Landscape and Urban Planning 48 (2000) 131142

Biodiversity concepts and urban ecosystems

Jean-Pierre L. Savarda,*, Philippe Clergeaub, Gwenaelle Mennechezb
b a Canadian Wildlife Service, 1141 Route de l'Eglise, P.O. Box 10100, Sainte-Foy (Que.), Canada G1V 4H5 INRA FAUNE SAUVAGE and UMR CNRS Ecobio, Avenue du General Leclerc, 35042 Rennes cedex, France

Received 10 January 1998; accepted 20 September 1999

Abstract The association of biodiversity and urban ecosystems has usually concerned the impact of urbanization on biodiversity. However, biodiversity concepts can easily be applied to the urban ecosystem itself. As more and more people live in cities, restoration, preservation and enhancement of biodiversity in urban areas become important. Concepts related to biodiversity management such as scale, hierarchy, species identity, species values, fragmentation, global approaches can be used to manage urban biodiversity. Application of these concepts in such articial ecosystems may yield important insights for the management of natural ecosystems. Birds are highly visible and quite sensitive to changes in habitat structure and composition. Bird species richness in urban ecosystems is inuenced both by local and landscape characteristics and a multiscale approach is essential to its proper management. Peoplewildlife conicts are an integral component of wildlife management in urban ecosystems and must be addressed. Enhancement of biodiversity in urban ecosystems can have a positive impact on the quality of life and education of urban dwellers and thus facilitate the preservation of biodiversity in natural ecosystems. # 2000 Elsevier Science B.V. All rights reserved.
Keywords: Biodiversity concepts; Urban ecosystems; Birds in cities

1. Introduction In the last decade, biodiversity concerns have been in the forefront of conservation efforts worldwide (Environment Canada, 1994; UNEP, 1995). Biodiversity has been dened in various ways (McNeely et al., 1990; Salwasser, 1990) but the term has generally been used in a very comprehensive manner meaning the variability of life (composition, structure and function). Biodiversity can be represented as an interlocked hierarchy of elements on several levels of biological organization (Noss, 1992). Since the term
*

Corresponding author.

`biodiversity' transcends all levels of life from genes to communities and all spatial and temporal scales (Noss, 1990; Savard, 1994), it has generated a lot of confusion and misunderstanding (West, 1993; Lautenschlager, 1997). However, when understood and used properly biodiversity concepts can provide a useful framework for conservation efforts (Savard, 1994). Urban ecosystems have usually been examined in terms of their impact on biodiversity (Middleton, 1994; Wackernagel and Rees, 1996). However, while the impact of a city on adjacent ecosystems can be signicant (Douglas, 1983), much can be learned by applying biodiversity concepts to the urban ecosystem

0169-2046/00/$20.00 # 2000 Elsevier Science B.V. All rights reserved. PII: S 0 1 6 9 - 2 0 4 6 ( 0 0 ) 0 0 0 3 7 - 2

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itself. Enhancement of biodiversity in urban ecosystems can be quite important as some evidences suggest that personal exposure to natural things in everyday life is a major determinant of sensitivity to environmental issues (Sebba, 1991; Rohde and Kendle, 1994). As the proportion of urban residents increases every year worldwide, the nature of urban ecosystems would become increasingly important in shaping people's views about natural ecosystems. Also, from an ecological perspective, urban ecosystems are highly dynamic (Gilbert, 1989; Adams, 1994) and can provide useful insights into the management of biodiversity in other ecosystems. They are characterized by a high level of heterogeneity often organized along gradients extending from the surrounding landscape to the town centre (McDonnell and Pickett, 1990; McDonnell et al., 1993). Biodiversity concerns related to urban ecosystems can be divided into three major groups: (1) those related to the impact of the city itself on adjacent ecosystems; (2) those dealing with how to maximize biodiversity within the urban ecosystem and (3) those related to the management of undesirable species within the ecosystem. While species diversity and abundance are often related to the quality of urban life (Adams, 1994; Middleton, 1994) the overabundance of some species can be at times undesirable (Cooper, 1987; Clergeau et al., 1996). This paper deals with concerns related to the urban ecosystem itself using birds as a target group. Birds are quite sensitive to changes in habitat structure and composition and are therefore excellent indicators of changes and stresses in the urban ecosystem (Savard and Falls, 1982; Clergeau et al., 1998). We start by reviewing some important concepts related to biodiversity preservation. We then review the importance of local and landscape attributes in inuencing the abundance, diversity and distribution of birds in urban ecosystems and nally, we identify approaches and activities than can enhance urban bird diversity. 2. Important concepts related to biodiversity 2.1. Hierarchy of scales Life is structured in a hierarchical fashion starting with cells forming individuals, which regroup into

