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Journal of Applied
Ecology 2001
38, 1122 1134

Are urban bird communities influenced by the bird


diversity of adjacent landscapes?

Blackwell Science Ltd

PHILIPPE CLERGEAU*, JUKKA JOKIMKI and


JEAN-PIERRE L. SAVARD
*INRA SCRIBE and UMR EcoBio, avenue Gnral Leclerc, 35 042 Rennes, Cedex, France; Arctic Centre,
University of Lapland, PO Box 122, 96101 Rovaniemi, Finland; and Canadian Wildlife Service, 1141 Rte de lglise,
PO Box 10100, Sainte-Foy, Qubec GIV4H5, Canada

Summary
1. The species diversity of adjacent landscapes influences the conservation or restoration of several animal groups in urban areas, but the effect on birds is unclear. To address
this question, we compared bird species richness (BSR) and community composition
between periurban (area surrounding the town) and urban (suburban and centre areas)
landscapes across three spatial scales.
2. At a large biogeographical scale (temperate and boreal climatic zone), relationships
between the BSR of urban areas and their surrounding landscapes were examined in a
meta-analysis of 18 published studies. In general, BSR was negatively correlated with
latitude and urbanization. The BSR of suburban and centre landscapes correlated
positively with the BSR of periurban landscapes. However, latitudinal effects were also
involved, as BSR in urban and periurban landscapes declined as town latitude increased.
Similarity indices were low (50%) between periurban and centre bird communities.
3. At a regional scale, we assessed winter bird data from several towns within three
regions of temperate and boreal countries (western France, northern Finland and eastern
Canada). The type of periurban landscape, number of inhabitants and town diameter did
not affect BSR. BSR was similar between the cities of a given biogeographical area. Bird
communities were more similar between similar habitat types of different cities than
between different habitats of the same city.
4. At a local scale, we tested the influence of proximity to the periurban landscape on
BSR in parks of western French towns of different size. Neither BSR nor community
similarity changed in relation to the distance of the park from the periurban landscape.
5. Guild composition according to diet and feeding habitat did not vary between urban
and periurban locations at regional or local scales.
6. We conclude that, at regional and local scales, urban bird communities are independent of the bird diversity of adjacent landscapes, and that local features are more
important than surrounding landscapes in determining BSR. Whatever the biodiversity
quality of the periurban landscape, site-specific actions such as shrub and tree planting,
water restoration and increasing vegetation diversity can change bird diversity in towns
and improve the quality of humanwildlife contacts.
Key-words: biodiversity, biogeography, bird species richness, landscape, urban planning,
urban system.
Journal of Applied Ecology (2001) 38, 11221134
Introduction
Processes associated with urbanization are one of the
major causes of landscape change and represent an

2001 British
Ecological Society

Correspondence: Philippe Clergeau, INRA SCRIBE and UMR


EcoBio, avenue Gnral Leclerc, 35 042 Rennes, Cedex, France
(e-mail clergeau@univ-rennes1.fr).

important threat to biodiversity (Wilcox & Murphy


1985). Urban planners need better information about
the factors affecting the distribution of species and
structure of communities in order to create or maintain
biodiversity in urban areas. Conservation or restoration efforts related to urban wildlife focus on limiting
artificial habitat, developing citizen participation in
wildlife conservation, improving the quality of life of

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1123
Urban bird
diversity and
landscape context

2001 British
Ecological Society,
Journal of Applied
Ecology, 38,
1122 1134

urban dwellers, and educating them about ecological


concepts (Gilbert 1989; Adams 1994; Niemel 1999).
The interest of urban residents for their immediate
environment is becoming increasingly important.
Additionally, their decisions on ecological issues at
regional or national levels are based largely on their
perception of nature at local levels (Michelson 1970;
Middleton 1994). Therefore, urban wildlife diversity
can influence significantly the management of biodiversity at regional, national and even global scales
(Owen 1978; Hadidian et al. 1997).
In general, human activities have produced similar
ecological structures in urban areas even in different
biogeographical regions. The response of birds to these
environmental changes could lead to the dominance
of bird communities by a few very abundant species
(Bezzel 1985). This, in turn, might lead to the general
hypothesis that urbanization causes uniform bird
communities in urban areas (Jokimki et al. 1996). A
town can be an original ecosystem with its own characteristics and species (Davis & Glick 1978; Bezzel
1985).
A few ecological studies have been conducted on the
relationships between urban (artificial built-up area)
and periurban (area adjacent to the town) landscapes
and on the conservation of biodiversity in urban ecosystems. Data on dispersion, colonization and settlement of several wildlife groups such as mammals
(Harris 1977; Dickman & Doncaster 1989), insects
(Owen 1978) and lizards (Germaine 1995) have clearly
underlined the effect of proximity to non-urban source
populations in urban biological conservation. Could
this relationship hold also for birds in urban landscapes, i.e. could periurban diversity influence bird species richness (BSR) in towns?
The composition of urban flora and fauna may be
determined by the biotic and abiotic factors associated
with the available species pool in the region (BhningGaese 1997; Roy, Hill & Rothery 1999). Surrounding
habitat features may affect bird community structures
(Andrn 1994; Kubes & Fuchs 1998; Jokimki 1999),
and bird diversity in urban areas has been linked to
periurban landscapes (Siegfried 1968; Jones 1983;
Munyenyembe, Harris & Hone 1989). However, Erz
(1966) suggested that bird species do not colonize
newly urbanized areas from surrounding countryside,
but immigrate from already urbanized populations.
Thompson, Greenwood & Greenway (1993) compared
garden birds between several European countries and
found that BSR in urban, suburban and rural gardens
was similar in countries of northern Europe, but not in
western or southern Europe.
These opposite results might depend on the geographical location or size of the town, but also on the
spatial scale used in these studies. The landscape and
community structures are scale-dependent and different factors are involved at each ecological organization
level (Forman & Godron 1986; ONeill 1989; BhningGaese 1997; Baillie et al. 2000).

