associated with buildings and residential
gardens), and intermediate community (that shared some of these
habitats characteristics) (Baker et al., 2010). By characterizing bird
communities we can increase our understanding of the interac-
tions between birds and various urban habitats they use. Also,
we can provide further insights on the effects that environmental
variables have on different bird species, an improvement over cal-
culating only species richness and diversity. Such environmental
variables, which might affect fitness of birds, may include physi-
cal habitat diversity, for example – structures that resemble cliffs
(high rising buildings), structures that provide nesting sites that
are naturally limited (i.e. holes) in illumination poles and traffic
lights, exotic vegetation with new feeding and nesting opportuni-
ties, etc.
Several urban factors are known to affect these indices. For
example, in a review article, Goddard, Dougill, and Benton (2010)
discuss mechanisms for encouraging ‘wildlife-friendly’ manage-
ment of collections of gardens across scales from the neighborhood
to the city, and one of their assumptions is that garden size pos-
itively affects species richness. They based this assumption on
previous studies (e.g. Daniels & Kirkpatrick, 2006) that found that
bird species richness in Australia was positively related to garden
size within the range of 50–1600 m2 . Garden location in relation to
city center also affects bird species richness and total abundance:
bird species richness decreases from natural or rural areas to city
centers (Bino et al., 2008) while bird total abundance increases
(Blair, 1999). Urbanization level around gardens or parks affects
187
bird species composition: in sites that are surrounded by simi-
lar urbanization levels bird species composition is similar. Level
of urbanization in the surrounding area can be more important
than site size and plant structure in determining bird commu-
nity composition (Huste & Boulinier, 2011). Urban environments
can cause an increase of total bird richness and abundance. For
example, in Britain it was found that these two indices increased
over a wide range of household densities and then declined at
greater household densities (Taratalos et al., 2007). However, the
decline occurred at house densities below the one required in
new developments, indicating the difficulty in maintaining a bal-
ance between biodiversity conservation and urban planning. Birds
are also sensitive to landscape composition and configuration
(Pellissier, Cohenb, Boulayb, & Clergeau, 2012).
The level of urbanization may have a differential effect on
indigenous, migrant and invading species. On one hand, in
countries with medium to high precipitation, low urbanization
level may be synonymous with high plant density, thus “inviting”
indigenous birds and arboreal migrants. Indigenous birds may also
be attracted to native vegetation that grows spontaneously in such
areas. On the other hand, high urbanization level may be accompa-
nied by more opportunities for invading species that are attracted
to garbage. It may also provide more opportunities to invading
urban exploiters that prefer to nest in holes (i.e. the Indian mynah
Acridotheres tristis and the Rose-ringed Parakeet Psittacula krameri)
or on ledges in buildings (i.e. laughing dove Streptopelia senegalen-
sis) that in Israel are abundant in areas of high urbanization level.
Studies that report a positive relation between bird richness and
urban environment explain this outcome by the presence of urban
exploiters and alien bird species that thrive in urban areas (Chace &
Walsh, 2006), and a recent review (Lowry, Lill, & Wong, 2013) points
out that species that have greater behavioral flexibility to the new
selection pressures presented by cities should have greater suc-
cess in urban habitats. Other studies characterize species that favor
urban development as generalist, which feed on plant material and
nest above the ground (Evans, Chamberlain, Hatchwell, Gregory,
& Gaston, 2011). Similarly in Israel, Kark, Iwaniuk, Schalimtzek,
and Banker (2007) found that being successful in more urbanized
environments depends on a combination of traits, including diet,
sociality, sedentariness and preferred nesting sites. They report that
ground nesters are rarely seen in dense city areas, because humans
or domesticated animals threaten them. However, nesting above
the ground is also not always successful in urban areas, and exper-
imental studies in Finland indicate that predation affected birds in
the city, and nest predation was higher in the town center than
in the less urbanized area of detached houses (Huhta, Jokimäki, &
Rahko, 1999; Jokimaki & Huhta, 2000). Most of the nests in the town
center were destroyed by avian predators.
Plant composition in urban gardens is another important fac-
tor that affects birds’ communities: High coverage of shrubs and
tall trees were found to be important for native forest birds in Tas-
mania (Daniels & Kirkpatrick, 2006). Adult trees and heterogenic
plants layers were positively correlated with high bird diversity
in urban open areas in the U.S.A (Mason et al., 2007) and Sweden
(Mortberg & Wallentinus, 2000). Tree cover was found to be impor-
tant also for species richness and bird abundance of migrating birds
in Mexico (MacGregor-Fors, Morales-Perez, & Schondube, 2010). In
summary, several factors have been found to affect bird presence
and bird community composition in gardens. The most important
of them are plant composition in gardens, the proportion of tree and
shrub cover, garden size, the degree of urbanization, and predation
risk. However, the effect of plant composition and garden spatial
structure on bird communities might not be the same in different
parts of the world. In Israel, this question has not been yet inves-