populations, which in turn form species, who end up as communities. Biodiversity concerns can occur at any level of organization. Levels of biological organization often correspond to specic spatial and temporal scales and must be addressed at their appropriate scale. Some concerns may focus on managing a population of geese in an urban park, others on increasing the number of bird species in the entire city. It is important when dealing with a concept as general as biodiversity to identify at which organizational level we want to act, or which group of organisms we target. In this case, we conne ourselves to birds but the same argument could be applied to plants, insects or mammals. In fact, biodiversity includes all forms of life. As biodiversity is expressed in a multitude of spatial scales, it is essential to specify the scale(s) of interest when using the term biodiversity (Savard, 1994). Failure to do so will quickly generate confusion and misunderstanding, since it is frequent to nd different and even contradictory results on different scales (Wiens, 1989). A multi-scale approach is thus necessary to properly address biodiversity questions. Scales are not independent from one another but linked in a hierarchical way (Allen and Starr, 1982) so the effects of an action at a given scale must be considered on higher and lower scales (Savard, 1994). Within urban ecosystems, actions taken to preserve or enhance biodiversity should be so at scales ranging from individual plants to the entire city itself and even its surrounding areas (Table 1). Temporal scales are also important to consider. Bird abundance and diversity in urban ecosystems vary with the seasons and even from year to year (Savard, 1978). The use of an urban park by birds varies according to the time of day causing bird diversity and abundance to uctuate throughout the day. When discussing and studying bird diversity it is also important to specify the type of diversity that interests us. For example, a park may have only two breeding species but be used for feeding by 10 species during the summer and ve during the winter. The hierarchical approach emphasizes logical and functional linkages among scales. Because biodiversity transcends several spatial and temporal scales as well as several organizational levels (from genes to communities), its management should be incorporated into a hierarchical decision system (Rice, 1992) as in some species management (Clergeau, 1995). In such a

Table 1 Various actions related to increasing bird diversity and abundance in urban ecosystem in relation to spatial scales Scale Adjacent landscapes Management level Regional Government or equivalent Planning Design Design parks and green corridors to optimize their use by birds; design roadways to minimize fragmentation of natural areas, Management Plant vegetation and restore habitats to improve parks and corridors; promote management options that favor bird diversity Reduce lighting of building at night during migration periods; manage waste to mini mize bird problems; plant vegetation in parks, green corridors and along streets. Enhance or restore vegetation in industrial and commercial areas; reduce use of herbicides and pesticides; plantation of shrubs, fruit trees, conifers; modify building structures to avoid bird problems. Install and maintain nest boxes and bird feeders; avoid use of pesticides and herbi cides; manage to reduce vulnerability of birds to cats; retain safe snags.

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City

Zone landscape use; identify and protect important natural areas for birds; identify green corridors linking the city to natural areas; create regional parks. Municipal Government Extend green corridors within the city; identify important areas for birds within the city; identify important bird species.

Local authorities City sectors (industrial, commercial, residential, recreational)

Individual lots

Owner

Shape, structure and size up corridors to optimize bird use; design park to increase bird abundance and diversity; design Building to reduce bird collisions. Establish vegetation objectives. Design type, structure and distribution Interact with higher levels to insure of vegetation to favor birds; insure optimal location of natural areas, parks building architecture compatible and corridors; create management with birds. plan for parks that will preserve and enhance bird diversity. Landscape property to attract birds: Select type of vegetation most coordinate with neighbors to compatible with birds; design maximize vegetation volume. nesting and feeding structures.