Our main goal in this study was to evaluate the


relative importance of local and regional landscapes
in shaping the structure of urban bird communities.
An effect of periurban diversity on urban BSR would
stress the need to incorporate landscape consideration
in managing communities of urban birds, whereas a
lack of strong relationships would emphasize local
actions. We used a multi-scale approach to investigate
the relationship on bird diversity between urban and
periurban landscapes. At a large biogeographical scale,
we compared bird community indices of bird studies
from towns in temperate and boreal climatic zones.
At a regional scale, we assessed the level of variability
in the avifauna of several cities within similar geographical areas. At a local scale, we compared the
avifauna of different habitats within a single town.
As life histories of species also constrain the spatial
scales at which resources can be exploited (Hansen &
Urban 1992), we analysed our data in more detail by
using groups of species (guilds) in our regional and
local scale analyses.

Methods

To look at relationships between urban areas and their
surrounding landscape, we reviewed studies where breeding BSR had been estimated similarly in both urban
and periurban landscapes. We excluded some studies
where BSR was estimated by numerous observers
(Luniak 1990; Konstantinov 1996). We limited our
sample to studies conducted in temperate and boreal
regions. From about 50 papers on urban birds, we
found 18 studies that fulfilled our pre-selection criteria.
In all these studies, description of study sites allowed
the coherent definition of three landscape types representative of typical forms of urban development. We
characterized the urban area by two landscape types
that we called centre and suburb for simplification.
(i) The centre includes both the commercial downtown
area, i.e. historic centre of the city with less than 15% of
vegetated area and buildings with three to five stories,
and the residential sector adjacent to the commercial
area, where buildings have in general one to three stories and vegetated open areas cover between 20% and
40% of the area. (ii) The suburb consists of residential
areas with single-family detached houses, large apartment building complexes, parks and cemeteries, and of
large office or supermarket sites with lawns and parking areas. The height and disposition of houses and
buildings vary considerably and vegetated areas cover
up to 70% of this sector. (iii) The third landscape, the
periurban sector, includes the various landscapes adjacent to the town, up to 10 km from the city centre for
large cities. It mainly comprises managed rural features
such as crops and cattle farming, but also includes
some leisure sites such as golf courses and parks, and
natural sites such as woodlots and lakes. In some

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P. Clergeau,
J. Jokimki &
J.-P.L . Savard

cases, towns are surrounded mainly by natural landscapes like forest or heath.
For all selected studies (Table 1), we classified surveyed sites into these three landscape types. For periurban landscapes, we extracted separately, based on the
previous remarks, data for managed (dominated by
agriculture or leisure sites) and natural (dominated by
forest or heath) landscapes when possible. In some
studies, we selected only one year of results because
other years did not involve the same study sites. Two
main types of census were represented: absolute
methods (spot-mapping or strip census), used in the
majority of publications, and relative methods (fixed
point counts on transect lines or not), used in only
three papers.

2001 British
Ecological Society,
Journal of Applied
Ecology, 38,
11221134

We analysed winter bird data from the towns of three


regions: (i) western France (between 47 and 49 north),
with a temperate climate and numerous cities surrounded by agricultural landscapes (P. Clergeau & G.
Mennechez, unpublished data); (ii) northern Finland,
which belongs to the northern coniferous forest biome
(between 65 and 67 north), with a cold climate and
small cities surrounded mainly by forests (J. Jokimki
& M. Kaisanlahti, unpublished data); and (iii) eastern
Canada (between 45 and 47 north), with a cold
climate and big cities surrounded by a large river and
forests (Clergeau et al. 1998). We compared BSR and
similarity indices (see below) between different types
of (i) suburban development in Europe and (ii) urban
parks in Canada. Because seasonal differences may
modify community composition and structure, our
regional scale results from the winter period may not be
directly comparable with large and local biogeographical scale results collected during the breeding period.
We compiled data in winter (January and February
1999) on birds from five cities in a small area
(40 000 km2) of the eastern part of Brittany (western
France) and Lapland (northern Finland). These towns
differed in size and types of periurban landscapes
(Table 2). In each city, we identified two traditional
suburban landscapes typical in their structure: blocks
of large apartment buildings and areas of single-family
detached houses. All sites were approximately 30 ha in
size and were surveyed using a single visit, i.e. a zigzag
walk through the site taking 60 min (Jokimki et al.
1996). The mapping method of Svensson (1974) used in
these studies reduced many problems associated with
counting birds in urban areas (DeGraaf, Geis & Healy
1991). The validity of this quick technique has been
confirmed (Jokimki & Suhonen 1998). All birds were
counted, except individuals flying over that did not stay
within the study site.
In addition, we compiled data on wintering birds
from the parks of four cities in the St Lawrence Valley,
southern Quebec (eastern Canada) (Table 2). We
selected data from two types of parks according to their

structure (Morneau et al. 1996): managed parks with


some trees or wooded portions (< 20%) and large lawns
(more than 50%), and more natural parks with forest or
wetland biotopes and with lawn area < 50%. To avoid
size effects (Opdam, Rijsdijk & Hustings 1985), as
parks varied from 03 ha to 117 ha (Morneau et al. 1996),
we selected only parks between 4 and 10 ha in size.


