Environmental Pollution 159 (2011) 1974e1983

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Environmental Pollution
journal homepage: www.elsevier.com/locate/envpol

Review

Novel urban ecosystems, biodiversity, and conservation

Ingo Kowarik
Department of Ecology, Technische Universität Berlin, Rothenburgstr. 12, D 12165 Berlin, Germany

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

Article history: With increasing urbanization the importance of cities for biodiversity conservation grows. This
Received 22 December 2010 paper reviews the ways in which biodiversity is affected by urbanization and discusses the conse-
Received in revised form quences of different conservation approaches. Cities can be richer in plant species, including in
11 February 2011
native species, than rural areas. Alien species can lead to both homogenization and differentiation
Accepted 13 February 2011
among urban regions. Urban habitats can harbor self-sustaining populations of rare and endangered
native species, but cannot replace the complete functionality of (semi-)natural remnants.
Keywords:
While many conservation approaches tend to focus on such relict habitats and native species in
Biodiversity loss
Biological invasions
urban settings, this paper argues for a paradigm shift towards considering the whole range of urban
Exotic species ecosystems. Although conservation attitudes may be challenged by the novelty of some urban
Nature conservation ecosystems, which are often linked to high numbers of nonnative species, it is promising to consider
Ecosystem services their associated ecosystem services, social benefits, and possible contribution to biodiversity
conservation.
Ó 2011 Elsevier Ltd. All rights reserved.

1. Cities and biodiversityda major challenge
Biodiversity conservation emerged as a field of international
policies in the 2nd half of the 20th century, culminating in the
Convention on Biological Diversity (CBD), which entered into force
in 1993. Although urban areas globally cover only about 3% of the
earth’s land surface (MEA, 2005), urban growth imposes major
challenges to biodiversity conservation as it has already induced
a profound transformation at the landscape level and is regarded as
a major threat to biodiversity (Antrop, 2004; Hansen et al., 2005).
Cities are not dispersed randomly at the global scale, but are often
located in biodiversity hotspots. As the human population grows
extraordinarily, especially in developing countries (MEA, 2005;
United Nations, 2008), so do conflicts with conservation aims
(Cincotta et al., 2000; Araújo, 2003; Liu et al., 2003; Kühn et al.,
2004; Zhao et al., 2006).
While urban growth accelerates in many regions, stimulating
urban sprawl and leading to the formation of “megacities” (Kraas,
2008), divergent trends have been observed in regions of
economic decline. Here, the phenomenon of shrinking cities is
associated with an enhanced emergence of “wild” ecosystems in
urban-industrial areas which well illustrate the ecological potential
of urban regions (Kowarik and Körner, 2005; Langner and
Endlicher, 2007).
Urban ecology has a long history, mainly in Europe, and has
developed rapidly over the last few decades as illustrated by
E-mail address: kowarik@tu-berlin.de.
Sukopp’s (2002) review and the papers in Marzluff et al. (2008).
There is an increasing body of evidence that urban land uses effect
profound changes in all environmental components and that
humans are the main drivers of change (Sukopp et al., 1979; Gilbert,
1989; Pickett et al., 2001; Alberti et al., 2003), thus leading to the
idea of addressing cities as “socioecosystems” (Grimm et al., 2008).
Yet, a better understanding of urban biodiversity patterns requires
insights into the functioning and interaction of environmental and
social drivers.
Urban growth changes landscapes and related biodiversity
patterns at the city level, the wildland (rural)eurban interface and,
because of the ecological footprint of cities, even at regional to
global scales (Grimm et al., 2008). Understanding, assessing, and
enhancing urban biodiversity is of paramount importance, from
both conservation and social perspectives. With urbanization as
a major global trend, the question of whether, and to what extent,
species of animals and plants can survive in urban settings becomes
increasingly vital. Conserving and enhancing urban biodiversity
also has unique implications for human well-being, public health,
and for making citizens aware of the importance of biodiversity
conservation as the majority of people globally will experience
“nature” and related ecosystem services primarily within the urban
fabric (Gilbert, 1989; McKinney, 2002; Miller and Hobbs, 2002;
Miller, 2005; Goddard et al., 2010).
As many convincing arguments already exist regarding the need
to prevent biodiversity losses in (semi-)natural landscapes that are
affected by urban growth (e.g., Hansen et al., 2005), the focus is
here on biodiversity patterns and related conservation approaches
within cities. This paper first reviews evidence on how biodiversity

0269-7491/$ e see front matter Ó 2011 Elsevier Ltd. All rights reserved.
doi:10.1016/j.envpol.2011.02.022
 I. Kowarik / Environmental Pollution 159 (2011) 1974e1983 1975

responds to urbanization in space and time and to what extent
patterns of urban biodiversity differ from those of nonurban areas.
Then, some conceptual ideas are addressed that may be fruitful for
stimulating a diversification of urban conservation approaches,
covering the whole array of urban nature components.
2. Biotic responses to urbanization
A wealth of urban studies addresses biotic responses to urban-
ization, although care must be taken with generalizing their results.
Most studies are from Europe and North America, while other
regions are still underrepresented. Biodiversity patterns have been
addressed at different scales and with a variety of approaches to
differentiate urban from nonurban systems. This hampers
comparisons between different studies, but some general insights
into the changes in urban species composition and underlying
mechanisms are nevertheless feasible.
Humans govern urban biodiversity directly by habitat loss,
habitat fragmentation and the introduction of new species, and
indirectly by changing urban climate, soils, hydrology, and
biogeochemical cycles (Table 1). Beyond physical alterations to the
urban environment, socioeconomic activities directly affect biodi-
versity patterns by setting filters for species selection and dispersal
(Table 2). Both types of mechanisms interact, affect the overall
species richness in urban settings and lead to compositional
changes in urban biota.
2.1. Species richness
Variation in species richness and composition has been abun-
dantly analyzed along gradients from the urban center to the rural

Table 1
Habitat-related parameters of cities, biodiversity responses, and proposed underlying mechanisms illustrated with examples.

Driver of change Biodiversity response Underlying mechanisms and examples

Habitat Increase in overall (plant)
heterogeneity species richness
Habitat Negative impacts on species
fragmentation that rely on large habitats,
decline in species with low
dispersal ability, increase in
species with long distance
dispersal ability
Invasion of isolated natural
remnants by introduced
species
Short-term evolutionary
adaptation
Loss and degradation Loss or decline in native
of (semi-)natural species with a high habitat
habitats specialization
Emergence of novel Habitat shift of common
urban habitats generalist native species
from semi-natural to urban
sites
Habitat shift in rare
native species
Increase in introduced
species
Elevated levels of Changes in species
nutrients, pollutants, composition and
and soil pH; reduced preference for selected
availability of water species traits
Urban heat island effects Increase in species which
are sensitive to deep
frost and tolerate heat
and drought stress; at
the species scale, changes
in phenology, arrival time
of migrant birds
Habitat disturbance Increase in short-lived
species and in introduced
species
Reduced establishment
of self-sustaining
populations
Evolutionary processes
High numbers of small patches with strongly contrasting environmental conditions can be used by an
array of native and introduced plant species with starkly different ecological demands
(Sukopp et al., 1979; Gilbert, 1989; Pysek, 1993; among many others).
Larger habitats support larger populations of birds and other animal groups (Evans et al., 2009; cf.
Goddard et al., 2010). Less mobile species such as carabid beetles incapable of flight decrease in urban
habitats (Niemelä and Kotze, 2009). The ability to move in the urban matrix can, however, counteract
effects of habitat fragmentation (Goddard et al., 2010). Bee species rely more on quality than
connectedness of habitats (Kearns and Oliveras, 2009), and habitat quality of trees for arthropods
depends more on origin and size of the trees than on tree connectedness (Yasuda and Koike, 2009).
Urban floras have more species that are moved by humans, animals, or water (Knapp et al., 2008b).
Human-mediated dispersal can enhance nonnative species, which are, e.g., more often moved by traffic
over long distances than are native species (von der Lippe and Kowarik, 2007).
Urban forest remnants are subject to enhanced plant invasions due to increased edge effects and
propagule pressure from surrounding urban matrix (e.g., Cadenasso and Pickett, 2001; Borgmann and
Rodewald, 2005; Duguay et al., 2007)
Rapid evolution in the herb Crepis sancta towards lower dispersal types in highly fragmented urban
habitats to prevent high costs of dispersal has been demonstrated (Cheptou et al., 2008).
Decline of less common or rare species that are dependent on wet and/or nutrient-poor habitats, such as
fens, bogs, nutrient-poor grasslands due to changes in hydrology and N-enrichment in urban areas
(Preston, 2000; DeCandido, 2004; Knapp et al., 2010). Also, species confined to areas of traditional
agricultural practice decline, including natives and many archaeophytes (Chocholousková and Pysek, 2003;
Knapp et al., 2010).
A large subset of native plant species can colonize urban habitats even when subject to high disturbance
levels (about two-thirds in Berlin; Kowarik, 1990), mostly species with a broad ecological niche such as
the grass Calamagostis epigejos (Rebele and Lehmann, 2001).
Some rare species colonize novel urban-industrial sites as these mimic environmental conditions of
natural habitats (Eversham et al., 1996), e.g., plant species from highly disturbed floodplains on gravel
on industrial sites (Rebele and Dettmar, 1996) or grassland species on old green roofs
(Dvorak and Volder, 2010).
Urban sites with profoundly changed soils, climate, and hydrology often show an increased proportion
of introduced species pre-adapted to the novel site conditions that mimic those of their original areas
(Kowarik, 1995; Pysek, 1998; Pickett et al., 2001; Sukopp and Wurzel, 2003).
Species, often nonnatives, preferring nitrogen-rich, warm and dry habitats are overrepresented in cities
(Wittig and Durwen, 1982; Knapp et al., 2009). Urban floras show increased proportions of plants with
scleromorphic or succulent leaves as a response to drought (Knapp et al., 2008b). Increased pH values due to
emissions of a cement plant enhanced seedling establishment in the shrub Mahonia aquifolium (Auge, 1997).
Climate warming enhances plant invasions (Nobis et al., 2009; Walther et al., 2009), and urban heat
island effects may anticipate associated effects. Climate chamber experiments revealed an increased
growth in the tree Ailanthus altissima as a response to increased temperatures (Kowarik and Säumel,
2007). Several studies illustrate an expansion of the growing season and phenological changes in urban
areas, e.g. in bud burst or flowering (White et al., 2002; Lu et al., 2006; Neil et al., 2010). Some migrant
bird species arrive earlier in spring (Biadun et al., 2009).
Habitat disturbance enhances annual and biennial species in urban regions (Chocholousková and Pysek,
2003; Knapp et al., 2008b). With increasing levels of human-mediated habitat disturbance, the number
of introduced species usually increases (Kowarik, 1995).
Previously abundant species tend to become rarer in urban settings (Chocholousková and Pysek, 2003).
With increasing habitat transformation and disturbance, the number of species with self-sustaining
populations decreases (Kowarik and von der Lippe unpubl.)
Evolution of novel Oenothera species from introduced North American ornamentals in Europe on urban
sites and along traffic corridors (Cleland, 1972). Evolution of plant ecotypes with heavy metal tolerance
on industrial sites (Ernst and Joosse-van Damme, 1983). Adaptation of birdsong to urban noise
(Mockford and Marshall, 2009).
1976 I. Kowarik / Environmental Pollution 159 (2011) 1974e1983

