Urban Ecology

This is the urban century. For the first time the majority of people live in towns
and cities. Understanding how people influence, and are influenced by, the
‘green’ component of these environments is therefore of enormous significance.
Providing an overview of the essentials of urban ecology, the book begins by
covering the vital background concepts of the urbanisation process and the effect that
it can have on ecosystem functions and services. Later sections are devoted to
examining how species respond to urbanisation, the many facets of human–ecology
interactions, and the issues surrounding urban planning and the provision of urban
green spaces.
Drawing on examples from urban settlements around the world, it highlights the
progress to date in this burgeoning field, as well as the challenges that lie ahead.

Kevin j. Gaston is Professor of Biodiversity and Conservation in the Department

of Animal and Plant Sciences, University of Sheffield. He has 20 years of research
experience in environmental science, conducting studies both in the UK and overseas
on a wide range of topics, including conservation prioritisation, ecosystem services,
invasive species, people–wildlife interactions, and urban ecology.
Ecological Reviews
s e r i e s e d i t o r Hefin Jones Cardiff University, UK
series editorial board
Mark Bradford Yale University, USA
David Burslem University of Aberdeen, UK
Lindsay Haddon British Ecological Society, UK
Sue Hartley University of Sussex, UK
Mark Hunter University of Michigan, USA
Heikki Setala University of Helsinki, Finland
Phillip Warren University of Sheffield, UK

Ecological Reviews publishes books at the cutting edge of modern ecology,

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Urban Ecology
Edited by

KEVIN J. GASTON
University of Sheffield
cambridge university press
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Cambridge University Press
The Edinburgh Building, Cambridge CB2 8RU, UK

Published in the United States of America by

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Information on this title: www.cambridge.org/9780521760973

# British Ecological Society 2010

This publication is in copyright. Subject to statutory exception

and to the provisions of relevant collective licensing agreements,
no reproduction of any part may take place without
the written permission of Cambridge University Press.

First published 2010

Printed in the United Kingdom at the University Press, Cambridge

A catalogue record for this publication is available from the British Library

Library of Congress Cataloging-in-Publication Data

Urban ecology / [edited by] Kevin J. Gaston.
p. cm. – (Ecological reviews)
ISBN 978-0-521-76097-3 (Hardback) – ISBN 978-0-521-74349-5 (pbk.)
1. Urban ecology (Sociology) 2. City planning–Economic aspects. 3. Sustainable
development. I. Gaston, Kevin J. II. Title. III. Series.
HT241.U7243 2010
307.76–dc22
2010031323

ISBN 978-0-521-76097-3 Hardback

ISBN 978-0-521-74349-5 Paperback

Cambridge University Press has no responsibility for the persistence or

accuracy of URLs for external or third-party internet websites referred to
in this publication, and does not guarantee that any content on such
websites is, or will remain, accurate or appropriate.
Contents

List of contributors page vii

Preface xi

1 Urban ecology 1
Kevin J. Gaston
2 Urbanisation 10
Kevin J. Gaston
3 Urban environments and ecosystem functions 35
Kevin J. Gaston, Zoe G. Davies and Jill L. Edmondson
4 Individual species and urbanisation 53
Karl L. Evans
5 Species diversity and urbanisation: patterns, drivers
and implications 88
Gary W. Luck and Lisa T. Smallbone
6 Urbanisation and alien invasion 120
Stefan Klotz and Ingolf Kühn
7 Interactions between people and nature
in urban environments 134
Richard A. Fuller and Katherine N. Irvine
8 Urban ecology and human social organisation 172
Paige S. Warren, Sharon L. Harlan, Christopher Boone,
Susannah B. Lerman, Eyal Shochat and Ann P. Kinzig
9 Urban ecology and human health and wellbeing 202
Jo Barton and Jules Pretty
10 Bringing cities alive: the importance of urban green spaces
for people and biodiversity 230
Jon Sadler, Adam Bates, James Hale and Philip James
vi CONTENTS

11 Integrating nature values in urban planning and design 261

Robbert Snep and Paul Opdam
12 Urban futures 287
Michael L. McKinney

Index 309
Contributors

JO BARTON KARL L. EVANS

Centre for Environment & Society, Department of Animal and
Department of Biological Sciences, Plant Sciences,
University of Essex, Colchester, University of Sheffield,
Essex CO4 3SQ , UK Sheffield S10 2TN, UK