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system, higher levels constrain to various degrees the actions taken at lower levels. In urban areas for example, city regulations limit what individuals can or cannot do. It is imperative to adopt a hierarchical approach to properly manage biodiversity across scales. Such an approach inuences local actions whenever those actions have undesirable effects on larger scales. Constraints may include mandatory coordination across scales and levels. For example, a grassland park in a city may have only ve breeding bird species, three of which are specic to that park. Local bird diversity of the park could be doubled or tripled by converting this grassland park into a wooded one. However, this would result in the loss of three species for the city as a whole. So, in this case, an increase in local diversity would result in a decrease in diversity at the scale of the city. Obviously, a choice has to be made here, as to which scale we want to give priority to. It doesn't necessarily have to be the greater scale as long as efforts are coordinated across the scales. Since urbanized areas already function within a hierarchical decision system, it should be relatively easy to extend to biodiversity management. In the example described here, there could be a general policy on the scale of the city to preserve and enhance the unique features of some parks. Once those features are identied and protected, a general goal of increased species richness can then be applied to all parks. Food webs and trophic relationships illustrate well the hierarchical organization of life and stress an other important point to consider when managing biodiversity: all species are not equal. 2.2. Species One important aspect in dealing with biodiversity is that not all species are equal. Species vary in size, shape, abundance, distribution, trophic position, ecological function, feeding habits and desirability. Therefore, diversity indices which assume all species as being equal are of little use for properly managing biodiversity (Savard, 1994). Some species may play important roles in the community, so their absence would signicantly affect several other species. Identication of these species is difcult since their role may be seasonal or even habitat specic (Bond, 1993). It is important to identify desirable species. For exam-

ple, in the context of an objective to increase bird people interactions, bird abundance would be more important than bird diversity. Conspicuous species would be preferred to furtive ones, as would species yielding positive interactions. Other species-related concepts used in biodiversity conservation are the concept of umbrella species and agship species (Hunter, 1990). Umbrella species are at the top of food chains or with large home ranges so that by protecting these species we protect all species on which they depend or the species with similar requirements but smaller home ranges. For example, Merlin (Falco columbarius) could be considered as an umbrella species in several North American cities as it feeds on songbirds and nests in tall trees. An abundance of small birds is therefore essential for the well being of Merlin in urban areas. Because House Sparrow (Passer domesticus) and European Starling (Sturnus vulgaris) often represent over 70% of the birds in urban areas, they are crucial as a prey base for the maintenance of some birds of prey in urban ecosystems. This concept is like the coarse lter approach mentioned further. Flagship species are charismatic species which attract attention and which can be used to galvanize public support for conservation efforts. A good example of a charismatic species is the Peregrine Falcon (F. peregrinus) whose nesting in cities has attracted considerable attention and greatly facilitated conservation efforts (Cade and Bird, 1990). An other example is the White Stork (Ciconia ciconia) in Europe which nest on chimneys (Cramp et al., 1977). Birds can also be excellent indicators of various aspects of urban ecosystems (biodiversity; vegetation structure; contaminants). In this respect, Rock Dove (Columbea livia) has been used successfully to monitor heavy metal contamination along urbanization gradients (Drasch et al., 1987). In urban ecosystems bird species also vary in terms of how they are perceived by people (Brown et al., 1979; Penland, 1987). Some species such as European Starlings, House Sparrow and Rock Doves are often appreciated at low density but are sometimes perceived as a nuisance at high densities. Peoplewildlife interactions are crucial in urban ecosystems (Brown et al., 1979) and any enhancement of biodiversity must take them into consideration (Michelson, 1970; Gilbert, 1989; Clergeau et al., 1997).

