Occurrence of breeding bird species in urban parks: Effects of park structure and broad-scale variables
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Urban Ecosystems, 3, 21–34, 1999
°
c 2000 Kluwer Academic Publishers. Manufactured in The Netherlands.
Occurrence of breeding bird species in urban parks:
Effects of park structure and broad-scale variables
JUKKA JOKIMÄKI jukka.jokimaki@urova.fi
Arctic Centre, University of Lapland, P.O. Box 122, FIN-96101 Rovaniemi, Finland
Abstract. Occurrences of breeding bird species in 54 urban parks were investigated in the city of Oulu in
northern Finland. Park area, human activity, habitat, and landscape structure within a 9-ha square surrounding
the study park were related to the bird species richness and occurrence of individual bird species. A total of 22
species was observed. The area of the park explained 39% of the variance of species richness. Seven species
(wheatear [Oenanthe oenanthe], common rosefinch [Carpodacus erythrinus], garden warbler [Sylvia borin],
lesser whitethroat [Sylvia curruca], linnet [Acanthis cannabina], redpoll [Carduelis flammea], and yellowhammer
[Emberiza citrinella]) were not detected in parks of22 JOKIMÄKI Furthermore, urban habitats are complicated by the many disturbance factors associated with human activity. These “nonnatural” factors may play an additional role in regulating animal populations. Unfortunately, most bird–habitat relationship studies have been made in natural ecosystems, while urban areas have been largely ignored (but see Gavareski, 1976; Luniak, 1981; Tilghman, 1987). To promote biodiversity in urban areas, research is needed to develop landscape planning and management methods (White, 1994). In general, urbanization decreases bird species diversity and richness but increases the total density of birds (Bezzel, 1985; Hohtola, 1978; Jokimäki, 1992; Lancaster and Rees, 1979). Studies of bird communities in urban parks have shown that parks are considerably richer in bird species diversity and richness than other urban habitats (Hadidian et al., 1997; Jokimäki and Suhonen, 1993; Tilghman, 1987). Thus it is important to know the factors that restrict the occurrence of bird species in urban parks and how we can maintain or even increase the diversity of birds in these urban patches. In this paper I examine relationships among area, small- and broad-scale landscape structure, human activity, and bird assemblages in urban parks. The purpose of this study was to determine which factors determine species richness and the occurrence of breeding bird species in urban parks. Methods Study area This study was conducted in the city of Oulu (65◦ 010 N, 25◦ 280 E) in northern Finland. The human population of the town at the end of 1994 was 106,419, and the average human population density in the main areas of the city varied between 500 and 2000 per km2 . The city covers about 3547 ha of land. All study parks were bounded by roads, cleared open areas, or buildings. The average size of the study parks was 1.2 ha (range 0.1–7.0 ha) and they covered a total of 64.1 ha (N = 54). The study parks were grouped in three size categories; ≤0.5 ha (N = 23), 0.75–1 ha (N = 19), and 1.5–7 ha (N = 12). The number of managed parks (i.e., parks that have shrub and small tree plantations, hedges, etc., and that are continuously tended as regards lawn mowing and shrub clipping by gardeners) was 31, and the number of unmanaged parks (i.e., green areas with more natural vegetation; grasses, shrubs, and trees) was 23. In species–area studies, the role of habitat structure is often neglected. Normally, parks in the centers of town are small and usually managed, whereas those on the periphery are larger and usually less groomed (Gilbert, 1989). I selected the study parks so that the effects of park size, location of the park, and habitat structure could be separated; i.e., I selected parks of different sizes and management levels (managed/unmanaged) from the center of the town to the private house area. Because the total number of parks found in the town center was small (N = 28), I included all of them in the study. Thus randomization was not necessary. Bird census Parks were surveyed from 5 to 27 June 1995. This census period corresponds well with the breeding season of the most bird species in the study area. In northern Finland, birds
