Biol. Rev. (2010), 85, pp. 643667.

643
doi: 10.1111/j.1469-185X.2010.00121.x

A conceptual framework for the colonisation

of urban areas: the blackbird Turdus merula
as a case study
Karl L. Evans , Ben J. Hatchwell, Mark Parnell and Kevin J. Gaston
Department of Animal and Plant Sciences, University of Sheffield, Sheffield S10 2TN, UK

(Received 12 December 2009; revised 15 December 2009; accepted 3 January 2010)

ABSTRACT

Despite increasing interest in urban ecology the factors limiting the colonisation of towns and cities by species from rural
areas are poorly understood. This is largely due to the lack of a detailed conceptual framework for this urbanisation
process, and of sufficient case studies. Here, we develop such a framework. This draws upon a wide range of ecological
and evolutionary theory and the increasing number of studies of how the markedly divergent conditions in urban
and rural areas influence the traits of urban populations and the structure of urban assemblages. We illustrate the
importance of this framework by compiling a detailed case study of spatial and temporal variation in the urbanisation
of the blackbird Turdus merula. Our framework identifies three separate stages in the urbanisation process: (i) arrival, (ii)
adjustment, and (iii) spread. The rate of progress through each stage is influenced by environmental factors, especially
human attitudes and socio-economic factors that determine the history of urban development and the quality of urban
habitats, and by species ecological and life-history traits. Some traits can positively influence progression through one
stage, but delay progression through another. Rigorous assessment of the factors influencing urbanisation should thus
ideally pay attention to the different stages. Urbanisation has some similarities to invasion of exotic species, but the two
clearly differ. Invasion concerns geographic range expansion that is external to the species original geographic range,
whilst urbanisation typically relates to filling gaps within a species original range. This process is exemplified by the
blackbird which is now one of the commonest urban bird species throughout its Western Palearctic range. This is in
stark contrast to the situation 150 years ago when the species was principally confined to forest. Blackbird urbanisation
was first recorded in Germany in 1820, yet some European cities still lack urban blackbirds. This is especially so in
the east, where urbanisation has spread more slowly than in the west. The timing of blackbird urbanisation exhibits a
marked spatial pattern, with latitude and longitude explaining 76% of the variation. This strong spatial pattern contrasts
with the weaker spatial pattern in timing of urbanisation exhibited by the woodpigeon Columba palumbus (with location
explaining 39% of the variation), and with the very weak spatial pattern in timing of black-billed magpie Pica pica
urbanisation (in which location explains 12% of the variation). Strong spatial patterns in timing of urbanisation are more
compatible with the leap-frog urbanisation model, in which urban adapted or imprinted birds colonise other towns and
cities, than with the independent urbanisation model, in which urban colonisation events occur independently of each
other. Spatial patterns in isolation do not, however, confirm one particular model. Factors relating to the arrival and
adjustment stages appear particularly likely to have influenced the timing of blackbird urbanisation. Spatial variation
in the occurrence of urban populations and the timing of their establishment creates opportunities to assess the factors
regulating urbanisation rates, and how the composition of urban assemblages develops as a result. These are major
issues for urban ecology.

Key words: adaptation, blackbird, Columba palumbus, corvids, magpie, Pica pica, Turdus merula, range expansion, species
traits, urbanisation, woodpigeon

* Address for correspondence: (Tel: 0114 2220125; Fax: 0114 2220002; E-mail: karl.evans@sheffield.ac.uk)

Biological Reviews 85 (2010) 643667 2010 The Authors. Journal compilation 2010 Cambridge Philosophical Society
644 K. L. Evans and others

CONTENTS
I. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 644
II. The three stages of the urbanisation process . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 645
(1) Arrival . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 645
(2) Adjustment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 646
(a) Environmental factors . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 646
(b) Mechanisms of adjustment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 647
(c) Species traits . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 648
(3) Spread . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 648
(4) Positive and negative feedbacks between stages . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 648
III. The blackbird Turdus merula as a case study . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 650
(1) Data compilation and assessment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 651
(2) Spatial and temporal patterns . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 651
(3) Independent or leap-frog urbanisation? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 654
IV. Applying the conceptual framework to the blackbird . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 654
(1) Stage 1 - arrival . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 654
(2) Stage 2 - adjustment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 655
(3) Stage 3 - spread . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 657
V. Variation in urbanisation - the blackbird in a broader context . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 657
(1) What proportion of Turdus species are urbanised? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 657
(2) Spatial and temporal patterns in urbanisation in other species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 658
(a) Turdus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 658
(b) Corvus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 659
(c) Columba . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 660
(d) Other genera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 661
VI. Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 661
VII. Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 662
VIII. References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 662
IX. Supporting Information . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 667

I. INTRODUCTION
The world is becoming increasingly dominated by concrete
and other hard surfaces, with about 4% of the terrestrial
area currently urbanised (UNDP, 2000; UN, 2008). This
percentage is much higher in some areas, such as western
Europe, in which urban land comprises approximately 10%
of Britain (Haines-Young et al., 2000; Fuller et al., 2002) and
12% of the Netherlands (CBS, 2003). At a global scale,
urbanisation is the fastest growing land use, and half of
the worlds human population now resides in urban areas
(UN, 2008).
Interest in the ecological effects of urbanisation was rather
limited until recently, but is now growing rapidly. Topics cur-
rently under investigation include (i) how urbanisation influ-
ences habitat quality and other environmental conditions
(e.g. Grimm et al., 2008; Evans, Newson & Gaston, 2009d), (ii)
describing variation in biodiversity along gradients in human
population density or other measures of urbanisation (e.g.
Blair, 1996; Marzluff, 2001; Evans et al., 2006; Evans, Green-
wood & Gaston, 2007; Tratalos et al., 2007), (iii) how species
ecological and life history traits influence their ability to
colonise and survive in urban areas (e.g. Lim & Sodhi, 2004;
Croci, Butet & Clergeau, 2008; Kark et al., 2007; Thompson
& McCarthy, 2008), and (iv) how these traits respond to the
different environmental conditions occurring in urban areas
and thus differ from those exhibited by rural conspecifics (e.g.
Marzluff, 2001; Yeh, Hauber & Price, 2007; Chamberlain
et al., 2009; Evans et al., 2009b,c; reviewed by Evans, in press).
These research agendas have clear connections to a num-
ber of well-established fields including conservation biology,
macroecology, island biogeography and invasion biology.
Whilst invasion biology and urban ecology both concern
changes in species distributions and adaptation to novel envi-
ronments there is a notable contrast between the two. Broadly
speaking, invasion biology concerns processes external to
a species original geographic range, whilst colonisation of
urban areas generally relates to processes internal to a species
range, i.e. filling gaps within range boundaries. There is a lack
of a clear conceptual framework for this internal range expan-
sion process, which is not helped by the paucity of case studies.
Here we first develop such a framework, and then provide a
detailed case study focusing on spatial and temporal variation
in the colonisation of urban areas by the blackbird Turdus
merula (hereafter referred to as the blackbird), which across
much of its geographic range is one of the commonest avian
species in urban areas. This is in stark contrast to the situa-
tion 150 years ago, when it was regarded as a forest specialist
(Luniak, Mulsow & Walasz, 1990). In so doing we compare
the spatial patterns in the occurrence and timing of urbani-
sation exhibited by the blackbird with that of other species.

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A conceptual framework for the colonisation of urban areas 645

II. THE THREE STAGES OF THE URBANISATION
PROCESS
How species become urbanised (i.e. colonise urban areas) is
poorly understood and has received rather little attention in
the ecological literature. There has been some exploration
of the traits that characterise synanthropic species and
urban adapters, primarily focusing on birds (Lim & Sodhi,
2004; Chace & Walsh, 2006; McKinney, 2006; Bonier,
Martin & Wingfield 2007; Kark et al., 2007; for plants see
Thompson & McCarthy, 2008). However, few studies have
attempted to describe how such species initially become
established in urban areas and subsequently expand their
urban populations (but see Vuorisalo et al., 2003; Yeh &
Price, 2004; Rutz, 2008).
An understanding of the urbanisation process has been
hindered by the lack of discrimination between each of its
three distinctive stages. These are rarely explicitly recognised
in the urbanisation literature (but see Marzluff, 2005) despite
much focus being given to their equivalents in invasion
biology (e.g. Williamson, 1996; Kolar & Lodge, 2001; Puth
& Post, 2005). The three stages of the urban colonisation
process are (1) arrival in urban areas, (2) adjustment to the
urban environment, and (3) spread within urban areas and
to neighbouring towns and cities (Fig. 1). Whilst urbanisation
is essentially a continuous process it can usefully be divided
into these three stages because it can succeed or fail at each
stage, with the outcome depending on numerous factors that
are stage specific. These factors are environmental ones and
species ecological and life-history traits. The environmental
factors primarily relate to the spatial configuration and
development history of urban areas and the quality of urban
habitats; they are thus dependent on human attitudes and
socio-economics. Notably, some species traits may have
opposing effects on the rate of progression through different
stages.
(1) Arrival
This stage concerns the initial dispersal to an urban area.
The importance of dispersal in determining species richness is
firmly recognised by island biogeography theory (MacArthur
& Wilson, 1967), and its importance in determining
urban biodiversity has received some initial discussion
(Marzluff, 2005).
The probability of a species initially arriving in an urban
area will increase with both the proportion of its geographic
range that is urbanised, and the time available for this process,
i.e. the history of urban development (Fig. 1). Concerning
species traits, a high population density in surrounding rural
habitats will increase the numerical probability of dispersal
to urban areas. Indeed, annual intraspecific variation in
the frequency of avian species occurring in gardens in the
United Kingdom, most of which are urban, is positively

Fig. 1. Schematic representation of the three stages of the urbanisation process, and the environmental and species characteristics
that increase the probability of each stage being successful. Socio-economics and human attitudes play a major role in shaping the
environmental characteristics, but have been excluded for simplicity. Some factors influence more than one stage, but can have
opposing effects on different stages. See Section II for more details.