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2.3. Habitat fragmentation and habitat quality Habitat fragmentation can be extreme within urban ecosystems. Its effect occurs at all spatial scales and affects all organisms. Fragments of natural vegetation may be too small or even too isolated to support some species. In a park, shrubs abundance and distribution will inuence the presence and abundance of bird species nesting in shrubs whereas park distribution, size and abundance in a city will affect bird diversity. Vegetation corridors linking urban green areas between themselves and/or with rural habitats are important to maintain and enhance urban biodiversity (Flink and Searns, 1993). They facilitate movements and ensure colonization of isolated natural areas (Clergeau, 1998). Streams form natural corridors in urban areas and should be managed as such. In Toronto, natural ravines within the city have been preserved and act as corridors between parks. Wellvegetated residential areas constitute aerial corridors through their tree canopy. Such corridors are especially useful for migrating birds which use them extensively as they provide food and protection against aerial predators (Savard, 1978). Greenways act not only as movement corridors (Clergeau and Burel, 1997) but can also provide breeding habitats for several edge species (Noss, 1993). Like for most ecosystems, our knowledge of urban biodiversity and its distribution is limited especially to small organisms. In view of that lack of knowledge a coarse lter approach is indicated where we seek to protect natural habitats within cities and by doing so protect most of the species associated with that habitat. For example, an increase in the volume and diversity of vegetation in a city would increase bird abundance and diversity and such an approach can be applied to all spatial scales (Table 1). The coarse grain approach does not protect all species and needs to be complemented by a ner approach dealing with specic species (Hunter, 1990). This is most crucial in urban ecosystems where habitats are most fragmented and isolated. Protected fragments may lack some key resources needed by species we would like to retain (i.e. nest site, shelter, food, etc.). These concepts are closely related to biodiversity conservation (Savard, 1994) and can easily be applied to urban ecosystems. Biodiversity concepts are quite general and must be rened at manageable levels and

scales. Statements like `our goal is to enhance urban biodiversity' are not workable because of the encompassing nature of the word biodiversity. Something like `our goal is to increase bird diversity in urban parks' is more appropriate as it species the group of organisms, as well as the scale of action. It is essential to elaborate achievable goals. It is also important not to confound biodiversity and species diversity as they are quite different concepts. Species diversity is only a small portion of biodiversity and deals mostly with species richness, sometimes incorporating abundance as well. Biodiversity includes all forms of life, as well as structural and functional aspects. 3. Importance of local and landscape attributes for bird abundance and biodiversity Three species have adapted particularly well to urban ecosystems and have colonized them worldwide: the House Sparrow, the European Starling and the Rock Dove. These species dominate the urban avifauna of most cities and are usually quite dominant in the most urbanized part of a city (Table 2). Although often neglected by ornithologists, these species constitute the core of urban avifauna and their role is quite crucial, especially in the more densely populated areas. Species richness tends to be higher in the more vegetated areas and lowest in the downtown areas, and this throughout all studies (Table 2). New residential developments with little or no original vegetation have a low diversity of birds, but depending on the architectural structure of their buildings, can sometimes support high densities of birds. Bird diversity typically increases as vegetation develops (Hohtola, 1978; Luniak, 1994). Urban woodlands are an important component of the urban landscape in terms of bird species diversity. The larger the woodland, the more species it supports (Sasvari, 1984; Tilghman, 1987a). For similar size woodlots, spatial heterogeneity, complex vertical structure and diverse species composition of vegetation are all associated with higher bird species richness (Cavareski, 1976; Tilghman, 1987b). Composition of the native component of urban bird communities varies according to the geographical location of the city (Table 2), its landscape context and the amount of natural vegetation contained in the city.