We tested the neighbourhood effect of periurban
landscapes using a similar habitat type located along a
distance gradient from periurban areas. In three cities
of different sizes in western France, Angers, Rennes
and Nantes (Table 2), we chose one park in the town
centre, one in the suburbs and one at the edge of town:
the distance between town centre parks and potential
species sources (periurban area) increased with the size
of the town. We selected 410-ha parks to avoid size
effects; in the large parks of town edges, we chose only
an isolated part of about 10 ha. All selected sites presented similar structures, with old trees, lawns and
ponds (Clergeau 2000). BSR was determined during
the breeding season by the point-count method (Bibby,
Burgess & Hill 1992). Two survey stations were located
in each park, and all birds seen or heard were recorded
during a 20-min count between 07:00 hours and
09:00 hours. Each station was surveyed twice per
breeding season (May and June 1998 and 1999). A
more detailed description of the study sites is available
from Clergeau (2000).


We used the cumulative number of bird species
obtained in each landscape type (centre, suburban and
periurban areas) as well as the mean number of species
per landscape type. The two methods gave similar
results, so we present here only the mean number of
species. In cases where the size of study plots differed,
we used the rarefaction method to estimate species
richness (Heck, van Belle & Simberloff 1975). Biogeographical data were first analysed by the general linear
model (GLM) univariate procedure (SPPS 1999). BSR
was the dependent variable and fixed factors included
the number of inhabitants (three groups: > 600 000,
n = 16; < 600 00060 000, n = 19; < 60 000, n = 15), latitude (two groups: > 50N, n = 19; others, n = 31), urbanization level (three groups: centre, n = 14; suburban,
n = 18; periurban, n = 18) and location in the old or
new world (two groups). The first model was constructed by using main effects of all independent variables and their interactions. After the second phase,
only significant variables (P < 005) were entered in the
model.
We used the Srensen similarity index (Jongman, ter
Braak & Tongeren 1995) to measure the similarity of
two assemblages. It is based on species numbers and
does not take species abundance into account:

City

Number of species (mean)

Authors

City (country)

Size
agglomerate
1000 inhab.

Weber (1972)

Vancouver (Canada)

1380

49 NW

Geis (1974)

Columbia (USA)

395

34 NW

Campbell & Dagg (1976)

Waterloo (USA)

80

43 NW

Marchetti (1976)

Marseille (France)

900

43 OW

Huhtalo & Jarvinen (1977)

Tornio (Finland)

20

66 OW

Savard (1978)

Toronto (Canada)

3425

44 NW

Hohtola (1978)

Kuopio (Finland)

80

63 OW

Lancaster & Rees (1979)

Vancouver (Canada)

1380

49 NW

Tatibouet (1981)

Lyon (France)

455

46 OW

Latitude
world part

Method
census /site
sample size
2 years*
16 census
10 ha
1 year (1972)*
1 census
8 97 ha
1 year*
7 census
25 ha
1 year
1 census
50 points
1 year*
4 census
30 95 ha
1 year*
5 census
10 ha
1 year*
3 census
2 17 ha
1 year*
> 5 census
8 ha
1 year
1 census
7 points

World part = old world, OW; new world, NW.


Study plots were not standardized and we have applied rarefaction diversity method (basis of 30 pairs).
ML, managed landscape; NL, natural landscape.

Similarity index

Centre data
(n sites)

Suburban data
(n sites)

Periurban data
(n sites)

Centre/
suburban

Suburban/
periurban

Centre/
periurban

14 (2)

275 (2)
275 (2) ML

056

111 (5)

161 (4)
161 (4) ML

068

069

058

13 (1)

25 (2)

31 (1)
31 (1) NL

043

071

036

9 (2)

28 (2)

373 (3)
365 (2) ML
39 (1) NL
94 (1)
94 (1) ML

052

078

040

063

063

057

122 (5)
88 (4) ML
261 NL
111 (1)
111 (1)ML

063

075

058

032

058

021

51 (7)

104 (1)

108 (1)

75 (6)

15 (2)

5 (3)

136 (7)

6 (1)

188 (4)

23 (1)
23 (1) NL

15 (2)

215 (2)

31 (3)
305 (2) ML
32 (1) NL

078

085

062

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diversity and
landscape context

2001 British
Ecological Society,
Journal of Applied
Ecology, 38,
1122 1134

Table 1. Selected studies in temperate and boreal regions where bird surveys were conducted in urban and periurban areas with the same method and in the same season. Managed landscape = areas where
agriculture or leisure sites predominate; Natural landscape = areas where forest or heath predominate. *Absolute method (mapping method, strip census); relative method (plot method, fixed point method on
transect line) (reviewed in Bibby, Burgess & Hill 1992)

City

Authors

City (country)

Bessinger & Osborn (1982)

Oxford (USA)

Jokimaki (1989)

Number of species (mean)


Size
agglomerate
1000 inhab.

Latitude
world part

115

52 NW

Rovanieni (Finland)

35

67 OW

Vauhkonen (1990)

Heinola (Finland)

15

67 OW

Sasvari (1990)

Budapest (Hungary)

2100

47 OW

Sodhi (1992)

Saskatoon (Canada)

200

52 NW

Rauhala (1994)

Kemi (Finland)

25

66 OW

Blair (1996)

Palo Alto (USA)

55

37 NW

Clergeau et al. (1998)

Rennes (France)

210

48 OW

Clergeau et al. (1998)

Quebec (Canada)

605

47 NW

Method
census /site
sample size
1 year*
> 10 census
6 ha
1 year*
1 census
30 ha
1 year (1990)*
2 census
50 420 ha
5 years*
4 census
50 ha
3 years*
6 census
9 ha
1 year
1 census
13 73 ha
2 years
12 census
16 points
1 year*
4 census
12 ha
1 year*
4 census
12 ha

World part = old world, OW; new world, NW.


