Professional Documents
Culture Documents
Journal of Applied
Ecology 2001
38, 1122 1134
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
1123
Urban bird
diversity and
landscape context
2001 British
Ecological Society,
Journal of Applied
Ecology, 38,
1122 1134
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
1124
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 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
Authors
City (country)
Size
agglomerate
1000 inhab.
Weber (1972)
Vancouver (Canada)
1380
49 NW
Geis (1974)
Columbia (USA)
395
34 NW
Waterloo (USA)
80
43 NW
Marchetti (1976)
Marseille (France)
900
43 OW
Tornio (Finland)
20
66 OW
Savard (1978)
Toronto (Canada)
3425
44 NW
Hohtola (1978)
Kuopio (Finland)
80
63 OW
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
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
1125
Urban bird
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)
Oxford (USA)
Jokimaki (1989)
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)
55
37 NW
Rennes (France)
210
48 OW
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
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)
1126
P. Clergeau,
J. Jokimki &
J.-P.L . Savard
2001 British
Ecological Society,
Journal of Applied
Ecology, 38,
11221134
Table 1. (contd )
1127
Urban bird
diversity and
landscape context
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
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
similarity index = 2c /( a + b )
40
30
20
10
0
10
15
20
25
30
35
40
1128
P. Clergeau,
J. Jokimki &
J.-P.L . Savard
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
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).
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.
Redon
17
21
Dinan
22
19
Fougres
18
17
St Malo
18
17
Rennes
21
20
070
095
Kemijrvi
4
9
Tornio
7
9
Kemi
7
11
Rovaniemi
4
9
Oulu
8
11
094
092
St Nicolas
9
6
Levis
4
8
Quebec
45
5
Montreal
4
10
276
043
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
2001 British
Ecological Society,
Journal of Applied
Ecology, 38,
1122 1134
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
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
061
068
069
065
069
074
Rennes
079
069
075
076
063
082
Nantes
071
070
068
078
074
079
073 009
072 007
Within town:
070 013
074 009
067 006
074 013
Park in centre:
Angers
22 km
30
20
10
0
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
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
1131
Urban bird
diversity and
landscape context
2001 British
Ecological Society,
Journal of Applied
Ecology, 38,
1122 1134
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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Received 3 March 2001; revision received 13 July 2001
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
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
1134
P. Clergeau,
J. Jokimki &
J.-P.L . Savard
Appendix (contd )
Name
Country*
Season census
Diet guild
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
2001 British
Ecological Society,
Journal of Applied
Ecology, 38,
11221134