tigated in depth, and the only study that refers to this issue found
that high lawn coverage negatively affected bird species richness
188 Y. Paker et al. / Landscape and Urban Planning 122 (2014) 186–195
(Shwartz, Shirley, & Kark, 2008). We hypothesized that high urban
garden variability (i.e. spatial structure of vegetation within the
garden) positively affects bird species diversity, while uniform gar-
dens support low bird diversity. Thus, the aims of this study were:
(1) to explore the influence that plant species richness and gar-
den spatial structure might have on bird species richness, diversity
and community structure in urban gardens in Israel; (2) to iden-
tify plant composition and garden spatial structure that will attract
many native bird species; (3) to test whether migrating birds can
use urban green spaces in Israel as their wintering grounds.
2. Methods
2.1. Study area
We investigated bird communities in 25 public gardens in the
cities of Tel-Aviv and nearby Holon, Israel (32◦ 02 N, 34◦ 47 E; high-
est point above sea level is 62 m). Tel Aviv and the cities around it
create the biggest metropolis in Israel, which lies next to the eastern
Mediterranean Sea on a low plateau. We selected gardens of about
similar size (0.96–2.44 ha) but different in plant composition. This
range of medium-sized gardens was chosen because larger parks
are relatively few in the city and smaller gardens are limited a pri-
ori in their plant diversity. The gardens differ in vegetation: for
example, some were characterized by monoculture trees, others
contained a relatively large lawn, while the rest had diverse shrub
areas. Gardens were scattered around the cities, including north-
ern and southern neighborhoods (Fig. 1). The minimal distance
between gardens was 200 m.
Fig. 1. Layout of the 25 public gardens which were included in the study, within
the cities of Tel-Aviv (its city center is indicated) and nearby Holon (southern to
Tel-Aviv), Israel. Map data: Google earth; Data SIO, NOAA, U.S. Navy, NGA, GEBCO;
image@2013 DigitalGlobe.
2.2. Bird surveys
Following Bibby, Hill, and Mustoe (2000) we used fixed-radius
point counts, a useful method when the habitat around the point
counts is important for the study. In each garden we selected two
stationary observation points, except one garden that had only one
suitable point, and therefore total number of points was 49. The
distance between the two points varied (30–100 m), depending on
the structure of the garden, to ensure that the observed areas will
not overlap. Birds were counted around each point in half circle
plots with a radius of 30 m. In each point count, at the begin-
ning of each observation, the observer sat down quietly for 5 min,
and then registered the birds for additional 10 min. Birds that flew
over the garden were not included, except for species that forage
while flying (i.e. Falco tinnunculus; see Daniels & Kirkpatrick, 2006).
Each point count was visited a total of 21–25 times: twice monthly
visits during autumn and spring migrations (August–October and
March–May, respectively), plus a monthly visit during the rest of
the year. There was a single exception from this pattern, in one
point count which totalled in only 13 monthly visits due to techni-
cal problems. All observations were made by a single observer (YP),
during 3 h from sunrise, between January 2008 and May 2009. We
also noted birds utilizing plant species for food and recorded forag-
ing activities whenever we observed nectar feeding from flowers,
fruit feeding (whole or parts), and gleaning of insects from foliage
and bark.
We recorded four types of bird species: residents, migrants,
aliens, and urban exploiters. Residents are species that stay year-
round in Israel, while migrants arrive to Israel in autumn or spring,
and either pass through on their way to their final destination, or
stay for the winter or summer. Aliens are species that are not part of
the local avifauna and became established in natural or semi natural
ecosystem or habitat elsewhere. They are agents of change, threat-
ening native biological diversity (Westphal, Browne, MacKinnon,
& Noble, 2008). Urban exploiters are species that are dominant
in highly urbanized surroundings, usually social and sedentary,
often aliens. Their typical characteristic is that they thrive as urban
commensals to the point that they become dependent on urban
resources (reviewed in Kark et al., 2007).
2.3. Vegetation and other environmental variables
Vegetation structure was evaluated around each point count, at
half circular plots, within a radius of 50 m from the observer. Trees
and shrubs species were counted in each plot and we also recorded
whether they were natives or aliens. Height of plants was divided
into three groups: low (under 1.5 m), medium (1.6–5 m), and high
(above 5 m). Annual plants were excluded. Trees, shrubs and lawn
coverage were calculated using aerial photos in ArcView 9.3. Cover-
age area of each height group was calculated with ArcView 9.3. We
recorded the existence of water source and irrigation in the gardens.
Number of cats, dogs and people that entered the observed areas
during observations was recorded by using the same protocol as
described above for birds’ counts. We also noted total vegetation
coverage 100 m around each garden by using aerial photographs
and GIS software.
2.4. Data analysis
Bird species diversity was calculated for each garden using
Shannon-Wiener index. Bird species richness was compared
between diverse and uniform gardens by calculating, for each gar-
den type, a rarefaction curve in EstimateS 8.20. The estimator for
species richness was Jacknife 1, which calculates species accumu-
lation (Symonds & Johnson, 2008). Definition of diverse or uniform
gardens was based on the number of trees and shrubs species in
 Y. Paker et al. / Landscape and Urban Planning 122 (2014) 186–195 189