Study plots were not standardized and we have applied rarefaction diversity method (basis of 30 pairs).
ML, managed landscape; NL, natural landscape.

Similarity index

Centre data
(n sites)

Suburban data
(n sites)

Periurban data
(n sites)

Centre/
suburban

Suburban/
periurban

Centre/
periurban

185 (2)

275 (2)
275 (2) NL

056

5 (1)

105 (4)

119 (10)
119 (10) ML

044

064

024

99 (1)

122 (1)

164 (1)
164 (1) NL

067

083

058

24 (3)

16 (6)

249 (7)
233 (3) ML
26 (4) NL
157 (6)
157 (6) ML

047

163 (2)
163 (2) ML

065

067

077

237 (3)
25 (2) ML
21 (1) NL
27 (2)
27 (2) ML

030

059

023

082

081

079

25 (2)
25 (2) NL

067

064

057

106 (1)

96 (1)

7 (1)

145 (2)

203 (3)

24 (3)

19 (3)

243 (4)

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P. Clergeau,
J. Jokimki &
J.-P.L . Savard

2001 British
Ecological Society,
Journal of Applied
Ecology, 38,
11221134
Table 1. (contd )

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landscape context

Table 2. Principal characteristics of towns studied in France, Finland and Canada


Size
City

Inhabitants

Periurban landscape
Diameter (km)

Major component

Secondary component

France (Brittany)
Redon
Dinan
Fougres
St Malo
Rennes

11 000
16 000
28 000
46 000
200 000

2
2
3
4
7

Bocage*
Bocage
Open field
Littoral
Open field

Wetland
Woodland
Bocage
Open field
Bocage

France (Other)
Angers
Nantes

140 000
260 000

44
12

Bocage
Open field

Woodland
Bocage

Finland (Lapland)
Kemijrvi
Tornio
Kemi
Rovaniemi
Oulu

10 000
20 000
25 000
35 000
105 000

25
25
5
5
85

Forest
Littoral
Littoral
Forest
Littoral

Lake side
Farming
Farming
River side
Farming

7500
40 000
600 000
1010 000

15
25
15
30

Farming
Farming
Forest
Farming

Forest
Forest
Farming
Forest

Canada (Quebec, all river side)


St Nicolas
Levis
Quebec
Montreal

*Hedgerow network landscape with grasslands and some farming.

where c is the number of species shared by the two


sites, and a and b the total number of species at each
site. Values of this index vary from 0 to 1; 0 indicates
that assemblages differ totally, and 1 that they are
identical.
To analyse potential ecological variations, we compared biological traits of bird species. First, we defined
a diet guild from the main diet of each species: insect
feeder; seed and vegetation feeder; omnivore; carnivore
(Clergeau et al. 1998). Secondly, we defined each species according to its main feeding habitat: aquatic; tree
dwelling; shrubs; meadows; other. Classification of
each species observed in France, Finland and Canada
is presented in the Appendix. Data were analysed by
comparing the number of species at regional and local
scales for each guild and each town using chi-squares.
Non-parametric tests were performed using correction for ties (two-tailed). All mean values are given with
standard errors (95% confidence limits).

Results

2001 British
Ecological Society,
Journal of Applied
Ecology, 38,
1122 1134

According to the GLM univariate model using all independent variables and their interactions, only the level
of urbanization (F = 1485, d.f. = 2, 27, P < 0001) and
latitude (F = 655, d.f. = 1, 27, P = 0016) significantly
affected BSR. None of the interaction terms was significant (P > 005). When we dropped non-significant

variables from the model, both urbanization level


(F = 1348, d.f. = 2, 46, P < 0001) and latitude
(F = 1651, d.f. = 1, 46, P < 0001) were significant,
explaining together 555% of the variation in BSR.
BSR of periurban landscapes was positively correlated with the BSR of suburban (Spearman rs = 075,
n = 18, P = 0002) and town centre landscapes (rs =
058, n = 14, P = 004), but the correlation between
BSR of suburban and centre landscapes was not significant (rs = 046, n = 14, P = 010). Relationships
between suburban and periurban BSR according to
types of periurban landscapes (managed or natural)
corroborated the correlation: significant regression
coefficients and similar slopes were obtained with managed and natural landscapes (Fig. 1). BSR decreased with
latitude both in suburban and periurban landscapes,

BSR in suburban landscape

similarity index = 2c /( a + b )

40

With periurban managed landscape


With periurban natural landscape

30
20
10
0
10

15

20

25

30

35

40

BSR in periurban landscape

Fig. 1. Bird species richness (BSR) in suburban landscape


is correlated with BSR observed in natural (y = 512 +
057x, r = 066, t = 420, P = 0002) or in managed
( y = 553 + 054x, r = 058, t = 372, P = 0004) periurban
landscapes (area adjacent to the town). Data from 18 selected
studies in temperate and boreal regions.

1128
P. Clergeau,
J. Jokimki &
J.-P.L . Savard

BSR in suburban landscape BSR in periurban landscape

JPE_666.fm Page 1128 Thursday, September 20, 2001 2:50 PM

P < 0003). In four cases in northern countries


(Finland and Canada), BSR did not follow this trend,
being greater in suburban than periurban landscapes
(Table 1).
Highest similarity values were observed between
periurban and suburban landscapes (067 021) and
lowest between periurban and centre landscapes
(050 038); similarity values between suburban and
centre landscapes (058 033) were intermediate. Bird
communities were more similar between periurban
and suburban landscapes than between periurban and
centre landscapes (MannWhitney U = 450, n = 26,
P = 001). Similarity between centre and suburban
landscapes did not differ from others (U = 725, n = 26,
P = 017; U = 570, n = 26, P = 016, respectively).
Similarity indices between communities were not influenced by latitude (n = 13, 13, 16, respectively, all rs < 02
and P > 04).