BIRDS IN URBAN PARKS: A LANDSCAPE ECOLOGICAL VIEW 23 seldom breed twice during one year. I used a mapping census method of breeding land birds (Koskimies and Väisänen, 1991) to determine the breeding presence of individual bird species. All surveys were made between 4 and 7 A.M. (i.e., before most people go their workplaces, etc.) during weekdays, in good weather only. Thus both the time of day and the day of the week were controlled in surveys. Each park was thoroughly and systematically searched twice, and the breeding territories of species were located. A survey did not consist of a single route through the park but rather a zig-zag walk through the park. The time spent in a park for each census was about 10 minutes. In larger parks (≥1.5 ha) the census time was 5 min longer than in smaller parks (≤1.0 ha). Because of the small size of the parks, it is assumed that practically all territory holders were detected. Bird species were considered to be breeders if they exhibited territorial behavior (e.g., singing or warning) on both visits or if breeding behavior (e.g., nesting activities, feeding nestlings, or fledglings) was recorded on any visit. Overflying birds and obvious nonresidents were not included in the analysis. Typically, the single-visit method detects about 60% of the breeding pairs and 90% of the species in forested areas of boreal zones (Järvinen and Lokki, 1978). I anticipated that the census efficiency in my study area may have been even higher because of the simple habitat structure and the short and synchronous breeding season (see Helle, 1986; Järvinen and Lokki, 1978). Latin names of bird species detected are shown in Table 1. Measured variables Area, human activity, and landscape structure at small and broad scales were determined for each study park (Table 2). Recreational activity was measured by counting the num- ber of people visiting each park at midday (11:00–13:00) during weekdays for a period of 5 minutes. Thus both the time of day and the day of the week were controlled in countings. Censuses were carried out in good weather only. Since the count was done only once in each park, and time spent in each park was relatively short (5 min), the mea- sured variable must be seen as indicative. The area of the park was measured from the city map; other habitat variables within the park boundary were measured in the field. Five vegetation sampling points, each with a diameter of 6 m (28 m2 ), were used to de- scribe the structure of each park. One sampling point was situated in the middle of the park, and the four others were situated at a distance of 15 m from the central point to- wards the four main compass directions. I calculated the number of pines (Pinus sylvestris), spruces (Picea abies), and deciduous shrubs (height 2–5 m, >5–10 m, >10–15 m, and >15 m) within each sampling point. Values for shrubs and trees were converted to densities (average number of shrubs or trees/ha). The tree species and foliage height diversity (Shannon formula (Begon et al., 1990)) were calculated from the tree species frequency series. The height of the domi- nant trees (m) in each park was measured using a hypsometer. I also counted the number of nest boxes in each park. Because of their small sizes (28 m2 ), vegetation plots were not intended to describe park vegetation structure in detail. However, since each study park included only managed or unmanaged habitats, each was homogeneous in respect to its management status and general habitat structure. Therefore it is supposed that the sampling design used gives an adequate estimate of the general structure of the study parks.
24 JOKIMÄKI
Table 1. Proportion of the breeding occurrence (%) of different bird species in different park size classes and in
the whole study area, and the total number of pairs.
Total Total
≤0.5 ha ≥0.75–1 ha ≥1.5–7 ha occurence number
Bird species (N = 23) (N = 19) (N = 12) (%) of pairs
Chaffinch Fringilla coelebs (TD) 30 63 50 46 29
Great tit Parus major (CD) 30 53 59 44 24
Willow warbler Phylloscopus trochilus (GD) 13 42 59 41 31
Fieldfare Turdus pilaris (TD) 26 32 50 33 38
Hooded crow Corvus corone cornix (T-) 9 37 50 28 15
Blue tit Parus caeruleus (CD) 19 21 67 26 14
Greenfinch Carduelis chloris (TD) 19 42 33 26 14
Pied flycatcher Ficedula hypoleuca (CD) 13 21 42 22 14
Pied wagtail Motacilla alba (GO) 17 21 25 20 11
Magpie Pica pica (T-) 22 5 33 19 10
Spotted flycatcher Muscicapa striata (CC) 9 21 25 17 8
Redstart Phoenicurus phoenicurus (CD) 4 16 25 13 7
Rook Corvus frugilegus (T-) 9 5 17 9 32
Siskin Carduelis spinus (SC) 4 5 17 7 4
Redwing Turdus iliacus (SD) 9 0 8 6 3
Wheatear Oenanthe oenanthe (GO) 0 0 25 6 3
Rosenfinch Carpodacus erythrinus (SD) 0 5 8 4 2
Garden warbler Sylvia borin (SD) 0 5 0 2 1
Lesser whitethroat Sylvia curruca (SD) 0 5 0 2 1
Linnet Acanthis cannabina (SD) 0 0 8 2 1
Redpoll Carduelis flammea (SD) 0 0 8 2 1
Yellowhammer Emberiza citrinella (GO) 0 0 8 2 1
The letters after the bird species indicate the breeding habits of the species according to von Haartman et al.