them: uniform gardens contained about 4 species of trees and 4 Table 1

Characteristics of the 25 gardens included in the study.
species of shrubs; diverse gardens contained about 20 species of
trees and 20 species of shrubs. Garden Garden size % Lawn Number of tree Number of
The relations between 14 environmental variables and bird (ha) coverage species shrub species
species richness and diversity were tested using stepwise linear Aliens Natives Aliens Natives
regressions in SPSS. Our study is the first in Israel to look at the
1 0.96 44 14 1 19 1
effect of garden structure and plants’ composition on birds. Hence, 2 1.00 0 7 1 3 3
we preferred not make any a priori predictions (by using a specific 3 1.01 54 11 0 9 1
a priori model which builds upon the literature), because such pre- 4 1.07 55 9 2 13 2
dictions would have been based on studies that had been carried 5 1.14 0 20 2 5 2
6 1.17 5 2 0 5 0
out in other ecosystems, with different climates, flora and fauna 7 1.21 33 11 1 13 1
(e.g. northern Europe, Britain, Australia). Rather, we chose to adopt 8 1.21 0 8 8 12 7
stepwise linear regressions as an initial investigative approach to 9 1.32 55 14 2 2 1
discern and sort the important variables which may affect usage 10 1.32 32 16 2 14 1
11 1.37 34 16 3 22 2
by birds, and exclude those that are not. The 14 variables were
12 1.43 31 8 1 15 2
as follows: garden size; richness of tree species; richness of shrub 13 1.50 58 13 2 24 2
species; coverage of trees; coverage of shrubs; coverage of high 14 1.54 0 7 1 0 0
plants; coverage of medium plants; coverage of low plants; lawn 15 1.55 81 2 1 2 1
coverage; number of cats; number of dogs; number of people; pres- 16 1.57 0 12 18 8 28
17 1.65 45 11 0 20 1
ence of irrigation; and presence of water source. Our hypothesis
18 1.68 0 3 2 0 0
was that the first five variables will positively affect bird species 19 1.72 0 2 19 1 47
richness and diversity. We also expected that the number of cats, 20 1.74 33 16 3 17 2
dogs and people would be negatively related to bird species rich- 21 1.76 0 6 0 0 1
22 2.00 50 9 5 19 3
ness. We hypothesized that the rest of the variables will be less
23 2.02 37 14 4 2 1
important in that respect. Some variables (shrub species richness, 24 2.40 0 4 1 2 0
cat and people presence, coverage of high and medium plants and 25 2.44 1 29 11 11 2
coverage of shrubs) were not normally distributed and therefore we
preformed square root transformations to normalize them. Other
environmental variables (trees, low plants and lawn coverage, tree
species richness, presence of dogs, garden size) were normally dis-
and the rest (60%) were migrants (including wintering birds and
tributed, and binary variables (water source, irrigation) were not
summer breeders). Nine species that are defined as urban exploiters
transformed. The two dependent variables (bird species richness
or alien species (marked in bold in Appendix A) composed 54% of all
and diversity) were normally distributed. The effect of the envi-
individuals seen. Additional 13 bird species that were observed in
ronmental variables on bird community structure was tested using
the gardens incidentally and outside the set observation periods
canonical correspondence analysis (CCA) in CANOCO.
are presented in Appendix B. Sixty-two percent of these latter
In order to compare the attraction of native and alien trees to
13 species belonged to the Passeriformes order; and only one
native or alien bird species, we summed the number of observations
predator species was observed – the resident common buzzard.
of feeding activities (obtaining nectar, eating fruit, foliage and bark
In addition, Appendix B shows that 77% of the species that were
gleaning) that were preformed by alien or native birds on native
observed incidentally and outside the set observation periods were
or aliens trees, during the observations periods. We then used chi-
migrants.
squared tests in order to assess whether native and alien birds select
native or alien trees. In addition, for the most utilized tree species
we divided the number of incidents observed by the number of
point counts that each tree species was present, in order to stan- 3.3. Effect of environmental variables on bird species richness and
dardized our estimate of species use of trees by their proportionate diversity
representation across sites, to enable a comparison between rare
and abundant plant species. Of the 14 environmental variables tested for their effect on
bird species richness and diversity (see Section 2 for details),
only one – shrub species richness – was found to have a signifi-
3. Results cant positive effect on bird species diversity (P = 0.001; a = 0.889,
b = 0.149, R2 = 0.678, df = 24, F = 48.358). None of the 14 environ-
3.1. Gardens mental variables had any effect on bird species richness (Appendix
C).
The gardens in our study ranged in size, from 0.96 to 2.44 ha, We compared the seven most diverse gardens vs. the six most
with a mean of 1.5 ha ± 0.8 (n = 25). Most (70%) of the gardens had uniform ones, by using a rarefaction test that included a total of 59
a lawn, covering 5–88% of their size, with a mean of 25.9% ± 5.0 bird species and 3403 individuals (Fig. 2). Although the two garden
(n = 25). Alien plants dominated the gardens: in most gardens (84%) types show a similar pattern, there is a consisted tendency of higher