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Table 2 Proportion of introduced species (%) in breeding bird communities of urban areas in relation to level of urbanizationa Location Downtown area Residential area Vegetation rare Vegetation abundant 9297 (5) [710] 73 (1) [19] 76 (1) [10] 97 (1) [6] 95 (1) [] 8085 (4) [916] 41 (1) [14] 7282 (6) [715] 2752 (3) [2325] 2643 (3) [2123] 57 (1) [9] 5257 (2) [1417] 60 (1) [] 24 (1) [27] 1234 (3) [1119] Vegetation very abundant 73 (1) [12] 34 (1) [31] 25 (1) [26] 23 (1) [16] 3139 (3) [1624] 0 (1) [25] 848 (4) [818] Semi-natural area Reference

Toronto, Canada Quebec, Canada Rennes, France Vancouver, Canada Blacksburg, Virginia Washington DC Warsarw, Poland Tornio, Finland Kuopio, Finland
a

9498 (2)b [55]c 68 (1) [13] 52 (1) [10] 6679 (3) [35]

4354 (2) [2122] 48 (1) [27] 34 (1) [30] 34 (1) [14] 9 (1) [21] 38 (1) [] 451 (l) [23] 033 (4) [614]

Savard (1978) Clergeau et al. (1998) Clergeau et al. (1998) Weber (1972) Lucid (1974) Williamson (1974) Luniak (1994) Huhtalo and Jarvinen (1977) Hohtola (1978)

Introduced species include European Starling, House Sparrow and Rock Dove. Although not introduced in European cities they have been regrouped there for comparison purposes. b Range of % of introduced species; (number of areas sampled). c [Number of species (range)].

Urban ecosystems are quite similar worldwide in terms of structure, functions and constraints. They differ in terms of their geographical location, their size and the type of landscape they modify. The landscape surrounding a city greatly inuences plant and wildlife species that will be found within the new articial ecosystem. It is essential to consider landscape factors in the management of urban biodiversity. For example, a high breeding density of European Starlings in a residential area can be due in part to the presence of extensive lawn surface in neighboring

areas or parks where birds feed. Similarly, the presence of large numbers of gulls in a sector can be due to the presence of nearby open water or of a breeding colony in surrounding areas. The reproductive success of birds often varies with the type of habitat selected. We found such a pattern in the reproductive parameters of European Starlings that varied from low in the downtown area to high in periurban sectors (Table 3). Thus, some sectors of the city may act as a source of starlings while others may not sustain viable populations. Such variation in repro-

Table 3 Reproductive parameters of European Starling in Rennes in relation to the urbanization gradient Urban center Laying date Clutch size Number of eggs hatching 6 April1 day (n7) 5.00.0 (n6) 2.82.0 (n6) Peripheric areas 4 April2 days (n14) 5.60.9 (n14) 5.00.8 (n15) Countryside 1st April2 days (n12) 6.00.5 (n10) 5.70.5 (n10)

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ductive success between urban habitats has been reported for several species (Nuorteva, 1971; Smith and Gilbert, 1984). The presence of urban populations of starlings may play a role in attracting large roosting populations of rural starlings. Understanding of urbanrural movements and their cause is essential to properly manage these populations (Clergeau, 1981). The impact of urbanization on a given landscape is partially a function of the original composition of the landscape. Urbanization will increase bird diversity of simple landscapes such as desertic areas or grassland areas by creating new habitats and supporting exotic species of birds (Emlen, 1974). In diversied landscapes that support an initial high diversity of habitats, a decrease in bird diversity is likely to occur as urbanization becomes more prevalent, eliminating unique natural habitats (Batten, 1972). Landscape factors such as the composition and arrangement of habitat types around a local area have signicant impacts on bird populations of that local area and must be taken into account when managing the local area (i.e. a park, a residential area, etc.). Landscape context is especially important in complex landscapes such as found in urban ecosystems (Dunning et al., 1992). 4. Approaches to enhance urban biodiversity There is no single best starting point when dealing with biodiversity. Local actions as well as regional actions are equally important on their respective scale. In urban areas home-owners can take various actions on the scale of their lot. It is important to realize that it is the concerted efforts at various scales that produce the best results. It is essential that home-owners realise that their own local action can contribute to a larger collective effort that would culminate in the creation of a real biological corridor that facilitates the movements of several species throughout the city. Such collective efforts proved quite powerful in Belgium (Laurence and Palmaerts, 1991). Urban planners can act on a larger scale, either in neighborhoods or in the entire city. Urban growth must now take into consideration the creation of large recreational zones which often, upon request of citizens, must remain as natural as possible. These zones must integrate