(1963–72) (T = nesting mainly in trees, C = in cavities, G = on ground, and S = in shrubs) and the main habitat
type (D = deciduous forest species, C = coniferous forest species, O = open area species).
Six broad-scale landscape variables were measured from the city map (Table 2). I es-
timated the proportion of the different habitat types (parks, built-up areas, open areas)
within a 9-ha square around the study parks and calculated the diversity of the surrounding
landscape by the Shannon formula. Three different variables were measured to describe
the level of isolation. These measurements described the distances from the study parks to
the nearest woods or parks and the proportion of the park area adjacent to the study parks
(Table 2).
Data analyses
Variables with nonnormal distributions were either log or arcsin transformed. Analyses were
carried out on breeding bird species richness and the occurrence of individual bird species.BIRDS IN URBAN PARKS: A LANDSCAPE ECOLOGICAL VIEW 25
Table 2. Variables used in logistic regression analysis.
Variables within the park boundary
Recreational activity (Mean number of people/park/visit) (RECREATION)
Area of park (ha) (AREA)
Density of shrubs (/ha) log-transformed (SHRUBS)
% of trees >2–5 m of the total density of trees (TREE 1)
% of trees >5–10 m of the total density of trees (TREE 2)
% of trees >10–15 m of the total density of trees (TREE 3)
% of trees >15 m of the total density of trees (TREE4)
% of deciduous trees of the total density of trees (DECITREE)
Total density of trees (/ha) (TREEDEN)
Height of the dominant trees (m) (TREEHEI)
Tree species diversity (Shannon index) (TREEDIV)
Foliage height diversity of the tree cover (Shannon index) (FHD)
Total number of nest boxes (NBOXES)
Broad-scale variables
Total length of streets within a 9-ha square surrounding the study park (STREETS)
% of block of flats area within a 9-ha square surrounding the study park (FLATS)
Isolation I (Distance to the nearest wood or park of >2.2 ha, m, log-transformed) (ISO1)
Isolation II (Distance to the nearest wood or park >9 ha, m) (ISO2)
Isolation III (Proportion of park area within a 9-ha square surrounding the studied park) (ISO3)
Diversity of the surrounding landscape within a 9-ha square surrounding the park (LDIV)
The variation in species richness was analysed by forward-stepwise multiple regression
analysis, and all breeding species were included in the analysis. The existence of individual
bird species (presence/absence) was modeled using a forward-stepwise logistic regression
analysis (see Trexler and Travis, 1993), since most of the bird species were absent, or only
one pair of the species bred in the study park. Species occurring in fewer than 10% of the
parks were omitted from that analysis because of the small sample size (see Table 1). The
dependent variable, presence or absence of each species in each park, was scored as 1 or 0,
respectively. The significance of each variable included in the models is based on the Wald
test (Hosmer and Lemeshow, 1989). The best models were not selected on purely statistical
grounds. I included only ecologically meaningful variables in the models. Therefore some
variables with an individual significance slightly less than the conventionally accepted
p = 0.05 level were included in some models (Table 3).
Results
A total of 22 bird species bred in the study parks (N = 54) of Oulu (Table 1). The
average number of breeding bird species per study park was lower in small-sized parks
(x̄ = 2.1, SD = 2.6) than in large-sized (x̄ = 5.9, SD = 3.7) or medium-sized parks26 JOKIMÄKI
Table 3. Logistic regression models for the individual bird species. The abbreviation of each variable is
given in Table 1. Negative coefficients are indicated with -. The significance of each coefficient is based
on Wald test (∗∗ P < 0.01, ∗ P < 0.05 and ◦ P < 0.10). The “adequacy” of each model is assessed with
the fit of its predictions to data (goodness-of-fit, G 2 ). Common names of bird species are given in Table 1.
Species Variables G2 P
Fringilla coelebs No model
Phylloscopus trochilus FLATS(-)∗ 6.1 0.0132
Corvus corene cornix FLATS(-)∗ 6.2 0.0132
Ficedula hypoleuca NBOXES∗∗ , FLATS(-)∗ , TREEDIV◦ 25.2BIRDS IN URBAN PARKS: A LANDSCAPE ECOLOGICAL VIEW 27
Table 4. Distribution of most abundant birds (≥10 pairs) in relation to patch size with expected pair numbers
in parentheses.