Table 2
Examples of socioeconomic factors and related biodiversity responses in urban regions, and proposed underlying mechanisms with examples.

Factor Biodiversity response Underlying mechanisms and examples

Income Increased number Wealthy neighborhoods can be associated with more intense landscaping and planting of exotic species
of planted and escaped (“luxury effect,” Hope et al., 2003). Newly naturalized plants were found more frequently in richer suburbs
introduced species (Sullivan et al., 2004). At the same time, vacant lots, often associated with less wealthy neighborhoods,
also appear to enhance the presence of alien species (Gulezian and Nyberg, 2010).
Availability of Escape from cultivation Species that have been available for longer periods on the market are more likely to have escaped from
ornamentals cultivation, likely due to a higher propagule pressure (Dehnen-Schmutz et al., 2007). The consistent supply
on the market of semi-hardy plants in garden centers provides seed sources for species that may escape subsequent to
climate warming (Niinemets and Peñuelas, 2008).
Species selection Escape from cultivation, The plant diversity in neighborhoods depends on socioeconomic and cultural characteristics
for landscaping changes in species and (Kinzig et al., 2005). The similarity of adjacent front gardens suggests a kind of social control (neighbor
plant trait composition ‘mimicry’ effect, Zmyslony and Gagnon, 1998). The number of planted individuals drives the propagule
pressure and by this the probability of naturalization (Krivanek et al., 2006; Lockwood et al., 2005). High
numbers of planted nonnative woody ornamentals lead to increased proportions of shrubs and trees in
urban floras (Sullivan et al., 2004; Knapp et al., 2009; Zhao et al., 2010). As species with shiny fruits are
often preferred by gardeners, animal-dispersed species are overrepresented in urban regions (Knapp
et al., 2009, 2010).
Management of Regeneration of ornamentals Management practices reduce threats to the survival of planted species, and thereby enhance their
urban green and side-effects for spontaneously opportunities to escape from cultivation (Mack, 2000; Kowarik, 2005b). Continued maintenance over
spaces growing species long periods may foster the persistence of ornamentals in historical green spaces over periods of more
than a century, thereby providing evidence of former gardening styles (Bakker and Boeve, 1985).
Recent gardening styles were found to explain the occurrence of native species and archaeophytes
(Brunzel et al., 2009).
Feeding practices Changes in bird species Bird feeding in urban gardens affects assemblages of urban birds (Fuller et al., 2008)
composition
Human mobility Preferential dispersal of Household mobility in urbanized settlements explains the increase in alien species (neophytes) over time
selected species (groups) better than habitat-related factors (Niggemann et al., 2009). Vehicles move seeds of nonnative species over
longer distances than seeds of native species (von der Lippe and Kowarik, 2007). Humans disperse species
attached to shoes and clothes (Mount and Pickering, 2009; Wichmann et al., 2009).

hinterlands of cities (McDonnell and Hahs, 2008). The observed
patterns differ conspicuously across different groups of taxa. Birds,
butterflies, carabid beetles, lichens, and mosses usually increase in
species richness along urban-to-rural gradients (e.g., Seaward,
1982; Blair, 1999; Marzluff, 2001; Niemelä et al., 2002; Clergeau
et al., 2006). The comparative study by Niemelä and Kotze (2009)
illustrates some general trends. Most of the eight cities they
studied around the world had more carabid beetles in rural
compared to urban and suburban areas; this can by explained, at
least partly, by habitat preferences, species traits, and the history of
species introductions. Forest specialist species and species inca-
pable of flight tend to be more common in suburban and rural
zones, while specialists of open habitats and species capable of
flight predominate in the urban core. However, deviations from
these general patters occur in some cities, showing peaks of species
richness in suburban plots or missing differences between urban
and rural sample sites. Generally, less mobile species seem to be
sensitive to fragmentation, which usually increases towards urban
centers, while mobile species such as birds or bees are more
affected by habitat features (e.g., Marzluff, 2001; Angold et al.,
2006; Croci et al., 2008; Kearns and Oliveras, 2009).
Several European and North American studies revealed striking
differences in vascular plant species richness compared with many
groups of animals: cities appear to be hot spots of plant species
richness. Numbers of plant species and communities increase with
the size of the city and the human population (Klotz, 1990; Pysek,
1993), and cities usually have higher numbers of plant species
compared with rural surroundings (Haeupler, 1975; McKinney,
2002; Hope et al., 2003; Kühn et al., 2004; Wania et al., 2006;
Knapp et al., 2009). In one of the first gradient studies, Kunick
(1982) found plant species richness to peak in the transition zone
between densely built-up inner-city areas of Berlin and the adja-
cent areas.
The high species richness of urban floras has often been
explained by (1) the high habitat heterogeneity of urban areas
resulting in a patchwork of (semi-)natural remnants and novel
urban habitats offering something to match the strongly differing
habitat requirements of many species, and (2) the high number of
introduced species (Sukopp and Werner, 1983; Gilbert, 1989;
Kowarik, 1990; Pysek, 1993; McKinney, 2002). A third line of argu-
ments addresses socioeconomic factors as drivers of urban biodi-
versity (Alberti et al., 2003; Kinzig et al., 2005; Essl et al., 2011a).
Tables 1 and 2 offer examples to illustrate some of the underlying
mechanisms, allowing us to infer processes from urban biodiversity
patterns.
2.2. Native species
Grid cell comparisons of the floristic assemblages of rural and
urban areas in Germany revealed significantly higher numbers of
both introduced and native species in cells with prevailing urban
land use than in rural. Even endangered native species had higher
species numbers in urbanized grid cells (Kühn et al., 2004). The
high plant species richness of cities is thus not exclusively due to
introduced species. Geological heterogeneity, as a proxy for natural
landscape heterogeneity, was the most important factor for
explaining the number of native species while city effects were
more essential for introduced species. The authors concluded that
cities, at least in Germany, have been preferably established at
rivers and along steep environmental gradients, which made them
naturally rich in species. It thus remains an open question whether
these species occur despite or because of urban conditions.
Further comparisons between the floristic assemblages of
urbanized and rural areas in Germany revealed some general
insights beyond species numbers: the floras of urbanized areas offer
a reduced phylogenetic diversity with a few species-rich lineages
(Knapp et al., 2008a), and significant changes in species traits of
urban plant species compared with those from rural areas (Knapp
et al., 2008b, 2009). For example, urban regions harbor fewer
species with hygromorphic leaves, but more species with scle-
romorphic or succulent leaves and species that are indicators
of nutrient-rich, warm, and dry conditions, in other words, species
 I. Kowarik / Environmental Pollution 159 (2011) 1974e1983
that are well adapted to changed urban environments (see Table 1
for further examples).
A few studies on temporal changes in urban floras over periods
of more than 100 years suggest an increase in total species number
over time, but also marked changes in species composition. In the
city of Plzen  , Czech Republic, the floristic richness of the
surroundings decreased over a period of about 130 years, most
likely due to intensified agricultural practice, while the species
number in the city increased surprisingly (Chocholousková and
Pysek, 2003). In the city of Halle, Germany, urbanization caused
a considerable species turnover of 22% over a period of 320 years
with the major changes occurring in the 20th century. Over that
period the total species number increased, including that of native
species, but different species groups significantly declined, espe-
cially species inclined to wet or nutrient-poor habitats (Knapp et al.,
2010). A decrease in wetland species has been also reported in New
York, Rome, and Beijing (DeCandido et al., 2007; Wang et al., 2007;
Ceschin et al., 2009; Zhao et al., 2010).
Analyses of species composition at the scale of grid cells with
prevailing urban land use or of total urban regions offer essential
insights but provide only limited information at the habitat scale as
the data usually represent highly heterogeneous areas. Integrating
habitat features into analyses of large floristic data sets helps to
identify the role of individual land use types for species richness,
e.g., archaeological areas or parks in Rome (Celesti-Grapow et al.,
2006). Analyses at the habitat scale in Berlin showed that about
two-thirds (63%) of native plant species were able to colonize sites
subject to high levels of human-mediated disturbances, whereas
the remaining species were confined to more natural sites
(Kowarik, 1990). In Beijing, a quarter to almost a third of all plant
species, which had been assessed as endangered or protected at the
regional or national scale, also occur in urban and suburban parts of
the city, but a third of all species that occur outside of the built-up
areas of Beijing was not able to thrive in more urbanized areas of
the city (Wang et al., 2007). The historical analysis of the flora of
Plzen indicates further limitations for plant species to survive in
urban settings. The proportional representation of common
species decreased over the period of the last 130 years, because
species that were once common became rare. This suggests that
many species can occur in cities, but with only small populations
(Chocholousková and Pysek, 2003). Consistently, rare species
declined with increasing human population density in Britain
(Thompson and Jones, 1999).
The presence of many species in urban settings is not necessarily
a proxy, however, for the occurrence of self-sustaining populations.
Unfortunately, there are only a few studies addressing whether
cities act as “ecological traps,” i.e., if species are able to reproduce
successfully on urban land or if their populations there depend on
a continued influx from nonurban areas. Leston and Rodewald
(2006), for example, tested the ecological traps hypothesis for
a bird, the northern cardinal, and demonstrated the species’s
successful regeneration in urban forest patches. The analyses of
the species assemblages of more or less urbanized grid cells in
Germany suggest that rare native species generally decline, leading
to a homogenization of native urban floras (Kühn et al., 2004). An
expert assessment of the establishment status of all plant species
occurring in Berlin reveals a likely important underlying habitat-
related response (Kowarik and von der Lippe unpubl.): native
species have a decreasing chance of establishing self-sustaining
populations on urban man-made sites that are subject to higher
levels of disturbance. Some endangered species are established on
such sites, but the majority of Red List species are confined to more
natural conditions.
All of these findings suggest that habitat changes associated with
urban land uses function as key filters in urban species composition.
1977
Obviously some species benefit from urban conditions while others
experience decline. This insight has led to different categorizations
to express the affinity of a species for urban conditions. The first was
probably by the Danish plant geographer Joakim Schouw (1823)
who described “Plantae urbanae” in the early 19th century. Wittig
et al. (1985) differentiated “urbanophileous” and “urbanophobic”
species; Blair (2001) described “urban avoiders,” “urban adapters,”
and “urban exploiters”; and Hill et al. (2002) discussed “urban
specialists.”
2.3. Nonnative species
Cities are outstanding points of entry as well as foci for the
secondary release of introduced species, with trade, traffic, and
horticulture as most prominent dispersal pathways (Hodkinson
and Thompson, 1997; Dehnen-Schmutz et al., 2007; Kowarik and
von der Lippe, 2007). Early urban invasion studies demonstrated
the importance of human-mediated activities for changes in the
urban species pool. In his seminal work, Flora of Montpellier,
Southern France, Albert Thellung (1912) assessed the relative
importance of different dispersal vectors by which nonnative
species are incorporated into the urban flora. His work reveals that
more species were accidentally introduced, for example, through
the import of wool or with ballast material, but that deliberately
introduced species, mostly ornamentals, had a higher chance of
establishing self-sustaining populations.
Urban land use in general and other parameters such as cover of
impervious surfaces, distance to city center, human population size,
and density are positively related to the abundance of introduced
species (Pysek, 1998; Roy et al., 1999; Hill et al., 2002; Celesti-
Grapow et al., 2006; Wania et al., 2006; Godefroid and Koedam,
2007; Nobis et al., 2009). These parameters serve as proxies for
two different types of mechanisms associated with urban land use:
(1) changes in urban environmental conditions and (2) human
activities that drive species selection and management of urban
green spaces.
Many studies have demonstrated cities as hotspots of alien plant
species. Pysek (1998) reported an average proportion of 40% alien
species in the urban floras of 54 European cities, ranging from 20 to
60% and including 25% neophytes (post-1500 introductions) and
15% archaeophytes (pre-1500 introductions) on average. Clemants
and Moore (2003) found a similar range for some North Amer-
ican cities (19e46%) with an average percentage of nonnative
species of 35%. For the flora of the built-up area of Beijing, Zhao
et al. (2010) report a 53% proportion of introduced species. The
large ranges in introduced species may reflect differences among
cities but may be also due to different approaches to determining
the reference area for urban floras or in lacking knowledge on the
origin of species. In Korea, for example, it is difficult to differentiate
early immigrants from China from native species, which would
explain the relatively low proportion of nonnatives (17%) in the
urban flora of Chonju (Zerbe et al., 2004).
Historical analyses of urban floras revealed a strong increase in
neophytes over the last 100 years and over even longer periods of
time (Godefroid, 2001; Chocholousková and Pysek, 2003;
DeCandido, 2004; Knapp et al., 2010). In contrast, archaeophytes
tend to decline, likely due to their association with traditional
agricultural practices (Preston et al., 2004; Pysek et al., 2005).
However, most alien species are generally less abundant than
native species (Williamson and Fitter, 1996; Hulme, 2008; Knapp
et al., 2009), thus leading to differentiation among urban regions
(Kühn and Klotz, 2006). Nonnative species are most prevalent in
early successional stages on urban wasteland (Kowarik, 1995; Pysek
et al., 2004; Muratet et al., 2007), in the garden flora (Loram et al.,
2008), and on naturally disturbed sites such as floodplains (Maskell
1978 I. Kowarik / Environmental Pollution 159 (2011) 1974e1983