social and ecological considerations to avoid conicts. The overall thinking about green spaces, corridors, wildlife and people is common to all cities but solutions vary greatly depending on local, physical, social and ecological constraints. Whatever the scale of intervention, one important aspect is to evaluate the impact of a given action on smaller and larger scales in order to avoid unwanted results. A rst approach for urban planners would be to inventory resources within the city (natural area, lake, riverF F F) and their organization (unique structures, corridors) and to assess the abundance, distribution and possible concerns about these resources. Once resources have been identied on the scale of the city itself, strategies can be established for preserving and using them whenever necessary. Such strategies should account for local constraints and assets. Local actions will develop around those constraints or opportunities and address specic objectives. For example, trees may be planted in a residential area to increase vegetation volume and thus increase insect and bird diversity. In some countries where the weather is harsh, bird feeders may be encouraged or even maintained in parks to enhance local diversity. Similarly, nest boxes may be erected for particular species, etc. Another important aspect to consider is the landscape context. This is important, as the maintenance of some resources within the city may depend on what is happening outside the city. For example, a stream originating outside the city will be affected by neighboring areas, or an urban park, by neighboring residential areas. Table 4 lists a multi-scale approach that reects the considerations given earlier. Some activities to enhance urban bird diversity are worth mentioning here: 4.1. Plantation of trees and shrubs Bird diversity in urban areas tends to be proportional to the existing volume of vegetation (Emlen, 1974; Lancaster and Rees, 1979). During migration, numbers and diversity of migrating birds within the city are also proportional to vegetation abundance (Savard, 1978). Tree species selection is also important. Conifer trees provide nesting cover for several species of birds and are heavily used for cover in winter (Savard, 1978). Fruit trees attract frugivorous birds, especially in winter. For example, in some

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Table 4 A multi-scale approach: to enhance bird abundance and diversity in the city we may want to 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. Examine the country side surrounding the city and secure or restore important bird habitats that may act as a source of birds for the city; establish a greenbelt around the city. Identify and consolidate vegetation corridors linking these areas to the city and link parks whenever possible; make use of natural streams and right of ways. Increase the volume and diversity of vegetation in the city (along streets, right of ways, industrial, commercial and residential sectors). Exploit and enhance the features of some parks that may attract particular species of birds. Increase the structural diversity of vegetation in natural and recreational parks of the city. Plant conifers and fruit trees to provide cover and food for birds; promote the night blackout of tall buildings during bird migration and encourage architecture that minimizes bird collisions. Erect special nesting structure for cavity or cliff nesting species (nest boxes for owls, chimneys for swifts, nesting platforms for falcons or storks). Encourage homeowners to manage their property for birds, to restrain their pets and to minimize the use of pesticides and herbicides. Distribute guidelines as to the proper planting of trees and shrubs to attract birds. Maintain bird feeders, nest boxes, bird baths in backyards.

Canadian cities, plantation of Mountain Ash (Sorbus americana) along city streets have attracted large numbers of wintering Bohemian Waxwings (Bombycilla garrulus) which in turn provided food for wintering Merlin which increased in numbers within the city (Oliphant and Haug, 1985). Several bird species nest and forage in shrubs so that addition of shrub thickets to an area would increase bird species diversity. 4.2. Provision of articial nesting structures Several bird species frequenting urban ecosystems are cavity or cliff nesters and are often limited by availability of nesting structures. Reproductive success and breeding densities of several of these species are sometimes greater in urban ecosystems than in their natural habitats (Smith and Gilbert, 1984). Articial chimneys have been used to attract nesting swifts (Apus sp.) and Peregrine Falcons have successfully nested on articial structures in cities (Cade and Bird, 1990). Architectural design of buildings can provide nesting sites for House Sparrow, Starlings and Rock Dove, which is not always a desirable thing. Careful design of buildings may avoid costly problems. 4.3. Provision of bird feeders Bird feeders are quite efcient in increasing local bird diversity especially in winter. Species' attraction depends on the type of food found in feeders, their location and the type of surrounding habitat. However, feeding of birds may attract undesirable species or