we recorded more alien tree species than native ones. Similarly, bird species richness as well as a higher rate of observing new bird
most gardens (76%) had more alien shrub species than native ones. species in diverse gardens, compared with uniform ones. This indi-
Table 1 presents these parameters for each of the 25 gardens. cates that with the same observational effort one might find more
new bird species during revisits to diverse gardens, in comparison
3.2. Observed bird species with uniform ones. In addition, bird species diversity (expressed
by Shannon index) changed seasonally: it was significantly higher
A total of 65 bird species (14,278 individuals) were recorded during spring and fall due to the presence of migrating bird species
from all 25 gardens (Appendix A). Forty percent (82% of total num- (Fig. 3; Kruskal–Wallis: H(3) = 34.14986, N = 1192; p < 0.010, multi-
ber of individuals) of bird species were residents (including aliens), ple comparisons of mean ranks of all groups).
190 Y. Paker et al. / Landscape and Urban Planning 122 (2014) 186–195
Fig. 2. Bird species richness as function of number of observations in diverse gardens
() and in uniform ones ().
3.4. Effect of environmental variables on bird community
structure
We also tested the effect of the 14 environmental variables that
were specified above on bird community structure. The relation-
ship of each of these variables with the two axes of the ordination
is shown in Fig. 4. The CANOCO analysis revealed that the Eigen val-
ues of first (horizontal) axis is 0.402; second (vertical) axis – 0.151.
p = 0.002, F = 7.752; Cumulative percentage variance of species-
environment relation for the two axes = 59.2. It can be seen that
along the 1st (horizontal) axis in Fig. 4, most bird species are located
at the right-hand part of the axis (squares 2 and 3), where shrub and
tree species richness is high, indicating that these variables have a
positive effect on bird community structure; fewer bird species are
located at the left-hand part of this axis (squares 1 and 4), where
presence of people and dogs is high, indicating that these variables
Fig. 3. Seasonal effect on bird species diversity (expressed by Shannon index).
Fig. 4. Effect of environmental variables (long arrows) on bird community structure.
The direction of each arrow (environmental variable) points to an increase of that
particular variable. The longer the arrow, the stronger the effect that a particular
variable had on bird community structure. Bird species that are located near each
other were influenced in a similar manner by the different environmental variables.
For example, Saxicola torquata (Saxi tor) and Sturnus burmannicus (Stur bur) are
located where lawn coverage (p lawn) is high. The ordination is divided into four
squares (1–4), to assist in generally defining each one of them by the variables within
it, and to identify the bird species in it. Environmental variables that are written in
codes are as follows; those with correlation >0.5 with one or two axes are significant
for determining scatter of bird species and are indicated in bold: low = low plant
coverage; s tre – tree species richness; s shr – shrub species richness; med –
medium plant coverage; p shr – shrub coverage; pond – water source; irrigati –
irrigation; p law – lawn coverage; siz – garden size; p tre – tree coverage; high
– high plant coverage. Names and details of bird species that are represented by
triangles and written in codes appear in Appendix D; Due to lack of space, names
of bird species which are included in groups A–C (each of these groups is circled by
a line) appear in Appendix E. Bird species that were seen in less than 5% of count
points were excluded from this analysis.
have a negative effect on bird community structure. Along the 2nd
(vertical) axis in Fig. 4, fewer bird species are located at the upper
part, where lawn coverage is high, or at the lower part, where tree
coverage is high, and from this we infer that these variables nega-
tively affect bird community; most bird species are located around
the middle part, where tree and lawn coverage are low or medium,
indicating that these variables contribute to bird community struc-
ture.
3.5. Tree species that are attractive for birds
In the 25 observed gardens, alien trees compromised 72% of the
total trees counted (1659 out of total of 2306 trees). Respectively,
native trees compromised 28% of the total trees counted (647 trees).
Our data show that native birds (including three native species
that may be categorized as urban exploiters) made a total of 742
visits for feeding activities to trees, out of which 467 visits were
to alien trees. This number of visits to alien trees is less than the
expected one (as calculated by 742 total visits × 0.72 proportion of
alien tress = 534 expected visits). In addition, native birds made 275
visits to native trees, which is higher than expected (as calculated
by 742 total visits × 0.28 proportion of native trees = 208 expected
visits). Analysis of these data revealed that native birds select to
forage on native trees (Chi-square = 29.988, df = 1, p = 0.001). Alien
birds made 65 visits to alien trees for feeding activities, and only
 Y. Paker et al. / Landscape and Urban Planning 122 (2014) 186–195 191