Patch size classes
≤0.5 ha ≥0.75–1 ha ≥1.5–7 ha
(Tot. area 9.1 ha) (Tot. area 18.0 ha) (Tot. area 37 ha) χ 2 (d f = 2)
Fringilla coelebs 7(4) 14(8) 8(17) 10.77∗∗
Parus major 7(3) 19(7) 7(14) 8.75
Phylloscopus trochilus 4(4) 14(9) 16(11) 4.59
Turdus pilaris 12(5) 16(11) 10(22) 17.21∗∗∗
Corvus corone cornix 2(2) 7(4) 6(9) 2.66
Parus caeruleus 2(2) 4(4) 8(8) 0.002
Carduelis chloris 2(2) 8(4) 4(8) 6.27
Ficedula hypoleuca 4(2) 4(4) 6(8) 2.57
Motacilla alba 4(2) 4(3) 3(6) 5.86
Pica pica 5(1) 1(3) 4(6) 10.74∗∗
Corvus frugilegus 1(5) 4(9) 21(18) 1.77
Expected number of pairs proportional to area in each patch size class, significance levels used: ∗∗ P < 0.01
and ∗∗∗ P < 0.001. Common names of bird are given in Table 1.
positive relationship between the smallest park area in a which a species occurred and dis-
tance from the town center (Spearman Rank correlation, r S = 0.64, p < 0.01, N = 22
bird species).
Distribution of the most abundant species (≥10 pairs) was proportional to area in each
size class for 8 out of the 11 species (Table 4). The chaffinch, fieldfare, and magpie were
disproportionally more abundant in the smaller than in the larger parks (Table 4).
Logistic models of individual bird species
From the logistic regressions, 9 of the 12 most common bird species had variables that
significantly affected their distribution, and a total of 13 variables (5 broad-scale variables
and 8 variables within the park boundaries) out of the 19 were entered in the models
(Table 3). The proportion of blocks of flats within a 9-ha square negatively influenced the
breeding presence of the willow warbler, hooded crow, spotted flycatcher, and pied fly-
catcher. The proportion of park area within the 9-ha square surrounding the park positively
influenced the breeding presence of the fieldfare. The total length of streets within a 9-ha
square surrounding the study park influenced negatively, whereas the distance to the nearest
park over 9 ha influenced positively the distribution of the spotted flycatcher. Diversity of
the surrounding landscape within the 9-ha square surrounding the study park negatively
influenced the breeding occurrence of the great tit.
Park area had a positive influence on the breeding presence of the blue tit and the green-
finch. The proportion of trees 5 to 10 m high was the most important variable influencing28 JOKIMÄKI
the presence of the great tit. For the fieldfare, the height of the dominant trees was the most
important variable. The total density of shrubs and the proportion of trees over 15 m high
negatively influenced the breeding presence of the greenfinch. The total density of trees
was the only variable that influenced the presence of the wagtail. The number of nest boxes
was the most important variable for the pied flycatcher. Tree species diversity influenced
positively the breeding presence of the pied flycatcher and the spotted flycatcher. The num-
ber of people visiting the parks was not a significant variable in any of the models (Table 3).
Birds in managed vs. unmanaged parks
Bird species richness per park did not differ between the managed (x̄ = 3.3, SD = 2.6,
N = 31) and the unmanaged parks (x̄ = 4.2, SD = 3.6. N = 23) (Mann–Whitney U = 32.3,
p > 0.05). The willow warbler (χ12 = 4.1, p < 0.05) and magpie (χ12 = 3.8, p = 0.05)
were the only bird species out of 22 species for which there were significant differences in
managed and unmanaged parks, both species were observed more often in unmanaged than
in managed parks.
Recreational activity and the proportion of deciduous trees were higher in managed than
in unmanaged parks, whereas the density of shrubs and trees was greater in unmanaged
than in managed parks (Table 5).
Table 5. Habitat and landscape structure of managed and unmanaged parks.