ADAM BATES RICHARD A. FULLER

School of Geography, The Ecology Centre,
Earth and Environmental Sciences, The University of Queensland,
Birmingham University, St Lucia, Qld 4072, Australia
Birmingham B15 2TT, UK
KEVIN J. GASTON
CHRISTOPHER BOONE Biodiversity and Macroecology
School of Human Evolution Group,
and Social Change, Department of Animal and Plant
Arizona State University, Sciences,
Tempe, AZ 85287, USA University of Sheffield,
Sheffield S10 2TN, UK
ZOE G. DAVIES
Biodiversity and Macroecology Group, JAMES HALE
Department of Animal and Plant School of Geography,
Sciences, Earth and Environmental Sciences,
University of Sheffield, Birmingham University,
Sheffield S10 2TN, UK Birmingham B15 2TT, UK

JILL L. EDMONDSON SHARON L. HARLAN

Biodiversity and Macroecology Group, School of Human Evolution
Department of Animal and Plant and Social Change,
Sciences, University of Sheffield, Arizona State University,
Sheffield S10 2TN, UK Tempe, AZ 85287, USA
viii LIST OF CONTRIBUTORS

KATHERINE N. IRVINE PO Box 789, Albury,

Institute of Energy and Sustainable NSW 2640, Australia
Development,
De Montfort University, M I C H A E L L . MC K I N N E Y
The Gateway, Leicester LE1 9BH, UK Department of Earth & Planetary
Sciences,
PHILIP JAMES University of Tennessee,
School of Environment and Life Knoxville,
Sciences, TN 37916, USA
University of Salford, Salford,
Greater Manchester M5 4WT, UK PAUL OPDAM
Department of Land Use Planning &
ANN P. KINZIG Alterra Landscape Center,
School of Life Sciences, Wageningen University Center,
Arizona State University, PO Box 47, 6700 AA Wageningen,
Tempe, AZ 85287, USA The Netherlands

STEFAN KLOTZ JULES PRETTY

Helmholtz Centre for Environmental Centre for Environment & Society,
Research – UFZ, Department of Biological Sciences,
Department of Community Ecology, University of Essex, Colchester,
Theodor-Lieser-Str. 4, 06120 Halle, Essex CO4 3SQ , UK
Germany
JON SADLER
I N G O L F K Ü H N School of Geography,
Helmholtz Centre for Environmental Earth and Environmental Sciences,
Research – UFZ, Birmingham University,
Department of Community Ecology, Birmingham B15 2TT, UK
Theodor-Lieser-Str. 4, 06120 Halle,
Germany EYAL SHOCHAT
Global Institute of Sustainability,
SUSANNAH B. LERMAN Arizona State University,
Graduate Program in Organismic Tempe, AZ 85287, USA
and Evolutionary Biology,
University of Massachusetts-Amherst, LISA T. SMALLBONE
Amherst, MA 01003, USA Institute for Land, Water and
Society,
GARY W. LUCK Charles Sturt University,
Institute for Land, Water and Society, PO Box 789, Albury,
Charles Sturt University, NSW 2640 Australia
 LIST OF CONTRIBUTORS ix

ROBBERT SNEP PAIGE S. WARREN

Department of Land Use Planning & Department of Natural Resources
Alterra Landscape Center, Conservation,
Wageningen University Center, University of Massachusetts-Amherst,
PO Box 47, 6700 AA Wageningen, Amherst, MA 01003, USA
The Netherlands
Preface

Recent years have witnessed an extraordinary explosion of interest in the field

of urban ecology. Not only has the academic ecological community decided
that it is both an inherently interesting and practically significant topic of
study, but there has also been a growing realisation in other quarters that an
improved understanding of the ecology of urban areas may be vital for major
issues such as human health and wellbeing.
This volume is intended to provide a broad overview of the field of urban
ecology, particularly for those wanting to find out what it is essentially about,
either as part of their general ecological education or perhaps with a view to
begin working in this area. Some of the chapter topics could be equally
relevant to almost any ecosystem (although the chapter contents would vary),
others particularly reflect the intimate links between people and ecological
pattern and process in urban areas. The book opens with chapters addressing
what urban ecology is (Chapter 1) and some of the essentials of the urbanisa-
tion process (Chapter 2) and of ecosystem function in urban areas (Chapter 3).
It moves on to consider how individual species (Chapter 4), species richness
(Chapter 5) and alien species (Chapter 6) respond to urbanisation. The next
three chapters tackle various facets of human–ecology interactions in urban
environments, respectively those between people and wildlife (Chapter 7),
between human social organisation and ecology (Chapter 8), and between
human health and wellbeing and ecology (Chapter 9). Two chapters then follow
that focus on urban green spaces and urban planning (Chapters 10 and 11).
The book concludes with consideration of the future of urban systems and
urban ecology (Chapter 12).
I am grateful to the British Ecological Society for the invitation to put this
volume together, and to all of the authors for producing and revising their
chapters amidst many competing pressures on their time. Other contributors
and a number of additional people kindly provided helpful comments on
individual chapters. Thanks are also due to P. Johnson for all her assistance.
CHAPTER ONE

Urban ecology
KEVIN J. GASTON

What is urban ecology?