Biological Reviews 85 (2010) 643667 2010 The Authors. Journal compilation 2010 Cambridge Philosophical Society
646
correlated with the nature of their population trends in
the wider countryside (Cannon et al., 2005). In addition,
if rural densities are high relative to the environmental
carrying capacity, selection pressures that favour dispersal to
unoccupied areas, such as urban ones, may increase (Herzig,
1995; Loxdale & Lushai, 1999; Lambin, Aars & Piertney,
2001). These selection pressures take numerous forms, but
may be related to deteriorating quality of rural environments;
for example, a succession of severe winters and increased
persecution are associated with the arrival of goshawks
Accipiter gentilis in Hamburg (Rutz, 2008). Similarly, a rapid
increase in the urban population of Australian white ibis
Threskiornis molucca in Sydney in recent decades is associated
with a reduction in the extent of their inland wetland habitat
due to drought, dam construction and agricultural water
extraction (J. Martin, unpublished data).
The successful introduction of species to urban areas
within their natural range indicates that a lack of dispersal,
rather than insufficient habitat, can limit a species
occurrence in urban areas (Cade & Bird, 1990; Martell,
Englund & Tordoff, 2002). Indeed, long-term studies
document a gradual increase in avian species richness in
urban areas as the time available for species to colonise
increases. In some cases this is largely in response to
successional changes in vegetation structure (Vale & Vale,
1976), but in others it is a consequence of colonisation events
that are independent of changes in habitat quality (Abs &
Bergen, 2008). It thus seems likely that the arrival stage can
frequently limit the structure of urban assemblages.
(2) Adjustment
Following arrival in an urban area an individual must cope
with a markedly different environment than its former
rural one. The adjustment stage concerns the processes
through which this adjustment occurs. In this section we
first summarise the characteristics of urban areas, and
thus describe the novel environment to which urbanising
species must adjust. We then outline the processes that
can generate the required adjustments, and conclude by
reviewing examples of the adjustments exhibited by urban
populations.
(a) Environmental factors
Broadly speaking urban and rural environments differ in
seven ways. First, habitat modification is marked in urban
areas, with the magnitude of change often exceeding that
resulting from rural land-uses, such as logging and farming.
Urban centres are characterised by highly fragmented
habitat patches with a simple habitat structure and very
low habitat diversity, although at small spatial scales
habitat diversity may be markedly higher in suburban areas
(Marzluff & Ewing, 2001; Grimm et al., 2008; Evans et al.,
2009d). The magnitude of this divergence between urban
and rural habitats is clearly regulated by human attitudes
and socio-economic factors that determine: (i) the level of
supplementary resource provision, such as avian nest sites
Biological Reviews 85 (2010) 643667 2010 The Authors. Journal compilation 2010 Cambridge Philosophical Society
K. L. Evans and others
and food, which can be an important influence on urban
assemblages (Fuller et al., 2008; Davies et al., 2009; Evans
et al., 2009d), (ii) levels of predation by humans (Vuorisalo
et al., 2003) and their commensals, such as domestic cats
(Baker et al., 2008; Sims et al., 2008), and (iii) the location
of urban areas, which is likely to be non-random with
regard to their ecological situations (e.g. towns and cities
are more likely to be located in areas of relatively high
primary productivity, close to permanent water supplies, or
at sites whose topography promotes defence), and which
could increase the divergence between urban and rural
habitats.
The second major difference between rural and urban
areas is that the latter are characterised by elevated
temperatures and modified rainfall patterns (Lowry, 1998;
Arnfield, 2003; Zhou et al., 2004; Collier, 2006), and thus
tend to have longer growing seasons and altered patterns of
vegetation phenology (White et al., 2002; Zhang et al., 2004;
Neil & Wu, 2006). Third, towns and cities are frequently
more polluted than rural areas, with regard to chemicals,
night-time lights and noise (Sharp, 2002; Small, Pozzi &
Elvidge, 2005; Grimm et al., 2008).
Fourth, native species richness is typically very low in
city centres, although at local scales the richness of some
taxa, such as birds, may peak in suburban areas (Clergeau,
Jokimaki & Savard, 2001; Marzluff, 2001; Tratalos et al.,
2007; Grimm et al., 2008). Fifth, urban areas often contain
increased numbers of exotic species, especially of plants
(Blair, 2001; McKinney, 2006; Smart et al., 2006; de Victor
et al., 2007; Grimm et al., 2008). Sixth, species population
densities typically differ markedly between urban and rural
areas; whilst most species occur at lower densities in urban
areas a significant, but small, proportion of species reach
much higher densities in towns and cities (Tratalos et al.,
2007; Evans et al., 2009d). Seventh, these alterations in
assemblage composition may change the nature of species
interactions and have a major influence on the quality of
urban habitat types, e.g. pollination rates may be reduced due
to a relative lack of insect pollinators (Cheptou & Avendano,
2006). Similarly, urbanisation can dramatically influence
exposure to natural enemies, i.e. predators, pathogens
and parasites, and competitors. There is support for the
contrasting hypotheses that for birds urban areas provide
safe nesting environments (Gering & Blair, 1999; Kosinski,
2001; Newhouse, Marra & Johnson, 2008), and that predator
densities and thus predation rates are higher in urban areas
(Thorington & Bowman, 2003; Jokimaki et al., 2005; Beck
& Heinshon, 2006; Marzluff et al., 2007; Sims et al., 2008),
and meta-analysis shows that no general trends have yet
emerged with regard to the impact of urbanisation on rates
of predation on birds nests (Chamberlain et al., 2009). The
nature of general patterns in the prevalence and intensity of
parasites and pathogens is equally uncertain, and probably
differs markedly with the identity of the parasite/pathogen,
its vector and host (Bradley & Altizer, 2007; Gosselink et al.,
2007; Bradley, Gibbs & Altizer, 2008; Fokidis, Greiner &
Deviche, 2008; Evans et al., 2009c; reviewed by Evans, in
A conceptual framework for the colonisation of urban areas
press). Competitors can also be viewed as natural enemies.
Interspecific competition can structure urban asssemblages
[plants - Ogawa & Mototani (1985); reptiles - Petren &
Case (1996); Ptilinopus fruit doves in New Guinea - Bell
(1986); corvids - Lehmann (2002); sparrows - Bannerman &
Bannerman (1965)], and higher densities of some generalists
in urban areas may increase the intensity of both intra-
and inter-specific competition (Tratalos et al., 2007; Grimm
et al., 2008).
In combination, the above seven factors will determine
the magnitude of contrast between urban and rural
environments, and thus the requirement for adjustment.
For some species urban features may be sufficiently similar
to those of ancestral rural areas to limit the requirement
for adjustment. For example, ruderal plant species that are
adapted to early successional vegetation stages tend to do
particularly well in urban environments due to frequent
anthropogenic disturbance of soils (Gilbert, 1989). More
specifically, plants adapted to growing on rocky surfaces, e.g.
Sedum species, frequently grow on similar surfaces in cities,
such as roof tiles and concrete. Similarly, the breeding sites
of urban peregrines Falco peregrinus, tower blocks and other
tall buildings, have been considered analogous to the cliff
ledges used by rural peregrines (Cade & Bird, 1990).
(b) Mechanisms of adjustment
Given these marked differences between urban and rural
areas it is highly likely that almost all species which colonise
towns and cities from rural sites will require some adjustment
to the novel urban environment. Phenotypic plasticity and
genetic adaptation are both likely to contribute to this
adjustment. In some cases phenotypic plasticity may be
all that is required for the establishment of an urban
population, in which case progress through the adjustment
stage will be rapid. It is more likely, however, that optimal
urban phenotypes will require genetic adaptation. The
requirement for genetic adaptation may arise for three
reasons (Price, Qvarnstrom & Irwin, 2003; West-Eberhard,
2005; Ghalambor et al., 2007). First, even if plasticity results
in an adaptive phenotype it may do so in a more costly
manner than that possible with genetic adaptation, resulting
in selection pressures that ultimately generate genetic change.
Second, plastic responses to the new environment may
approach, but fall short of, the optimum urban phenotype,
resulting in directional selection for extreme phenotypes
which generates genetic adaptation (i.e. genetic assimilation;
Waddington, 1961). Third, the plastic responses induced by
arrival in novel environments, such as urban areas, may
generate additional selection pressures that drive genetic
adaptation of other traits.
Marked intraspecific trait divergence between rural and
urban conspecific populations is increasingly documented
for a wide range of morphological, physiological and
behavioural traits (e.g. Slabbekoorn & den Boer-Visser,
2006; Angilletta et al., 2007; Evans et al., 2009b,c; reviewed
by Evans, in press). These studies provide evidence regarding
the magnitude to which selection pressures differ between
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647
urban and rural environments, but their usefulness in
assessing the relative roles of plasticity and genetic change
in generating trait divergence is limited as most studies
have focused on describing patterns rather than assessing
their causal mechanisms, few determine the adaptive value
of trait divergence in urban areas, and almost none has
distinguished between genetic adaptation and epigenetic
effects. Addressing these limitations will be a major challenge
for urban ecology in the future.
There is, however, some evidence that trait divergence
between urban and rural populations is attributable to
phenotypic plasticity. The frequently documented tendency
for urban birds to have louder and/or higher pitched
songs than rural conspecifics (which may be an adaptive
response to noise pollution in cities, and the low frequency
of anthropogenic noise, but see Nemeth & Brumm, 2009)
appears to have arisen primarily through phenotypic
plasticity (Brumm, 2004; Slabbekoorn & Ripmeester, 2008;
Bermudez-Cuamatzin et al., 2009; Mockford & Marshall,
2009). Vegetation phenology and the timing of avian
reproduction typically occur earlier in urban areas than
rural ones (Zhang et al., 2004; Chamberlain et al., 2009), and
this seems likely to be a plastic response as phenological
advancement in response to climate change is typically
driven by phenotypic plasticity (Brommer et al., 2005;
Brommer, Rattiste & Wilson, 2008; Charmantier et al.,
2008). Furthermore, experiments indicate that the early
reproduction of urban blackbirds, relative to rural ones, is
a plastic response (Partecke, vant Hof & Gwinner, 2004).
Urban birds typically have shorter flight initiation distances
following human disturbance than their rural counterparts
(Cooke, 1980; Campbell, 2006; Mller, 2008). Individual
Eurasian woodpigeons Columba palumbus (hereafter termed
woodpigeon) change their reactions to disturbance when
foraging in urban and rural areas, indicating that at least
in this species changes are a consequence of a flexible
strategy (Tomiaojc, 1976). Plasticity thus appears to be
a major mechanism through which adjustments to urban
environments arise, but it is plausible that the ability to
exhibit plastic responses is under genetic control (Price et al.,
2003; West-Eberhard, 2005; Ghalambor et al., 2007).
Turning to genetic adaptation, experiments in which
urban and rural individuals were raised in identical
conditions are increasingly suggesting that adjustment to
urban areas can have a genetic basis. This is the case for the
smaller body size of urban house sparrows Passer domesticus
(Liker et al., 2008) and increased sedentariness and reduced
stress response of urban blackbird populations (Partecke,
Schwabl & Gwinner, 2006; Partecke & Gwinner, 2007),
although epigenetic effects cannot be ruled out in either of
these studies. The clearest example of genetic adaptation to
urban environments concerns the annual plant Chamaecrista
fasciculate. This species engages in mutualistic interactions
with ants; the plant provides ants with food from extra-
floral nectaries, and the ants feed on herbivorous insects
that attack the plants. Urban plants have significantly fewer
extra-floral nectaries than rural ones as there are fewer
648
herbivorous insects in urban areas. Experiments indicate that
this trait divergence between urban and rural populations
arises entirely from genetic adaptation, and not plasticity or
epigenetic effects (Rios, Marquis & Flunker, 2008).
Species will adapt rapidly to urban environments in
those, probably relatively few, cases where phenotypic
plasticity is sufficient for complete adaptation. Plasticity will
probably also facilitate most other urbanisation events by
increasing the ability to persist in urban environments, whilst
further beneficial adaptation occurs through genetic change.
However, much additional work is required to assess the
extent to which such genetic adaptation facilitates adjustment
to urban environments.
(c) Species traits
Ecologically flexible species are more likely than specialists to
exhibit sufficient plasticity to cope with the novel urban envi-
ronment. Indeed, generalists appear to contribute dispro-
portionately to urban assemblages (Bonier et al., 2007; Kark
et al., 2007; K.L. Evans, D.E. Chamberlain, B.J. Hatchwell,
R.D. Gregory & K.J. Gaston, unpublished data). Behavioural
flexibility may also play a role in adjustment to urban areas.
This trait is linked to relative brain size in birds and mam-
mals, which can predict a species ability to invade areas
outside their natural range, and their response to rapid envi-
ronmental change (Shultz et al., 2005; Sol et al., 2005). So far,
however, studies have failed to find an association between
relative brain size and avian species responses to urban
areas (Kark et al., 2007; K.L. Evans, D.E. Chamberlain, B.J.
Hatchwell, R.D. Gregory & K.J. Gaston, unpublished data).
When adjustment requires genetic adaptation three factors
will determine the rate at which this genetic change occurs.
The first concerns life-history traits that are associated with
rapid rates of evolution, the most prominent ones are sexual
reproduction and fast reproductive rates; notably, both of
these traits facilitate adaptation in invasive exotic species
(Nikolaev et al., 2007; Barrett, Colautti & Eckert, 2008).
Other life-history traits may promote faster evolution, for
example intense sexual selection may do so by increasing
mutation rates, but the evidence for this effect is more
equivocal (Ellegren, 2007; Petrie & Roberts, 2007).
Second, the magnitude of genetic diversity limits the
capacity of populations to adapt to novel conditions (Pujol &
Pannell, 2008). Initial reduction in genetic diversity is likely
to occur as a consequence of founder effects associated with
colonisation of urban areas by a small number of individuals.
To assess the scale of this effect, we compiled data on the rel-
ative genetic diversity of urban and rural populations. These
data certainly indicate that genetic diversity is more limited
in many, but not all, urban populations compared to nearby
rural ones (Table 1). Mutation and immigration from rural
populations may subsequently enable urban populations to
recover genetic diversity, at least partially. This has been
documented for the red fox Vulpes vulpes (Wandeler et al.,
2003). By contrast, urban blackbird populations appear to
become increasingly isolated from nearby rural birds over
time; this pattern may arise if urban birds that disperse to
Biological Reviews 85 (2010) 643667 2010 The Authors. Journal compilation 2010 Cambridge Philosophical Society
K. L. Evans and others
rural areas have reduced fitness relative to urban individuals
that remain in an urban area (Evans et al., 2009a).
The third factor that may regulate the rate of genetic
adaptation to urban areas is gene flow, which can prevent
genetic adaptation to local conditions by swamping evolving
locally adaptive genotypes (Postma & Van Noordwijk, 2005;
Rasanen & Hendry, 2008). To assess the potential for this
in the specific context of urban populations we compiled
data on gene flow between urban and rural populations.
Table 1 shows that such populations are rarely panmictic,
but gene flow is frequently marked, suggesting that there
is often the potential for genetic adaptation to urban
environments to be hindered by gene flow from rural
populations. However, divergent selection pressures can
enable local genetic adaptations to arise despite strong gene
flow (e.g. Garant et al., 2005; Senar et al., 2006), and barriers
to gene flow can evolve rapidly in sympatric populations (i.e.
within dozens of generations; Hendry, Nosil & Rieseberg,
2007). The extent to which gene flow from rural populations
prevents local adaptation to urban environments is thus
uncertain, and assessing the magnitude of this effect is another
major challenge for urban ecology.
(3) Spread
The third and final stage of urbanisation concerns the spread
of populations that have adjusted to urban environments;
broadly speaking, this spread may arise through two
mechanisms. First, through independent urbanisation events,
thus limiting this final stage to spread within an urban area.
Alternatively, urbanisation may follow a leap-frog model
in which urban environments are colonised by individuals
from other urban populations that disperse over or through
rural environments. This could occur because individuals
are adapted to urban habitats, and/or because they are
imprinted on them (Davis, 2008); such a leap-frog model is
more likely to generate a strong spatial pattern in timing of
urbanisation than independent urbanisation events.
In the leap-frog model the spread of urbanised populations
will partly be determined by the spatial configuration of
urban areas, with faster rates of spread expected where
those areas are close together. Species traits influencing
rates of dispersal are likely to play a greater role in
determining the rate of spread of urbanised populations
in leap-frog urbanisation events than independent ones.
Leap-frog urbanisation is also more likely to occur in species
that require genetic change to adjust to urban areas; this is
because the genetic adaptations will already be present in
the colonising individuals from urban populations, but not
in those that originate from rural populations.
(4) Positive and negative feedbacks between stages
Factors influencing the three urbanisation stages may be
correlated with each other. This is perhaps most obviously
the case for environmental factors. Regions with a high
proportion of urbanised land are likely to have longer his-
tories of urban development, and shorter distances between
A conceptual framework for the colonisation of urban areas 649

Table 1. Changes in genetic diversity of urban populations relative to nearby rural ones. A, number of alleles; AR , allelic richness;
Ha, number of haplotypes; H, Shannons diversity index; HaD , haplotype diversity; Ho , observed heterozygozity; GD, Neis gene
diversity; GI, mean number of genotypes per individual; H, Shannons diversity index; PB, percentage of polymorphic bands; PL,
percentage of polymorphic loci. Genetic divergence is measured by FST , and its related metric PT , Neis genetic distance DS (Nei
1978), and the Nei and Li distance coefficient DNL (Nei & Li 1979). Ns, not significant

Change in urban
Group Species Region genetic diversity Genetic divergence Source
Plants the moss Leptodon smithii Italy Ha -36%, PL -20% FST 0.37 Spagnuolo et al. (2007)
McGillivray Grevillea Australia A +95%, Ho +27% n/a Roberts et al. (2007)
macleayana
common dandelion USA GI -41% n/a Keane et al. (2005)
Taraxacum officinale
broad-leaved Scotland A -10%, Ho -29% n/a Hollingsworth & Dickson
helleborine Epipactis (1997)
helleborine
rue-leaved saxifrage Germany ns (GD, H, PB) PT 0.30 Reisch (2007)
Saxifraga tridactlites
downy yellow violet USA ns (H, PL) DNL 0.45 Culley et al. (2007)
Viola pubescens
Invertebrates small white butterfly Japan ns (PL) n/a Takami et al. (2004)
Pieris rapae
the butterfly P. melete Japan ns (PL) n/a
the ground beetle UK ns (AR , Ho ) n/a Desender et al. (2005)
Pterostichus madidus
Belgium ns (AR , Ho ) n/a
the ground beetle Abax UK AR -15%, Ho -88% n/a
ater
Belgium ns (AR , Ho ) n/a
Amphibians the frog Leptodactylus GD -46%, PB -32% DS 0.21 Aruda & Morielle-Versuta
ocellatus (2008)
the frog L. fuscus GD -10%, PB -8% DS 0.04
the frog L. podicipinus GD +50%, PB +37% DS 0.04
the frog L. labyrinthicus GD +11%, PB +6% DS 0.06
common toad Bufo bufo UK A -22%, Ho -15% n/a Hitchings & Beebee (1998)
common frog Rana UK A -10%, Ho -25% n/a Hitchings & Beebee (1997)
temporaria
red-backed salamander Canada A -29%, AR -39%, FST 0.19 Noel et al. (2007)
Plethodon cinerus Ho -34%
Reptiles caissaca viper Bothrops Brazil PL -28% FST 0.19 Dutra et al. (2008)
moojeni
Blandings turtle USA PB -18% n/a Rubin et al. (2001)
Emydoidea blandingii
Birds dark-eyed junco Junco USA AR -41%, Ho -21% n/a Rasner et al. (2004)
hyemalis
blackbird Turdus merula Western AR -3% to -20%, FST 0.00 to 0.04, DS Evans et al. (2009a)
Palearctic Ho +8% to -11%, 0.00 to 0.07,
depending on city depending on city
Eurasian kestrel Falco Poland AR -10%, Ho -8% FST 0.05 Rutkowski et al. (2006)
tinnunculus
Mammals large Japenese field Japan Ha -77%, HaD - 71% FST 0.49 Hirota et al. (2004)
mouse Apodemus
speciosus
red fox Vulpes vulpes Switzerland A -14%, Ho -17% FST 0.04 Wandeler et al. (2003)

urban areas, than less developed regions; all these factors
will positively influence both the arrival and spread stages
of urbanisation (Fig. 1). Positive correlations may also exist
between species traits, for example, species with high pop-
ulation densities are more likely to arrive in urban areas
(Fig. 1) because such species will have a larger number of
dispersive offspring. These abundant species also tend to
have large geographic ranges (Gaston, Blackburn & Lawton,
1997; Blackburn, Cassey & Gaston, 2006), which will further
increase the probability of arrival in urban areas as larger
ranges increase the probability of a species range encompass-
ing towns and cities, and these species tend to have greater
dispersal ability (Gaston, 2003). Species with high population
densities also tend to use resources that are abundant in the

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650 K. L. Evans and others

environment (Gregory & Gaston, 2000), increasing the prob-
ability that they will occur in urban areas, thus reducing the
difference between urban and rural areas and the amount of
adjustment to the former that is required (Fig. 1).
Species traits that influence the urbanisation process
may, however, have opposing effects on the three stages
of urbanisation. For example, high dispersal rates promote
arrival in urban areas (Fig. 1), but may hinder adjustment
for two reasons. First, selection pressure in urban areas
can favour reduced dispersal (Cheptou et al., 2008). Second,
species with strong dispersal abilities will experience marked
gene flow between urban and rural populations that may
hinder the evolution of locally adaptive genotypes (see
Section II. 2c). The rate of progression though each stage
of the urbanisation process may thus correlate negatively.
The presence of such complex interactions between traits
that influence different stages of the urbanisation process
emphasise the importance, when sufficient data are available,
of considering each stage independently when assessing
which ecological or life-history traits promote urbanisation.
III. THE BLACKBIRD TURDUS MERULA
AS A CASE STUDY
A failure to recognise the three separate stages of the
urbanisation process has undoubtedly hindered studies of
urban colonisation, but the lack of detailed case studies is
also important. Such studies are difficult to conduct because
the arrival of many species in urban areas is unlikely to have
been consistently recorded throughout a region. For native
species this is no doubt partly because the process is internal
to the geographic range. Thus an urbanising species will
be familiar to observers from rural areas, perhaps reducing
the likelihood of its initial occurrence in urban areas being
documented. The speed at which urbanisation occurs may
also have an influence on the probability of the process being
recorded, with gradual urbanisation events being less likely
to be noted.
The blackbird is a widespread Palearctic thrush (Fig. 2).
It originally inhabited forests, but is now common in urban
areas throughout much, but not all, of its geographic range
(Luniak et al., 1990). Due to the species conspicuousness
and ease of identification there is a wealth of literature
documenting its arrival in urban areas; it thus provides a
notable exception to the dearth of detailed data on the timing
of urbanisation events. Whilst there have been attempts to
review this literature (Heyder, 1955; Luniak et al., 1990),
much of it remains obscure and highly dispersed, and no
formal analyses of spatial and temporal patterns have been
conducted. Here we synthesise the available information to
provide a formal case study documenting the spatial and
temporal spread of urban populations across much of a
species range. We use these data to analyze spatial patterns
in the timing of urbanisation, to record the rate at which
the habit of occupying urban areas has spread through the
species range, and apply our conceptual framework of the

Fig. 2. The distribution of subspecies in each of the four species in the Eurasian blackbird species complex: (i) Eurasian blackbird:
E At Turdus merula aterrimus, E Az T. m. azorensis, E Ca T. m. cabrerae, E In T. m. intermedius, E Ma T. m. mauritanicus, E Me T. m.
merula, E Sy T. m. syriacus; (ii) Tibetan blackbird: T Ma Turdus maximus maximus; (iii) Chinese blackbird: C Ma Turdus mandarinus
mandarinus, C So T. m. sowerbyi; and (iv) Indian Blackbird I Bo Turdus simillimus bourdilloni, I Ki T. s. kinisii, I Ni T. s. nigropileus and
I Si T. s. simillimus. The position of Australasia, in the inset, has been moved for presentation purposes.