concentrate too many birds into a small area. Whereas feeding of small seed eating birds is generally a positive activity, feeding of large birds in parks or other urban areas often results in negative impacts on the habitat and/or generates peoplewildlife conicts. 4.4. Regulating human behavior Urban problems related to birds are usually due in part to human behavior. For example, improper storage of human waste may attract large numbers of scavenging birds. Architectural features of buildings often encourage nesting of undesirable species. Free roaming by cats and dogs is an important source of disturbance and predation on birds. Some Canadian cities have regulations restricting free roaming of cats and dogs which greatly reduces their impact on wildlife. 4.5. Creation, restoration and management of natural areas Parks or other green spaces within urban areas can act as a source of birds for neighboring residential areas. Bird diversity observed in backyards would likely be higher in properties adjacent to a natural area. Size and shape of green areas are important in determining the number of species they attract. These patches of natural habitats can greatly enhance local bird diversity (Cavareski, 1976; Tilghman, 1987a, b). Riparian areas along streams provide important corridors for wildlife in urban areas. Preservation of the

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vegetative integrity of these natural greenways can signicantly enhance urban biodiversity. Degraded urban habitats can be restored and contribute to local biodiversity, as well as to the development of restoration techniques for natural ecosystems (Berger, 1988). 5. Management of peoplewildlife conicts Urban ecosystems are highly dynamic and in constant evolution. Like in natural ecosystems, wildlife communities uctuate and evolve. New species invade urban areas (Morneau et al., 1999) and some species explode in abundance creating conicts with people (Cooper, 1987). These conicts need to be properly addressed and they pose quite a challenge as they are often caused by inadequate human behavior. Furthermore, since the dynamics of urban ecosystems are still poorly documented, management actions often have unexpected results. Pigeons, gulls and starlings sometimes are a nuisance in towns. Several American cities also have serious problems with urban geese that have exploded in numbers within urban parks creating serious habitat degradation problems (Nelson and Oetting, 1981; Conover, 1987; Breault and McKelvey, 1991). Like urban biodiversity, management of urban bird problems requires a multi-scale approach as well as a sociological component. Unfortunately little research has been done so far on human perception and appreciation of urban wildlife (Brown et al., 1979; Lemoine and Sauvage, 1996). 6. Conclusions Principles used for managing or enhancing biodiversity can be applied to urban ecosystems. Because of the highly dynamic nature of urban ecosystems a small effort in management can have a great effect on bird abundance and diversity. As we have shown, actions are required on several spatial scales simultaneously and local as well as landscape features are important to consider. Although we limited our paper to birds and emphasized species richness, which is only a small component of biodiversity. Urban ecosystems can be of high value to a variety of other organisms or to several other aspects of biodiversity (population structure, genetic diversity). For example, rare plants or

rare forms of plants can be cultivated in backyards, maintaining a source of genetic variability; owers that attract butteries can be grown in parks and even private lots so that buttery diversity increases; creation of ponds and wetlands in urban parks attracts a variety of aquatic organisms, etc. Whatever the group of organisms considered, biodiversity concepts apply. Enhancement of urban biodiversity can also have direct economic impacts as residential properties adjacent to greenways or urban parks have a higher market value than similar non-adjacent properties (Hammer et al., 1974; King et al., 1991, see Luttik this issue). It is essential to incorporate a sociological component to biodiversity management in urban areas. No other ecosystems support such high densities of people so that desires and perceptions of urban residents must form an integral part of biodiversity management in urban ecosystems. These aspects need to be better quantied. Research priorities include a greater understanding of peoplewildlife interactions. Such research will require a joint approach by sociologists and ecologists and is crucial in the quest for sustainable urban biodiversity. Finally, another important research priority is a greater understanding of the importance of landscape features in local areas and of the dynamics of exchanges between urban and rural areas. The ecology and population dynamics of urban birds need also to be better quantied. Enhancement of biodiversity in urban ecosystems, if well done, can have a signicant and positive impact on the quality of life and education of the increasingly growing urban population and thus, indirectly facilitate the preservation of biodiversity in natural ecosystems. Acknowledgements We would like to thank G. Falardeau for his assistance. This work was supported by the Canadian Wildlife Service and the `Centre national de recherche scientique, Universite de Rennes I'.