Real ecologists study wild and natural places. At least, that has been the view
which has dominated the field of ecology for much of its existence. Some
lone voices argued that this was a narrow, unrealistic and unhelpful perspec-
tive, but their arguments were easily ignored by the majority as they focused
on their supposedly pristine field sites. This was always something of an
irony, as surely there was never any real doubt that less wild and natural
areas equally functioned as ecosystems, containing the same essential com-
ponents (e.g. water, soil, microorganisms, plants, animals), and being subject
to much the same processes (e.g. carbon, nutrient and water cycles). Thus,
towards the close of the twentieth century and the dawning of the twenty-
first, this viewpoint changed radically. Growing numbers of studies focused
on the patterns and processes structuring systems that had been heavily
modified, or were essentially created, by human activities. This change was
particularly evident with respect to the ecology of urban areas, with numer-
ous research studies, commentaries and reviews being published (e.g. Gilbert
1989; McDonnell & Pickett 1993; Grimm et al. 2000, 2008a, 2008b; Marzluff
et al. 2001, 2008; Pickett et al. 2001; Kelcey & Rheinwald 2005; Forman 2008).
Indeed, not only did ecologists begin to study urban systems in earnest, but
urban ecology also began to influence the study of ecology more widely. In
short, urban ecology came of age.

The rise of urban ecology

Ecology has variously been described as the scientific study of the processes
determining the abundance and distribution of organisms, of the interactions
between organisms, of the interactions between organisms and the environ-
ment, and of the flows of energy and materials through ecosystems. Urban
ecology is quite simply therefore the study of these issues within urban systems.

Urban Ecology, ed. Kevin J. Gaston. Published by Cambridge University Press.

# British Ecological Society 2010.
2 K. J. GASTON

The rise of urban ecology has arguably been fuelled by seven things. First,
there has been mounting recognition that much of the world is now covered by
human-dominated ecosystems, and that humans are integral to any general
models and understanding of ecosystems. Urban systems are simply at one
extreme of the spectrum from areas which are entirely uninfluenced by human
activities (now very scarce, if existing at all) through to those which are
predominantly shaped by interactions with people.
Second, a growing proportion of the Earth and its human population has
become urbanised, and in more and more regions almost everybody has come
to live in towns and cities. Thus, an understanding of the ecology of urban
systems has become much more obviously relevant to people’s lives and to
solving the environmental problems that they face. This has resulted in par-
ticular in attention being given to the ecosystem services (the benefits humans
obtain from ecosystems) associated with urban areas, the influence of different
forms of urban development and management on them, and how these ser-
vices can best be maintained and improved (e.g. Bolund & Hunhammar 1999;
Alberti 2005).
Third, urban areas have been studied more closely in order to elucidate the
connections between urban and rural landscapes. Indeed, urban areas are
disproportionately responsible for many of the pressures that more natural
ecosystems elsewhere experience. In large part as a consequence of the high
proportion of the human population that they account for, urban systems
make major local, regional and global demands for resources and for waste
assimilation (e.g. Rees 1992; Folke et al. 1997; Millennium Ecosystem Assess-
ment 2005; Bagliani et al. 2008). They are thus the primary drivers of habitat
loss and fragmentation (particularly through demands for food and timber)
and the multitude of consequences that follow, are the principal source
of greenhouse gas emissions (and hence have significant implications for
climate change) and many other atmospheric and aquatic pollutants, and
are major sources for biological invasions (Grimm et al. 2008a; Trusilova &
Churkina 2008).
Fourth, the potential significance for people’s health and wellbeing of their
having direct interactions with the natural world has become progressively
apparent (e.g. Ulrich et al. 1991; Kaplan 1995; Health Council of the Nether-
lands 2004; Maas et al. 2006; Fuller et al. 2007). Because such interactions are
often at their most sparse and least frequent in urban ecosystems, these areas
have also become the places in which this significance is most easily demon-
strated, in which observational studies and experiments can most readily be
conducted, in which a need to understand how these can be improved becomes
most apparent, and in which actions to do so are most pressing.
Fifth, the ecology of urban areas is rather different from that of other
systems, owing to the pervasive influence that human activities have on
 URBAN ECOLOGY 3