Biological Reviews 85 (2010) 643667 2010 The Authors. Journal compilation 2010 Cambridge Philosophical Society
A conceptual framework for the colonisation of urban areas
urbanisation process to these data to discuss the underlying
factors driving the urbanisation process in relation to its three
different stages.
(1) Data compilation and assessment
Data on blackbird urbanisation dates were obtained
using five methods. First, we consulted material in the
Alexander Library at the Edward Grey Institute of Field
Ornithology, Oxford University; this is arguably the best
ornithological library in the world relating to the focal
region. For each country within the range of the blackbird
we searched country-specific reprint collections, regional
handbooks and local ornithological journals. Second, we
consulted all papers in our research groups collection
on urban ecology and urbanisation. Third, we referred
to the citations in papers obtained using the above
methods. Fourth, we corresponded with local ornithologists.
Finally, using Web of Knowledge (WoK) we conducted
searches using the terms Turdus* AND urban* and
Turdus* AND town*; equivalent searches were conducted
on the Ornithological Worldwide Literature (OWL) database
(http://egizoosrv.zoo.ox.ac.uk/OWL/default.htm), which
focuses on the grey literature.
Following del Hoyo, Elliott & Christie (2005) we recognize
seven sub-species of Turdus merula (Fig. 2), although it is
not always possible to delimit precisely the geographic
range boundaries of the continental subspecies due to
clinal variation (Cramp, 1988). Three taxa have recently
been recognized as distinct species but were previously
considered to be conspecific with T. merula, these are Tibetan
blackbird T. maximus and Indian blackbird T. simillimus (split
from T. merula by both Clement et al., 2000 and del Hoyo
et al., 2005) and Chinese blackbird T. mandarinus (which is
regarded as a separate species by Clement et al., 2000).
Of these only T. mandarinus is urbanised (del Hoyo et al.,
2005; A. Clements, personal communication; J. Hornskov,
personal communication), but we do not consider these taxa
in the rest of this paper.
In total we collated over 120 references to blackbird
urbanisation. We use the names of geopolitical units and
towns/cities as listed in the Times Comprehensive Atlas of the
World (2000). Latitude and longitude for each location were
obtained from the Google Earth mapping service. When
urbanisation dates referred to a specific region, rather than
a town or city, we used the latitude and longitude of the
town closest to the geographic centre of the focal region. The
pattern of urbanisation in each geopolitical unit is described
in Appendix A.
The literature tended to state a range of possible
urbanisation dates. In statistical analyses we used the mid-
point of any such ranges, but discarded data when the date
range exceeded 50 years; although in the vast majority of
cases the date range was much smaller than this. The extent of
genetic differentiation and gene flow between each subspecies
of the blackbird is uncertain; indeed, many avian subspecies
described from the Western Palearctic do not represent
independent genetic units (Zink, 2004; Phillimore & Owens,
Biological Reviews 85 (2010) 643667 2010 The Authors. Journal compilation 2010 Cambridge Philosophical Society
651
2006). We thus conduct analyses using both the entire dataset
(N = 81 locations), and data restricted to the nominate
subspecies (N = 68 locations). Data were insufficient to
conduct separate analyses for other blackbird sub-species.
Sample sizes are smaller than those used to produce country
summaries (Appendix A) due to the repetition of data from
different sources, and exclusion of data with insufficiently
precise urbanisation dates.
A strong spatial pattern in the timing of urbanisation is
less likely to arise if urbanisation occurs independently than
if it occurred in a leap-frog manner (Section II. 3). We thus
assessed the strength of any spatial pattern by regressing
urbanisation date against latitude and longitude using linear,
quadratic and cubic terms for each predictor. Within our
data latitude and longitude were weak predictors of each
other (all data r 2 = 8%; nominate subspecies r 2 = 10%), thus
analyses are not influenced strongly by colinearity. We used
an information theoretic approach to model building, and
followed standard protocols (Burnham & Anderson, 2002;
Johnson & Omland, 2004). We constructed all possible
models given our set of predictor variables, and using
Akaike Information Criteria (AIC) values we calculated each
models weight, i.e. the probability that it provides the most
parsimonious fit to the data. The smallest number of models
whose cumulative weights summed to 0.95 was included in
the 95% confidence set of models, and model averaging was
conducted to assess the influence of predictor variables on
urbanisation date. All analyses were conducted using SAS
(v. 9.1).
We compared the rate of spread of urban blackbird
populations, i.e. internal range expansion, to the rate
of documented avian range expansions that have been
external to a species original geographic range. To facilitate
discussion of the environmental factors determining the rate
of spread of blackbird urbanisation we also assessed regional
variation in its magnitude. A formal analysis of rate of spread
assumes that urban birds moved throughout the species
range from a single locality. This assumption is most likely to
be valid for blackbirds if data are restricted to a single sub-
species. The first records of urbanisation by the nominate
race are from Bamberg and Erlangen (Germany) in 1820,
which are located 34 km apart. We calculated the annual rate
of spread from the mid-point between these two localities to
every urban population of T. merula merula for which we could
estimate the timing of urbanisation. We then calculated the
overall mean rate of spread, and assessed if this differed for
localities to the east and west of the initial urbanisation event.
(2) Spatial and temporal patterns
There is a strong spatial pattern in the timing of blackbird
urbanisation (Fig. 3). When data from all sub-species were
pooled together three models were retained in the 95%
confidence set (Table 2). Model averaging revealed that
total explanatory power was high (r 2 = 76%). Urbanisation
date was not strongly related to latitude (model averaged
partial r 2 = 7%; Fig. 4A), but was strongly related to
longitude in a positive decelerating manner (model averaged
652 K. L. Evans and others

Fig. 3. Timing of urbanisation in the Eurasian blackbird Turdus merula. Each panel represents a different time period (A) 18201849,
(B) 18501879, (C) 18801909, (D) 19101939, and (E) 1940 onwards. Circles represent urban locations for which data are
available. Black circles indicate the presence of urban blackbirds, and grey ones their absence. In E a grey circle represents the
absence of urban blackbirds by 2000. The position of Iceland, in the inset, has been adjusted for presentation purposes.

Biological Reviews 85 (2010) 643667 2010 The Authors. Journal compilation 2010 Cambridge Philosophical Society
A conceptual framework for the colonisation of urban areas 653

Table 2. Results of multiple regression analyses of spatial patterns in the urbanisation dates of blackbird Turdus merula, magpie Pica
pica, and woodpigeon Columba palumbus. The linear, quadratic and cubic terms of latitude (lat) and longitude (long) were used as
predictors. Model selection followed an information theoretic approach, and for each response variable we present all models in the
95% confidence set

lat lat2 lat3 long long2 long3 model weight model r 2 (%) partial r 2 lat (%) partial r 2 long (%)
T. merula all populations + + 0.88 76.15 7.34 41.50
+ + + 0.04 76.65 7.84 41.08
+ + 0.03 72.81 4.00 72.73
T. m. merula only + + + 0.37 73.28 5.58 24.99
+ + 0.22 71.03 10.36 22.74
+ + 0.17 71.39 3.69 33.95
+ + 0.10 73.27 5.76 24.98
+ + 0.04 71.36 3.85 33.92
+ + 0.02 67.45 6.78 30.01
+ + 0.02 74.07 6.23 25.78
Pica pica + 0.65 13.45 0.00 13.45
+ 0.20 13.49 0.04 13.45
0.12 0.02 0.02 0.00
Columba palumbus + + + 0.40 39.85 7.35 35.20
+ + 0.18 36.15 3.65 35.38
+ + + + 0.16 39.86 7.36 35.21
+ + + 0.07 36.17 3.66 35.40
+ 0.06 34.93 2.43 34.17
+ + + + 0.06 45.13 7.56 40.48
+ 0.02 34.96 2.46 34.20

(A) (B)

(C) (D)

Fig. 4. Relationships between known dates of Eurasian blackbird Turdus merula urbanisation across the species range and (A) latitude
and (B) longitude, and for only the nominate sub-species with (C) latitude and (D) longitude. Note the variation in the scale of the x
axes.

Biological Reviews 85 (2010) 643667 2010 The Authors. Journal compilation 2010 Cambridge Philosophical Society
654
partial r 2 = 43%; Fig. 4B). When restricting data to the
nominate subspecies seven models were retained in the
95% confidence set (Table 2). Model averaging revealed
that total explanatory power was high (r 2 = 72%). Latitude
(partial r 2 = 6%; Fig. 4C) explained less of the variation in
urbanisation date than longitude (partial r 2 = 27%; Fig. 4D),
which was related to urbanisation date in a more linear
manner than when data from all subspecies were used.
Blackbirds were thus urbanised earlier in the west of their
range than in the east, and to a lesser extent were urbanised
earlier in the south than in the north. In nine countries
occupied by the nominate subspecies the timing of blackbird
urbanisation is closely associated with an increase in the
species geographic range or population size in the wider
landscape (Appendix A). When these data are excluded
from analyses the timing of urbanisation remains strongly
associated with longitude (model averaged partial r 2 = 49%),
and poorly associated with latitude (model averaged partial
r 2 = 0.4%).
The overall mean rate of spread of blackbird urbanisation
within the range of the nominate subspecies was 7.7
0.7 km per year (mean S.E.M). Spread was faster to
the west (10.1 1.7 km per year, N = 23) than to the east
(6.5 0.6 km, N = 42); the 84% confidence intervals of these
estimates do not overlap and the difference is thus statistically
significant at = 0.05 (Payton, Greenstone & Schenker,
2003). This pattern remains when data from countries in
which the blackbird has experienced more general range
expansions are excluded from analyses (westerly spread
10.0 2.0 km per year, N = 20; easterly spread 5.3
0.7 km per year, N = 32; again 84% confidence intervals do
not overlap).
(3) Independent or leap-frog urbanisation?
The strong spatial pattern in the timing of blackbird
urbanisation is predicted by the leap-frog urbanisation model
in which individuals from an established urban population
colonise other built-up areas either because they are adapted
to such areas, or because they are imprinted on them.
However, spatial patterns in the timing of urbanisation
cannot, in isolation, confirm a particular urbanisation
model, and it is important to consider other evidence. The
introduction of rural blackbirds to previously uncolonised
urban areas failed in Bialystok and Olsztyn (north-east
Poland), whilst the introduction of urban blackbirds to other
uncolonised cities, Lublin in south-east Poland and Kiev in
Ukraine, succeeded (Grachyk et al., 1975; Graczyk, 1982).
This suggests that urban blackbirds, relative to rural ones,
are more likely to colonise built-up areas. Indeed, urban
blackbirds exhibit some adjustments to towns and cities that
probably arise, at least in part, from genetic adaptation
(Partecke et al., 2006; Partecke & Gwinner, 2007). The
genetic basis of such urban adaptations may explain why
introductions of rural blackbirds to urban environments
failed; however, an effect of climate cannot be ruled out,
as the failed introductions were also to colder environments
than the successful ones.
Biological Reviews 85 (2010) 643667 2010 The Authors. Journal compilation 2010 Cambridge Philosophical Society
K. L. Evans and others
In some regions, urbanisation was probably independent
from that in Europe. The most likely candidates
for populations exhibiting independent urbanisation are
T. merula azorensis and T. merula cabrerae, subspecies that
are confined to Atlantic islands (the Azores, Canaries and
Madeira) that are outside the normal wintering range of other
blackbird populations. Another strong case for independent
urbanisation is the north-west African subspecies, T. merula
mauritanicus. Urban blackbirds were common in this region
within 70 years of the first urbanisation events being
documented in Europe, at localities approximately 1,700 km
away. The mean adult and natal dispersal distances of
blackbirds are 3.2 km and 3.3 km respectively (Paradis et al.,
1998). Whilst dispersal over much longer distances occurs,
it is a rare event. Paradis et al. (1998) recorded a maximum
dispersal distance of 300 km between breeding sites for adults
(N = 1,800), and 350 km for first-year birds (N = 2,200).
Given the very small number of urban blackbirds in Europe
at the start of the urbanisation process it is highly unlikely
that sufficient individuals could have colonised north-west
Africa and started the urbanisation process there. Indeed,
the mean rate of spread to the west, i.e. in the direction of
Tunisia, from the first urbanisation event is just 10 km per
year. A much faster rate of spread (approximately 25 km
per year) would be required if urbanised blackbirds from the
site of the first documented urbanisation events in Germany
were to have reached Tunisia at the time of blackbird
urbanisation in that country. Therefore, in combination,
the data strongly suggest that within the range of T. merula
urbanisation happened independently in north-west Africa
and Europe. Urbanisation events in other subspecies (i.e.
T. merula aterrimus, intermedius and syriacus) would also be
independent if these taxa are genetically distinct from the
nominate subspecies. There is no evidence for this, however,
and many avian subspecies that have been described from
Europe are not genetically distinct (Zink, 2004; Phillimore &
Owens, 2006).
Therefore, despite the strong spatial signal in the
urbanisation of blackbirds across their Western Palearctic
range, independent colonisation of urban areas has almost
certainly occurred in parts of that range. Furthermore,
analysis of the genetic structure of paired urban and rural
blackbird populations does confirm that urbanisation has
often occurred through multiple independent colonisations
of towns and cities from nearby rural populations (Evans
et al., 2009a).
IV. APPLYING THE CONCEPTUAL
FRAMEWORK TO THE BLACKBIRD
(1) Stage 1 - arrival
Throughout Europe, cities existed long before blackbirds
and other avian species became urbanised. For example,
London occupied 2 km2 in 1300, over 4 km2 by 1673, and
12.5 km2 by 1819 (Shepherd, 1300; Blome, 1673; Howgego,
A conceptual framework for the colonisation of urban areas
1978; Fitter, 1990), yet blackbirds did not become common in
English city centre gardens until the 1920s and 1930s (Simms,
1978; de Laet, 2000). Below we discuss the mechanisms
that may have contributed to the long time lag between
the development of urban areas and their colonisation by
blackbirds.
The first stage of urbanisation is arrival in urban areas
(Fig. 1). In much of Europe blackbirds appear to have been
reasonably abundant in rural areas for hundreds of years
before urbanisation (Appendix A), and it is unlikely that a
lack of dispersal into built-up areas limited the potential for
urbanisation to commence. However, this is not the case in
northern and eastern Europe where a marked increase in the
size of blackbird populations and their range has occurred in
nine countries (Belarus, Denmark, Estonia, Finland, Iceland,
Latvia, Lithuania, Norway and Sweden). This process started
in Denmark and Norway approximately 150 years ago, and
spread eastwards and northwards; blackbirds are currently
much commoner throughout Scandinavia, the Baltic states
and Iceland than previously. This range expansion is linked
to climatic amelioration and habitat conversion (Jarvinen
& Vaisanen, 1979; Burton, 1995; Newton, 2003). Also,
in Denmark the Game Act of 1849 prohibited catching
blackbirds and the species population increase commenced
shortly after the acts introduction (Lppenthin, 1967). In
eight of the nine countries (Belarus is the exception) in which
blackbirds have markedly increased their range, urbanisation
has occurred within two decades of the range expansion. This
suggests that a delayed arrival in urban areas, due to a small
number of potential colonisers in the wider landscape, may
have slowed the urbanisation process in north-east Europe.
It is notable, however, that the strong spatial pattern in the
timing of urbanisation remains even when data from these
nine countries are excluded from analyses.
Other Turdus species also exhibit a close similarity in the
timing of range expansion and urbanisation, including the
fieldfare T. pilaris (Section V. 2a) and kurrichane thrush
T. libonyana (Vernon, 1997). More generally, urbanisation
of the woodpigeon, magpie, sparrowhawk Accipiter nisus and
peregrine in the UK is associated with population increases,
often following recovery from persecution and accidental
poisoning (Newton, 1986; Marchant et al., 1990; Ratcliffe,
1993; Wilson, 1993). Similarly, the number of goshawk
observations in Hamburg and surrounding rural areas
increased simultaneously (Rutz, 2008), and urbanisation
of the black woodpecker Dryocopus martius in Hungary is
associated with range expansion (Gorman, 1998). It is thus
plausible that in many cases the initial arrival stage of the
urbanisation process may be a response to high population
densities in rural habitats increasing the number of potential
colonisers, and possibly the selection pressure to colonise
other areas (Fig. 1). This process has some similarities to the
buffer effect (Brown, 1969), in which a species colonises new
habitats when the preferred ones are saturated. However,
the higher densities of blackbirds, and some other species,
in urban areas than rural ones (Tratalos et al., 2007; Evans
et al., 2009d), and the lack of a significant difference in
Biological Reviews 85 (2010) 643667 2010 The Authors. Journal compilation 2010 Cambridge Philosophical Society
655
annual productivity between urban and rural blackbird
populations (Chamberlain et al., 2009), suggests that urban
habitats are currently no less preferable than rural ones. This
was probably also true in the past as there are historical
reports of high blackbird densities in urban areas (Heyder,
1955; Luniak et al., 1990).
Blackbirds frequently first arrived in urban areas in the
winter (e.g. Glaser, 1871; Fallon, 1875; le Rabe, 1886;
Collett, 1921; Heyder, 1955; Graczyk, 1959; Rabose et al.,
1995). A similar situation still applies; for example, blackbirds
occur in the centre of Montpellier (France) in winter, but
do not yet breed there. Other avian studies also document
that breeding in towns is preceded by winter occurrences,
examples include the hill pigeon Columba rupestris (del Hoyo,
Elliott & Sargatal, 1997), dark-eyed junco Junco hyemalis
(Yeh, 2004), and mallard Anas platyrhynchos (Pulliainen, 1963).
Counter examples are available, such as the woodpigeon
(Tomiaojc, 1976), but at least in temperate regions the typical
first step in avian urbanisation may be wintering in urban
areas. Temperate urban areas are likely to be particularly
advantageous in winter, compared to rural ones, due to
higher temperatures, and human activities that increase food
availability. The latter may occur directly through provision
at bird feeders, or indirectly by activities such as planting of
trees that produce berries or fruit, and clearing areas of snow
that facilitates foraging by ground-feeding birds.
Many blackbirds in northern and eastern Europe are
migratory, and this proportion was even higher in the past
than is the case today (Rivalan et al., 2007; van Vliet, Musters
& ter Keurs, 2009). The small number of resident blackbirds
in north-east Europe, prior to urbanisation, may thus have
further limited the number of individuals that could have
undertaken the initial urbanisation step by wintering in
local towns. It is also possible that urbanisation began
in central Europe due to the combination of a sufficient
number of resident blackbirds that could start the process,
and sufficiently cold winters to provide a greater selective
advantage in doing so compared to warmer regions elsewhere
within the species range.
(2) Stage 2 - adjustment
The lag between the development of urban areas and
blackbird urbanisation may also have arisen if time is
required for blackbirds to adjust to urban areas. In areas of
Europe where its arrival is unlikely to have been limiting this
time-lag may have occurred because of factors that influence
the rate of adjustment to urban environments (Fig. 1). First,
urban environments are dynamic and their characteristics
may have changed over time in a manner that favoured
blackbirds, and thus increased the extent to which they were
pre-adapted to urban areas (Fig. 1). Second, urbanisation
may have required genetically determined adaptations that
took time to evolve (Fig. 1). These mechanisms are not
mutually exclusive and we consider each in turn.
The dynamic nature of towns and cities means that habitat
availability and quality may have improved over time. In
1895 blackbirds were rarely present in Neisse (Germany)
656
but numbers increased soon after the establishment of parks,
and by 1906 the species was a common breeder (Kollibay,
1906). Blackbirds reach higher densities in urban areas with
more green space and a greater proportion of trees and
shrubs, although the strength of such relationships appears
to be scale dependent (Karlsson & Kallander, 1977; Mason,
2003). Increased availability of bushy vegetation also appears
to promote recruitment into the breeding population, and
such areas are preferred as breeding territories (Fernandez-
Juricic & Telleria, 1999; Wysocki et al., 2004). It is thus highly
likely that changes in the nature of some urban areas would
have increased their suitability for blackbirds. For example, in
Britain many parks were created in urban areas between the
1830s and early 1900s (Jordan, 1994; Taylor, 1995), which
broadly coincides with when the species became urbanised
(Appendix A).
The extent to which such changes were essential for
blackbird urbanisation is, however, debatable. Historical
descriptions of urban areas suggest that suitable habitat for
breeding blackbirds was present many decades before they
colonised cities. For example, Aeneas Sylvius Piccolomini
described the presence of rich flower gardens and orchards
in the German towns of Erfurt, Frankfurt, Nuremberg and
Augsburg in 1485 (Landau & Schneider, 1928). Similarly,
large parks in central London, such as St. Jamess Park and
Hyde Park, were broadly similar in the early 1700s to their
appearance today (Fitter, 1990). The successful introduction
of blackbirds to urban environments which previously lacked
them, such as Poznan and Kiev (Grachik et al., 1975;
Graczyk, 1982), is clear evidence that urbanisation does
not always occur as soon as conditions are suitable. In
addition, the blackbird and song thrush Turdus philomelos
have similar habitat requirements (their densities in urban
areas of the United Kingdom are positively correlated; Evans
et al., 2009d), but song thrushes are urbanised in Moscow
and blackbirds are not. This strongly suggests that a lack
of suitable habitat has not prevented the urbanisation of
blackbirds in some regions. A greening of urban areas
also appears unlikely to explain the timing of urbanisation
in some woodpigeon populations. Both the blackbird and
woodpigeon are originally woodland species, and their
densities in urban areas of Britain are strongly correlated
(Evans et al., 2009d), suggesting that they have similar habitat
preferences in cities. However, the blackbird colonised
Munich 90 years before woodpigeons did, suggesting that
suitable habitat was present for the latter species many
years prior to their arrival. Similarly, woodpigeons arrived
in Prague and Vienna respectively 55 and 60 years after
blackbirds. Therefore, although it is probable that improved
habitat conditions may have partly facilitated urbanisation
in some cases, this seems unlikely to have been a general
factor.
Another factor contributing to the dynamic nature of
urban areas concerns changes in human activities related
to wildlife. It is possible that anthropogenic supplementary
feeding facilitated urbanisation in some regions, as is thought
to be the case for mallard wintering in urban areas in Finland
Biological Reviews 85 (2010) 643667 2010 The Authors. Journal compilation 2010 Cambridge Philosophical Society
K. L. Evans and others
(Pulliainen, 1963). One British study has demonstrated that in
urban areas the total breeding density of all bird species that
take supplementary food, including the blackbird, increases
with the density of feeding stations (Fuller et al., 2008).
However, it is uncertain if such patterns are causal as they
may arise because people provide more food where there
are more birds. Unfortunately, it is not possible to correlate
the timing of blackbird urbanisation with when feeding of
garden birds started because insufficient data are generally
available [although it commenced in Finland at the end of the
19th Century (Vuorisalo, Lahtinen & Laaksonen, 2001), i.e.
a few decades prior to blackbird urbanisation]. Moreover,
the provision of supplementary food is unlikely to have
been a consistently important factor in promoting blackbird
urbanisation as this activity is still rare in many regions of
eastern Europe that contain urbanised populations.
Concerning less positive human attitudes, the lack of
urban blackbirds in south-east France has previously been
attributed to the high levels of persecution of Turdus species
in this region (Ingram, 1926). Other authors have also linked
urbanisation to a change in levels of human persecution,
albeit in combination with habitat alteration. For example,
Holloway (2002) states that at the end of the 19th Century
a switch from using suburban gardens for growing food
to planting of ornamental shrubs and plants that provided
better nesting sites was associated with reduced persecution,
and probably allowed blackbirds to become tamer, and thus
to start adapting to urban environments. Similarly, in the
1930s in Belgium, the blackbird was much commoner in
urban gardens with shrubs than in village gardens where
vegetables were grown, perhaps due to improved habitat
structure and reduced persecution (Morbach, 1939). Rural
persecution is certainly associated with the occurrence of
urbanised populations of other species (Section II. 1), but
there is insufficient evidence to suggest that the timing of
blackbird urbanisation is a consequence of changes in human
attitudes.
The second suite of factors influencing the rate of progress
through the adjustment stage of urbanisation concerns the
extent to which genetic adaptation is required, and the rate
at which such adaptations can arise. Avian body size has a
strong genetic basis (Glazier, 2002), and some populations of
urban blackbirds are morphologically divergent from their
rural counterparts (Gregoire, 2003). However, blackbirds do
not appear to require any specific morphological adaptations
to become established in urban areas (Evans et al., 2009b).
By contrast, urban blackbirds have a reduced corticosterone
stress response than rural ones, and rearing urban and
rural individuals in a shared environment suggests that these
differences have a genetic basis (Partecke et al., 2006). Also,
a number of anecdotal reports state that urban blackbirds
are more sedentary than rural ones (Bruch et al., 1978;
Luniak, 2004). Experimental studies have also found reduced
migratory restlessness in urban blackbirds relative to forest
ones (Graczyk, 1961), and stable isotope analysis of migratory
behaviour of paired urban and rural blackbird populations
indicates that the former are less migratory (K.L. Evans,
A conceptual framework for the colonisation of urban areas
J. Newton, K.J. Gaston, S. Sharp, A. McGowan & B.J.
Hatchwell unpublished data). There is good evidence from
many passerines that such changes in migratory strategy
require genetic change (Berthold et al., 1992), and rearing
experiments suggest that increased sedentariness in urban
blackbirds partly has a genetic basis (Partecke & Gwinner,
2007). Therefore, at least some of the changes documented
in urban blackbird populations probably require genetic
adaptation, but early maternal effects cannot be excluded
(Partecke et al., 2006; Partecke & Gwinner, 2007). Whether
such changes were requirements for urbanisation to succeed,
or if they arose following urbanisation is unknown. It is
thus not possible to conclude that the need for genetic
adaptation has contributed to the time required for blackbirds
to adjust to urban areas, but such effects are certainly
possible.
(3) Stage 3 - spread
We provide the first estimate of the rate of avian range
expansion that is internal to a species geographic range, i.e.
colonisation of unoccupied areas within a species original
range. Assuming a leap-frog model we find that the mean rate
of spread of urban blackbirds, 8 km per annum, is very similar
to the median value documented for avian range expansions
that are external to species geographical ranges (6 km per
annum, Table 3). This similarity may arise because both
internal and external range expansion involve dispersal and
adaptation to novel areas with different ecological conditions
and may thus be limited by similar processes. As in studies
of external range expansion we find geographical variation
in the rate of spread. Urbanisation of T. m. merula has been
more rapid in the west of its range than in the east. Part of
this variation may be due to delayed arrival in eastern urban
areas. This is unlikely to be a consequence of the species
former rarity in parts of this region, because the rate of
spread is still slower in the east when such areas are removed
from the analysis. Spatial variation in migratory strategies
(Section IV. 1) may also contribute to variation in the rate
of spread, and the less urbanised nature of this region may
have further slowed the urbanisation process by reducing the
number of focal points at which urbanisation could occur
(Fig. 1).
V. VARIATION IN URBANISATION - THE
BLACKBIRD IN A BROADER CONTEXT
We place our analysis of blackbird urbanisation in a wider
context in two ways. First, we assess the extent to which
other Turdus species are urbanised, and compare this to the
proportion of urbanised species in other avian genera at
a global scale. We use these data to facilitate a discussion
of the environmental and biological traits that influence
the probability of urbanisation. Second, in order to assess
whether the patterns we describe for the blackbird can be
considered general, we assess the extent to which other
Biological Reviews 85 (2010) 643667 2010 The Authors. Journal compilation 2010 Cambridge Philosophical Society
657
urbanised species exhibit spatial variation in the occurrence
and timing of their establishment.
(1) What proportion of Turdus species are
urbanised?
It is a common perception that avian genera differ in the
extent to which they are urbanised, but there has been
little formal assessment of variation in the proportion of
urbanised species in different genera, and of whether this
is consistent across faunal regions. Here, we conduct a
preliminary assessment of these issues. We compare the
Turdus genus to Corvus, which is often considered to be
one of the most highly urbanised avian genera (Marzluff,
2001), and Columba which contains one of the most highly
urbanised bird species in the world, the feral pigeon
C. livia.
Data on habitat use by each species of Turdus, Corvus and
Columba in their native ranges were obtained by consulting
recently published guides to each genus (Turdus: Clement
et al., 2000; del Hoyo et al., 2005; Corvus: Madge & Burn,
1994; Columba: del Hoyo et al., 1997; Gibbs, Barnes &
Cox, 2001). We also searched Web of Knowledge (WoK)
and the Ornithological Worldwide Literature (OWL) database
using the terms Turdus* AND urban* and Turdus* AND
town* and their equivalents for Corvus and Columba. We
classified species as being fully urbanised if their habitat
descriptions included towns, cities, urban or suburban
areas, and being partly urbanised if they occurred in large
gardens, villages, or on the edge of towns, cities or urban
areas.
Globally, there was marked variation in the proportion
of each genus that was urbanized. Urbanisation was most
prevalent in Corvus and least common in Columba (Table 4).
Whilst this lends some support to the assertion that Corvus is
amongst the most highly urbanised of avian genera (Marzluff,
2001), the genus does not contain significantly more fully
urbanised species than the other taxa ( 2 = 1.50, P =
0.47), and the trend for Corvus to be more urbanised still
only tended towards significance when partially and fully
urbanised species were combined ( 2 = 4.91, P = 0.09).
It is thus perhaps more appropriate to consider Corvus as
just one example of a highly urbanised genus. Corvids
are particularly intelligent, as evidenced by their relatively
large brain size, and the high rate at which they acquire
novel feeding adaptations (Emery, 2006). This behavioural
flexibility may be one factor contributing to their success in
urban areas (Fig. 1).
The three genera differed with respect to which geographic
region contained the greatest proportion of urbanised species
(Table 4). The clearest example of this is that the Indo-Malay
region supports the greatest proportion of fully urbanised
Corvus species (50%), but that none of its ten Turdus species or
its six Columba species are fully urbanised. The mechanisms
driving this variation are uncertain, but variation among
genera in the spatial congruence between species geographic
ranges and urban areas may contribute to the pattern.
Indeed, most Turdus species in the Indo-Malay region occur
658 K. L. Evans and others