40

r s = 049; P = 004

30
20
10
0
30

35

40

45

50

55

60

65

70

40
30

r s = 056; P = 002

20
10
0
30

35

40

45

50

55

60

65

70

BSR in town centre

40
30

r s = 004; P = 088

20
10
0
30

35

40

45

50

55

60

65

70

Latitude
Fig. 2. Bird species richness (BSR) both in periurban and in
suburban landscapes are correlated with latitude, but BSR in
town centre is independent of latitude (Spearman rs).

but not in town centres (Fig. 2). In northern Europe,


the number of species in each landscape (centre:
82 57; suburban: 113 31; periurban: 130 62)
was half that in southern Europe (148 111, 244
53, 301 107, respectively).
BSR decreased from periurban (215 154) and
suburban landscapes (169 114) to centre landscapes (102 95) (n = 14, 14, 18 pairs, all Z < 292,

In Brittany, similar wintering BSR were found in


blocks of apartment building (1920 380) and singlehouse areas (1880 313; Z = 067, n = 10, P = 050).
BSR did not vary between cities (Table 3) nor was
influenced by the number of inhabitants (below and
above 26 000 inhabitants), town diameter (below and
above 35 km) or the type of periurban landscape
(bocage/others) (Table 2) (MannWhitney, all n = 10,
P > 024).
In Lapland, BSR was lower in areas with blocks
of apartment building (600 328) than in singlehouse areas (980 192; Z = 203, n = 10, P = 004;
Table 3). BSR did not vary between cities, nor was
influenced by the number of inhabitants, town diameter or the type of periurban landscape (forest/
littoral) (MannWhitney, all n = 10, P > 007). In
pooled data, BSR was lower in Lapland than in Brittany (U = 0001, n = 20, P = 0008).
In Quebec, BSR did not differ between parks with
lawn occupying > 50% (550 404) and parks with

Table 3. Number of wintering bird species in suburban landscapes (two housing types in Europe; two park structures in Canada)
of towns varying in terms of size and periurban landscapes (see Table 2). Chi-squares on matrix towns/landscape types per region
did not show significant variation according to towns
Towns

2001 British
Ecological Society,
Journal of Applied
Ecology, 38,
11221134

Chi-square

d.f.

Brittany (western France)


Housing type
Block of apartment buildings
Single-family detached houses

Redon
17
21

Dinan
22
19

Fougres
18
17

St Malo
18
17

Rennes
21
20

070

095

Lapland (northern Finland)


Housing type
Block of apartment buildings
Single-family detached houses

Kemijrvi
4
9

Tornio
7
9

Kemi
7
11

Rovaniemi
4
9

Oulu
8
11

094

092

Quebec (eastern Canada)


Park type
With lawn > 50%
With lawn < 50%

St Nicolas
9
6

Levis
4
8

Quebec
45
5

Montreal
4
10

276

043

JPE_666.fm Page 1129 Thursday, September 20, 2001 2:50 PM

1129
Urban bird
diversity and
landscape context

Table 4. Similarity indices within towns and between towns for single-family detached house/blocks of apartment building areas
(Brittany and Lapland) and between towns for parks more or less managed (Quebec)
Mean SE

Towns
Brittany (western France)
Redon
Dinan
Fougres
St Malo
Rennes

Redon
068
072
074
063
063

Dinan
075
059
070
075
070

Fougres
084
078
080
078
062

St Malo
068
072
077
080
067

Rennes
068
072
081
070
078

Kemijrvi
062
073
073
075
067

Tornio
078
075
086
073
080

Kemi
080
090
078
055
093

Rovaniemi
100
078
080
046
050

Oulu
080
080
091
080
074

St Nicolas
024
050
057
040

Levis
013
040
043
032

Quebec
012
067
057
033

Montreal
025
057
067
050

Single-house: 075 011


Within town: 073 016
Blocks: 069 011

Lapland (northern Finland)


Kemijrvi
Tornio
Kemi
Rovaniemi
Oulu

Single-house: 084 015


Within town: 067 012
Blocks: 072 026

Quebec (eastern Canada)


St Nicolas
Levis
Quebec
Montreal

2001 British
Ecological Society,
Journal of Applied
Ecology, 38,
1122 1134

lawn occupying < 50% (805 199; Z = 109, n = 8,


P = 027) and BSR did not vary between cities
(Table 3). Also in this region, neither the size of town
nor its periurban landscape could explain BSR variations (MannWhitney, all n = 8, P > 022).
In Brittany, bird community similarity between
blocks of apartment building and single-house areas
within the same town was about the same as that
between similar habitats in different towns (Mann
Whitney, n = 15, 15, 25, all P > 027; Table 4). In Lapland, within-town similarity in bird communities was
lower than that between bird communities of singlehouse areas of different towns (U = 0001, n = 15,
P = 0002) and similar to that between areas of blocks
of apartment buildings of different towns (U = 215,
n = 15, P = 067; Table 4). Bird communities of singlehouse landscapes were more similar than those of
blocks (U = 205, n = 20, P = 002). In Quebec, park
similarity indices did not differ between different towns
and between different parks with lawn < 50% or with
lawn > 50% (n = 10, 10, 12, all P > 091; Table 4).
Therefore, in the three regions, similarity indices were
not significantly greater within town than within habitat types. In pairwise comparisons, similarity indices
did not vary with size of towns (number of inhabitants,
town diameter) or types of periurban landscapes
(n total = 16, 25, 25, all P > 008).
The BSR of different diet and feeding habitat guilds
did not vary between different towns in Brittany or
Lapland (chi-square test, d.f. = 8, 12, all P > 079). In
Brittany, the number of species per category did not
vary with location (Wilcoxon test, all n = 10, P > 018)