et al., 2006), but can also dominate pioneer forests on derelict land
(Kowarik, 2005a).
A few studies demonstrated that many successful introduced
species are well adapted to drier, warmer, and highly disturbed
urban habitats (Table 1) and are often preferentially enhanced by
human activities (Table 2). However, it is challenging for further
research to disentangle these two mechanisms as they may often
interact, as illustrated by the example of Ailanthus altissima. This
tree, native to China, is virtually confined to urban centers in central
Europe, which was originally explained as being due to urban heat
island effectsdbut which could also simply reflect planting
patterns, which in turn drive propagule pressure and subsequent
invasion processes. Recent climate chamber experiments revealed,
however, that increased temperatures actually enhance the growth
of this tree (Kowarik and Säumel, 2007).
3. Novel urban ecosystems and the four natures approach
Profound human-mediated changes in natural settings can lead
to lasting deviations of ecosystem dynamics and thereby to a shift
in ecosystem types. Such developments were recognized early
on in intensively used European landscapes (e.g., Bernatsky, 1904;
von Hornstein, 1950), and conceptually assessed as a change in
“potential natural vegetation” (Tüxen, 1956). In a seminal paper,
Hobbs et al. (2006) addressed the result of such changes as “novel
ecosystems,” which are characterized by two features: major
human-induced changes in the abiotic environment and significant
changes in the species pool due to local extinctions or introduction
of novel biotic elements, with the potential for changes in
ecosystem functioning.
The examples given by Hobbs et al. (2006) mostly consider
ecosystem processes subsequent to the abandonment of agricul-
tural land and the degradation of forests owing to species invasions.
Surprisingly, the authors do not mention urban systems although
many of these certainly match the cited definition criteria of novel
ecosystems. As reviewed above, species assemblages in urban
settings are profoundly changed, both by the decline of rare native
species and the introduction of alien species. In addition, changes
in urban climate, hydrology, and soils, with associated biotic
feedback loops, heavily affect ecosystem functioning, as reviewed
by Alberti (2005).
Although cities as a whole can easily be seen as novel systems
contrasting with rural surroundings, scaling down to the habitat
level shows significant differences. From a historical viewpoint,
urban regions usually present a mosaic of fragmented habitats that
differ conspicuously in their history and their pace and level of
transformation from pristine to urban ecosystems. With a simple
landscape matrix, Table 3 illustrates major types of ecosystems that
can occur within urban areas, reflecting different human-mediated
transformation stages, triggered by urbanization. These range from
pristine remnants to agriculturally and horticulturally shaped
systems to distinctive urban-industrial ecosystems that emerge on
severely changed sites, mostly after habitat destruction.
As an invitation to consider the whole range of ecosystems in
urban cities, these ecosystem types can be conceptualized simply as
different kinds of nature that reflect the transformation of pristine
environmental conditions due to urbanization (“the four natures
approach”; Kowarik, 2005a). In terms of urban novelty, emerging
ecosystems on previously built-up areas or heavily changed urban
land are novel (“nature of the fourth kind”), as are some horticul-
tural systems (“nature of the third kind”) that have been estab-
lished, e.g., on landfills with artificial substrates or as green roofs on
the top of buildings. Both novel urban ecosystems and remnants of
pristine or agricultural systems are influenced by urbanization,
either directly by land use or indirectly by effects resulting from the
urban matrix. As an example of the latter, forests in urban areas
often show an increased influx of introduced species and higher
decomposition and nitrification rates compared with rural forest
stands (McDonnell et al., 1997; Borgmann and Rodewald, 2005).
The historical perspective that underlies the matrix in Table 3
illustrates fundamental differences in landscape history that are
also vital for conservation issues as several studies have disclosed
the general importance of habitat continuity for maintaining
species assemblages including key species for conservation (Hermy
et al., 1999; Vähä-Piikkiö et al., 2004; Johansson et al., 2008). Hence,
rare and endangered grassland and forest species are believed to
occur preferentially in settings of nature of the first kinddor
second kinddthan in the other types. In the face of increasing
intensity of agricultural and silvicultural practices in many regions
(e.g., Henle et al., 2008) the conservation value of such relict
habitats in urban regions increases, and a few studies document
that urban-industrial wastelands, parks and gardens, and even
green roofs can also harbor many species (e.g., Rebele and Dettmar,
1996; Dvorak and Volder, 2010; Goddard et al., 2010). One
outstanding example is an old green roof on a filtration plant in
Zurich, Switzerland, which provides a habitat for 170 plant species
including 9 rare or endangered species of orchids (Chrisman, 2005).
4. Nature conservation in urban areas
In face of an accelerating transformation of rural landscapes by
urbanization the need to protect remaining remnants of (semi-)
natural habitats increases. Yet in the urban millennium, the
importance of biodiversity conservation within cities generally
grows as well. Early conceptual papers on urban nature conserva-
tion (e.g., Auhagen and Sukopp, 1983) as well as more recent papers
highlight two main reasons for enhancing conservation activities in
urban settings: (1) as a contribution to the conservation of biodi-
versity and, (2) to enable the social functions and other ecosystem
services that are associated with urban nature (e.g., McKinney,
2002; Miller and Hobbs, 2002; Dunn et al., 2006; Tzoulas et al.,
2007; Goddard et al., 2010).