Table 3. Mean rates of spread for avian range expansions external to species original ranges. Data include natural range expansions
and those following introduction to non-native locations. When a range of rates is given this represents geographical variation,
for bronzed cowbird and great-tailed grackle it also includes temporal variation. In some species estimates vary between studies.
Additional papers describe or map range changes for other species, but no attempt has been made to calculate rates of change from
such data

Species Exotic Location Rate km year1 Source

Bewicks wren Thryomanes bewickii no North America 0.4 Hitch & Leberg (2007)
red kite Milvus milvus no Wales 0.5 Lensink (1997)
osprey Pandion haliaetus no Scotland 0.9 Lensink (1997)
tree sparrow Passer montanus yes North America 1 Williamson (1996)
summer tanager Piranga rubra no North America 1.6 Hitch & Leberg (2007)
marsh harrier Circus aeruginosus no England 2 Lensink (1997)
carolina chickadee Poecile carolinensis no North America 2.1 Hitch & Leberg (2007)
yellow-billed sapsucker Sphyrapicus varius no North America 2.2 Hitch & Leberg (2007)
buzzard Buteo buteo no UK & Holland 0.32.3 Lensink (1997)
Egyptian goose Alopochen aegyptiacus yes Holland 3 Lensink (1998)
sparrowhawk Accipiter nisus no UK & Holland 2.43.1 Lensink (1997)
blue-winged warbler Vermivora pinus no North America 3.3 Hitch & Leberg (2007)
bobolink Dolichonyx oryzivorous no North America 3.4 Hitch & Leberg (2007)
yellow-rumped warbler Dendroica coronata no North America 3.5 Hitch & Leberg (2007)
hooded warbler Wilsonia citrina no North America 4.4 Hitch & Leberg (2007)
peregrine Falco peregrinus no UK 1.14.5 Lensink (1997)
goshawk Accipiter gentiles no UK & Holland 1.54.5 Lensink (1997)
tree sparrow Passer montanus yes Australia 5 Blakers et al. (1984)
fish crow Corvus ossifragus no North America 5.1 Hitch & Leberg (2007)
willow flycatcher Empidonax trailii no North America 5.2 Hitch & Leberg (2007)
Swainsons thrush Catharus ustulatus no North America 5.4 Hitch & Leberg (2007)
Kentucky warbler Opornis formosus no North America 5.7 Hitch & Leberg (2007)
white-winged crossbill Loxia leucoptera no North America 6.1 Hitch & Leberg (2007)
Bachmans sparrow Aimophila aestivalis no North America 6.8 Hitch & Leberg (2007)
pine siskin Carduelis pinus no North America 7.1 Hitch & Leberg (2007)
alder flycatcher Empidonax alnorum no North America 7.4 Hitch & Leberg (2007)
golden-winged warbler Vermivora chrysoptera no North America 8.4 Hitch & Leberg (2007)
Inca dove Columbina inca no North America 9.8 Hitch & Leberg (2007)
black-billed cuckoo Coccyzus erythropthalmus no North America 10.6 Hitch & Leberg (2007)
blue-grey gnatcatcher Polioptila caerulea no North America 12.1 Hitch & Leberg (2007)
bronzed cowbird Molothrus aenus no America 6.213.8 Kostecke et al. (2004)
serin Serinus serinus no Europe 16.5 Margalef (1974)
yellow-breasted bunting Emberiza aureola no Europe 16.5 Timofeeff-Ressovsky (1940)
house sparrow P. domesticus yes North America 16.8 van den Bosch et al. (1992)
Brewers blackbird Euphagus cyanocephalus no America 17.5 Stepney & Power (1973)
house finch Carpodacus mexicanus yes North America 18.9 Mundinger & Hope (1983)
house finch Carpodacus mexicanus yes North America 20.8 Veit & Lewis (1996)
great-tailed grackle Quiscalus mexicanus no North America 12.722.6 Wehtje (2003)
house sparrow Passer domesticus no Europe 27.9 van den Bosch et al. (1992)
shiny cowbird Molothrus bonariensis no Caribbean 8.330.6 Post & Wiley (1977)
serin Serinus serinus no Europe 40 Olsson (1971)
collared dove Streptopelia decaoto no Europe 43.7 van den Bosch et al. (1992)
starling Sturnus vulgaris yes North America 91.6 van den Bosch et al. (1992)
cattle egret Bubuculus ibis no Americas 93.3 Kuyt (1971), Rodriguez- Mata (1973)
house sparrow Passer domesticus yes Australia 6100 Blakers et al. (1984)
cattle egret Bubuculus ibis no Americas 106.2 van den Bosch et al. (1992)
starling Sturnus vulgaris yes North America 200 Grosholz (1996)

in the extreme north where urban areas are rare, thus limiting
the potential for species to become urbanised, whilst most
Corvus species in this region have large geographic ranges
that overlap with many cities. This variation emphasises the
potential importance of the history of urban development in
determining the potential for a species to become urbanised
(Fig. 1).
(2) Spatial and temporal patterns in urbanisation in
other species
(a) Turdus
In many Turdus species urban populations do not occur
throughout a species geographic range. For example,
the mistle thrush T. viscivorus is absent as an urban

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A conceptual framework for the colonisation of urban areas 659

Table 4. Regional variation in the proportion of partly and fully urbanised Turdus, Corvus and Columba species in their native ranges.
The summed number of species in each region is greater than the global total as some species occur in more than one region. Data
are primarily from Madge & Burn (1994) for Corvus; from Clement et al. (2000) and del Hoyo et al. (2005) for Turdus; and from del
Hoyo et al. (1997) and Gibbs et al. (2001) for Columba. Biogeographic realms were defined following Olson et al. (2001)

Turdus Corvus Columba

% fully or % fully or % fully or
%fully partly % fully partly % fully partly
# species urbanised urbanised # species urbanised urbanised # species urbanised urbanised
Global 70 29 41 46 33 57 50 20 28
Afrotropics 9 33 44 6 17 33 13 15 39
Australasia 1 0 100 16 25 38 3 33 67
Indo-Malay 10 0 20 4 50 75 6 0 17
Nearctic 1 100 100 7 43 57 1 100 100
Neotropics 33 30 48 4 25 75 17 18 18
Oceania 0 n/a n/a 2 0 0 0
Palearctic 19 26 32 11 36 82 12 25 25

breeding species in many towns in eastern Europe, including
St. Petersburg (Russia; Khrabryi, 2005) and Lublin (Poland;
Biaduo, 2005), and is rare in many other towns in this
region (e.g. Warsaw, Poland; Luniak, 2005). By contrast, it
frequently occurs in urban areas in western Europe, including
Belgium (Weiserbs & Jacob, 2005) and the United Kingdom
(Mason, 2000; Evans et al. 2009d). Similarly, the song thrush
T. philomelos is absent from Warsaw (Luniak, 2005), yet is
an urban breeder in towns to the east (e.g. St. Petersburg,
Khrabryi, 2005; Moscow, Konstantinov & Zakharov, 2005;
Lublin, Biaduo, 2005), and the west (e.g. in Hamburg:
Mulsow, 2005; in Brussels: Weiserbs & Jacob, 2005; and
throughout the United Kingdom: Mason, 2000; Evans et al.,
2009d). Species also vary in their timing of urbanisation,
for example the song thrush colonised German cities in the
1930s, several decades later than in the United Kingdom
(Fitter, 1990).
More detailed data on the timing of urbanisation are
available for the fieldfare, which has only recently become
urbanised in parts of its range. It colonised Krakow
(south-east Poland) in the 1960s (Harmata, 1979), and Lublin
(south-east Poland) in the late 1970s, when it started to breed
in parks and cemeteries before moving to more residential
areas of the city in the early 1990s where it is now established
(Biaduo, 2005). Fieldfares began breeding in inner Warsaw
(central Poland) in 1975 when there were less than 10 pairs,
but it has since become more numerous with 80100 pairs
in 1985 and 250350 pairs in 1990 (Luniak et al.,1990).
Fieldfares became urbanised in the Vosges region (north-
east France) in the late 1970s and early 1980s (Sueur &
Sueur-Bellart, 1983). These examples of recent urbanisation
by the fieldfare are against a background of a marked south-
westwards expansion in its breeding range between the 1800s
and 1960s (Lubcke & Furrer, 1985).
Away from Europe there are numerous other examples
of Turdus species being urbanised in only part of their
geographic range. Such variation is exhibited by at least
seven of the 33 Turdus species occurring in the Neotropics
(unless stated otherwise data are from Clement et al., 2000).
The rufous-bellied thrush T. rufiventris is only urbanised in
the south of its range, occurring in the centre of cities such
as Rio de Janerio (Brazil) and Buenos Aires (Argentina).
The pale-breasted thrush T. leucomelas frequents parklands
and gardens in some areas, including suburbs of Caracas,
Venezuela, and has recently colonised urban areas in Rio
Grande do Sul (southern Brazil; Bencke & Grillo, 1995).
The creamy-bellied thrush T. amaurochalinus occurs in parks
and large gardens, including around large cities, just in the
southern parts of its South American range. Similarly, the
great thrush T. fuscater has become common in gardens in
large cities, such as Quito (Ecuador) and la Paz (Bolivia),
in the last few decades but is not urbanised elsewhere. In
Mexico, the rufous-backed thrush T. rufopalliatus colonised
Mexico City and Oaxaca City during the 1980s and 1990s,
and was first noted in San Miguel de Allende in the late
1990s. It is uncertain if this species has become urbanised
through natural range expansion or establishment of escaped
cage birds, but the data clearly demonstrate the potential for
it to thrive in urban areas in some but not all parts of its
range (Howell & Webb, 1995; Brooks, 1999).
In the Afrotropics the olive thrush T. olivaceus is urbanised
in South Africa, but not in the more northern parts of
its range (Urban, Fry & Keith, 1997). The taxonomy of
this species is, however, complex and the sub-species in
the non-urbanised section of the range may be specifically
distinct from the taxa in the urbanised section of the range
(Bowie et al., 2005). Similarly, the Palearctic Tickells thrush
T. unicolor is a rural bird in most of its range, but is partially
urbanised in some sections, such as the Kashmir region of
the Indian sub-continent (Clement et al., 2000).
(b) Corvus
Corvus species also exhibit temporal variation in their
occurrence in urban areas. In Australia, the Torresian crow
C. orru was rarely seen in urban areas prior to the 1950s, but
is now ubiquitous (Low, 2002; Everding & Jones, 2006). The
jungle crow C. macrorhynchos became much more urbanised