Managed (N = 31) Unmanaged (N = 23)
x̂ SD x̂ SD U p
RECREATION (people/park/visit) 4.1 5.4 1.2 2.3 175.0BIRDS IN URBAN PARKS: A LANDSCAPE ECOLOGICAL VIEW 29 Discussion Species richness and composition In the study area, species richness was strongly associated with woodlot area, as shown in many other studies (e.g., Blake and Karr, 1987; Lussenhop, 1977). However, other factors, including the height of the dominant trees and the density of small-sized trees, also had a positive effect on bird species richness (see also Freemark and Merriam, 1986; Gavareski, 1976; Tilghman, 1987). The total number of breeding species (22) in this study area compares well with results from Central Europe, where the species richness varied from 5 to 21 in parks of similar sizes (see Sasvari (1984) and references therein). Despite being in a northern latitude, most of the breeding species in my study area were the same as those found in Central European parks: see Biadun, 1994 (Lublin); Luniak, 1981 (Warsaw); Mulsow, 1982 (Hamburg); Sasvari, 1984 (Budapest); Sukopp et al., 1982 (Berlin); Tomialojc, 1970 (Lenica); Tomialocj and Profus, 1977 (Wroclaw). One reason for the similarity of species composition between different cities may be that urban areas are evolutionarily young habitats. Only a few species are adapted to life in cities, and these dominant bird species are mainly the same in Central and Northern Europe: see also Huhtalo and Järvinen, 1977; Jokimäki et al., 1996; Luniak, 1990; Thompson et al., 1993. House sparrows (Passer domesticus) and feral pigeons (Columba livia domestica), which might be expected to compete for nesting places with park birds, were not nesting in the parks of this study (see also Lussenhop (1977) but for opposite results see Tilghman (1987)). However, sparrows and pigeons quite commonly foraged in the managed parks of the town center (personal observation). Some bird species (woodpigeon Columba palumbus, song thrush Turdus philomelos, chiffchaff Phylloscopus collybita, robin Erithacus rubecula, and jay Garrulus glandarius) were absent in this study area although these species are quite common in urban areas of Central Europe (see references in earlier paragraph) and in the forests surrounding the study area (Väisänen et al., 1998). Central European cities are older than northern cities, and perhaps the northern populations of those bird species have not yet adapted to life in cities in the North. Another reason might be that the park and landscape structure differ between Central European cities and northern ones. It is also possible that in northern Finland, there is still a lot of habitat for these species outside urban areas, in contrast to Central Europe, where birds may be forced into urban areas because other habitats have been decimated. Most of the bird species in my urban study area were deciduous forest species (see also Jokimäki and Suhonen, 1998; Suhonen and Jokimäki, 1988). This was understandable, since deciduous trees and shrubs are favored over conifers in urban planning in Finland (Jokimäki, 1996) and in many other parts of Europe (Gilbert, 1989). Ground-nesting birds were poor colonizers of urban parks (see also Gavareski, 1976; Luniak, 1981; Tomialojc, 1970; Tomilojc and Profus, 1977), whereas cavity-nesting species were among the most common bird species (this study, see also Suhonen and Jokimäki, 1988). Lack of suitable nest sites with concealment vegetation or nest predation pressure may limit the occurrence of ground-nesters (Jokimäki, 1996). However, cavity-nesters may benefit from the large numbers of nest boxes in urban parks (Luniak, 1992).
30 JOKIMÄKI Influence of park area Park area was the most important predictor of the number of breeding bird species in this study. One reason for this observation may be that in the smallest parks, the minimum area requirements of the individual bird species are not fulfilled. However, the minimum area requirements of bird species in urban parks are seldom discussed (but see Luniak, 1981). In this study area many of the bird species (32%) avoided small parks (
BIRDS IN URBAN PARKS: A LANDSCAPE ECOLOGICAL VIEW 31 decline in abundance with increases in built structure proportion. All these species were affected negatively by the proportion of blocks of flats (FLATS) within 9 ha of the park. A large number of adjacent buildings may negatively affect both birds (Tilghman, 1987) and mammals (Dickman, 1987). This may be due to the reduced amount of vegetation cover in areas around the park. Also, the impacts of trampling may be greater in parks with a high building density in the immediate vicinity. However, human activity was not higher in parks surrounded by high building density in this study. Human activity may influence breeding success of birds (Osborne and Osborne, 1980; Tomialocj, 1978; Tomialojc and Profus, 1977). It has been suggested that recreational use of urban parks is one of the main features affecting bird species in urban woodlots (Luniak, 1982; van der Zande et al., 1984). However, in this study the presence of people in parks did not affect the breeding occurrence of birds (see also Luniak, 1981; Tilghman, 1987). Perhaps the intensity of the recreational use was not so heavy in the parks, or human activity is of only minor importance