Table 3
Different types of ecosystems within a city area resulting from landscape transformation due to urbanization. These types can be conceptualized as different kinds of nature to
illustrate fundamental differences in landscape history (adapted from Kowarik, 2005a).

Type of ecosystem History Prevailing level Type of

of transformation nature
Pristine (e.g., forests, wetlands) Remnants of natural ecosystems Low 1
Agricultural (e.g., grassland, fields) Remnants of man-made ecosystems, Medium 2
resulting from early habitat transformation
Horticultural (e.g., parks, gardens) Transformed remnants or newly Medium to high 3
established after habitat destruction
Urban-industrial (e.g., vacant lots, Emerging after habitat destruction High 4
industrial sites, transport corridors)
 I. Kowarik / Environmental Pollution 159 (2011) 1974e1983
4.1. Urban contribution to biodiversity conservation
As reviewed above, cities are often richer in plant species than
their rural surroundings, but this does not necessarily hold for
other taxonomic groups. Urban areas usually harbor profoundly
changed species assemblages with novel mixtures of native and
introduced species, although the native species are not necessarily
fewer in number and abundance than introduced species (Kühn
et al., 2004). Analyses of changes in species numbers and compo-
sition at the city level are highly important for a better under-
standing of biodiversity responses to urbanization. It should be
kept in mind, however, that species numbers and related diversity
indices have limited relevance for conservation issues (Rodda,
1993; Prendergast and Eversham, 1997), although they are
frequently used as value-connoted biodiversity indicators (Heink
and Kowarik, 2010).
For assessing the contribution of cities to biodiversity conser-
vation, the key question is instead whether speciesdin particular,
rare and endangered speciesdnot only occur on urban land but
can also establish self-sustaining populations. There is a great need
for further research as only a few studies address this crucial
question (e.g., Leston and Rodewald, 2006). Results on the estab-
lishment of plant species in (semi-)natural relict habitats and on
transformed urban sites in Berlin suggest that some Red List
species can indeed establish self-sustaining populations on man-
made urban sites, whereas the majority of endangered species rely
on relicts of pristine ecosystems (Kowarik and von der Lippe,
unpubl.). These findings correspond well with results from several
studies at the city level, revealing that, in general, rare native
species became rarer or extinct in urban areas, especially when
bound to wet and/or nutrient-poor sites (e.g., DeCandido, 2004;
Knapp et al., 2010). Studies at the habitat scale, however, offer
prominent exceptions to this trend, for example for grassland
species in historical parks (Maurer et al., 2000) or insect species in
gardens (Goddard et al., 2010). Some farmland birds that are
subject to a strong decline in many agricultural landscapes
(BirdLife International, 2004) are able to establish self-sustaining
populations in inner-city airports and on early successional urban
wasteland in Berlin (Ökologie and Planung unpubl.; Meffert,
Marzluff and Dziock, unpubl.).
The big picture thus suggests that urban areas cannot substitute
the functioning of natural ecosystems as habitats for a broad array
of species, including rare habitat specialists or species that are
highly sensitive to fragmentation. This conclusion strongly
supports strategies to protect (semi-)natural remnants in urban
regions and to minimize negative impacts of new urban growth on
biological and environmental resources. However, there is an
increasing body of evidence that individual urban habitat types
across the whole spectrum of the four natures (cf. Table 3) can add
a significant contribution to biodiversity conservation, and further
research at the habitat and population scales is strongly encouraged
to elucidate urban habitat functions in detail.
At the regional scale, the relative urban contribution to the
conservation of plant and animal species is expected to depend on
the environmental qualities of the particular surroundings. The
function of urban areas for biodiversity conservation is likely minor
in cities that are embedded in more natural settings, but could be
highly significant when cities are surrounded by intensively used
and biologically impoverished landscapes. Within cities, intensity of
urban land use, habitat continuity, and management practices are
likely crucial factors that determine biological richness. These factors
can also explain differences between cities, or within cities between
neighborhoods, with different histories and different cultural norms
and legacies, which can have an impact on the acceptance of natural
processes within urban areas.
1979
4.2. Linking biodiversity with social functions
Research from environmental psychology and environmental
medicine has shown that the exposure of people to natural systems
positively affects human well-being and health. Contact with green
space speeds up the recovery from stress (Kaplan and Kaplan, 1989),
enhances the development of children (Kahn and Kellert, 2002),
could mitigate juvenile hyperactivity disorder (Kuo and Taylor,
2004), and is generally associated with better health (Mitchell and
Popham, 2008). Such results strongly support urban conservation
strategies under the motto “conservation where people live and
work” (Miller and Hobbs, 2002).
One could argue that many ecosystem services in urban regions
depend on the spatial distribution and abundance of biomass rather
than on the level of species richness or the occurrence of particular
species. Yet biodiversity does matter as a study from England shows.
In the urban green spaces of Sheffield, psychological benefits
increased with the species richness of parks, and green space users
were able to perceive species richness more or less accurately
depending on the taxonomic group in question. The degree of
psychological benefit was positively correlated with species rich-
ness of plants and to a lesser extent of birds. Plant variety correlated
with the ability of people to reflect and bird variety with people’s
emotional attachment (Fuller et al., 2007).
While the study by Fuller et al. (2007) demonstrated a clear link
between biodiversity and social benefits of urban nature, Miller
(2005) found an increasing “extinction of experience” with natural
systems in the human population living in biologically impoverished
urban areas. This would contribute to inequity in urban life and
strongly emphasizes the need for city-wide conservation strategies
beyond the traditional focus on nature reserves. Better opportunities
for urban residents to interact with nature are also thought to foster
a wider interest in nature conservation issues and could thus
enhance conservation activities at and beyond the city scale (Miller,
2005; Dunn et al., 2006).
4.3. Considering the whole range of urban nature
Existing conservation strategies mostly aim at preserving native
ecosystems by curbing urban growth and resulting habitat losses,
preserving more or less transformed relicts of natural habitats
within cities or restoring native species in urban habitats. These
approaches are highly important and necessary as specific urban
ecosystems are not expected to substitute the whole functioning of
(semi-)natural systems for biodiversity conservation (see above).
Nevertheless the question arises as to whether traditional conser-
vation approaches need to be supplemented as they do not cover
the whole range of urban nature.
Although not systematically analyzed thus far, most papers on
urban conservation approaches appear to be biased to some extent
by conservation, or restoration, efforts that prioritize relicts of
pristine (“native”) ecosystems or native species in urban regions
and tend to neglect other urban systems. There is a simple argu-
ment for a paradigm shift towards considering the whole range of
urban nature: an exclusive focus on “native nature” risks ignoring
the benefits of the other aspects of urban nature emerging on
profoundly altered sites or developing on sites shaped by horti-
culture (cf. Table 3).
These novel urban ecosystems may challenge the perception of
conservationists in different ways. First, one must consider
whether ecosystems that represent profound human-induced
changes in natural systems have value or whether they are per
se to be negatively viewed because they diverge from natural
settings. The paper by Hobbs et al. (2006) provides an important
contribution to this discussion as it makes clear that such systems
1980 I. Kowarik / Environmental Pollution 159 (2011) 1974e1983

Table 4
Risks often attributed to nonnative species in urban regions.

Risk Explanation/Comment
Spread of introduced Cities and other settlements are prominent points of entry for nonnative species and decreasing numbers of exotics along urban-to-rural
species into the gradients suggest the functioning of settlements as invasion foci (Sullivan et al., 2005; Vidra and Shear, 2008; Gavier-Pizarro et al., 2010;
adjacent countryside Essl et al., 2011b). A prominent vector is dispersal by birds (Catling and Porebski, 1994; Seidling, 1999), and also traffic and moving
water can move introduced species outside of cities (Truscott et al., 2006; von der Lippe and Kowarik, 2008; Säumel and Kowarik, 2010).
However, comparisons over time in Britain revealed no general trend of spreading out of urban habitats into the wider countryside
(Botham et al., 2009). In Plzen , the numbers of alien species have increased in the city, but not in the surroundings over a period of more
than 100 years (Chocholousková and Pysek, 2003). As numerous other human activities directly contribute to the establishment of
nonnative species in near-natural surroundings, either through planting, seeding, or deposition of garden waste (Hodkinson and
Thompson, 1997; Kowarik, 2003), it remains challenging to disentangle and quantify invasion processes that rely on urban propagule
sources from those that depend on the release of introduced species outside of cities.
Displacement of native The high proportion of alien species in urban regions often coincides with a decline in native species. There is not necessarily a causal
species relationship between the two processes, however, as introduced species are often passengers rather than drivers of change (MacDougall
and Turkington, 2005; Didham et al., 2005). Some studies show that numbers of natives and nonnatives increase together in urban areas
(Celesti-Grapow et al., 2006; Kühn and Klotz, 2006). Introduced species may, but need not necessarily, interact with existing native
species. It is thus a challenge for impact studies to disclose a causal role of introduced species in the decline of native species on urban land.
Biotic homogenization There is some concern that the high numbers of nonnative species in urban regions contribute to a global homogenization of biodiversity
patterns as the same suite of nonnative species can be found in many cities (McKinney, 2006). For German cities, Kühn and Klotz (2006)
revealed homogenizing effects of urbanization for native plant species assemblages and pre-1500 alien plant species (archaeophytes) while
post-1500 alien species (neophytes) led to a differentiation of urban floras at local to regional scales. In contrast to European cities,
nonnative species appear to be associated with losses in beta-diversity in North American cities, likely due to the presence of
European archaeophytes (La Sorte et al., 2008).
Human health risks Many native species impose health risks, e.g., species with an allergen potential, but few exotic species add novel risks. Exceptions include
phytodermatitis associated with the former ornamental herb Heracleum mantegazzianum (Jaspersen-Schib et al., 1996) or enhanced allergy
risks due to North American Ambrosia species in Europe (Taramarcaz et al., 2005).
Impacts on higher As more animal species feed on native than on nonnative plant species, the cultivation of the latter is believed to negatively affect
trophic levels higher trophic levels. Studies on the relationship of nonnative plant species with wildlife in gardens yielded contrary results indicating that
other factors such as the structural heterogeneity of gardens may be more important than the origin of garden plants
(cf. Goddard et al., 2010).
Maintenance costs The prolific reproduction and vigorous growth of some introduced species in urban habitats may increase management costs for green
spaces and sometimes even for maintaining built infrastructure. Examples include Fallopia species (Child and Wade, 2000) or the tree
Ailanthus altissima (Celesti-Grapow and Blasi, 2004).
Emission of VOCs Many conifers and broad-leaved evergreen species emit biogenic volatile organic compounds (VOCs), which can increase summer smog
in cities. In regions with low numbers of native evergreen species, the planting of introduced evergreen trees or shrubs may reduce air
quality. However, some important native species, such as many oaks, also emit VOCs (Niinemets and Peñuelas, 2008).
Impacts on ecosystem Alien species may affect ecosystem services by many ways (Charles and Dukes, 2007), but also positive effects have been found. In
services urban forests in Florida, invasive trees were most apt at sequestering CO2 (Escobedo et al., 2010). Assessing the cultural services
provided by introduced species is still challenging.