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660 K. L. Evans and others

in Japan between the 1970s and 1990s, probably in response
to changes in waste disposal practices that increased food
availability (Ueta et al., 2003). Similarly, the hooded crow
Corvus cornix is becoming more urbanised in many European
regions. The species first bred in Finnish cities in the earlier
half of the 20th Century, but remained at low population
densities until the 1960s (Vuorisalo et al., 2003), and also
became urbanised in other European cities at a similar time
(e.g. Berlin; Lehmann, 2002). This contrasts with a much
older association with towns and cities further east in its
range (Madge & Burn, 1994).
We are not aware of sufficiently large compilations of data
on timing of urbanisation in any Corvus species for assessing
spatial patterns in urbanisation, but such a compilation is
available for another corvid, the magpie (Jerzak, 2001).
We analysed the spatial pattern of urbanisation using the
methods described above for the blackbird (Section III. 1),
but longitude was rescaled to facilitate the use of polynomial
terms (thus the most westerly location, Dublin, had a
longitude of zero rather than -6, i.e. 6 W). There is little
association between latitude and longitude in these data (r 2 =
0.5%), thus colinearity is not a concern. Magpie urbanisation
date exhibits a limited spatial pattern (model average r 2 =
12%; Table 2), with latitude having a negligible influence on
urbanisation date (model averaged partial r 2 < 1%; Table 2,
Fig. 5A), and a slight trend for more recent urbanisation
in eastern Europe (longitude: model averaged partial r2 =
12%; Fig. 5B). The very weak spatial signal in these data
suggests that this species has become urbanised in a largely
independent manner, preventing an analysis of rates of
spread of urban populations.
(c) Columba
Even the archetypal urban bird, the rock pigeon C. livia, is
not fully urbanised throughout its range. This species started
to colonise cities in Turkmenistan and Uzbekistan only in
the early 1980s (Obukhova, 2001). Similarly, in the 1980s
the population of C. livia breeding in Dushanbe (Tajikistan)
fed almost entirely on crops located 1015 km outside the
city, rather than on food sources within the city (Obukhova,
2001). The hill pigeon Columba rupestris started to invade
towns in the early 1900s, initially to feed during the winter,
with regular breeding in urban areas occurring in the second
half of the century; in some regions over 90% of this species
population now occurs in urban areas (del Hoyo et al., 1997).
A number of other Columba species are urbanised just in

(A) (B)

(C) (D)

Fig. 5. Relationships between known dates of magpie Pica pica urbanisation and (A) latitude, and (B) longitude (data from Jerzak
2001), and of woodpigeon Columba palumbus urbanisation with (C) latitude, and (D) longitude (data from Tomiaojc 1976) in Europe.
The open symbols represent the raw data, and the grey line the model averaged predicted values from the models described in
Table 2. To facilitate the use of polynomial terms in the magpie data longitude was rescaled so that the western-most location,
Dublin, had a longitude of zero rather than 6 W.