Park lawn > 50%: 040 051


Within town: 043 029
Park lawn < 50%: 043 019

but, in Lapland, seed and vegetation feeders (species


noted g) were more abundant in single-house areas
(340 214) than in blocks of apartment buildings
(120 255; Z = 202, n = 10, P = 004). In the feeding
habitat guild, tree-dwelling species were more numerous in Lapland single-house areas (520 163) than in
areas of blocks of apartment buildings (240 297;
Z = 202, n = 10, P = 004). No other differences were
observed (P > 005). The Canadian study did not
reveal any variation between sites (Wilcoxon test, diet
guild: all n = 8, P > 007; feeding habitat guild: all
n = 8, P > 008) and corroborated a lack of difference
between towns (chi-square, all d.f. = 6, P > 042).


Using French towns of different sizes, we tested
whether BSR in urban parks was related to the distance
of the park from the periurban landscape, i.e. the distance that birds would have to travel across the urban
matrix. BSR varied between years (more species in
1998, U = 1450, n = 18, P = 002) but BSR of parks
located in the centre landscape (1998: 2633 206;
1999: 2367 299), suburban landscape (1998: 2533
113; 1999: 2133 599) or just at the edge of town
(1998: 2567 147; 1999: 2300 588) did not vary
with the diameter of the town (1998: 2 = 135, d.f. = 4,
P = 086; 1999: 2 = 105, d.f. = 4, P = 090). Thus the
distance from the periurban landscape did not affect
park BSR (Fig. 3).
Similarity indices between the different habitats of a
town did not differ from those between parks in town

JPE_666.fm Page 1130 Thursday, September 20, 2001 2:50 PM

1130
P. Clergeau,
J. Jokimki &
J.-P.L . Savard

Table 5. Similarity indices within French towns and between towns for park in centre/park just in fringe; in first line 1998, in
second line 1999. n km = distance between park in centre and park just in fringe
Mean SE

Angers (22 km)


Rennes (35 km)
Nantes (6 km)

Angers
061
068
069
065
069
074

Rennes
079
069
075
076
063
082

Nantes
071
070
068
078
074
079

Park just in fringe:

073 009
072 007

Within town:

070 013
074 009
067 006
074 013

Park in centre:

Angers
22 km

30
20
10
0

Bird species richness

Centre

Suburban

Town edge

Rennes
35 km

30
20
10
0
Centre

Suburban

Town edge

Nantes
6 km

30
20
10
0
Centre

Suburban

Town edge

Park location
Fig. 3. Bird species richness (breeding species) count in
similar parks (4 10 ha parks with old trees, lawns and ponds)
of three towns of western France (km = distance between the
town edge and the town centre). 1998, grey bars; 1999, dark
bars. All chi-square tests not significant.

2001 British
Ecological Society,
Journal of Applied
Ecology, 38,
11221134

centres or between parks just at the edge of different


towns (all U > 012, n = 6, P > 034; Table 5). Similarity
indices did not vary between years (U = 2650, n = 18,
P = 022). BSR of diet and habitat guilds did not vary
between park locations in 1998 (2, all d.f. = 4, P > 063
and all d.f. = 8, P > 091, respectively) and in 1999 (all
d.f. = 4, P > 073 and all d.f. = 8, P > 084, respectively).
At a local scale, distance from the periurban landscape did not affect park BSR, community similarity or
guild composition.

Discussion
Our results indicated that BSR in urban areas was independent of the adjacent landscapes of study towns
across different spatial scales. In addition, the low similarity values observed between periurban and urban
bird communities supported this conclusion. Using
several studies from the northern hemisphere, we found
positive correlations in BSR between suburban and
periurban landscapes. However, this was mostly due to
a latitudinal effect on BSR rather than a relationship
between birds of urban and periurban landscapes; BSR
in urban and periurban landscapes increases or
decreases together with town latitude. These results are
coherent with classical decreasing trends in BSR from
south to north, e.g. from 273 regular breeding bird species in France to 235 in Finland (Yeatman-Berthelot
& Jarry 1994; Visnen, Lammi & Koskimies 1998).
This latitudinal relationship did not always persist at
the regional scale of the country. Highly urban areas
tended to have similar BSR independent of their geographical location, as obtained by Jokimki et al.
(1996). Similarity indices obtained from our selected
studies indicate that a large part of the bird community
is similar throughout the urbanization gradient, but
also that in a town centre about 50% of the avifauna is
independent of periurban species (mean similarity
index between centre and periurban = 050). Interestingly in Finland and Canada, some BSR was higher in
suburban than in periurban landscapes. This indicates
that intermediate level disturbances caused by urbanization (Blair 1996), and also food supplementation
given by humans in some habitats (see our Finland
results on seed-eater functional group; Jokimki &
Suhonen 1998), might be beneficial for birds in northern conditions. Our analysis at the regional scale corroborates the hypothesis that BSR does not vary
greatly within the city as a result of the types of periurban landscapes. Sites with similar structures had
similar BSR, independent of the size of the city or the
type of periurban landscapes, as Blondel et al. (1984)
have suggested. Bird communities were more similar
between cities for a given habitat than within cities
between different habitats. Our local scale analysis,
which compared similar types of sites, parks with trees
and lawns, at different distances from the source, did

JPE_666.fm Page 1131 Thursday, September 20, 2001 2:50 PM

1131
Urban bird
diversity and
landscape context

2001 British
Ecological Society,
Journal of Applied
Ecology, 38,
1122 1134

not reveal any difference in BSR and similarity data.