ecosystem structure and function. This makes them particularly interesting

subjects of ecological study because it means on the one hand that models
cannot necessarily simply be borrowed from more traditional ecology and
applied successfully to urban systems, and on the other hand that the develop-
ment of models appropriate for urban ecology may enrich ecological research
at large, especially as the human influence on ecological systems rises more
generally.
Sixth, and more methodologically, urban ecology has been fuelled by
the recognition that an understanding of urban ecosystems requires an
approach that draws on multiple disciplines, and by a generally more
positive and constructive attitude to interdisciplinary research than has
typified much of the recent history of academic discourse. Studies of urban
ecosystems require inputs from social, behavioural and economic sciences,
as they are profoundly shaped by human activities, attitudes and choices
(McIntyre et al. 2000; Alberti et al. 2003; Hope et al. 2003; Kinzig et al. 2005;
Grove et al. 2006).
One final stimulus to the study of urban ecology has been provided by the
discovery, which at the time was surprising, that in many regions of the world
areas that have experienced urban development often coincide with those that
support high native species richness and endemism (Kerr & Currie 1995;
Balmford et al. 2001; Chown et al. 2003; Gaston & Evans 2004; Gaston 2005).
Conservation biology has thus had to engage not simply with generic issues of
human population growth, but more particularly with understanding just
where that growth is taking place, what its consequences are, and how to
maintain habitats and species of conservation concern in areas with high
human densities.

Ecology in and of urban areas

Urban ecology can be viewed from two, closely related, perspectives (Grimm
et al. 2000; Pickett et al. 2001). The first addresses ecology in urban areas. It is
closely allied to traditional ecology, in that it typically investigates ecological
patterns and processes in urban areas in much the same way as they might be
examined in any other environment. That said, increasingly there is a much
greater socioeconomic component, acknowledging the dominance of human
activities in shaping urban ecosystems.
Studies in urban areas tend to be of four broad types: (i) comparison of
different land-use types within an urban setting; (ii) comparison of an urban
area with a nearby ‘natural’ area; (iii) monitoring of an urban area through
time; and (iv) gradient analysis (McIntyre et al. 2000). A particularly dominant
theme has been the study of rural–urban gradients, in which typically the
ecological characteristics of sites are compared along a, usually spatial, tran-
sect of increasing urbanisation (e.g. McDonnell & Pickett 1990; Blair 1996;
4 K. J. GASTON

Guntenspergen & Levenson 1997; McDonnell et al. 1997; Pouyat et al. 1997;
Carreiro et al. 1999; Niemelä et al. 2002). However, having resulted in many
valuable insights, this dominance is fading with a greater diversification of
approaches. Indeed, even studies treading this same route now seem to be
considering multi-dimensional environmental gradients more frequently.
Temporal patterns have received far less attention in urban environments
than have spatial patterns, largely because of a lack of appropriate data,
although this is also beginning to change (e.g. Zapparoli 1997; Morneau
et al. 1999; Chocholoušková & Pyšek 2003; Puth & Burns 2009).
The second perspective on urban ecology is a more system-oriented
approach, the ecology of urban areas. The distinction from ecology in urban
areas is not so much one of methodology per se, but rather something of a
conceptual shift. The ecology of urban areas concerns how those areas
function as aggregated wholes. There has thus been a growing number of
studies particularly of aspects of the ecology of entire cities (e.g. Hadidian
et al. 1997; Kent et al. 1999; Grimm et al. 2000; Turner 2003; Pickett &
Cadenasso 2006; Davies et al. 2008; Walton 2008). A logical extension of such
work is then the comparison of the ecologies of different cities, and this is
indeed developing rapidly, predominantly in relation to patterns of land
cover and assemblage composition (e.g. Pyšek 1998; Turner et al. 2004; La
Sorte et al. 2007).
Neither the ecology in urban areas nor the ecology of urban areas perspective
has particular precedence, and knowledge will advance most swiftly when both
are employed. An analogy might perhaps usefully be drawn between local
ecology and macroecology (Gaston & Blackburn 2000). Local studies have dom-
inated ecology for much of its existence and have contributed much to under-
standing of the dynamics of populations, communities and ecosystems.
However, a great deal remains unexplained without also taking a much
broader view, and placing local study sites in the context of landscapes and
the processes that shape the spatial flows of individuals and materials. It is
when the two views are combined that the greatest insights and predictive
power can be achieved. Likewise, the combination of studies of ecology in and
ecology of urban areas is likely to bring about the most rapid advances in
understanding, particularly because of the inherent complexity of urban
systems.
To date, urban ecology has predominantly been an observational science,
given the practical constraints on conducting field experiments posed by issues
of land ownership and access, security from interference and the spatial
variation of urban landscapes. However, some small-scale experiments have
been conducted (e.g. Denys & Schmidt 1998; Gaston et al. 2005), and there may
be considerable potential to do so at much larger scales through collaboration
with urban designers (Felson & Pickett 2005).
 URBAN ECOLOGY 5