Biological Reviews 85 (2010) 643667 2010 The Authors. Journal compilation 2010 Cambridge Philosophical Society
A conceptual framework for the colonisation of urban areas
portions of their ranges, examples include the band-tailed
pigeon C. fasciata (urbanised in parts of the USA, but not
South America), picazuro pigeon C. picazuro (urbanised in
south-east Brazil and Argentina, but not further north), and
spot-winged pigeon C. maculosa (urbanised in Bolivia and
Peru, but not further south; all data from Gibbs et al., 2001).
The woodpigeon also exhibits much spatial and temporal
variation in the development of urban populations. Whilst
Tomiaojc (1976) documents this urbanisation process there
has been no formal assessment of spatial patterns in the
timing of these urbanisation events. Using the methods we
applied to the blackbird (Section III.1), we found a moderate
spatial pattern in the timing of woodpigeon urbanisation
in Europe (model average r 2 = 39%; Table 2; again there
is little colinearity in longitude and latitude in these data,
r 2 = 1.4%). Urbanisation exhibited a complex but weak
relationship with latitude (model averaged partial r 2 = 6%;
Fig 5C), but was more recent in the east (model averaged
partial r 2 = 35%, Fig. 5D). The strength of the spatial signal
in these data is sufficiently weak to prevent an analysis of rates
of spread, and suggests that leap-frog urbanisation events are
unlikely to be frequent in this species, although they cannot
be entirely excluded.
(d) Other genera
Spatial variation in the occurrence of urban populations has
been recorded in many other avian genera. Towns and cities
have been colonised by the merlin Falco columbarius in the
Nearctic (Sodhi et al., 1992), wallcreeper Tichodroma muraria
in Tibet (Gantlett & Millington, 1992), black woodpecker
in Hungary (Gorman, 1998), dunnock Prunella modularis in
the United Kingdom (Vogel & Tuomenpuro, 1997), lesser
whitethroat Sylvia curruca in eastern Europe (Bijlsma & Saris,
1997), northern wheatear Oenanthe oenanthe in parts of north-
east Europe (Walasz & Mielczarek, 1992), and common
redstart Phoenicurus phoenicurus and thrush nightingale Luscinia
luscinia in parts of eastern Europe (Lykov, Avilova & Beme,
2009); in other parts of their geographic ranges these species
are absent from urban areas.
A number of other bird species also show temporal
variation in their occurrence in urban areas. The following
are just some of the examples from the European avifauna:
long-tailed tit Aegithalos caudatus has recently colonised such
habitats in Europe (Nilsson, 1997); goshawks Accipiter gentilis
moved into urban parks in some European countries,
including Latvia, Germany and The Netherlands, at the
end of the 20th Century, but are absent from many
European cities (Wurlefs, 1994; Bijlsma & Sulkava, 1997); the
greenfinch Carduelis chloris has become increasingly common
in many European towns and villages since the 1950s (Gil-
Delgado & Newton, 1997); the Eurasian jay Garrulus glandarius
colonised urban areas in western and central Europe from
the 1930s, Latvia in the 1960s, Ukraine in the 1970s and
more recently in Russia (Bejcek & Gorban, 1997).
Spatial and temporal variation in urbanisation patterns
are thus probably more general phenomena than is typically
recognised. Indeed, this variation may hinder large-scale
Biological Reviews 85 (2010) 643667 2010 The Authors. Journal compilation 2010 Cambridge Philosophical Society
661
studies that seek to explain interspecific variation in
species responses to urban areas. Rather than viewing the
composition of urban assemblages as fixed, it is perhaps
more appropriate to consider them as a dynamic suite of
species that is subject to continuing colonisation events, as
well as local extinction. The detailed case study of spatial and
temporal patterns we present for the blackbird enables us
to assess how underlying factors related to each of the three
stages of the urbanisation process drives the development of
urban populations. Additional case studies are required from
a diverse range of taxonomic groups and regions to assess
fully which aspects of our conceptual framework have the
greatest influence on the development of urban assemblages.
VI. CONCLUSIONS
(1) Studies of range expansions have focused on those that
are external to a species original range, rather than
internal processes in which gaps within the current
range are filled. The urbanisation of a species is
a prime example of such internal range expansion.
However, despite a recent explosion of interest in the
ecological effects of urbanisation, a detailed theoretical
framework for studies of biotic colonisation of urban
areas has previously not been available. We provide
such a framework.
(2) There are two main potential mechanisms through
which a species may colonise urban areas. First,
colonisation may arise independently in a number
of locations. Alternatively, leap-frog urbanisation may
occur in which individuals first adapt to an urban
environment in a single location and then disperse to
other towns and cities.
(3) Each of these types of urbanisation progresses through
three stages: (i) arrival, (ii) adjustment, and (iii) spread.
The rate of progress through these stages will vary
with environmental factors relating to the spatial
configuration and development of urban areas (and
which are influenced by human attitudes and socio-
economic factors), species ecological and life-history
traits and whether urbanisation occurs independently
or in a leap-frog manner. These factors may be
interrelated in ways that can synergistically promote
progress through different stages or facilitate progress
through one stage yet delay progress through another.
(4) We use the framework that we have developed
to inform discussion of avian urbanisation events,
focusing primarily on the blackbird Turdus merula. This
is currently one of the most widely urbanised avian
species within its Western Palearctic range, but there
is marked spatial variation in the occurrence of urban
populations and the timing of their establishment,
which varies by over 150 years. The occurrence of
such variation is likely to be a general pattern in other
taxa and regions.
662
(5) The blackbird appears to be unusual in that the timing
of urbanisation exhibits a strong spatial signature,
such a signature is much less marked in the magpie
and woodpigeon. Whilst the blackbirds strong spatial
pattern is suggestive of leap-frog urbanisation other
evidence suggests that many urbanisation events are
likely to have arisen in an independent manner.
(6) The overall rate of spread of urban blackbird
populations is similar to the mean rate of spread
of avian range expansions that are external to species
original ranges. This suggests that internal and external
range expansions are governed by similar processes as
each involves the colonisation of novel environments.
(7) Blackbird urbanisation has generally occurred more
recently in eastern Europe, and the rate of spread
of urban populations is also slower in this region.
This appears likely to have arisen from reduced
opportunities for arrival in urban areas. The former
rarity of blackbirds in parts of this region, an
increased migratory tendency limiting the extent
to which individuals moved into urban areas in
winter (which appears to be a precursor of complete
urbanisation in the blackbird and other species), and
the reduced number of urban developments may all
have contributed to limiting the opportunities for
arrival in urban areas in eastern Europe. The extent to
which urbanisation requires novel genetic adaptations
remains uncertain, but these may also have been slower
to arise in more migratory eastern populations. In some
areas urbanisation may also have been promoted by
reduced persecution and increased habitat suitability
in urban areas, but these appear not to have generally
influenced the timing of blackbird urbanisation.
(8) Additional case studies are required before generali-
sations can be made regarding the factors promoting
progression through each of the three key stages of
this process. It is clear, however, that the composition
of urban biotas is not fixed. Rather their composition
can exhibit great variability, and additional species are
likely to be urbanised in the future, especially given
the unprecedented growth in the number and extent
of urban areas.
VII. ACKNOWLEDGEMENTS
We are grateful to M. Luniak for his earlier work
that promoted investigations into blackbird urbanisa-
tion. C. Benskin, B.M. Braschler, J. Elts, E. Garcia-
Berthou, A. Gregoire, K. Hammerschmidt, R.C. Hatchwell,
J. Hornskov, Z. Karpowicz, Y. Kolbeinsson, E. Leibak,
V. Lummaa, A. Nagy, K. Reinhardt, C.I.S. Salas, V.H.
Salas, L.C. Salzar, M. Simeoni, S. Svazas and D. Wysocki
helped with translations and supplying information on urban-
isation patterns; J. Marzluff provided assistance. This work
Biological Reviews 85 (2010) 643667 2010 The Authors. Journal compilation 2010 Cambridge Philosophical Society
K. L. Evans and others
was funded by the Natural Environment Research Coun-
cil. K.J.G. holds a Royal Society-Wolfson Research Merit
Award.
VIII. REFERENCES
Abs, M. & Bergen, F. (2008). A long term survey of the avifauna in an urban park.
In Urban ecology: an international perspective on the interaction between humans and nature (eds.
J. M. Marzluff, E. Shulenberger, W. Endlicher, M. Alberti, G. Bradley,
C. Ryan, U. Simon and C. Zumbrunnen), pp. 373376. New York: Springer.
Adamian, M. S. & Klem, D. (1999). Handbook of the birds of Armenia. American
University of Armenia, Yerevan, Republic of Armenia.
Akimov, I. A. & Kostyuhin, V. (1994). Urban zoology in the Ukraine: a brief review
of the vertebrates. Memorabilia Zoologica 49, 283288.
Alexander, H. G. (1927). The birds of Latium, Italy. Ibis 1927, 245283.
Andrews, I. J. (1995). The birds of the Hashemite Kingdom of Jordan. Musselburgh:
published privately by I.J. Andrews.
Angilletta, M. J., Wilson, R. S., Niehaus, A. C., Sears, M. W., Navas, C. A. &
Ribeiro, P. L. (2007). Urban physiology: city ants possess high heat tolerance. PLoS
One 2, e258.
Arnfield, A. J. (2003). Two decades of urban climate research: A review of turbulence,
exchanges of energy and water, and the urban heat island. International Journal of
Climatology 23, 126.
Aruda, M. P. & Morielle-Versuta, E. (2008). Cytogenetic and random amplified
polymorphic DNA analysis of Leptodactylus species from rural and urban
environments. Genetics and Molecular Research 7, 161176.
Baker, P. J., Molony, S. E., Stone, E., Cuthill, I. C. & Harris, S. (2008). Cats
about town: is predation by freeranging pet cats Felis catus likely to affect urban
bird populations? Ibis 150, 8699.
Baldamus, E. (1880). Der Wurzburger Amselprozess und die Amsel. Frankfurt Am Main:
Morgenstern.
Bannerman, D. A. (1963). Birds of the Atlantic Islands, vol. 1. A history of the birds of the
Canary Islands and of the Salvages. Edinburgh: Oliver and Boyd.
Bannerman, D. A. & Bannerman, W. M. (1965). Birds of the Atlantic islands, vol. 2.
A history of the birds of Madeira, the Desertas and the Porto Santo Islands. Edinburgh: Oliver
and Boyd.
Bannerman, D. A. & Bannerman, M. (1966). Birds of the Atlantic Islands, vol. 3. A history
of the birds of the Azores. Edinburgh: Oliver and Boyd.
Barrett, G., Silcocks, A., Barry, S., Cunningham, R. & Pulter, R. (2003).
The new atlas of Australian birds. Victoria: Royal Australian Ornithologists Union.
Barrett, S. C. H., Colautti, R. I. & Eckert, C. G. (2008). Plant reproductive
systems and evolution during biological invasion. Molecular Ecology 17, 373383.
Beck, N. R. & Heinsohn, R. (2006). Group composition and reproductive success of
cooperatively breeding white-winged choughs (Corcorax melanorhamphos) in urban and
nonurban habitat. Austral Ecology 31, 588596.
Bejcek, V. & Gorban, I. (1997). Jay Garrulus glandarius. In: The EBCC atlas of European
breeding birds: their distribution and abundance. (eds. W. J. M. Hagemeijer and M.
J. Blair), pp. 670671. London: T & AD Poyser.
Bell, H. (1986). Occupation of urban habitats by birds in Papua New Guinea.
Proceedings of the Western Foundation of Vertebrate Zoology 3, 148.
Bencke, G. A. & Grillo, H. C. Z. (1995). Range expansion of the Palebreasted
Thrush Turdus leucomelas (Aves, Turdidae) in Rio Grande do Sul, Brazil. Iheringia,
Serie Zoologia 79, 175176.
Benoit, L. (1840). Ornitologia Siciliana. Messina.
BermudezCuamatzin, E., RosChelen, A. A., Gil, D. & Garcia, C. M.
(2009). Strategies of song adaptation to urban noise in the house finch: syllable
pitch plasticity or differential syllable use? Behaviour 146, 12691286.
Berthold, P., Helbig, A. J., Mohr, G. & Querner, U. (1992). Rapid microevolu-
tion of migratory behaviour in a wild bird species. Nature 360, 668670.
Biadun, V. (2005). Lublin. In Birds in European cities. (ed. J. G. Kelcey and
G. Rheinwald), pp. 171196. St. Katharinen: Ginster Verlag.
Bijlsma, R. G., Hustings, F. & Camphuysen, K. C. J. (2001). Avifauna van Nederland,
vol. 2. Haarlem-Stichting Uitgeverij van der KNNV; GMB Uitgeverij.
Bijlsma, R. G. & Saris, F. (1997). Lesser Whitethroat Sylvia curruca. In: The EBCC atlas
of European breeding birds: their distribution and abundance. (eds. W. J. M. Hagemeijer
and M. J. Blair), pp. 594595. London: T & AD Poyser.
Bijlsma, R. G. & Sulkava, S. (1997). Goshawk Accipiter gentiles. In: The EBCC atlas of
European breeding birds: their distribution and abundance. (eds. W. J. M. Hagemeijer and
M. J. Blair), pp. 154155. London: T & AD Poyser.
Blackburn, T. M., Cassey, P. & Gaston, K. J. (2006). Variations on a theme:
sources of heterogeneity in the form of the interspecific relationship between
abundance and distribution. Journal of Animal Ecology 75, 14261439.
Blair, R. B. (1996). Land use and avian species diversity along an urban gradient.
Ecological Applications 6, 506519.
A conceptual framework for the colonisation of urban areas
Blair, R. B. (2001). Birds and butterflies along urban gradients in two ecoregions
of the United States: is urbanisation creating a homogenous fauna? In: Biotic
homogenization. (eds. M. L. McKinney and J. L. Lockwood), pp. 3356. New York:
Plenum Publishers.
Blakers, M., Davies, S. J. J. F. & Reilly, P. N. (1984). The atlas of Australian birds.
Melbourne: Melbourne University Press.
Blome, R. (1673). Map of London. (available at http://en.wikipedia. org/wiki/
History of London).
Bodenheimer, F. S. (1935). Animal Life in Palestine. Jersualeum: Mayer.
Bonier, F., Martin, P. R. & Wingfield, J. C. (2007). Urban birds have broader
environmental tolerance. Biology Letters 6, 670673.
Bowie, R. C. K., Voelker, G., Fjeldsa , J., Lens, L., Hackett, S. J. &
Crowe, T. M. (2005). Systematics of the olive thrush Turdus olivaceus species complex
with reference to the taxonomic status of the endangered Taita thrush T. helleri.
Journal of Avian Biology 36, 391404.
Bradley, C. A. & Altizer, S. (2007). Urbanisation and the ecology of wildlife
diseases. Trends in Ecology and Evolution 22, 95102.
Bradley, C. A., Gibbs, S. E. J. & Altizer, S. (2008). Urban land use predicts West
Nile Virus exposure in songbirds. Ecological Applications 18, 10831092.
Brommer, J. E., Merila, J., Sheldon, B. C. & Gustafsson, L. (2005). Natural
selection and genetic variation for reproductive reaction norms in a wild bird
population. Evolution 59, 13621371.
Brommer, J. E., Rattiste, K. & Wilson, A. J. (2008). Exploring plasticity in the
wild: laying datetemperature reaction norms in the common gull Larus canus.
Proceedings of the Royal Society of London, Series B 22, 687693.
Brooks, D. M. (1999). Rufous-backed Thrush Turdus rufopalliatus in Guanajuato,
Mexico. Cotinga 12, 71.
Brown, J. L. (1969). The buffer effect and productivity in tit populations. American
Naturalist 103, 347354.
Bruch, A., Elvers, H., Pohl, C., Westphal, D. & Witt, K. (1978). Die vogel in
Berlin (west) eine ubersicht. Berlin: Ornithologischer Bericht fur Berlin.
Brumm, H. (2004). The impact of environmental noise on song amplitude in a
territorial bird. Journal of Animal Ecology 73, 434440.
Bull, P. C., Gaze, P. D. & Robertson, C. J. R. (1985). The atlas of bird distribution in
New Zealand. Wellington: Ornithological Society of New Zealand.
Burnham, K. P. & Anderson, D. R. (2002). Model selection and multimodel inference: a
practical information-theoretic approach. New York: Springer.
Burton, J. F. (1995). Birds and climate change. London: Christopher Helm.
Cade, T. J. & Bird, D. M. (1990). Peregrine falcons, Falco peregrinus, nesting in an
urbanenvironment - a review. Canadian Field Naturalist 104, 209218.
Campbell, O. (2006). Urban parks as shared spaces? The utility of alert distances as
indicators of avian tolerance of humans in Stirling, Scotland. Area 38, 301311.
Cannon, A. R., Chamberlain, D. E., Toms, M. P., Hatchwell, B. J. & Gas-
ton, K. J. (2005). Trends in the use of private gardens by wild birds in Great
Britain. Journal of Applied Ecology 42, 659671.
CBS (2003). Bodemgebruik in Nederland. Voorburg/Heerlen: Statistics Netherlands.
(Available at http://statline.cbs.nl)
Chace, J. F. & Walsh, J. J. (2006). Urban effects on native avifauna: a review.
Landscape and Urban Planning 74, 4649.
Chamberlain, D. E., Cannon, A. R., Toms, M. P., Leech, D. I., Hatchwell,
B. J. & Gaston, K. J. (2009). Avian productivity in urban landscapes: a review and
meta-analysis. Ibis 151, 118.
Charmantier, A., McCleery, R. H., Cole, L. R., Perrins, C., Kruuk, L. E. B.
& Sheldon, B. C. (2008). Adaptive phenotypic plasticity in response to climate
change in a wild bird population. Science 320, 800803.
Cheptou, P. O. & Avendano, L. G. V. (2006). Pollination processes and the Allee
effect in highly fragmented populations: consequences for the mating system in
urban environments. New Phytologist 172, 774783.
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 of the USA 105, 37963799.
Ciochia, V. (1992). Pasarile clocitoare din Romania Atlas. Bucharest: Editura Stiintifica.
Clement, P., Hathaway, R., Wilczur, J. & Byers, C. (2000). Thrushes. London:
Christopher Helm.
Clergeau, P., Jokimaki, J. & Savard, J. P. L. (2001). Are urban bird communities
influenced by the bird diversity of adjacent landscapes? Journal of Applied Ecology 38,
11221134.
Collett, R. (1921). Norges fugle. Kristiania: Forlagt Av H. Aschehoug and Co.
Collier, C. G. (2006). The impact of urban areas on weather. Quarterly Journal of the
Royal Meteorological Society 132, 125.
Cooke, A. S. (1980). Observations on how close certain passerine species will tolerate
an approaching human in rural and suburban areas. Biological Conservation 18, 8588.
Corti, U. A. (1959). Die brutvogel der deutschen und osterreichischen Alpenzone; die vogelwelt der
Alpen band 5. Chur: Verlag Bischofberger and Co.
Cramp, S. (ed.) (1988). The birds of the western Palearctic Vol 5: Tyrant flycatchers to thrushes.
Oxford: Oxford University Press.
Croci, S., Butet, A. & Clergeau, P. (2008). Does urbanisation filter birds on the
basis of their biological traits? The Condor 110, 223240.
Biological Reviews 85 (2010) 643667 2010 The Authors. Journal compilation 2010 Cambridge Philosophical Society
663
Culley, T. M., Sbita, S. J. & Wick, A. (2007). Population genetic effects of urban
habitat fragmentation in the perennial herb Viola pubescens (Violaceae) using ISSR
markers. Annals of Botany 100, 91100.
Davis, J. M. (2008). Patterns of variation in the influence of natal experience on
habitat choice Quarterly Review of Biology 83, 363380.
Davies, Z. G., Fuller R. A., Loram, A., Irvine, K. N., Sims, V. & Gaston, K. J.
(2009). A national scale inventory of resource provision for biodiversity within
domestic gardens. Biological Conservation 142, 761771.
Deby, J. (1845). Notes on the birds of Belgium. The Zoologist 3: 813816, 858862,
933934, 979981, 10191022, 10701072, 11311132 and 11871189.
De Kerville, H. G. (1890). Faune de la Normandie Fasc II Oiseaux. Paris: Extrait du
Bulletin de la Societe de Amis des Sciences Naturelles de Rouen.
De Laet, J. (2000). Over merels en andere kerels. Brussels: Vubpress.
De Victor, V., Julliard, R., Couvet, D., Lee, A. & Jiguet, F. (2007). Functional
homogenization effect of urbanisation on bird communities. Conservation Biology 21,
741751.
Del Hoyo, J., Elliott, A. & Christie, D. A. (eds) (2005). Handbook of the birds of the