are inevitable parts of contemporary landscapes and should stim-
ulate expanding conservation or restoration approaches. We might
question whether biotic settings should be expected to remain the
same while almost everything in the social, physical, and chemical
urban environment changes. As the answer is likely to be negative,
it could be concluded that novel urban ecosystems deserve more
attention from a conservation perspective due to their paramount
social functions and their possible contributions to biodiversity
conservation as discussed above.
A second challenge associated with novel urban ecosystems is
the prevalence of many nonnative species. Invasions by alien
species are widely accepted as a leading cause of biodiversity loss.
As Didham et al. (2005), however, argue, much of the evidence for
this contention is based on simple correlations between exotic
dominance and native species decline in degraded systems. This
specifically holds for many urban systems that are dominated by
alien species. The decline of previously occurring native species is
often due to habitat loss and transformation associated with urban
land uses, which in turn provide beneficial conditions for intro-
duced species. These introduced species are, as in other cases
(MacDougall and Turkington, 2005), often passengers rather than
drivers of change. Novel mixtures of alien and native species that
emerge after profound habitat transformation on urban land are
expected to be adapted to novel environmental conditionsdand in
some cases possibly better adapted than previous assemblages of
native species. It is thus worthwhile to consider the benefits linked
to nonnative species. Possible consequences may involve the inte-
gration of gardens and of novel urban-industrial ecosystems into
urban green systems and the strengthening of their conservation
functions (e.g., Kowarik and Körner, 2005; Goddard et al., 2010).
It is also true however that several risks are associated with
introduced species, as sketched in Table 4. Although many authors
consider the high proportions of nonnative species in urban envi-
ronments to be self-evident risks that lead to further invasions of
the countryside, to a displacement of native species, or to biotic
homogenization and other negative impacts, the big picture is
much more complex, and more research about the impacts of alien
species in urban settings is urgently needed. Risks certainly do exist
but are usually related to particular species and not to the whole
group of nonnative species. To avoid uncritical generalizations, it is
thus promising to reconsider risks with a case-by-case approach. As
a normative baseline for such assessments, the recently published
definition of environmental damages due to invasive species may
be helpful. Such damage is expected to occur when an alien species
evokes “a significant adverse effect on a biotic or abiotic conser-
vation resource that has an impact on (a) the value of the conser-
vation resource, (b) the conservation resource as an ecosystem
component, or (c) the sustainable use of the conservation resource”
(Bartz et al., 2010). In keeping with this definition, change due to
novel species is not considered to be detrimental when it is not
associated with significant negative impacts on other species or
resources with special conservation interest. However, a risk
assessment should also consider potential risks which may result,
for example, from the invasion of adjacent habitats.
In conclusion, I thus argue for combining well-established
strategies aiming to preserve (semi-)natural remnants and enhance
 I. Kowarik / Environmental Pollution 159 (2011) 1974e1983
native species in urban regions with approaches that acknowledge
the contribution of novel urban ecosystems and associated species
assemblages. These may be valued as an adaptation to severe
habitat transformation and may secure the provision of ecosystem
services in urban settings in an era of global change.
Acknowledgements
This paper is dedicated to Prof. Dr. Dr. h.c. Herbert Sukopp, the
major pioneer of urban ecology in Europe, at the occasion of his
80th birthday in November 2010. My thanks are due to Bill
Manning and Claudia Wiegand for stimulating this contribution, to
an anonymous reviewer for helpful comments, and to Kelaine
Vargas for improving my English.
References
Alberti, M., 2005. The effects of urban patterns on ecosystem function. International
Regional Science Review 28, 168e192.
Alberti, M., Marzluff, J.M., Shulenberger, E., Bradley, G., Ryan, C., Zumbrunnen, C.,
2003. Integrating humans into ecology: opportunities and challenges for
studying urban ecosystems. Bioscience 53, 1169e1179.
Angold, P.G., Sadler, J.P., Hill, M.O., Pullin, A., Rushton, S., Austin, K., Small, E.,
Wood, B., Wadsworth, R., Sanderson, R., Thompson, K., 2006. Biodiversity in
urban habitat patches. Science of the Total Environment 360, 196e204.
Antrop, M., 2004. Landscape change and the urbanization process in Europe.
Landscape and Urban Planning 67, 9e26.
Araújo, M.B., 2003. The coincidence of people and biodiversity in Europe. Global
Ecology and Biogeography 12, 5e12.
Auge, H., 1997. Biologische Invasionen: das Beispiel Mahonia aquifolium. In:
Feldmann, R., Henle, K., Auge, H., Flachowsky, J., Klotz, S., Krönert, R. (Eds.),
Regeneration und nachhaltige Landnutzung: Konzepte für belastete Regionen.
Springer, Berlin, pp. 124e129.
Auhagen, A., Sukopp, H., 1983. Ziel, Begründungen und Methoden des Natur-
schutzes im Rahmen der Stadtenwicklungspolitik von Berlin. Natur und Land-
schaft 58, 9e15.
Bakker, P.A., Boeve, E., 1985. Stinzenplanten. Uitgeverij Terra, Zutphen.
Bartz, R., Heink, U., Kowarik, I., 2010. Proposed definition of environmental damage
illustrated by the cases of genetically modified crops and invasive species.
Conservation Biology 24, 675e681.
Bernatsky, J., 1904. Anordnung der Formationen nach ihrer Beeinflussung seitens
der menschlichen Kultur und der Weidetiere. Engler’s Botanisches Jahrbuch 94,
1e8.
Biadun, W., Kitowski, I., Filipiuk, E., 2009. Trends in the arrival dates of spring
migrants in Lublin (E Poland). Acta Ornithologica 44, 89e94.
BirdLife International, 2004. Birds in the European Union: a status assessment,
Wageningen.
Blair, R.B., 1999. Birds and butterflies along an urban gradient: surrogate taxa for
assessing biodiversity? Ecological Applications 9, 164e170.
Blair, R.B., 2001. Birds and butterflies along urban gradients in two ecoregions of the
United States: is urbanization creating a homogenous fauna? In: McKin-
ney, M.L., Lockwood, J.L. (Eds.), Biotic Homogenization. Plenum Publishers, New
York, pp. 33e56.
Borgmann, K.L., Rodewald, A.D., 2005. Forest restoration in urbanizing landscapes:
interactions between land uses and exotic shrubs. Restoration Ecology 13,
334e340.
Botham, M.S., Rothery, P., Hulme, P.E., Hill, M.O., Preston, C.D., Roy, D.B., 2009. Do
urban areas act as foci for the spread of alien plant species? An assessment of
temporal trends in the UK. Diversity and Distributions 15, 338e345.
Brunzel, S., Fischer, S.F., Schneider, J., Jetzkowitz, J., Brandl, R., 2009. Neo- and
archaeophytes respond more strongly than natives to socio-economic mobility-
and disturbance patterns along an urban-rural gradient. Journal of Biogeog-
raphy 36, 835e844.
Cadenasso, M.L., Pickett, S.T.A., 2001. Effect of edge structure on the flux of species
into forest interiors. Conservation Biology 15, 91e97.
Catling, P.M., Porebski, Z.S., 1994. The history of invasion and current status of glossy
buckthorn, Rhamnus frangula, in southern Ontario. Canadian Field-Naturalist
108, 305e310.
Celesti-Grapow, L., Blasi, C., 2004. The role of alien and native weeds in the dete-
rioration of archaeological remains in Italy. Weed Technology 18, 1508e1513.
Celesti-Grapow, L., Pysek, P., Jarosík, V., Blasi, C., 2006. Determinants of native and
alien species richness in the flora of Rome. Diversity and Distributions 15,
490e501.
Ceschin, S., Salerno, G., Caneva, G., 2009. Multitemporal floristic analysis on a humid
area in Rome’s archaeological site as indicator for environmental change.
Environmental Monitoring and Assessment 149, 29e42.
Charles, H., Dukes, J.S., 2007. Impacts of invasive species on ecosystem services. In:
Nentwig, W. (Ed.), Biological Invasions. Ecological Studies, vol. 193. Springer,