Similarly, guild comparisons between sites did not
produce any significant results at either regional or
local scales. This lack of modification in community
structures supports the relatively small influence of
periurban landscapes on urban BSR.
It could be that individuals of some species emigrate
to a new town from other urbanized areas, as suggested
by Erz (1966). Thus, urban bird communities can be
composed of species not common in the surrounding
landscapes (Clergeau et al. 1998) and local rather than
regional scale factors play a more important role in
shaping the structure of urban bird communities. This
result agrees with Emlen (1974), who noted, in the
recent town of Tucson, that some species absent from
the periurban landscape live in the urban area. This
sort of independence has also been observed in several
population level bird studies (Havlin 1962; Erz 1966;
Coombs et al. 1981; Birkhead 1991).
Among studies involving distances from a species
source ( patches of native vegetation) to explain bird
populations in urban areas, only that of Munyenyembe, Harris & Hone (1989) obtained significant
results. However, they worked along a short distance
gradient (less than 15 km) and only differences within
02 km from the woodland area were significant. This
represents a classic fringe effect (De Graaf, Geis &
Healy 1991; Chou & Soret 1996), where proximity of
resources or habitat explains the presence of birds.
Our results suggest that, for a similar number of species in periurban areas, we can expect different BSR in
urban areas due mainly to differences in local features.
Thus, in an urban biodiversity perspective, the management of BSR would be best served by actions at a
local scale. For example, diversity and density of vegetation and habitat heterogeneity in urban sites increase
BSR (Davis & Glick 1978; Lancaster & Rees 1979;
Goldstein, Gross & DeGraaf 1986; Clergeau et al.
1998), affect bird assemblages (Savard 1978; Jokimki
et al. 1996) and in some cases decrease nest predation rate (Jokimki & Huhta 2000). Also, the age of a
residential area is known to influence directly BSR
through the development of its vegetation (Vale & Vale
1976; Hohtola 1978; Munyenyembe, Harris & Hone
1989). The size of each landscape element also influences BSR by fulfilling minimum area requirements of
species (Hohtola 1978; Tilghman 1987; Jokimki
1999). In addition, urban feeders can not only change
the density but also the composition of bird assemblages, as well as increase bird diversity (Jokimki &
Suhonen 1998; Morneau et al. 1999).
Our study reinforces a suggestion not always clearly
expressed in recent syntheses of urban ecology (Germaine et al. 1998; Jokimki 1999; Savard, Clergeau &
Mennechez 2000): whatever the biodiversity quality of
the periurban landscape, site-specific actions such as
shrub and tree planting, water restoration and increasing
vegetation diversity can change bird diversity in the town
and improve the quality of humanwildlife contacts.

Acknowledgements
We thank our research organizations that allowed
the development of a new ecological perspective, with
studies in urban systems. This work was partially
supported by grants from Ministre Franais de
lEnvironnement and the Canadian Wildlife Service.
The authors are grateful to F. Gilot, A. Bregeon,
G. Falardeau and M.L. Kaisanlahti-Jokimki for their
field works and for two anonymous referees for their
comments on the manuscript.

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Appendix
Birds observed in France (spring and winter census), Finland (winter census) and Canada (winter census)

2001 British
Ecological Society,
Journal of Applied
Ecology, 38,
1122 1134

Name

Country*

Season census

Diet guild

Feeding habitat guild

Accipiter striatus (Viellot)


Aegithalos caudatus (Linnaeus)
Anas platyrhynchos (Linnaeus)
Anas rubripes (Brewster)
Apus apus (Linnaeus)
Ardea cinerea (Linnaeus)
Bombycilla garrulus (Linnaeus)
Bonasa umbellus (Linnaeus)
Bucephala clangula (Linnaeus)
Cardinalis cardinalis (Linnaeus)
Carduelis cannabina (Linnaeus)
Carduelis carduelis (Linnaeus)
Carduelis chloris (Linnaeus)
Carduelis flammea (Linnaeus)
Carduelis pinus (Wilson)
Carduelis spinus (Linnaeus)
Carduelis tristis (Linnaeus)
Carpodacus mexicanus (Miller)
Carpodacus purpureus (Gmelin)
Certhia brachydactyla (Brehm)
Coccothraustes coccothraustes (L.)
Colaptes auratus (Linnaeus)
Columba livia dom. (Gmelin)
Columba palumbus (Linnaeus)
Corvus brachyrhynchos (Brehm)
Corvus corone (Linnaeus)
Corvus frugilegus(Linnaeus)
Corvus monedula (Linnaeus)
Cyanocitta cristata (Linnaeus)
Delichon urbica (Linnaeus)
Dendrocopos major (Linnaeus)
Emberiza cirlus (Linnaeus)
Emberiza citrinella (Linnaeus)
Erithacus rubecula (Linnaeus)
Falco sparverius (Linnaeus)
Fringilla coelebs (Linnaeus)
Fulica atra (Linnaeus)
Gallinula chloropus (Linnaeus)
Garrulus glandarius (Linnaeus)
Hippolais polyglotta ( Vieillot)
Hirundo rustica (Linnaeus)
Lanius excubitor (Linnaeus)
Larus argentatus (Pontoppidan)
Larus marinus (Linnaeus)
Larus ridibundus (Linnaeus)
Loxia curvirostra (Linnaeus)
Loxia leucoptera (Gmelin)
Luscinia megarhynchos (Brehm)
Mergus merganser (Linnaeus)