world volume 10: cuckoo-shrikes to thrushes. Barcelona: Lynx edicions.
Del Hoyo, J., Elliott, A. & Sargatal, J. (eds) (1997). Handbook of the birds of the
world volume 4: sandgrouse to cuckoos. Barcelona: Lynx edicions.
Desender, K., Small, E. C., Gaublomme, E. & Verdyck, P. (2005). Rural-urban
gradients and the population genetic structure of woodland ground beetles.
Conservation Genetics 6, 5162.
Dubois, A. (1888). Compte-rendu des observations ornithologiques faites en Belgique
pendant lannee 1886. Bulletin du Musee royal dHistoire naturelle de Belgique 4, 177210.
Dubois, P. J., Le Mareechal, P., Olioso, G. & Yesou, P. (2000). Inventaire des
Oiseaux de France: aviafaune de la France metropolitaine. Paris: Nathan.
Dutra, N. C. L., Telles, M. P. C., Dutra, D. L. & Silva, N. J. (2008). Genetic
diversity in populations of the viper Bothrops moojeni Hoge, 1966 in central Brazil
using RAPD markers. Genetics and Molecular Research 7, 603613.
Dybbro, T. (1978). Oversight over Danmarks Fugle. Forening: Dansk Ornithologisk.
Ebenman, B. & Karlsson, J. (1984). Urban Blackbirds from egg to independence.
Annales Zoologici Fennici 21, 249251.
Ellegren, H (2007). Characteristics, causes and evolutionary consequences of
malebiased mutation. Proceedings of the Royal Society of London, Series B 274, 110.
Emery, N. J. (2006). Cognitive ornithology: the evolution of avian intelligence.
Philosophical Transactions of the Royal Society of London, Series B 361, 2343.
Epprecht, W. (1946). Die Verbreitung der Amsel, Turdus m. merula L. zur. Brutzeit
in Zirich, 1946. Der Ornithologische Beobachter 43, 97105.
Evans, K. L. (In press). Urbanisation and individual species. In Urban ecology (ed. K.
J Gaston). Cambridge: Cambridge University Press.
Evans, K. L., Gaston, K. J., Frantz, A. C., Simeoni, M., Sharp, S. P.,
McGowan, A., Dawson, D. A., Walasz, K., Partecke, J., Burke, T. &
Hatchwell, B. J. (2009a). Independent colonisation of multiple urban centres
by a formerly forest specialist bird species. Proceedings of the Royal Society of London, Series
B 276, 24032410.
Evans, K. L., Gaston, K. J., Sharp, S. P., McGowan, A. & Hatchwell, B. J.
(2009b). The effect of urbanisation on avian morphology and latitudinal gradients
in body size. Oikos 118, 251259.
Evans, K. L., Gaston, K. J., Sharp, S. P., McGowan, A., Simeoni, M. &
Hatchwell, B. J. (2009c). Effects of urbanisation on disease risk and age ratios in
blackbird Turdus merula populations. Oikos 118, 774782.
Evans, K. L., Greenwood, J. J. D. & Gaston, K. J. (2007). The positive correlation
between avian species richness and human population density in Britain is not
attributable to sampling bias. Global Ecology and Biogeography 16, 300304.
Evans, K. L., Newson, S. E. & Gaston, K. J. (2009d ). Habitat influences on urban
avian assemblages. Ibis 151, 1939.
Evans, K. L., Van Rensburg, B. J., Gaston, K. J. & Chown, S. L. (2006). People,
species richness and human population growth. Global Ecology and Biogeography 15,
625636.
Everding, S. E. & Jones, D. N. (2006). Communal roosting in a suburban population
of Torresian crows (Corvus orru). Landscape and Urban Planning 74, 2133.
Fallon, F. (1875). Monographie des oiseaux de la Belgique. Paris: J. B. Bailliere et Fils.
Fatio, V. & Studer, T. (1908). Catalogue des Oiseaux de la Suisse. Geneve: Departement
federal de lInterieur.
Fernandez-Juricic, E. & Telleria, J. L. (1999). Recruitment patterns of blackbirds
Turdus merula in urban fragmented populations. Ardeola 46, 6170.
Ferragni, O. (1885). Avifauna Cremonese.
Fitter, R. S. R. (1990). Londons Natural History 2nd edition. London: Bloomsbury
Books.
Fokidis, H. B., Greiner, E. C. & Deviche, P. (2008). Interspecific variation in avian
blood parasites and haematology associated with urbanization in a desert habitat.
Journal of Avian Biology 39, 300310.
Friedrich, C. G. (1850). Naturgesch der Vogel Deutschlands.
Frieling, H. (1942). Grossstadtvogel Krieg Mensch Natur. Stuttgart: Franckhsche
Verlagshandlung.
Frionnet, C. (1925). Les Oiseaux de la Haute-Marne. Haute-Marne: Societe Sciences
Naturelles de Haute-Marne.
664
Fritsch, A. (1870). Naturgeschichte der Vogels Europas. Prague: Verlag des Verfassers.
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, 131137.
Fuller, R. M., Smith, G. M., Sanderson, J. M., Hill, R. A., Thompson, A. G.,
Cox, R., Brown, N. J., Clarke, R. T., Rothery, P. & Gerard, F. F. (2002).
Countryside survey 2000 module 7: land cover map 2000 final report. Monks Wood: Centre
for Ecology and Hydrology.
Gantlett, S. & Millington, R. (1992). Photo-forum: wallcreeper. Birding World 5,
271274.
Garant, D., Kruuk, L. E. B., Wilkin, T. A., McCleery, R. H. & Sheldon, B. C.
(2005). Evolution driven by differential dispersal within a wild bird population.
Nature 433, 6065.
Gaston, K. J. (2003). The structure and dynamics of geographic ranges. Oxford: Oxford
University Press
Gaston, K. J., Blackburn, T. M. & Lawton, J. H. (1997). Interspecific abundance
range size relationships: an appraisal of mechanisms. Journal of Animal Ecology 66,
579601.
Geister, I. (1995). Ornitoloski Atlas Slovenije. Ljubljana: DZS.
Gering, J. C. & Blair, R. B. (1999). Predation on artificial bird nests along an urban
gradient: predatory risk or relaxation in urban environments? Ecography 22, 532541.
Ghalambor, C. K., McKay, J. K., Carroll, S. P. & Reznick, D. N. (2007).
Adaptive versus nonadaptive phenotypic plasticity and the potential for
contemporary adaptation in new environments. Functional Ecology 21, 394407.
Gibbs, D., Barnes, E. & Cox, J. (2001). Pigeons and doves: a guide to the pigeons and doves
of the world. Robertsbridge: Pica Press.
Gilbert, O. L. (1989). The ecology of urban habitats. London: Chapman & Hall.
GilDelgado, J. A. & Newton, I. (1997). Greenfinch Carduelis chloris. In The EBCC
atlas of European breeding birds: their distribution and abundance. (eds. W. J. M. Hagemeijer
and M. J. Blair), pp. 712713. London: T & AD Poyser.
Glaser, L. (1871). Mitteilungen aus dem Tierleben in den Wormser Promenaden.
Der Zoologische Garten 12, 311.
Glazier, D.S. (2002). Resource-allocation rules and the heritability of traits. Evolution
56, 16961700.
Goodman, S. M. & Meininger, P. L. (1989). The Birds of Egypt. Oxford: Oxford
University Press.
Gorman, G. (1998). The spread of the Black Woodpecker in Europe - will it reach
Britain next? Birding World 11, 390395.
Gosselink, T. E., Van Deelen, T. R., Warner, R.E. & Mankin, P. C. (2007).
Survival and cause-specific mortality of red foxes in agricultural and urban areas of
Illinois. Journal of Wildlife Management 71, 18621873.
Graczyk, R., Fedorenko, A. P., Loskot, V. M. & Chuprin, S. L. (1975).
Introduction of the Turdus merula town population from Poznan into Kiev. Vestrik
Zoologii 3, 2932.
Graczyk, R. (1959). Badania nad wystepowaniem I stanem ilosciowym kosa (Turdus
merula L.) w Polsce. Ekologia Polska 7, 5582.
Graczyk, R. (1961). Studies on variability, biology and economic status of the
Blackbird, Turdus merula. Ekologia Polska 9, 453485.
Graczyk, R. (1982). Ecological and ethological aspects of the synanthropization of
birds. Memorabilia Zoologica 37, 7991.
Gregoire, A. (2003). Demographie et differentiation chez le merle noir Turdus merula: liens avec
lhabitat et les relations hotes-parsites. PhD Thesis, Universite de Bourgogne.
Gregory, R. D. & Gaston, K. J. (2000). Explanations of commonness and rarity in
British breeding birds: separating resource use and resource availability. Oikos 88,
515526.
Grimm, N. B., Faeth, S. H., Golubiewsk, N. E., Redman, C. L., Wu, J., Bai, X. &
Briggs, J. M. (2008). Global change and the ecology of cities. Science 319, 756760.
Grosholz, E. D. (1996). Contrasting rates of spread for introduced species in
terrestrial and marine environments. Ecology 77, 16801686.
Gudmundsson, F. (1951). The effects of the recent climatic changes on the birdlife of
Iceland. In Proceedings of the Xth International Ornithological Congress, June 1950 Uppsala.
(ed. S. Horstadius), pp 502514. Uppsala: Almquist and Wiksell.
Haartman, L., Hilden, O., Linkola, P., Suomalainen, P. & Tenovuo, R.
(1967). Pohjolan Linnut Varikuvin II. Helsinki: Kustannusosakeyhtio Otavan
Kirjapaino.
Haines-Young, R. H., Barr, C. J., Black, H. I. J., Briggs, D. J., Bunce, R. G. H.,
Clarke, R. T., Cooper, A., Dawson, F. H., Firbank, L. G., Fuller, R. M.,
Furse, M. T., Gillespie, M. K., Hill, R., Hornung, M., Howard, D. C.,
McCann, T., Morecroft, M. D., Petit, S., Sier, A. R. J., Smart, G. M.,
Stott, A. P., Stuart, R. C. & Watkins, J. W. (2000). Accounting for nature: assessing
habitats in the UK countryside. London: Department of the Environment, Transport
and the Regions.
Harmata, W. (1979). Wzrost liczby gniezdzacych sie kwiczoow Turdus pilaris w
Krakowie. Chronmy Przyrode Ojczysta 33, 6772.
Hendry, A. P., Nosil, P. & Rieseberg, L. H. (2007). The speed of ecological
speciation. Functional Ecology 21, 455464.
Herzig, A. L. (1995). Effects of population density on long-distance dispersal in the
goldenrod beetle Trirhabda virgata. Ecology 76, 20442054.
Biological Reviews 85 (2010) 643667 2010 The Authors. Journal compilation 2010 Cambridge Philosophical Society
K. L. Evans and others
Heyder, R. (1955). Hundert jahre gartenamsel. Beitrage Vogelk 4, 6481.
Hirota, T., Hirohata, T., Mashima, H., Satoh, T. & Obara, Y. (2004).
Population structure of the large Japanese field mouse, Apodemus speciosus (Rodentia:
Muridae), in suburban landscape, based on mitochondrial D-loop sequences.
Molecular Ecology 13, 32753282.
Hitch, A. T. & Leberg, P. L. (2007). Breeding distributions of North American bird
species moving north as a result of climate change. Conservation Biology 21, 534539.
Hitchings, S. P. & Beebee, T. J. C. (1997). Genetic substructuring as a result of
barriers to gene flow in urban Rana temporaria (common frog) populations: implications
for biodiversity conservation. Heredity 79, 117127.
Hitchings, S. P. & Beebee, T. J. C. (1998). Loss of genetic diversity and fitness
in Common Toad (Bufo bufo) populations isolated by inimical habitat. Journal of
Evolutionary Biology 11, 269283.
Hollingsworth, P. M. & Dickson, J. H. (1997). Genetic variation in rural and
urban populations of Epipactis helleborine (L) Crantz (Orchidaceae) in Britain. Botanical
Journal of the Linnean Society 123, 321331.
Holloway, S. (2002). The historical atlas of breeding birds in Britain and Ireland: 18751900.
London: A and C Black Publishers Ltd.
Homeyer, A. (1860). Die Amsel in Neu-Vorpommern und am Rhein. Journal of
Ornithology 8, 396397.
Hortling, I. (1929). Ornitologisk Handbok. Helsinki: Helsingfors-Brando.
Howell, S. N. G. & Webb, S. (1995). The birds of Mexico and northern central America.
Oxford: Oxford University Press.
Howgego, J. (1978). Printed maps of London circa 15531850. 2nd edition. London:
Dawson.
Hudec, K. (1983). Fauna ESSR Svazek 23 Ptaci Aves. Prague: Academia.
Ingram, C. (1926). The birds of the Riviera: being an account of the avifauna of the Cote dAzur
from the Esterel Mountains to the Italian Frontier. London: H.F. and G. Witherby.
Jarvinen, O. & Vaisanen, R. (1979). Climatic changes, habitat changes, and
competition: dynamics of geographical overlap in two pairs of congeneric bird
species in Finland. Oikos 33, 261271.
Jerzak, L. (2001). Synurbanization of the magpie in the Palearctic. In Avian ecology
and conservation in an urbanizing world. (ed. J. M. Marzluff, R. Bowman and
R. Donnelly), pp. 403426. Boston, Kluwer Academic Publishers.
Jokimaki, J., Kaisanlahti-Jokimaki, M. L., Sorace, A., Fernandez-Juricic, E.,
Rodriguez-Prieto, I. & Jimenez, M. D. (2005). Evaluation of the safe nesting
zone hypothesis across an urban gradient: a multi-scale study. Ecography 28, 5970.
Johnson, J. B. & Omland, K. S. (2004). Model selection in ecology and evolution.
Trends in Ecology and Evolution 19, 101108.
Jordan, H. (1994). Public parks, 18851914. Garden History 22, 85113.
Kalela, O. (1952). Changes in the geographic distribution of Finnish birds and
mammals in relation to recent changes in climate. Fennia 75, 3851.
Kark, S., Iwaniuk, A., Schalimtzek, A. & Banker, E. (2007). Living in the city:
can anyone become an urban exploiter? Journal of Biogeography 34, 638651.
Karlsson, J. & Kallander, H. (1977). Fluctuations and density of suburban
populations of the blackbird Turdus merula. Ornis Scandinavica 8, 139144.
Kayser, C. (1914). Beitrage zur Ornis von Preufsisch-Schlesien. Journal of Ornithology
62, 530556.
Keane, B., Collier, M. H. & Rogstad, S. H. (2005). Pollution and genetic structure
of North American populations of the common dandelion (Taraxacum officinale).
Environmental Monitoring and Assessment 105, 341357.
Kemner, N. A. (1910). Koltrastens hackplatser. Fauna och Flora 5, 9293.
Kentish, B. J., Dann, P. & Lowe, K. W. (1995). Breeding biology of the common
blackbird Turdus merula in Australia. Emu 95, 233244
Khrabryi, V. (2005). St. Petersburgh. In Birds in European cities. (ed. J. G. Kelcey and
G. Rheinwald), pp. 307334. St. Katharinen: Ginster Verlag.
Knodt, B. (1886). In des Ausschusses fur Beobachtungsstationen der Vogel
Deutschlands, pp. 322. Journal of Ornithology 34, 129388.
Kolar, C. S. & Lodge, D. M. (2001). Progress in invasion biology: predicting
invaders. Trends in Ecology and Evolution 16, 199204.
Kolazy, J. (1880). Die Amsel. Mittheilungen des Ornithologischen Vereins in Wien 4, 5763.
Kollibay, P. (1906). Die Vogel der Preuischen Provinz Schlesien. Korn: Breslau.
Konstantinov, V. M. & Zakharov, R. (2005). Moscow. In Birds in European cities.
(ed. J. G. Kelcey and G. Rheinwald), pp. 197214. St. Katharinen: Ginster
Verlag.
Kosinski, Z. (2001). Effects of urbanisation on nest site selection and nesting success
of the Greenfinch Carduelis chloris in Krotoszyn, Poland. Ornis Fennica 78, 175183.
Kostecke, R. M., Ellison, K. & Summers, S. G. (2004). Continued range
expansion by Bronzed Cowbirds in the southwestern United States. The Southwestern
Naturalist 49, 487492.
Kovshar, A. F. & Sujko, B. P. (1984). Biologische daten zur verstadterung der Amsel
(Turdus merula) in Alma-Ata, Kasachstan, USSR. Mitteilungen Zoologie Museum Berlin 8,
97105.
Kruger, C. (1940). Nordiske solsorters (Turdus m. merula) forekomst traek. Dansk
Ornitologisk Forenings Tidsskrift 34, 114153.
Kuhk, R. (1939). Die Vogel Mecklenburgs. Faunistische, tiergeographische und okologische
Untersuchungen im mecklenburgischen Raume. Gustrow: Verlag Opitz & Co.
Kumari, E. (1954). The birds of Estonia. Tallinn: Eesti Riiklik Kirjastus,.
A conceptual framework for the colonisation of urban areas
Kumari, E. (1970). Lintujen kaupunkilaistumisest ja kaupunkilintujen tottumuksista
Baltian maissa: Suomalais-eestilaisen ornitologikongressin esitelmat. Ornis Fennica
47, 143.
Kuyt, E. (1971). First record of the cattle egret in the northwestern territories. Canadian
Field Naturalist 86, 8384.
Lambin, X., Aars, J. & Piertney, S. B. (2001). Dispersal, intraspecific competition,
kin competition and kin facilitation: a review of the empirical evidence. In Dispersal
(eds. J. Clobert, E. Danchin, A. A. Dhondt and J. D. Nichols), pp. 110122.
Oxford: Oxford University Press.
Landau, P. & Schneider, C. (1928). Der deutsche garten. Erleben: Ein jahrtausend
Natur.
Lehmann, R. (2002). Population development, habitat use and interaction of Magpie
Pica pica and Hooded Crow Corvus corone cornix in Berlin. Vogelwelt 123, 213221.
Leibak, E. (1981). Blackbirds have become town birds. Estonian Nature 11, 737738.
Lensink, R. (1997). Range expansion of raptors in Britain and the Netherlands since
the 1960s: testing an individual-based diffusion model. Journal of Animal Ecology 66,
811826.
Lensink, R. (1998). Temporal and spatial expansion of the Egyptian goose Alopochen
aegyptiacus in the Netherlands, 196794. Journal of Biogeography 25, 251263.
Le Rabe, F. (1886). Catalogue Annote des Oiseaux qui vivent a etat sauvage dans le departement
de LYonne. Paris: Vicor Mason.
Le Sauvage, H. E. (1872). Catalogue methodique des Oiseaux du Calvados. Memoires
de la Societe Linneenne Normandie 6, 171219.
Liker, A., Papp, Z., Bokony, V. & Lendvai, A. Z. (2008). Lean birds in the city:
body size and condition of house sparrows along the urbanisation gradient. Journal
of Animal Ecology 77, 789795.
Lim, H. C. & Sodhi, N. S. (2004). Responses of avian guilds to urbanisation in a
tropical city. Landscape and Urban Planning 66, 199215.
Lonnberg, E. (1935). Svenska Faglars Flyttning. Stockholm: Hugo Gebers Forlag.
Lppenthin, B. (1967). Danske ynglefugle I fortid og nutid (Danish breeding birds past and
present). Oxford: Odense University Press.
Lowry, W. P. (1998). Urban effects on precipitation amount. Progress in Physical
Geography 22, 477520.
Loxdale, H. D. & Lushai, G. (1999). Slaves of the environment: the movement of
herbivorous insects in relation to their ecology and genotype. Philosophical Transactions
of the Royal Society of London, Series B 354, 14791495.
Low, T. (2002). The New Nature. Sydney: Penguin.
Lubcke, W. & Furrer, R. (1985). Die wacholderdrossel. Wittenberg: Neue Brehm-
Bucherei 569 Ziemsen.
Luniak, M. (2004). Synurbanizationadaptation of animal wildlife to urban
development. In Proceedings of the 4th international symposium on urban wildlife conservation.
(eds. Shaw, W. W., Harris, L. K. and Vandruff, L.), pp. 5055. Tucson.
Luniak, M. (2005). Warsaw. In Birds in European cities. (ed. J. G. Kelcey and
G. Rheinwald), pp. 389416. St. Katharinen: Ginster Verlag.
Luniak, M., Mulsow, R. & Walasz, K. (1990). Urbanisation of the European
Blackbirdexpansion and adaptations of urban population. In Urban ecological
studies in Central and eastern Europe; proceedings of the international symposium Warszawa-
Jablonna 2425th September 1986. (ed. Luniak, M.), pp. 187198. Warsaw, Polish
Academy of Sciences.
Lykov, E. L., Avilova, K. V. & Beme, I. R. (2009). Some aspects of the
synanthropization of nine species of the Turdidae family in Kaliningrad. Moscow
University Biological Sciences Bulletin 64, 8491.
Lynes, H. (1924). An ornithological visit to north-west Morocco (Spanish Province of
Yebala). Novitates Zoologicae 31, 49103.
Macarthur, R. H. & Wilson, E. O. (1967). The theory of island biogeography. Princeton:
Princeton University Press.
Madge, S. & Burn, H. (1994). Crows and jays: a guide to the crows, jays and magpies of the
world. London: A & C Black.
Marchant, J. H., Hudson, R., Carter, S. P. & Whittington, P. (1990). Population
trends in British breeding birds. Tring: British Trust for Ornithology.
Margalef, R. (1974). Ecologia. Barcelona: Omega Ediciones.
Marschall, A. F. G. & Von Pelzeln, A. (1882). Ornis Indobonensis. Wien: Verlag
von Georg Paul Faesy.
Martell, M. S., Englund, J. V. & Tordoff, H. B. (2002). An urban osprey
population established by translocation. Journal of Raptor Research 36, 9196.
Martin, P. L. (1884). Illustrierte Naturgeschichte der Thiere. Leipzig: F.U. Brockhaus.
Marzluff, J. M. (2001). Worldwide urbanization and its effects on birds. In Avian
ecology and conservation in an urbanizing world. (eds. J. M. Marzluff, R. Bowman and
R. Donnelly), pp. 1938. Boston: Kluwer Academic Publishers.
Marzluff, J. M. (2005). Island biogeography for an urbanizing world: how
extinction and colonization may determine biological diversity in human-dominated
landscapes. Urban Ecosystems 8, 157177.
Marzluff, J. M. & Ewing, K. (2001). Restoration of fragmented landscapes for
the conservation of birds: A general framework and specific recommendations for
urbanizing landscapes. Restoration Ecology 9, 280292.
Marzluff, J. M., Withey, J. C., Whittaker, K. A., Oleyar, M. D., Unfried,
T. M., Rullman, S. & Delap, J. (2007). Consequences of habitat utilization by nest
Biological Reviews 85 (2010) 643667 2010 The Authors. Journal compilation 2010 Cambridge Philosophical Society
665
predators and breeding songbirds across multiple scales in an urbanizing landscape.
The Condor 109, 516534.
Mason, C. F. (2000). Thrushes now largely restricted to the built environment in
eastern England. Diversity and Distributions 6, 189194.
Mason, C. F. (2003). Some correlates of density in an urban Blackbird Turdus merula
population. Bird Study 50, 185188.
McKinney, M. L. (2006). Urbanisation as a major cause of biotic homogenization.
Biological Conservation 127, 247260.
Melchers, M. & Daalder, R. (1996). De vogels van Amsterdam. Haarlem: Schuyt and
Co.
Meltofte, H. & Fjeldsa, J. (eds) (1989). Fuglene i Danmark, bd. 1 and 2. Copenhaggen:
Gyldendal.
Merom, H. (1960). Birds of Israel. Tel Aviv: Hakibbutz Hameuchad.
Mockford, E. J. & Marshall, R. C. (2009). Effects of urban noise on song and
response behaviour in great tits. Proceedings of the Royal Society of London, Series B 276,
29792985.
Mller, A. P. (1978). Nordjyllands Fugle - deres yngleudbredelse og traekforhold. Klampenborg:
Scandanavian Press Ltd.
Mller, A. P. (2008). Flight distances of urban birds, predation and selection for
urban life. Behavioral Ecology and Sociobiology 63, 6375.
Morbach, J. (1939). Vogel der Heimat Band 1 familie der raben, stare, pirole und finken. Druk:
Herausgegeben im Auftrage der LLPO.
Mudie, R. (1834). The feathered tribes of the British islands. London: Whittaker and Son.
Muller, W. (1887). Die Vogelfauna des Grossherzogthums Hessen. Journal of
Ornithology 35, 8691.
Mulsow, R. (2005). Brussels. In Birds in European cities. (ed. J. G. Kelcey and
G. Rheinwald), pp. 127152. St. Katharinen: Ginster Verlag.
Mundinger, P. & Hope, S. (1983). Expansion of the winter range of the house finch:
19471979. American Birds 36, 347353.
Nankinov, D. (1982). Pticite na grad Sofia. Ornithological information Bulletin 12, 1386.
Nei, M. & Li, W.-H. (1979). Mathematical model for studying genetic variation in
terms of restriction endonucleases. Proceedings of the National Academy of Sciences of the
USA 76, 52695273.
Neil, K. & Wu, J. (2006). Effects of urbanisation on plant flowering phenology:
A review. Urban Ecosystems 9, 243257.
Nemeth, E. & Brumm, H. (2009). Blackbirds sing higherpitched songs in cities:
adaptation to habitat acoustics or sideeffect of urbanisation. Animal Behaviour 78,
637641.
Newhouse, M. J., Marra, P. P. & Johnson, L. S. (2008). Reproductive success of
House Wrens in suburban and rural landscapes. Wilson Journal of Ornithology 120,
99104.
Newton, I. (1986). The sparrowhawk. Calton: Poyser.
Newton, I. (2003). The speciation and biogeography of birds. London: Academic Press.
Nikolaev, S. I., Montoya-Burgos, J. I., Popadin, K., Parand, L., Margulies,
E. H. & Antonarakis, S. E. (2007). Life-history traits drive the evolutionary rates
of mammalian coding and noncoding genomic elements. Proceedings of the National
Academy of Sciences of the USA 104, 2044320448.
Nilsson, J. a. (1997). Long-tailed Tit Aegithalos caudatus. In The EBCC atlas of
European breeding birds: their distribution and abundance. (eds. W. J. M. Hagemeijer
and M.J. Blair), pp. 630631. London: T & AD Poyser.
Noel, S., Ouellet, M., Galois, P. & Lapointe, F. J. (2007). Impact of urban
fragmentation on the genetic structure of the eastern red-backed salamander.
Conservation Genetics 8, 599606.
Obukhova, N. Y. (2001). Geographic variation of colour in the synanthropic blue
rock pigeon. Russian Journal of Genetics 37, 649658.
Oddi E. (1929). Ornitologia Italiana. Milano: Hoepli.
Ogawa, K. & Mototani, I. (1985). Invasion of introduced dandelions and survival
of the native ones in the Tokyo metropolitan area of Japan. Japanese Journal of Ecology
35, 443452.
Olson, D., Dinerstein, M. E., Wikramanayake, E. D., Burgess, N. D., Powell,
G. V. N., Underwood, E. C., DAmico, J. A., Itoua, I., Strand, H. E.,
Morrison, J. C., Loucks, C. J., Allnutt, T. F., Ricketts, T. H., Kura, Y.,
Lamoreux, J. F., Wettengel, W. W., Hedao, P. & Kassem, K. R. (2001).
Terrestrial ecoregions of the world: a new map of life on earth. BioScience 51,
933938.
Olsson, V. (1971). Studies of less familiar birds 165: Serin. British Birds 64, 213223.
Palmgren, P. (1930). Quantitative Untersuchungen iiber die Vogelfauna in den
Waldern Siidfinnlands mit besonderer Berucksichtigung Alands. Acta Zoologica Fennica
7, 1218.
Palomino, D. & Carrascal, L. M. (2005). Birds on novel island environments.
A case study with the urban avifauna of Tenerife (Canary Islands). Ecological Research
20, 611617.
Paradis, E., Baillie, S. R., Sutherland, W. J. & Gregory, R. D. (1998). Patterns
of natal and breeding dispersal in birds. Journal of Animal Ecology 67, 518536.
Parslow, J. (1973). Breeding Birds of Britain and Ireland. Berkhamstead: T. and A. D.
Poyser.
Partecke, J. & Gwinner, E. (2007). Increased sedentariness in European blackbirds
following urbanisation: a consequence of local adaptation? Ecology 88, 882890.
666
Partecke, J., Schwabl, I. & Gwinner, E. (2006). Stress and the city: urbanisation
and its effects on the stress physiology in European Blackbirds. Ecology 87,
19451952.
Partecke, J., Vant Hof, T. & Gwinner, E. (2004). Differences in the timing of
reproduction between urban and forest European Blackbirds (Turdus merula): result
of phenotypic flexibility or genetic differences? Proceedings of the Royal Society of London,
Series B 271, 19952001.
Patrikeev, M. (2004). The birds of Azerbaijan. Moscow: Russian Nature Press.
Payton, M. E., Greenstone, M. H. & Schenker, N. (2003). Overlapping
confidence intervals or standard error intervals: what do they mean in terms of
statistical significance? Journal of Insect Science 3, 34.
Pax, F. (1925). Wirbeltierfauna v. Schlesien. Berlin: Gebruder Borntraeger.
Petraitis, A. (1976). Ecology of birds on the Lithuanian SSR I: the results of ornithological
research in Lithuania. Vilnius: Institute of Zoology and Parasitology of the Academy of
Sciences of the Lithuanian SSR.
Petrie, M. & Roberts, G. (2007). Sexual selection and the evolution of evolvability.
Heredity 98, 198205.
Petren, K. & Case, T. J. (1996). An experimental demonstration of exploitation
competition in an ongoing invasion. Ecology 77, 118132.
Phillimore, A. B. & Owens, I. P. F. (2006). Are subspecies useful in evolutionary
and conservation biology? Proceedings of the Royal Society of London, Series B 273,
10491053.
Price, T. D., Qvarnstrom, A. & Irwin, D. E. (2003). The role of phenotypic
plasticity in driving genetic evolution. Proceedings of the Royal Society of London, Series B
270, 14331440.
Post, W. & Wiley, J. W. (1977). The Shiny Cowbird in the West Indies. The Condor
79, 119121.
Postma, E. & Van Noordwijk, A. J. (2005). Gene flow maintains a large genetic
difference in clutch size at a small spatial scale. Nature 433, 6568.
Pujol, B. & Pannell, J. R. (2008). Reduced responses to selection after species range
expansion. Science 321, 96.
Pulliainen, E. (1963). On the history, ecology and ethology of the Mallards (Anas
platyrhynchos L.) overwintering in Finland. Ornis Fennica 40, 4565.
Puth, L. M. & Post, D. M. (2005). Studying invasion: have we missed the boat?
Ecology Letters 8, 715721.
Rabose, D., De Wavrin, H., Tricot, J. & Van Der Elst, D. (1995). Atlas des oiseaux
nicheurs de Bruxelles. Liege: Aves.
Rasanen, K. & Hendry, A. P. (2008). Disentangling interactions between adaptive
divergence and gene flow when ecology drives diversification. Ecology Letters 11,
624636.
Rasner, C. A., Yeh, P. J., Eggert, L. S., Hunt, K. E., Woodruff, D. S. & Price,
T. D. (2004). Genetic and morphological evolution following a founder event in the
darkeyed junco, Junco hyemalis thurberi. Molecular Ecology 13, 671681.
Ratcliffe, D. (1993). Peregrine Falco peregrinus. In The new atlas of breeding birds in Britain
and Ireland: 19881991. (eds. D. W. Gibbons, J. B. Reid and R. A. Chapman),
pp. 124125. T and A.D. Poyser, London.
Reisch, C. (2007). Genetic structure of Saxifrage tridactylites (saxifragaceae) from natural
and man-made habitats. Conservation Genetics 8, 893902.
Rios, R. S., Marquis, R. J. & Flunker, J. C. (2008). Population variation in plant
traits associated with ant attraction and herbivory in Chamaecrista fasciculata
(Fabaceae). Oecologia 156, 577588.
Rivalan, P., Frederiksen, M., Los, G. & Julliard, R. (2007). Contrasting
responses of migration strategies in two European thrushes to climate change.
Global Change Biology 13, 275287.
Roberts, D. G., Ayre, D. J. & Whelan, R. J. (2007). Urban plants as genetic
reservoirs or threats to the integrity. Conservation Biology 21, 842852.
Rodriguez-Mata, J. (1973). Presencia de Bubulcus ibis en la Argentina. Neotropica 19,
5556.
Rubin, C. S., Warner, R. E., Bouzat, J. L. & Paige, K. N. (2001). Population
genetic structure of Blandings turtles (Emydoidea blandingii) in an urban landscape.
Biological Conservation 99, 323330.
Rutkowski, R., Rejt, L. & Szczuka, A. (2006). Analysis of microsatellite
polymorphism and genetic differentiation in urban and rural kestrels Falco tinnunculus
(L.). Polish Journal of Ecology 54, 473480.
Rutz, C. (2008). The establishment of an urban bird population. Journal of Animal
Ecology 77, 10081019.
Schinz, H. R. (1854). Naturgesch. u. Abbildungen des Menschen. Zurich.
Schlegel, H. (1860). De dieren van Nederland, Vogels. Haarlem: Kruseman.
Schnurre, O. (1921). Die Vogel der deutschen Kulturlandschaft. Marburg.
Senar, J. C., Borras, B., Cabrera, J., Cabrera, T. & Bjorklund, M. (2006).
Local differentiation in the presence of gene flow in the citril finch Serinus citronella.
Biology Letters 2, 8587.
Sharp, D. (2002). Silencing cities. Journal of Urban Health 79, 162163.
Shepherd, W. R. (1300). London 1300 Historical Atlas. Available at http://en.wikipedia.
org/wiki/ History of London.
Shirihai, H. (1996). The birds of Israel. San Diego: Academic Press.
Shmueli, E. (1962). Changes in the avifauna of the Sharon. Teva Va-Aretz 4, 201214.
Biological Reviews 85 (2010) 643667 2010 The Authors. Journal compilation 2010 Cambridge Philosophical Society
K. L. Evans and others
Shultz, S., Bradbury, R., Evans, K. L., Gregory, R. & Blackburn, T. M.
(2005). Brain size and resource specialisation predict long-term population trends in
British birds. Proceedings of the Royal Society of London, Series B. 272, 23052311.
Simms, E. (1978). British thrushes. London: New Naturalist.
Sims, V., Evans, K. L., Newson, S. E., Tratalos, J. A. & Gaston, K. J. (2008).
Avian assemblage structure and domestic cat densities in urban environments.
Diversity and Distributions 14, 387399.
Skarphedhinsson, K. H., Petursson, G. & Hilmarsson, J. O. (1994).
Utbreisla varpfugla a Suvesturlandi Konnun 19871992. Reykjavk: Fjolrit
Natturufrd istofnunar.
Slabbekoorn, H. & Den BoerVisser, A. (2006). Cities change the songs of birds.
Current Biology 16, 23262331.
Slabbekoorn, H. & Ripmeester, E. A. P. (2008). Birdsong and anthropogenic noise:
implications and applications for conservation. Molecular Ecology 17, 7283.
Sladen, A. G. L. (1919). Notes on birds observed in Palestine. Ibis 1919, 222250.
Small, C., Pozzi, F. & Elvidge, C. D. (2005). Spatial analysis of global urban extent
from DMSPOLS night lights. Remote Sensing of Environment 96, 277291.
Smart, S. M., Thompson, K., Marrs, R. H., Leduc, M. G., Maskell, L. C. &
Firbank, L. G. (2006). Biotic homogenization and changes in species diversity
across human-modified ecosystems. Philosophical Transactions of the Royal Society of
London, Series B 273, 26592665.
Sodhi, N. S., James, P. C., Warkentin, I. G. & Oliphant, L. W. (1992). Breeding
ecology of urban merlins (Falco columbarius). Canadian Journal of Zoology 70, 14771483.
Sol, D., Duncan, R. P., Blackburn, T. M., Cassey P. & Lefebvre, L. (2005). Big
brains, enhanced cognition, and response of birds to novel environments. Proceedings
of the National Academy of Sciences of the USA 102, 53065307.
Spagnuolo, V., Muscariello, L., Terracciano, S. & Giordano, S. (2007).
Molecular biodiversity in the moss Leptodon smithii (Neckeraceae) in relation to
habitat disturbance and fragmentation. Journal of Plant Research 120, 595604.
Spencer, R. (1975). Changes in the distribution of recoveries of ringed Blackbirds.
Bird Study 22, 177190.
Stepney, P. H. R. & Power, D. M. (1973). Analysis of the eastward breeding
expansion of Brewers Blackbird plus general aspects of avian expansions. Wilson
Bulletin 8, 462464.
Stolker, C. (18661872). Versuch einer Vogelfauna der Kantone St. Gallen und Appenzell.
Svergies Ornitologiska Forening.
Sueur, F. & Sueur-Bellart, M. (1983). Urban nesting of the Fieldfare (Turdus pilaris)
in the Vosges (France). Alauda 51, 312.
Svensson, S., Svensson, M. & Tjernberg, M. (1999). Svensk fagelatlas supplement 31.
Stockholm: Var Fogelvarld.
Takami, Y., Koshio, C., Ishii, M., Fujii, H., Hidaka, T. & Shimizu, I. (2004).
Genetic diversity and structure of urban populations of Pieris butterflies assessed
using amplified fragment length polymorphism. Molecular Ecology 13, 245258.
Taylor, H. A. (1995). Urban public parks, 18401900: design and meaning. Garden
History 23, 201221.
Thompson, K. & McCarthy, M. A. (2008). Traits of British alien and native urban
plants. Journal of Ecology 96, 853859.
Thomson, G. M. (1922). The naturalisation of animals and plants in New Zealand.
Cambridge.
Thorington, K. K. & Bowman, R. (2003). Predation rate on artificial nests increases
with human housing density in suburban habitats. Ecography 26, 188196.
Timofeeff-Ressovsky, N. W. (1940). Mutations and geographical variation. In The
new systematics (ed. J. Huxley), pp. 73136. Oxford: Clarendon Press.
Tischler, F. (1941). Die Vogel Ostpreuens und seiner Nachbargebiete. Berlin: Konigsberg.
Tomiaojc, L. (1976). The urban population of the woodpigeon Columba palumbus
Linnaeus, 1758, in Europeits origin, increase and distribution. Acta Zoologica
Cracoviensia 21, 585632.
Transehe, N. V. (1965). Die vogelwelt lettlands mit Berucksichtigung der Nachbargebiete.
Hannover: Verlag Harro von Hirschheydt.
Tratalos, J., Fuller, R. A., Evans, K. L., Davies, R. G., Newson, S. E. &
Greenwood, J. J. D. & Gaston, K. J. (2007). Bird densities are associated with
household densities. Global Change Biology 13, 16851695.
Tristram, H. B. (1884). The Survey of Western Palestine: The Fauna and Flora of Palestine
XXII. London: Palestine Exploration Fund.
Tucker, B. W. (1927). A contribution to the ornithology of Naples and the Phlegraean
fields, with notes on some other neighbouring localities. Ibis 1927, 87114.
Ueta, M., Kurosawa, R., Hamao, S., Kawachi, H. & Higuchi, H. (2003).
Population change of jungle crows in Tokyo. Global Environmental Research 7, 131137.
UN (2008). World urbanisation prospects: the 2007 revision. New York: United Nations.
(Available at www.un.org/esa/population/ publications/wup2007/2007WUP
Highlights web. pdf).
UNDP (2000). World Resources 20002001. Amsterdam: Elsevier Science.
Urban, E. K., Fry, H. C. & Keith, S. (1997). Birds of Africa vol. 5: thrushes to puffback
flycatchers. London: Academic Press.
Vale, T. R. & Vale, G. R. (1976). Suburban bird populations on west-central
California. Journal of Biogeography 3, 157165.
Van Den Bosch, F., Hengeveld, R. & Metz, J. A. J. (1992). Analysing the velocity
of animal range expansion. Journal of Biogeography 19, 135150.
A conceptual framework for the colonisation of urban areas
Van Havre, G. C. M. (1928). Les Oiseaux de la faune Belge, releve documente des especes
sauvages observees en belgique. Brussels: Mautrice Lamertin.
Van Vliet, J., Musters, C. J. M. & Ter Keurs, W. J. (2009). Changes in migration
behaviour of blackbirds Turdus merula from the Netherlands. Bird Study 56, 276281.
Veit, R. R. & Lewis, M. A. (1996). Dispersal, population growth and the Allee effect:
dynamics of the House Finch invasion of eastern North America. The American
Naturalist 148, 255274.
Verheyen, R. (1948). De zangvogels van Belgie. Brussels: Koninklijk Belgisch Instituut
voor Natuurwetenschappen.
Vernon, C. J. (1997). Kurrichane Thrush Turdus libonyana. In The atlas of
southern African birds Vol 2: Passerines. (eds. J. A. Harrison, D.G. Allan,
L. G. Underhill, M. Herremans, A. J. Tree, V. Parker and C. J. Brown),
pp. 488490. Johannesburg: BirdLife South Africa.
Viksne, J. (1983). Birds of Latvia, territorial distribution and number. Riga: Latvian SSR
Academy of Sciences Institute of Biology and Zinatne Publishing House.
Vlek, R. (2000). Amsterdamse vogelhistorie. Nieuwe vogelsoorten in de regio Groot-
Amsterdam door de eeuwen heen 12851999. De Gierzwaluw 38(S1), 192.
Vogel, R. L. & Tuomenpuro, J. (1997). Dunnock Prunella modularis. In The EBCC Atlas
of European Breeding Birds: their distribution and abundance. (eds W. J. M. Hagemeijer
and M. J. Blair), pp. 506507. London: T & AD Poyser.
Voigt, A. (1908). Deutsches Vogelleben. Leipzig: Teubner.
Von Hammling, J. & Schulz, K. (1911). Beobachtungen aus der Umgegend von
Posen. Journal of Ornithology 59, 384433.
Vuorisalo, T., Lahtinen, R. & Laaksonen, H. (2001). Urban biodiversity in local
newspapers: a historical perspective. Biodiversity and Conservation 10, 17391756.
Vuorisalo, T., Andersson, H., Hugg, T., Lahtinen, R., Laaksonen, H. &
Lehikoinen, E. (2003). Urban development from an avian perspective: causes
of hooded crow (Corvus corone cornix) urbanisation in two Finnish cities. Landscape and
Urban Planning 62, 6987.
Waddington, C. H. (1961). Genetic assimilation. Advances in Genetics 10, 257293.
Walasz, K. & Mielczarek, P. (1992). The atlas of breeding birds in Ma3opolska
19851991 (south-eastern Poland). Wroc3aw: Biologica Silesiae.
Wandeler, P., Funk, S. M., Largiader, C. R., Gloor, S. & Breitenmoser, U.
(2003). The city-fox phenomenon: genetic consequences of a recent colonisation of
urban habitat. Molecular Ecology 12, 647656.
Wehtje, W. (2003). The range expansion of the great-tailed grackle (Quiscalus mexicanus
Gmelin) in North America since 1880. Journal of Biogeography 30, 15931607.
Weiserbs, A. & Jacob, J. P. (2005). Brussels. In Birds in European Cities. (ed.
J. G. Kelcey and G. Rheinwald), pp. 81102. St. Katharinen: Ginster Verlag.
WestEberhard, M. J. (2005). Developmental plasticity and the origin of species
differences. Proceedings of the National Academy of Sciences of the USA 102, 65436549.
White, M. A., Nemani, R. R., Thornton, P. E. & Running, S. W. (2002). Satellite
evidence of phenological differences between urbanised and rural areas of the
eastern United States deciduous broadleaf forest. Ecosystems 5, 260273.
Williamson, M. H. (1996). Biological invasions. London: Chapman and Hall.
Wilson, J. (1993). Magpie Pica pica. In The new atlas of breeding Birds in Britain and Ireland:
19881991. (eds. D. W. Gibbons, J. B. Reid and R. A. Chapman), pp. 386387.
London: T and A.D. Poyser.
Biological Reviews 85 (2010) 643667 2010 The Authors. Journal compilation 2010 Cambridge Philosophical Society
667
Wurlefs, M. (1994). Entwicklung einer stadtischen Population des Habichats (Accipiter
gentiles) und die rolle de elster (Pica pica) im Nahrungsspektrum des habichts. Charadrius
30, 8293.
Wust, W. (1986). Avifauna Bavariae die Vogelwelt Bayerns m Wandel der Zeit Band II.
Munchen: Im Auftrag der Ornithologischen Gesellschaft in Bayern.
Wysocki, D., Adamowicz, J., Kosciow, R. & Smietana, P. (2004). The size of
breeding territory in an urban population of the Blackbird (Turdus merula) in Szczecin
(NW Poland). Ornis Fennica 81, 112.
Yeatman, L.J. (1971). Histoire des oiseaux deurope. Paris: Societe Ornithologique de
France.
Yeh, P. J. (2004). Rapid evolution of a sexually selected trait following population
establishment in a novel habitat. Evolution 58, 166174.
Yeh, P. J., Hauber, M. E. & Price, T. D. (2007). Alternative nesting behaviours
following colonisation of a novel environment by a passerine bird. Oikos 116,
14731480.
Yeh, P. J. & Price, T. D. (2004). Adaptive phenotypic plasticity and the successful
colonisation of a novel environment. American Naturalist 164, 531542.
Zhang, X., Friedl, M. A., Schaaf, C. B., Strahler, A. H. & Schneider, A.
(2004). The footprint of urban climates on vegetation phenology. Geophysical Research
Letters 31, L12209.
Zhou, L. M., Dickinson, R. E., Tian, Y. H., Fang, J. Y., Li, Q. X., Kaufmann,
R. K., Tucker, C. J. & Myneni, R. B. (2004). Evidence for a significant
urbanisation effect on climate in China. Proceedings of the National Academy of Sciences of
the USA 101, 95409544.
Zink, R. M. (2004). The role of subspecies in obscuring avian biological diversity and
misleading conservation policy. Proceedings of the Royal Society of London, Series B 271,
561564.
IX. SUPPORTING INFORMATION
Additional supporting information may be found in the
online version of this article.
Appendix A Summary of the timing of blackbird Turdus
merula urbanisation in each geopolitical unit.
Please note: Wiley-Blackwell are not responsible for the
content or functionality of any supporting materials supplied
by the authors. Any queries (other than missing material)
should be directed to the corresponding author for the
article.
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