16

14

Number of observations
12

10

Ziziphus spina-christi*

Quercus sober
Acacia raddiana*
Quercus ithaburensis*

Pistacia palaestina*

Quercus calliprinos*

Olea europaea*
Elaeagnus angustifolia*
Pyracantha coccinea

Brachychiton populneus

Ficus rubiginosa
Pinus pinea

Dalbergia sissoo

Ficus religiosa

Acacia saligna
Ficus sycomorus*
Tamarix aphylla*

Rhamnus alaternus*
Erythrina corallodendrum

Acer obtusifolium*
Eucalyptus camaldulensis
Bombax ceiba

Callistemon sp

Ficus microcarpa
Sapium sebiferum

Fig. 5. Number of observations of birds’ feeding activity on different tree species. Native trees are indicated by asterisks. Bird species are divided into natives and urban
exploiters (gray bars) and alien (black bars).

two visits to native ones. Similar analysis of these data revealed of habitat and therefore occupy any available greenspace. On the
that alien birds significantly selected to feed on alien trees than on other hand, we found that bird species diversity is positively related
native ones (Chi-square = 21.231, df = 1, p = 0.001). to shrubs species richness. We believe that this evaluation of birds’
In addition, in order to identify which specific alien and native composition is more reliable than bird species richness, as it takes
tree species were preferred by the birds, we counted the number of into account not only the number of species present but also their
observed feeding activities of all birds (native and alien) on different relative abundance. This latter finding is also in line with others
tree species (Fig. 5). It can be seen that the most frequently utilized who showed that presence of shrubs in urban gardens is important
tree species are a mixture of alien (Bombax ceiba and Callistemon for many forest birds (Bino et al., 2008; Daniels & Kirkpatrick, 2006).
sp) and native trees (Acaia raddiana and Quercus ithaburensis). Shrubs provide birds with partial cover from extreme weather
(heavy rain and scorching sunshine) and predators, as well as food
in the form of insects, seeds and fruits. Therefore, we recommend
4. Discussion
planting multiple shrubs species in order to create diverse gardens
that will have high bird species diversity.
Although urban public gardens are designed primarily for the
In addition, we hope that our study, which is the first in Israel
benefit of humans, their enrichment with bird species will further
to look at the effect of various garden variables on birds, will be
enhance human enjoyment and may also provide opportunity to
followed by future studies in that respect, perhaps also in other
learn about wildlife. In addition, the importance of greenspaces for
parts of the country. Accumulating information, based on a larger
urban birds is more and more understood and has been demon-
and broader range of studies, might enable future analyses to incor-
strated in many recent studies (e.g. MacGregor-Fors et al., 2010;
porate suites of variable in a modeling approach. Moreover, such
Carbo-Ramirez & Zuria, 2011). Urban gardens can provide suitable
future studies will pave the road to move from descriptive studies
habitats for many bird species, other than urban exploiter and alien
that characterize patterns to mechanistic and experimental stud-
species that are very common in cities, as was previously reported
ies of urban ecology. This direction had already been suggested by
(e.g. McKinney & Lockwood, 1999) and similarly found in our study,
Schochat, Warren, Faeth, McIntyre, and Hope (2006), who argued
where more than half of all birds counted were urban exploiters and
that understanding the relevant ecological processes operating in
alien species. Therefore, by understanding the effects that urban
cities is essential to the practice of conservation in an urbanizing
garden characteristics have on bird species richness, diversity and
world.
community structure, landscape designers can take this informa-
tion into account in order to design gardens that will attract various
bird species. In this respect, our study indicates a few points to
4.2. Spatial garden structure
consider.
Our results show that garden composition is important in shap-
4.1. Bird species richness and diversity ing bird community structure in urban areas. The majority of bird
species were found in gardens in which trees and shrubs species
It is known that species richness increases with the diversity of richness was high, and lawn and tree coverage was low or medium
habitats (Hortal, Triantis, Meiri, Thebault, & Sphenthourakis, 2009) (Fig. 4). Along the same line, high coverage of lawn or trees was
and urban birds are no exception (Evans, Newson, & Gaston, 2009; attractive only to a few bird species, some of which were urban
Pino, RodaÁ, Ribas, & Pons, 2000). However, we found that bird exploiters and alien species. In accordance with our finding, tree
species richness was not affected by any of the 14 environmental coverage was not correlated with bird species richness also in
variables that we tested. This outcome might indicate that in the another Israeli study, carried out in a different urban area (Bino
Israeli urban environment birds can be less selective in their choice et al., 2008). Unlike these results from Israel, MacGregor-Fors et al.
192 Y. Paker et al. / Landscape and Urban Planning 122 (2014) 186–195
(2010) report that in Mexico tree coverage had a positive rela-
tionship with species richness of migrating birds. Therefore, we
recommend avoiding large open lawns without trees or shrubs,
as well as dense monoculture woods, because these components
support only few bird species, and moreover – mostly aliens, like
the Common Myna (Acridotheres tristis), or urban exploiters, like
the Feral Pigeon (Columba livia). Instead, our recommendation is
to plant many trees and shrub species, and to balance between
open lawn space and close shrubberies or small thickets. As evi-
dent from Fig. 2, such diverse gardens support more bird species,
and therefore can contribute to the enjoyment that people can get
from visiting gardens in their neighborhood.
Enjoyment to people can be achieved also by the seasonal
change in bird species diversity that is caused by migrating sea-
sons (Fig. 3), enabling people to observe different species at various
times of the year. Moreover, the distinct seasonal pattern which
is reflected in Fig. 3 indicates the importance of urban gardens to
migrating birds. This conclusion is further supported by the fact
that 45% of the birds species recorded in the studied gardens were
migrants (see Appendix A), as well as 77% of the species that were
additionally observed (Appendix B).
4.3. Native vs. alien tree species
Although Daniels and Kirkpatrick (2006) found that it is pos-