compared to other variables. In this study, isolation of parks seems not to be an important factor for park birds (see also Tilghman, 1987), since the isolation variable only entered into the models of the fieldfare and the spotted flycatcher. Perhaps the degree of isolation in the study area was negligible relative to dispersal ability of birds, or my measurements of the level of isolation were not important. Influence of park structure Different bird species were associated with different vegetation variables. The positive relationship between tree height and the breeding occurrence of the fieldfare may be related to the selection of safe nesting places. Nests situated in the lower parts of trees may suffer from higher levels of predation or disturbance by humans than nests situated in the higher parts of trees. Birds in urban areas tend to place their nests higher than in areas with lower disturbance (e.g., Dhindsa et al., 1989). The number of nest boxes was the most important variable explaining the occurrence of the cavity-nesting pied flycatcher, but not for the other cavity-nesters such as Parus species, in the city parks. Many studies have shown that nesting success is higher in nest boxes than in natural cavities (e.g., Nilsson, 1984). The pied flycatcher may outcompete the Parus species for the proffered nest boxes. During the nest building and egg laying, great tits are absent from the nest for such long periods that pied flycatchers succeed in usurping their nests (Slagsvold, 1975). Several studies have demonstrated the positive relationship between diversity of vegetation (e.g., Sukopp et al., 1982), foliage height diversity (FHD) (e.g., MacArthur and MacArthur, 1961), and bird species diversity. However, I did not find any effects of FHD on the bird species richness, and the effect of the tree species diversity was negative. Abundance of the undergrowth and shrub layer have been observed to correlate positively with bird species richness and diversity in urban parks (Gavareski, 1976; Luniak, 1981; Tilghman, 1987). However, in this study, there was no impact of the shrub layer on bird species richness or any bird species. It could be that factors other than “natural” factors (e.g., FHD, shrub density) are more important for urban birds. For example, in this study
32 JOKIMÄKI three park bird species were observed to suffer from adjacent buildings, and many wintering bird species correlated positively to human population density and factors associated with humans (e.g., winter feeding) (Jokimäki and Suhonen, 1998). The willow warbler and the magpie occurred more often in unmanaged than in managed parks. For the ground-nesting willow warbler, the adequate shrub cover in unmanaged parks may decrease nesting failures caused by visually searching avian nest predators (Jokimäki, 1996). As corvids prefer to build their nests in coniferous trees in urban areas (Ilyichev et al., 1990), the magpie was more often observed in the pine-dominated, unmanaged parks than in the managed parks, where the proportion of deciduous trees was higher. Conclusions The results of this study demonstrate that factors other than “natural” factors (e.g., tree density, etc.) can also affect the distribution of some bird species in urban parks. Although park area was one of the best single predictors of species richness, my results indicate that bird species responses to urbanization are individualistic. Some species selected small parks (e.g., the magpie and the fieldfare), while others avoided those small areas (e.g., Sylvia and Emberiza species) and selected larger ones. The pied flycatcher benefited from the occurrence of nest boxes in the parks. A great variety of cavity-nesting bird species may colonize urban parks if different kinds of nest boxes are available. Most of the bird species detected in urban parks were deciduous forest species. This may be related to the fact that deciduous trees are favored over conifers in urban planning in Finland. To increase occurrence opportunities of coniferous forest species, coniferous tress must be used in park management, especially in northern latitudes where deciduous trees and shrubs drop their leaves in autumn. Bird species composition in urban parks also depends on factors on a larger scale than the individual’s immediate habitat (see also Jokimäki and Huhta, 1996; Wiens et al., 1987; Wiens, 1989). Consideration of both landscape level and park characteristics are necessary for effective design and management of urban areas. Without sufficient knowledge of the importance of matrix structure, evaluation of bird–habitat relationships may be misleading. Acknowledgments I thank Dr. P. Helle, Dr. E. Huhta, Dr. M. Mönkkönen, Dr. J. Suhonen, Dr. J. Tiainen, and two anonymous referees for commenting on the manuscript and M.-L. Kaisanlahti for help in the field. L. Weber Müller-Wille revised the grammar of this paper. The study was financially supported by the Finnish Cultural Foundation. References Andrén, H. (1994) Effects of habitat fragmentation on birds and mammals in landscapes with different proportions of suitable habitat: A review. Oikos 71, 355–366. Baeyens, G. (1981) Magpie breeding success and Carrion Crow interference. Ardea 69, 125–139.
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