Berlin, New York, pp. 217e237.
1981
Cheptou, P.-O., Carrue, O., Rouifed, S., Cantarel, A., 2008. Rapid evolution of seed
dispersal in an urban environment in the weed Crepis sancta. Proceedings of the
National Academy of Sciences USA 105, 3796e3799.
Child, L.E., Wade, P.M., 2000. The Japanese Knotweed Manual. Packard Publishing
Limited, Chichester.
Chocholousková, Z., Pysek, P., 2003. Changes in composition and structure of urban
flora over 120 years: a case study of the city of Plzen  . Flora 198, 366e376.
Chrisman, S. (Ed.), 2005. Green roofs. Ecological Design and Construction. Schiffer,
Atglen.
Cincotta, R.P., Wisnewski, J., Engelman, R., 2000. Human population in the biodi-
versity hotspots. Nature 404, 990e992.
Cleland, R.E., 1972. Oenothera: Cytogenetics and Evolution. Academic Press, New
York.
Clemants, S.E., Moore, G., 2003. Patterns of species richness in eight northeastern
United States cities. Urban Habitats 1, 1e16.
Clergeau, P., Croci, S., Jokimäki, J., Kaisanlathi-Jokimäki, M.-L., Dinetti, M., 2006.
Avifauna homogenisation by urbanisation: analysis at different European lati-
tudes. Biological Conservation 127, 336e344.
Croci, S., Butet, A., Georges, A., Aguejdad, R., Clergeau, P., 2008. Small urban
woodlands as biodiversity conservation hot-spot: a multi-taxon approach.
Landscape Ecology 23, 1171e1186.
DeCandido, R., 2004. Recent changes in plant species diversity in urban Pelham Bay
park, 1947e1998. Biological Conservation 120, 129e136.
DeCandido, R., Calvanese, N., Alvarez, R.V., Brown, M.I., Nelson, T.M., 2007. The
naturally occurring historical and extant flora of Central Park, New York City,
New York 1857e2007. Journal of the Torrey Botanical Society 134, 552e569.
Dehnen-Schmutz, K., Touza, J., Perrings, C., Williamson, M., 2007. A century of the
ornamental plant trade and its impact on invasion success. Diversity and
Distributions 13, 527e534.
Didham, R.K., Tylianakis, J.M., Hutchinson, M.A., Ewers, R.M., Gemell, N.J., 2005. Are
invasive species the drivers of ecological change? Trends in Ecology and
Evolution 20, 470e474.
Duguay, S., Eigenbrod, F., Fahrig, L., 2007. Effects of surrounding urbanization on
non-native flora in small forest patches. Landscape Ecology 22, 589e599.
Dunn, R.R., Gavin, M.C., Sanchez, M.C., Solomon, J.N., 2006. The pigeon paradox:
dependence of global conservation on urban nature. Conservation Biology 20,
1814e1816.
Dvorak, B., Volder, A., 2010. Green roof vegetation for North American ecoregions:
a literature review. Landscape and Urban Planning 96, 197e213.
Ernst, W.H.O., Joosse-van Damme, E.N.G., 1983. Umweltbelastungen durch Miner-
alstoffe e Biologische Effekte. VEB Gustav Fischer, Jena.
Escobedo, F., Varela, S., Zhao, M., Wagner, J., Zipperer, W., 2010. Analyzing the
efficacy of subtropical urban forests in offsetting carbon emissions from cities.
Environmental Science and Policy 13, 362e372.
Essl, F., Dullinger, S., Rabitsch, W., Hulme, P.E., Hülber, K., Jarosík, V., Kleinbauer, I.,
Krausmann, F., Kühn, I., Nentwig, W., Vilà, M., Genovesi, P., Gherardi, F.,
Desprez-Loustau, M.L., Roques, A., Pysek, P., 2011a. Socioeconomic legacy yields
an invasion debt. PNAS 108, 203e207.
Essl, F., Milasowszkyb, N., Dirnböck, T., 2011b. Plant invasions in temperate forests:
resistance or ephemeral phenomenon? Basic and Applied Ecology 12, 1e9.
Evans, K.L., Newson, S.E., Gaston, K.J., 2009. Habitat influences on urban avian
assemblages. Ibis 151, 19e39.
Eversham, B.C., Roy, D.B., Telfer, M.G., 1996. Urban, industrial and other manmade
sites as analogues of natural habitats for Carabidae. Annales Zoologica Fennici
33, 149e156.
Fuller, R.A., Irvine, K.N., Devine-Wright, P., Warren, P.H., Gaston, K.J., 2007.
Psychological benefits of greenspace increase with biodiversity. Biology Letters
3, 390e394.
Fuller, R.A., Warren, P.H., Armsworth, P.R., Barbosa, O., Gaston, K.J., 2008. Garden
bird feeding predicts the structure of urban avian assemblages. Diversity and
Distributions 14, 131e137.
Gavier-Pizarro, G.I., Radeloff, V.C., Stewart, S.I., Huebner, C.D., Keuler, N.S., 2010.
Rural housing is related to plant invasions in forests of southern Wisconsin,
USA. Landscape Ecology 25, 1505e1518.
Gilbert, O.L., 1989. Ecology of Urban Habitats. Chapman and Hall, London.
Goddard, M.A., Dougill, A.J.H., Benton, T.G., 2010. Scaling up from gardens: biodi-
versity conservation in urban environments. Trends in Ecology and Evolution
25, 90e98.
Godefroid, S., 2001. Temporal analysis of the Brussels flora as indicator for changing
environmental quality. Landscape and Urban Planning 52, 203e224.
Godefroid, S., Koedam, N., 2007. Urban plant species patterns are highly driven by
density and function of built-up areas. Landscape Ecology 22, 1227e1239.
Grimm, N.B., Faeth, S.H., Golubiewski, N.E., Redman, C.L., Wu, J., Bai, X., Briggs, J.M.,
2008. Global change and the ecology of cities. Science 319, 756e760.
Gulezian, P.Z., Nyberg, D.W., 2010. Distribution of invasive plants in a spatially
structured urban landscape. Landscape and Urban Planning 95, 161e168.
Haeupler, H., 1975. Statistische Auswertungen von Punktrasterkarten der
Gefäßpflanzenflora Süd-Niedersachsens. Scripta Geobotanica 8, 1e141.
Hansen, A.J., Knight, R.L., Marzluff, J.M., Powell, S., Brown, K., Gude, P.H., Jones, K.,
2005. Effects of exurban development on biodiversity: patterns, mechanisms,
and research needs. Ecological Applications 15, 1893e1905.
Heink, U., Kowarik, I., 2010. What criteria should be used to select biodiversity
indicators? Biodiversity and Conservation 19, 3769e3797.
Henle, K., Alard, D., Clitherow, J., Cobb, P., Firbank, L., Kull, T., McCracken, D.,
Moritz, R.F.A., Niemelä, J., Rebane, M., Wascher, D., Watt, A., Young, J., 2008.
1982 I. Kowarik / Environmental Pollution 159 (2011) 1974e1983
Identifying and managing the conflicts between agriculture and biodiversity
conservation in Europeea review. Agriculture, Ecosystems and Environment
124, 60e71.
Hermy, M., Honnay, O., Firbank, L., Grashof-Bokdam, C., Lawesson, J.E., 1999. An
ecological comparison between ancient and other forest plant species of
Europe, and the implications for forest conservation. Biological Conservation 91,
9e22.
Hill, M.O., Roy, D.B., Thompson, K., 2002. Hemeroby, urbanity and ruderality: bio-
indicators of disturbance and human impact. Journal of Applied Ecology 39,
708e720.
Hobbs, R.J., Arico, S., Aronson, J., Baron, J.S., Bridgewater, P., Cramer, V.A.,
Epstein, P.R., Ewel, J.J., Klink, C.A., Lugo, A.E., Norton, D., Ojima, D.,
Richardson, D.M., Sanderson, E.W., Valladares, F., Vilá, M., Zamora, R., Zobel, M.,
2006. Novel ecosystems: theoretical and management aspects of the new
ecological world order. Global Ecology and Biogeography 15, 1e7.
Hodkinson, D.J., Thompson, K., 1997. Plant dispersal: the role of man. Journal of
Applied Ecology 34, 1484e1496.
Hope, D., Gries, C., Zhu, W.X., Fagan, W.F., Redman, C.L., Grimm, N.B., Nelson, A.L.,
Martin, C., Kinzig, A., 2003. Socioeconomics drive urban plant diversity.
Proceedings of the National Academy of Sciences USA 100, 8788e8792.
von Hornstein, F., 1950. Theorie und Anwendung der Waldgeschichte. For-
stwissenschaftliches Centralblatt 69, 161e177.
Hulme, P.E., 2008. Contrasting alien and native plant speciesearea relationships:
the importance of spatial grain and extent. Global Ecology and Biogeography 17,
641e647.
Jaspersen-Schib, R., Theus, L., Guirguis-Oeschger, M., Gossweiler, B., Meier-Abt, P.J.,
1996. Acute poisonings with toxic giants in Switzerland between 1966 and
1994. Schweizer Medizinische Wochenschrift 126, 1085e1098.
Johansson, L.J., Hall, K., Prentice, H.C., Ihse, M., Reitalu, T., Sykes, M.T., Kindström, M.,
2008. Semi-natural grassland continuity, long-term land-use change and plant
species richness in an agricultural landscape in Öland, Sweden. Landscape and
Urban Planning 84, 200e211.
Kahn Jr., P.H., Kellert, S.R. (Eds.), 2002. Children and Nature: Psychological, Socio-
cultural, and Evolutionary Investigations. MIT Press, Cambridge, MA.
Kaplan, R., Kaplan, S., 1989. The Experience of Nature: A Psychological Perspective.
Cambridge University Press, Cambridge, MA.
Kearns, C.A., Oliveras, D.M., 2009. Environmental factors affecting bee diversity in
urban and remote grassland plots in Boulder, Colorado. Journal of Insect
Conservation 13, 655e665.
Kinzig, A.P., Warren, P., Martin, C., Hope, D., Katti, M., 2005. The effects of human