References
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Savard, J.-P.L., 1994. General concepts related to biodiversity. Biodiversity in Canada: A Science Assessment for Environment Canada. Environment Canada, Ottawa, pp. 940. Savard, J.-P.L., Falls, B.J., 1982. Inuence of habitat structure on the nesting height of birds in urban areas. Can. J. Zool. 59, 924 932. Sebba, R., 1991. The landscapes of childhood: the reection childhood's environment in adult memories and in children's attitudes. Environ. Behav. 23, 395422. Smith, D.G., Gilbert, R., 1984. Eastern Screech-Owl home range and use of suburban habitats in southern Connecticut. J. Field Ornith. 55, 322329. Tilghman, N.G., 1987a. Characteristics of urban woodland affecting breeding bird diversity and abundance. For. Ecol. Manage. 21, 163175. Tilghman, N.G., 1987b. Characteristics of urban woodlands affecting breeding bird diversity and abundance. Landscape and Urban Planning 14, 481495. UNEP, 1995. Global Biodiversity Assessment. Cambridge University Press, Cambridge, 1140 pp. Wackernagel, M., Rees, W., 1996. Our Ecological Footprint: Reducing Human Impact on the Earth. The New Catalyst Bioregional Series, New Society Publishers, 160 pp. Weber, W.C., 1972. Birds in cities: a study of populations, foraging ecology and nest-sites of urban birds. M.Sc. Thesis, University of British Columbia, Vancouver, 269 pp. West, N.E., 1993. Biodiversity of rangelands. J. Range Manage. 46, 213. Wiens, J.A., 1989. Spatial scaling in ecology. Funct. Ecol. 3, 385 397. Williamson, R.D., 1974. Birds in Washington DC. In: Noyes, J.H., Progulske, D.R. (Eds.), Wildlife in an Urbanizing Environment. Proceedings of a Symposium, Springeld, Mass. Cooperative Extension Serv., University of Massachusetts Press, Amherst, 79 November 1973. Jean-Pierre L. Savard is a Research Scientist for the Canadian Wildlife Service (CWS) in Quebec. He holds a bachelor's degree from Universite Laval, Quebec, an M.Sc. from the University of Toronto, Ontario, and a Ph.D. from the University of British Columbia. He spent 14 years with the CWS in British Columbia, where he studied the distribution and ecology of molting and wintering seaducks (scoters, goldeneyes, Harlequin Duck), the breeding ecology of Barrow's Goldeneye and the Eared Grebe, oldgrowth forest birds, and seabirds (Marbled Murrelet). His current research interests in Quebec include urban wildlife ecology, impact of forest practices on birds, bird survey techniques, seaduck ecology (breeding, staging, moulting and wintering) and biodiver sity concepts. He has adjunct status at several Quebec universities where he co-supervises graduate students. Philippe Clergeau is a wildlife biologist at `Institut National de Recherche Agronomique' in France. He had received his Ph.D. and his `Habilitation a Diriger des recherches' from Rennes University. He teaches wildlife management and biodiversity control in agronomic schools and universities, and he collaborates with a `Centre National de la Recherche Scientifique' team (UMR

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J.-P.L. Savard et al. / Landscape and Urban Planning 48 (2000) 131142 Gwenaelle Mennechez has just received her Ph.D. from Rennes University. She has studied behavior and fitness of European starlings in different landscapes (gradient of urbanisation). She has participated in research projects on biodiversity and avifauna in urban areas.

EcoBio) on landscape ecology. His research activities focus on the pest bird biology and management, and on biological relations between rural and urban areas. He is project leader for several Publications on relations between humans and animals in towns.