sible to have a close to full assemblage of small native woodland
birds in a garden comprised totally of introduced plant species, it
is often reported that native plant species have a positive effect on
native bird species (Chace & Walsh, 2006; Day, 1995). Our finding
is in line with the latter studies and shows that native bird species
preferred native trees for foraging. An opposite trend was found
for alien birds, which significantly selected to feed on alien trees
than on native ones. This differential selection can be explained by
the fact that most of the observed native birds are relatively small
species that collect insects (aphids and others) from foliage and
tree branches (e.g. Sylvia and Phylloscopus species), or nectar from
flowers (e.g. Nectarinia osea), or consume small fruits (e.g. Pycnono-
tus xanthopygos). These food types are abundant on native trees,
which are mostly of Mediterranean origin and therefore smaller
and more bush-like than the alien trees, providing also shelter that
the small birds need. On the other hand, alien birds consisted of
only a few species (mostly Myna and two Psittacula species), which
foraged on trees that provided them big flowers and much nectar
(e.g. Bombax ceiba, Callistemon sp., Erythrina corallodendrum; Fig. 5).
These alien trees are also usually taller than the native ones and
therefore are suitable for parakeets that spend much time in high
places.
One aim of garden designers should be to construct gardens that
will attract native birds, but not alien ones. This aim is important
for both educational (enhancing knowledge of local avifauna by
the urban population) and conservation (supporting local avifauna
while discouraging invading species) reasons. In the light of this,
our finding that native birds are attracted to native trees while
alien ones are attracted to alien trees is of considerable impor-
tance to garden planners. It indicates that garden planners should
prefer local trees over aliens in spite of the often more impressive
appearance (greater height and extensive foliage) of the latter. We
emphasize this recommendation because our data show that cur-
rently, the situation in the Israeli city of Tel-Aviv is the opposite –
alien plant species dominated the studied gardens.
4.4. Presence of people, dogs and cats
Previously, it has been shown that presence of people has a nega-
tive effect on birds (Boyle & Sampson, 1985; Evans et al., 2009). Fig. 4
shows a similar result, and in addition, it indicates that fewer bird
species were found in gardens where presence of dogs was high.
It seems unlikely to assume that this outcome is because dogs and
people tend to go to gardens that birds do not like (i.e. those with
more open space and lawns), as we found no significant correlation
between presence of dogs and lawn coverage. Rather, this probably
indicates a negative effect of people and dogs on bird community
structure. To minimize this effect we recommend creating areas in
gardens that will be inaccessible to people and dogs. This can be
done by planning trails in such a way that relatively large areas of
shrubbery will be far from them, and also by keeping dogs on leash,
as required by law in Israel.
Although we did not find a significant negative effect of cats’
presence on bird species richness, a recent study suggests that
cats are likely the single greatest source of mortality for birds and
mammals, estimating that in the US cats kill 1.4–3.7 billion birds
annually (Loss, Will, & Marra, 2013). Similar concern rises an Israeli
report (Brickner, 2003). Hence, we predicted that cats’ presence
would be negatively related to bird species richness. Our analysis
showed that the number of all the observed bird species but three
(the hooded crow Corvus corone, laughing dove Streptopelia sene-
galensis and the white wagtail Motacilla alba) declined in relation
to the number of cats present, but this relationship was not sig-
nificant. We attribute this result to the large proportion (14%) of
birds belonging to the three species that were insensitive to cats’
presence. Urban density of cats in Israel is very high. For example, a
study in the city of Jerusalem estimated a density of 2300 adult stray
cats per km2 (Mirmovitch, 1995) in comparison to 229 cat/km2 in
Bristol, UK (Baker, Bentley, Ansell, & Harris, 2005; see also Natoli,
Ferrari, Bolletti, & Pontier, 1999). There are several reasons for this.
First, in Israel organic waste is often not properly treated and food
remains are found near garbage containers, in yards and parks. Sec-
ond, feeding stray cats is a widespread phenomenon in Israeli cities
and villages, and people that provide food for dozens of cats are
rather common. Third, Israel has a mild climate and regularly-fed
cats easily survive the winter (Brickner-Braun, Geffen, & Yom-Tov,
2007). Therefore, in order to minimize cats’ numbers and avoid
future threats on bird community in urban gardens, we recom-
mend excluding potential food and water sources accessible to cats
in gardens, and prohibiting feeding cats in gardens.
4.5. Implications for urban landscape design
It has been previously shown that local factors (e.g. housing
density, structural complexity of vegetation) are more important
than regional ones (e.g. latitude, climate) in determining the species
richness of urban avian assemblages, indicating the importance of
management of urban sites for conservation (for example, reviewed
by Evans et al., 2009). Therefore, we would like to emphasize
the need for interdisciplinary studies conducted in collaboration
between landscape designers and zoologists. Such studies can yield
a better understanding of the required garden structure and com-
position for enhancing the number of animal species living in it.
For example, in the present study, zoological knowledge (catego-
rizations of birds and plants to native and alien ones) enabled us to
understand the attraction of native birds to native plants and alien
birds to alien trees. Such understanding can be passed on to collab-
orating landscape designers, who can then apply and implement
this knowledge while planning urban gardens.
In addition to the importance of natural areas within cities for
biodiversity conservation (Fuller, Tratalos, & Gaston, 2009), there
has been a growing recent awareness as to their effect on the well-
being of the human population (Berman, Jonides, & Kaplan, 2008;
Breuste, 2004; Bryant, 2006; Pretty et al., 2007). For example, Fuller,
Irvine, Devine-Wright, Warren, and Gaston (2007) found that sev-
eral psychological benefits increase with species richness of urban
green spaces. Moreover, that study showed that visitors to urban
 Y. Paker et al. / Landscape and Urban Planning 122 (2014) 186–195 193