socioeconomic status and cultural characteristics on urban patterns of biodi-
versity. Ecology and Society 10, 23. http://www.ecologyandsociety.org/vol10/
iss1/art23/.
Klotz, S., 1990. Species/area and species/inhabitants relations in European cities. In:
Sukopp, H., Hejny, S., Kowarik, I. (Eds.), Urban Ecology. SPB Academic Publisher,
The Hague, pp. 99e104.
Knapp, S., Kühn, I., Schweiger, O., Klotz, S., 2008a. Challenging urban species
diversity: contrasting phylogenetic patterns across plant functional groups in
Germany. Ecology Letters 11, 1054e1064.
Knapp, S., Kühn, I., Wittig, R., Ozinga, W.A., Poschlod, P., Klotz, S., 2008b. Urbani-
zation causes shifts in species’ trait state frequencies. Preslia 80, 375e388.
Knapp, S., Kühn, I., Bakker, J.P., Kleyer, M., Klotz, S., Ozinga, W.A., Poschlod, P.,
Thompson, K., Thuiller, W., Römermann, C., 2009. How species traits and
affinity to urban land use control large-scale species frequency. Diversity and
Distributions 15, 533e546.
Knapp, S., Kühn, I., Stolle, J., Klotz, S., 2010. Changes in the functional composition of
a central European urban flora over three centuries. Perspectives in Plant
Ecology, Evolution and Systematics 12, 235e244.
Kowarik, I., 1990. Some responses of flora and vegetation to urbanization in Central
Europe. In: Sukopp, H., Hejny, S., Kowarik, I. (Eds.), Urban Ecology. SPB
Academic Publisher, The Hague, pp. 45e74.
Kowarik, I., 1995. On the role of alien species in urban flora and vegetation. In: Pysek, P.,
Prach, K., Rejmánek, M., Wade, M. (Eds.), Plant Invasions: General Aspects and
Special Problems. SPB Academic Publishing, Amsterdam, pp. 85e103.
Kowarik, I., 2003. Human agency in biological invasions: secondary releases foster
naturalisation and population expansion of alien plant species. Biological
Invasions 5, 293e312.
Kowarik, I., 2005a. Wild urban woodlands: towards a conceptual framework. In:
Kowarik, I., Körner, S. (Eds.), Wild Urban Woodlands. New Perspectives for
Urban Forestry. Springer, Berlin, Heidelberg, pp. 1e32.
Kowarik, I., 2005b. Urban ornamentals escaped from cultivation. In: Gressel, J. (Ed.),
Crop Ferality and Volunteerism. CRC Press, Boca Raton, pp. 97e121.
Kowarik, I., Körner, S. (Eds.), 2005. Wild Urban Woodlands. New Perspectives for
Urban Forestry. Springer, Berlin, Heidelberg.
Kowarik, I., von der Lippe, M., 2007. Pathways in plant invasions. In: Nentwig, W.
(Ed.), Biological Invasions. Ecological Studies, vol. 193. Springer, Berlin, New
York, pp. 29e47.
Kowarik, I., Säumel, I., 2007. Biological flora of central Europe: Ailanthus altissima (Mill.)
Swingle. Perspectives in Plant Ecology, Evolution and Systematics 8, 207e237.
Kraas, F., 2008. Megacities as global risk areas. In: Marzluff, J., et al. (Eds.), Urban
Ecology. An International Perspective on the Interaction between Humans and
Nature. Springer, New York, pp. 583e596.
Krivánek, M., Pysek, P., Jarosík, V., 2006. Planting history and propagule pressure as
predictors of invasion by woody species in a temperate region. Conservation
Biology 20, 1487e1498.
Kunick, W., 1982. Zonierung des Stadtgebietes von Berlin (West). Ergebnisse flori-
stischer Untersuchungen. Landschaftsentwicklung und Umweltforschung 14,
1e164.
Kuo, F.E., Taylor, A.F., 2004. A potential natural treatment for attention-deficit/
hyperactivity disorder: evidence from a national study. American Journal of
Public Health 94, 1580e1586.
Kühn, I., Klotz, S., 2006. Urbanization and homogenization e comparing the floras
of urban and rural areas in Germany. Biological Conservation 127, 292e300.
Kühn, I., Brandl, R., Klotz, S., 2004. The flora of German cities is naturally species
rich. Evolutionary Ecology Research 6, 749e764.
La Sorte, F.A., McKinney, M.L., Pysek, P., Klotz, S., Rapson, G.L., Celesti-Grapow, L.,
Thompson, K., 2008. Distance decay of similarity among European urban floras:
the impact of anthropogenic activities on ß-diversity. Global Ecology and
Biogeography 17, 363e371.
Langner, M., Endlicher, W. (Eds.), 2007. Shrinking Cities: Effects on Urban Ecology
and Challenges for Urban Development. Peter Lang Publishing Group, Frankfurt
am Main.
Leston, L.F.V., Rodewald, A.D., 2006. Are urban forests ecological traps for under-
story birds? An examination using northern Cardinals. Biological Conservation
131, 566e574.
von der Lippe, M., Kowarik, I., 2007. Long-distance dispersal of plants by vehicles as
a driver of plant invasions. Conservation Biology 21, 986e996.
von der Lippe, M., Kowarik, I., 2008. Do cities export biodiversity? Traffic as
dispersal vector across urban-rural gradients. Diversity and Distributions 14,
18e25.
Liu, J., Daily, G.C., Ehrlich, P.R., Luck, G.W., 2003. Effects of household dynamics on
resource consumption and biodiversity. Nature 421, 530e533.
Lockwood, J.L., Cassey, P., Blackburn, T., 2005. The role of propagule pressure in
explaining species invasions. Trends in Ecology and Evolution 20, 223e228.
Loram, A., Thompson, K., Warren, P.H., Gaston, K.J., 2008. Urban domestic gardens
(XII): the richness and composition of the flora in five UK cities. Journal of
Vegetation Science 19, 321e330.
Lu, P.L., Yu, Q., Liu, J.D., Lee, X.H., 2006. Advance of tree-flowering dates in response
to urban climate change. Agricultural and Forest Meteorology 138, 120e131.
MEA (Millennium Ecosystem Assessment), 2005. Ecosystems and Human Well-
being: Current State and Trends: Findings of the Condition and Trends Working
Group. Island Press, Washington. http://www.millenniumassessment.org.
MacDougall, A.S., Turkington, R., 2005. Are invasive species the drivers or passen-
gers of change in degraded ecosystems? Ecology 86, 42e55.
Mack, R.N., 2000. Cultivation fosters plant naturalization by reducing environ-
mental stochasticity. Biological Invasions 2, 111e122.
Marzluff, J.M., 2001. Worldwide urbanization and its effects on birds. In:
Marzluff, J.M., Bowman, R., Donnelly, R. (Eds.), Avian Ecology in an Urbanizing
World. Kluwer, Norwell, pp. 19e47.
Marzluff, J.M., Shulenberger, E., Endlicher, W., Alberti, M., Bradley, G., Ryan, C.,
Simon, U., ZumBrunnen, C. (Eds.), 2008. Urban Ecology. An International
Perspective on the Interaction between Human and Nature. Springer, New York.
Maskell, L.C., Bullock, J.M., Smart, S.M., Thompson, K., Hulme, P.E., 2006. The
distribution and habitat associations of non-native plant species in urban
riparian habitats. Journal of Vegetation Science 17, 499e508.
Maurer, U., Peschel, T., Schmitz, S., 2000. The flora of selected urban land-use types
in Berlin and Potsdam with regard to nature conservation in cities. Landscape
and Urban Planning 46, 209e215.
McDonnell, M., Hahs, A., 2008. The use of gradient analysis studies in advancing our
understanding of the ecology of urbanizing landscapes: current status and
future directions. Landscape Ecology 23, 1143e1155.
McDonnell, M.J., Pickett, S.T.A., Groffman, P., Bohlen, P., Pouyat, R.V., Zipperer, W.C.,
Parmelee, R.W., Carreiro, M.M., Medley, K., 1997. Ecosystem processes along an
urban-to-rural gradient. Urban Ecosystems 1, 21e36.
McKinney, M.L., 2002. Urbanization, biodiversity, and conservation. Bioscience 52,
883e890.
McKinney, M.L., 2006. Urbanization as a major cause of biotic homogenization.
Biological Conservation 127, 247e260.
Miller, J.R., 2005. Biodiversity conservation and the extinction of experience. Trends
in Ecology and Evolution 20, 430e434.
Miller, J.R., Hobbs, R.J., 2002. Conservation where people live and work. Conserva-
tion Biology 16, 330e337.
Mitchell, R., Popham, F., 2008. Effect of exposure to natural environment on health
inequalities: an observational population study. Lancet 372, 1655e1660.
Mockford, E.J., Marshall, R.C., 2009. Effects of urban noise on song and
response behaviour in great tits. Proceedings of the Royal Society B 276,
2979e2985.
Mount, A., Pickering, C.M., 2009. Testing the capacity of clothing to act as a vector
for non-native seed in protected areas. Journal of Environmental Management
91, 168e179.
Muratet, A., Machon, N., Jiguet, F., Moret, J., Porcher, E., 2007. The role of urban
structures in the distribution of wasteland flora in the Greater Paris area,
France. Ecosystems 10, 661e671.
Neil, K.L., Landrum, L., Wu, J.G., 2010. Effects of urbanization on flowering
phenology in the metropolitan phoenix region of USA: findings from herbarium