Ca
Fr
Fr
Ca
Fr
Fr
Fi
Ca
Ca
Ca
Fr
Fr
Fr/Fi
Fi/Ca
Ca
Fr
Ca
Ca
Ca
Fr
Fr
Ca
Fr/Fi/Ca
Fr
Ca
Fr/Fi
Fr
Fr
Ca
Fr
Fr/Fi
Fr
Fr/Fi
Fr
Ca
Fr
Fr
Fr
Fr
Fr
Fr
Ca
Fr/Ca
Ca
Fr
Ca
Fi/Ca
Fr
Ca

wi
sp/wi
sp
wi
sp
sp
wi
wi
wi
wi
sp
sp/wi
wi
wi
wi
wi
wi
wi
wi
sp/wi
sp
wi
sp/wi
sp/wi
wi
sp/wi
sp
sp/wi
wi
sp
sp/wi
wi
wi
sp/wi
wi
sp/wi
sp
sp
sp/wi
sp
sp
wi
sp/wi
wi
wi
wi
wi
sp
wi

c
i
g
g
i
c
g
g
i
g
g
g
g
g
g
g
g
g
g
i
g
i
o
g
o
o
o
o
o
i
i
g
g
i
c
g
g
g
o
i
i
c
o
o
c
g
g
i
c

t
t
a
a
o
a
t
t
a
s
s
m
o
t
t
t
t
t
t
t
t
m
o
o
m
m
m
m
t
o
t
o
o
s
m
o
a
a
t
s
o
m
o
a
a
t
t
s
a

*Fr = France; Fi = Finland; Ca = Canada.


sp = spring; wi = winter.
i = insect feeder; g = seed and vegetation feeder; o = omnivorous; c = carnivorous.
a = aquatic habitat; t = tree dwelling habitat; s = shrub habitat; m = meadow habitat; o = other habitat.

JPE_666.fm Page 1134 Thursday, September 20, 2001 2:50 PM

1134
P. Clergeau,
J. Jokimki &
J.-P.L . Savard

Appendix (contd )
Name

Country*

Season census

Diet guild

Feeding habitat guild

Motacilla alba (Linnaeus)


Muscicapa sriata (Pallas)
Parus ater (Linnaeus)
Parus atricapillus (Linnaeus)
Parus caeruleus (Linnaeus)
Parus cristatus (Linnaeus)
Parus major (Linnaeus)
Parus montanus (Conrad)
Parus palustris (Linnaeus)
Passer domesticus (Linnaeus)
Phoenicurus ochruros (Gmelin)
Phylloscopus collybita (Vieillot)
Phylloscopus trochilus (Linnaeus)
Pica pica (Linnaeus)
Picoides pubescens (Linnaeus)
Picoides villosus (Linnaeus)
Picus viridis (Linnaeus)
Podiceps ruficollis (Pallas)
Prunella modularis (Linnaeus)
Pyrrhula pyrrhula (Linnaeus)
Regulus sp.
Serinus serinus (Linnaeus)
Sitta canadensis (Linnaeus)
Sitta carolinensis (Latham)
Sitta europea (Linnaeus)
Sterna hirundo (Linnaeus)
Streptopelia decaocto (Frivaldsky)
Strix aluco (Linnaeus)
Sturnus vulgaris (Linnaeus)
Sylvia atricapilla (Linnaeus)
Troglodytes troglodytes (Linnaeus)
Turdus iliacus (Linnaeus)
Turdus merula (Linnaeus)
Turdus migratorius (Linnaeus)
Turdus philomenos (Brehm)
Turdus viscivorus (Linnaeus)
Zenaida macroura (Linnaeus)

Fr
Fr
Fi
Ca
Fr/Fi
Fr
Fr/Fi
Fi
Fr
Fr/Fi/Ca
Fr
Fr
Fr
Fr/Fi
Ca
Ca
Fr
Fr
Fr
Fr/Fi
Fr
Fr
Ca
Ca
Fr
Fr
Fr
Fr
Fr/Ca
Fr
Fr
Fr
Fr
Ca
Fr
Fr
Ca

wi
sp
wi
wi
sp/wi
sp/wi
sp/wi
wi
sp/wi
sp/wi
sp
sp/wi
sp
sp/wi
wi
wi
sp/wi
sp
sp/wi
sp/wi
sp/wi
sp/wi
wi
wi
sp/wi
sp
sp/wi
sp
sp/wi
sp/wi
sp/wi
wi
sp/wi
wi
sp/wi
sp/wi
wi

i
i
i
i
i
i
i
i
i
o
i
i
i
o
i
i
i
c
i
g
i
g
i
i
i
c
g
c
o
i
i
i
i
i
i
i
g

o
s
t
t
t
t
t
t
t
o
o
s
s
m
t
t
m
a
s
t
t
t
t
t
t
a
o
t
m
s
s
m
m
s
m
m
o

*Fr = France; Fi = Finland; Ca = Canada.


sp = spring; wi = winter.
i = insect feeder; g = seed and vegetation feeder; o = omnivorous; c = carnivorous.
a = aquatic habitat; t = tree dwelling habitat; s = shrub habitat; m = meadow habitat; o = other habitat.

2001 British
Ecological Society,
Journal of Applied
Ecology, 38,
11221134

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