green spaces can perceive differences in the species richness of Table A.1
Bird species, incidence and total individuals observed at the 25 gardens during the
plants, and to a lesser extent of birds. These findings indicate the
study period (urban exploiters and alien species are indicated in bold).
potentially important role that urban gardens might play in pro-
viding people who live in cities with some experience of nature. Species Status Incidence (% Total
(R = resident; gardens out individuals
Therefore, we believe that studies such as the current one will
M = migrant) of total) observed
provide important tools and data to be applied and used for decision
making in planning urban landscape. The role of landscape design- White-spectacled Bulbul R 100 1535
(Pycnonotus
ers is to shape urban sphere in a way that supports the needs of
xanthopygos)
both human beings and other species. Hence, the importance of Hooded Crow (Corvus R 100 688
this work is in its multidisciplinary perspective. corone)
Great Tit (Parus major) R 100 527
House Sparrow (Passer R 96 2543
5. Conclusions domesticus)
Warbler (Phylloscopus sp.) M 100 459
Our results indicate that a diverse garden, composed of many Palestine Sunbird R 96 517
(Nectarinia osea)
shrub and tree species and a diverse spatial structure is attractive
Graceful Prinia (Prinia R 92 253
to many bird species. Accordingly, we recommend that when plan- gracilis)
ning new gardens, designers will avoid large open lawns and dense Lesser Whitethroat (Sylvia M 92 238
monoculture woods without undergrowth, because these environ- curruca)
ments are favorable only for some urban exploiters and alien bird Laughing Dove R 88 1012
(Streptopelia
species. Instead, we recommend that an urban garden will con- senegalensis)
sist of diverse shrubs and trees species. We found that native bird Eurasian Blackbird (Turdus R 92 405
species select native trees for foraging, while alien birds select merula)
to feed on alien trees. Therefore, garden designers should prefer Eurasian Jay (Garrulus R 92 177
glandarius)
native plants, which in Israel are relatively small and bush-like,
Blackcap (Sylvia atricapilla) M 88 277
and thus provide food and shelter for native birds throughout the White Wagtail (Motacilla M 80 262
year. On the other hand, we recommend reducing the proportion alba)
of alien trees, because they provide high habitats and food (big Common Myna R 84 631
flowers with much nectar) that the larger alien birds need. Our (Acridotheres tristis)
Eurasian Hoopoe (Upupa R 84 125
results also indicate that urban gardens are important to migrating epops)
birds. In addition, since we found a negative effect of people and Barn Swallow (Hirundo M 80 178
dogs on bird community structure, we recommend creating areas rustica)
of thickets and shrubberies that will have limited access to people Chaffinch (Fringilla coelebs) M 80 326
European Greenfinch R 72 202
and dogs. Finally, we would like to emphasize the need for inter-
(Carduelis chloris)
disciplinary studies conducted in collaboration between landscape European Robin (Erithacus M 72 86
designers and zoologists, in order to achieve a better understanding rubecula)
of the required garden structure and composition for enhancing the European Turtle-Dove M 80 85
enjoyment of people as well as the number of bird species living in (Streptopelia turtur)
Olivaceous Warbler M 80 76
it.
(Hippolais pallida)
Syrian Woodpecker R 72 67
Acknowledgements (Dendrocopos syriacus)
Common Swift (Apus M 68 191
apus)
We are very grateful to Ido Itzhaki for his continuous help and Eurasian Collared-Dove R 68 147
important advice on various aspects of the study. We would also (Streptopelia decaocto)
like to thank Ilana Gelenter for her help in the statistical analysis, Masked Shrike (Lanius M 72 41
nubicus)
Dan Eisikowitch for identifying plant species, Shay Barkan for his
Feral Pigeon (Columba R 60 2488
help with the figures, and the reviewers, whose comments greatly livia)
improved and crystallized our manuscript. The Research Fund of Spotted Flycatcher M 72 41
the Open University of Israel supported this study. (Muscicapa striata)
Rose-ringed Parakeet R 52 129
(Psittacula krameri)
Appendix A. Red-backed Shrike (Lanius M 56 35
collurio)
Sardinian Warbler (Sylvia R 44 74
See Table A.1.
melanocephala)
Black Redstart (Phoenicurus M 48 31
Appendix B. ochruros)
White-throated Kingfisher R 40 20
(Halcyon smyrnensis)
See Table B.1. Vinous-breasted Starling R 28 92
(Sturnus burmannicus)
Common Nightingale M 40 14
Appendix C.
(Luscinia megarhynchos)
Common Whitethroat M 36 26
See Table C.1. (Sylvia communis)
Collared Flycatcher M 36 17
(Ficedula albicollis)
Appendix D. European Goldfinch R 28 39
(Carduelis carduelis)
See Table D.1.
194 Y. Paker et al. / Landscape and Urban Planning 122 (2014) 186–195