records. Journal of Arid Environments 74, 440e444.
Niemelä, J., Kotze, D.J., 2009. Carabid beetle assemblages along urban to rural
gradients: a review. Landscape and Urban Planning 92, 65e71.
Niemelä, J., Kotze, D.J., Venn, S., Penev, L., Stoyanov, I., Spence, J., Hartley, D.,
de Oca, E.M., 2002. Carabid beetle assemblages (Coleoptera, Carabidae)
 I. Kowarik / Environmental Pollution 159 (2011) 1974e1983
across urbanerural gradients: an international comparison. Landscape
Ecology 17, 387e401.
Niggemann, M., Jetzkowitz, J., Brunzel, S., Wichmann, M.C., Bialozyt, R., 2009.
Distribution patterns of plants explained by human movement behaviour.
Ecological Modelling 220, 1339e1346.
Niinemets, U., Peñuelas, J., 2008. Gardening and urban landscaping: significant
players in global change. Trends in Plant Science 13, 60e65.
Nobis, M.P., Jaeger, J.A.G., Zimmermann, N.E., 2009. Neophyte species richness at the
landscape scale under urban sprawl and climate warming. Diversity and
Distributions 15, 928e939.
Pickett, S.T.A., Cadenasso, M.L., Grove, J.M., Nilon, C.H., Pouyat, R.V., Zipperer, W.C.,
Costanza, R., 2001. Urban ecological systems: linking terrestrial ecological,
physical, and socioeconomic components of metropolitan areas. Annual Review
of Ecology and Systematics 32, 127e157.
Prendergast, J.R., Eversham, B.C., 1997. Species richness covariance in higher
taxa: empirical tests of the biodiversity indicator concept. Ecography 20,
210e216.
Preston, C.D., 2000. Engulfed by suburbia or destroyed by the plough: the ecology of
extinction in Middlesex and Cambridgeshire. Watsonia 23, 59e81.
Preston, C.D., Pearman, D.A., Hall, A.R., 2004. Archaeophytes in Britain. Botanical
Journal of the Linnean Society 145, 57e294.
Pysek, P., 1993. Factors affecting the diversity of flora and vegetation in central
European settlements. Vegetatio 106, 89e100.
Pysek, P., 1998. Alien and native species in central European urban floras: a quan-
titative comparison. Journal of Biogeography 25, 155e163.
Pysek, P., Chocholousková, Z., Pysek, A., Jarosík, V., Chytrý, M., Tichý, L., 2004. Trends
in species diversity and composition of urban vegetation over three decades.
Journal of Vegetation Sciences 15, 781e788.
Pysek, P., Jarosík, V., Chytrý, M., Kropác, Z., Tichý, L., Wild, J., 2005. Alien plants in
temperate weed communities: prehistoric and recent invaders occupy different
habitats. Ecology 86, 772e785.
Rebele, F., Dettmar, J., 1996. Industriebrachen. Ökologie und Management. Ulmer,
Stuttgart.
Rebele, F., Lehmann, C., 2001. Biological flora of central Europe: Calamagrostis epi-
gejos (L.) Roth. Flora 196, 325e344.
Rodda, G.H., 1993. How to lie with biodiversity. Conservation Biology 7, 959e960.
Roy, D.B., Hill, M.O., Rothery, P., 1999. Effects of urban land cover on the local species
pool in Britain. Ecography 22, 507e515.
Schouw, J.F., 1823. Grundzüge der allgemeinen Pflanzengeographie. Berlin.
Seaward, M.R.D., 1982. Lichen ecology of changing urban environments. In:
Bornkamm, R., Lee, J.A., Seaward, M.R.D. (Eds.), Urban Ecology. Blackwell
Scientific Publications, Oxford, pp. 181e189.
Seidling, W., 1999. Spatial structures of a subspontaneous population of Taxus
baccata saplings. Flora 194, 439e451.
Sukopp, H., 2002. On the early history of urban ecology in Europe. Preslia 74,
373e393.
Sukopp, H., Blume, H.-P., Kunick, W., 1979. The soil, flora and vegetation of Berlins
waste lands. In: Laurie, I.E. (Ed.), Nature in Cities. John Wiley, Chichester, pp.
115e131.
Sukopp, H., Werner, P., 1983. Urban environments and vegetation. In: Holzner, W.,
Werger, M.J.A., Ikusima, I. (Eds.), Man’s Impact on Vegetation. Junk Publishers,
The Hague, pp. 247e260.
Sukopp, H., Wurzel, A., 2003. The effects of climate change on the vegetation of
central European cities. Urban Habitats 1, 65e86.
Sullivan, J.J., Williams, P.A., Cameron, E.K., Timmins, S.M., 2004. People and time
explain the distribution of naturalized plants in New Zealand. Weed Technology
18, 1330e1333.
Sullivan, J.J., Timmins, S.M., Williams, P.A., 2005. Movement of exotic plants into
coastal native forests from settlements in northern New Zealand. New Zealand
Journal of Ecology 29, 1e10.
1983
Säumel, I., Kowarik, I., 2010. Urban rivers as dispersal corridors for primarily wind-
dispersed invasive tree species. Landscape and Urban Planning 94, 244e249.
Taramarcaz, P., Lambelet, C., Clot, B., Keimer, C., Hauser, C., 2005. Ragweed
(Ambrosia) progression and its health risks: will Switzerland resist this inva-
sion? Swiss Medical Weekly 135, 538e548.
Thellung, A., 1912. La Flore Adventice de Montpellier. E. Le Maout, Cherbourg.
Thompson, K., Jones, A., 1999. Human population density and prediction of local
plant extinction in Britain. Conservation Biology 13, 185e189.
Truscott, A.M., Soulsby, C., Palmer, S.C.F., Newell, L., Hulme, P.E., 2006. The dispersal
characteristics of the invasive plant Mimulus guttatus and the ecological
significance of increased occurrence of high-flow events. Journal of Ecology 94,
1080e1091.
Tzoulas, K., Korpela, K., Venn, S., Yli-Pelkonen, V., Kazmierszak, A., Niemelä, J., et al.,
2007. Promoting ecosystem and human health in urban areas using green
Infrastructure: a literature review. Landscape and Urban Planning 81, 167e178.
Tüxen, R., 1956. Die heutige potentiell natürliche Vegetation als Gegenstand der
Vegetationskartierung. Angewandte Pflanzensoziologie 13, 5e42.
United Nations, 2008. World urbanization prospects. The 2007 Revision. Highlights.
United Nations, New York. http://www.un.org/esa/population/publications/
wup2007/2007WUP_Highlights_web.pdf.
Vidra, R.L., Shear, T.H., 2008. Thinking locally for urban forest restoration: a simple
method links exotic species invasion to local landscape structure. Restoration
Ecology 16, 217e220.
Vähä-Piikkiö, I., Kurtto, A., Hahkala, V., 2004. Preservation of indigenous vegetation
in urban areas. Landscape and Urban Planning 68, 357e370.
Walther, G.-R., Roques, A., Hulme, P.E., Sykes, M.T., Pysek, P., et al., 2009. Alien
species in a warmer world: risks and opportunities. Trends in Ecology and
Evolution 24, 686e693.
Wang, G., Jiang, G., Zhou, Y., Liu, Q., Ji, Y., Wang, S., Chen, S., Liu, H., 2007. Biodiversity
conservation in a fast-growing metropolitan area in China: a case study of plant
diversity in Beijing. Biodiversity and Conservation 16, 4025e4038.
Wania, A., Kühn, I., Klotz, S., 2006. Plant richness patterns in agricultural and urban
landscapes in central Germanydspatial gradients of species richness. Land-
scape and Urban Planning 75, 97e110.
White, M.A., Nemani, R.R., Thornton, P.E., Running, S.W., 2002. Satellite evidence of
phenological differences between urbanized and rural areas of the eastern
United States deciduous broadleaf forest. Ecosystems 5, 260e273.
Wichmann, M.C., Alexander, M.J., Soons, M.B., Galsworthy, S., Dunne, L., Gould, R.,
Fairfax, C., Niggemann, M., Hails, R.S., Bullock, J.M., 2009. Human-mediated
dispersal of seeds over long distances. Proceedings of the Royal Society B 276,
523e532.
Williamson, M.H., Fitter, A., 1996. The characters of successful invaders. Biological
Conservation 78, 163e170.
Wittig, R., Durwen, K.-J., 1982. Ecological indicator-value spectra of spontaneous
urban floras. In: Bornkamm, R., Lee, J.A., Seaward, M.R.D. (Eds.), Urban Ecology.
Blackwell Scientific Publications, Oxford, pp. 23e31.
Wittig, R., Diesing, D., Gödde, M., 1985. Urbanophob e urbanoneutral e urbanophil.
Das Verhalten der Arten gegenüber dem Lebensraum Stadt. Flora 177, 265e282.
Yasuda, M., Koike, F., 2009. The contribution of the bark of isolated trees as habitat
for ants in an urban landscape. Landscape and Urban Planning 92, 276e281.
Zerbe, S., Choi, I.-K., Kowarik, I., 2004. Characteristics and habitats of the non-native
plant species in the city of Cheon-ju (Korea). Ecological Research 19, 91e98.
Zhao, S.Q., Da, L.J., Tang, Z.Y., Fang, H.J., Song, K., Fang, J.Y., 2006. Ecological conse-
quences of rapid urban expansion: Shanghai, China. Frontiers in Ecology and
the Environment 4, 341e346.
Zhao, J., Oujang, Z., Zheng, H., Zhou, W., Wang, X., Xu, W., Ni, Y., 2010. Plant species
composition in green spaces within the built-up areas of Beijing. Plant Ecology.
doi:10.1007/s11258-009-9675-3.
Zmyslony, J., Gagnon, D., 1998. Residential management of urban front-yard land-
scape: a random process? Landscape and Urban Planning 40, 295e307.