Table A.1 (Continued ) Table C.1

Statistical results for testing the effect of 14 environmental variables on bird species
Species Status Incidence (% Total
richness. N = 25 for all variables.
(R = resident; gardens out individuals
M = migrant) of total) observed Variable Correlation P value
Thrush Nightingale (Luscinia M 32 14 Garden size 0.015 0.944
luscinia) Richness of tree species 0.157 0.453
Common Redstart M 28 13 Richness of shrub species 0.331 0.106
(Phoenicurus phoenicurus) Coverage of trees −0.213 0.307
Common Kestrel (Falco R 28 11 Coverage of shrubs 0.243 0.242
tinnunculus) Coverage of high plants −0.149 0.478
Eurasian Sparrowhawk M 24 10 Coverage of medium plants 0.264 0.203
(Accipiter nisus) Coverage of low plants 0.167 0.424
Song Thrush (Turdus M 28 7 Lawn coverage 0.187 0.372
philomelos) Number of cats −0.298 0.148
Spanish Sparrow (Passer M 20 45 Number of dogs −0.232 0.265
hispaniolensis) Number of people −0.222 0.287
Eurasian Wryneck (Jynx M 16 5 Presence of irrigation −0.080 0.709
torquilla) Presence of water source −0.100 0.641
Rueppell’s Warbler (Sylvia M 20 5
rueppelli)
European Bee-eater (Merops M 16 32 Table D.1
apiaster) Codes and names of bird species that are shown in Fig. 4.
Spur-winged Lapwing R 12 15
(Vanellus spinosus) Latin name Code Latin name Code
Monk parakeet (Myiopsitta R 12 14
Accipiter nisus Acci nisu Passer domesticus Pass dome
monachus)
Acridotheres tristis Acri tris Passer hispaniolensis Pass hisp
Orphean Warbler (Sylvia M 8 4
Carduelis chloris Card chlo Phoenicurus phoenicurus Phoe phoe
hortensis)
Carduelis spinus Card spin Prinia gracilis Prin grac
Eurasian Siskin (Carduelis M 12 22
Columba livia Colu livi Saxicola torquata Saxi torq
spinus)
Corvus corone Corv coro Streptopelia decaocto Stre deca
Stonechat (Saxicola torquata) M 8 7
Falco tinnunculus Falc tinn Streptopelia turtur Stre turt
Garden Warbler (Sylvia borin) M 8 4
Garrulus glandarius Garr glan Sturnus burmannicus Stur burm
Gray Wagtail (Motacilla M 8 3
Jynx torquilla Jynx torq Sylvia hortensis Sylv hort
cinerea)
Lanius collurio Lani collu Sylvia rueppelli Sylv ruep
Chukar (Alectoris chukar) R 4 2
Merops apiaster Mero apia Turdus merula Turd meru
Black-eared Wheatear M 4 2
Motacilla alba Mota alba Upupa epops Upup epop
(Oenanthe hispanica)
Motacilla cinerea Mota cine Vanellus spinosus Vane spin
Black-crowned Night-Heron R 4 1
(Nycticorax nycticorax)
Eurasian Hobby (Falco M 4 1
subbuteo) Appendix E.
Cretzschmar’s Bunting M 4 1
(Emberiza caesia)
Whinchat (Saxicola rubetra) M 4 1 See Table E.1.
Eurasian Golden-Oriole M 4 1
(Oriolus oriolus) Table E.1
European Pied Flycatcher M 4 1 Latin names of bird species in groups A–C that are shown in Fig. 4.
(Ficedula hypoleuca)
Bluethroat (Luscinia svecica) M 4 1 A B C
Eurasian Thick-knee R 4 1 Apus apus Erithacus rubecula Carduelis carduelis
(Burhinus oedicnemus) Halcyon smyrnensis Hippolais pallida Dendrocopos syriacus
Wood Warbler (Phylloscopus M 4 1 Hirundo rustica Muscicapa striata Ficedula albicollis
sibilatrix) Lanius nubicus Nectarinia osea Fringilla coelebs
Common Cuckoo (Cuculus M 4 1 Luscinia luscinia Phoenicurus ochruros Myiopsitta monachus
canorus) Luscinia megarhynchos Phylloscopus sp. Parus major
Streptopelia senegalensis Psittacula krameri Sylvia borin
Turdus philomelos Pycnonotus xanthopygos
Sylvia atricapilla
Table B.1 Sylvia communis
Bird species that were observed in gardens incidentally and outside the set obser- Sylvia curruca
vation periods. Sylvia melanocephala

Species Status (R = resident;

M = migrant)

Barred Warbler (Sylvia nisoria) M

Cattle Egret (Bubulcus ibis)
Common Buzzard (Buteo buteo)
Common Kingfisher (Alcedo atthis)
Common starling (Strunus vulgaris)
Common Quail (Conturnix conturnix)
Fieldfare (Turdus pilaris)
Great Spotted Cuckoo (Clamator glandarius)
Lesser Gray Shrike (Lanius minor)
Olive-tree Warbler (Hippolais olivetorum)
Sedge Warbler (Acrocephalus schoenobaenus)
Tree Pipit (Anthus trivialis)
Meadow Pipit (Anthus pratensis)
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