‘Wicked’ problems
More so than in research into many other ecosystems, studies in urban ecology
are almost invariably conducted against the background of a concern to under-
stand the consequences of particular human-wrought changes to the environ-
ment. Whether the focus be a particular species, habitat or ecosystem process,
and whatever motivation drove the original investigator, such studies are apt
to be interpreted in terms of the successes or failures of past urban planning,
how the structures that resulted can best be exploited or modified, and how
future planning should be conducted. A measure of the maturity of the field of
urban ecology will be the extent to which these considerations are realistic, not
only about the complexity of urban ecosystems, but also about the formidable
array of pressures, constraints and compromises, and the interactions thereof,
that shape the planning process and contribute to (without being able entirely
to dictate) the resultant emergent urban systems. As the Royal Commission on
Environmental Pollution (2007, p. 5) states, urban systems ‘have evolved in
particular ways as agglomerations of people; accretions of buildings and roads;
infrastructures for water and energy supply and the removal of sewage and
waste; public and private spaces; places of business and residence; locations for
the production and consumption of goods and services; facilities for entertain-
ment, education and health; and so forth’.
It has been argued that urban environmental management presents a classic
case of a ‘wicked problem’ (Royal Commission on Environmental Pollution
2007); ‘wicked’ in the sense of nasty or vicious, rather than passing any
ethical judgement. Rittel and Webber (1973) observed that the kinds of societal
problems that planners deal with are intrinsically different from archetypal
problems in science in that they are ill-defined and cannot be definitively
solved. Their characterisation of the wickedness of planning problems provides
valuable context for studies in urban ecology, but arguably also extends to
some of the issues which those studies themselves set out to address: (i) there is
no definitive formulation of a wicked problem – the process of formulating the
problem and of conceiving the solution are essentially the same; (ii) wicked
problems have no stopping rule – because there is no definitive formulation
of the problem there is no point at which the solution has been found;
(iii) solutions to wicked problems are not true-or-false, but good-or-bad – the
particular solutions are likely to depend on who provides them; (iv) there is no
immediate and no ultimate test of a solution to a wicked problem – any
solution is likely to have many consequences, some of which will be unin-
tended and unexpected, which play out over potentially long periods of time;
(v) every solution to a wicked problem is a ‘one-shot operation’; because there is
no opportunity to learn by trial and error, every attempt counts significantly –
this is because planning actions are seldom reversible but have so many conse-
quences; (vi) wicked problems do not have an enumerable (or an exhaustively
6 K. J. GASTON

describable) set of potential solutions, nor is there a well-described set of

permissible operations that may be incorporated into the plan – the set
of potential solutions and the extent to which they are permissible will depend
on who provides them; (vii) every wicked problem is essentially unique – there
are always particularities to a problem which may override the commonalities
with other problems; (viii) each wicked problem can be considered a symptom
of another problem; (ix) the existence of a discrepancy representing a wicked
problem can be explained in numerous ways – the choice of explanation
determines the nature of the problem’s resolution; and (x) the planner has
no right to be wrong – the objective is to improve a situation.

Prospects
Despite its relatively late emergence as an important component of the eco-
logical research agenda, understanding of urban ecology has developed rap-
idly. The numbers of papers, books, symposia and conferences dedicated to the
topic continue on steeply rising trajectories. Contributions appear increasingly
frequently in high-profile general science journals as well as more subject-
specific ones and, perhaps more tellingly, key findings are more regularly
being cross-referenced in other fields of ecological and environmental science.
All of this reflects a vibrant and fast moving research agenda.
Of course, much work remains to be done in urban ecology. Almost all of the
subsequent, more detailed, chapters in this volume highlight questions for
which answers are urgently required, and important topics that have remained
poorly explored or are as yet unexplored. More generically, obvious issues for
future work include (i) the ecology of towns and cities in developing countries;
(ii) the relationship between the ecology of towns and cities and how urban
areas grow; (iii) improved understanding of the role of history and culture in
shaping the ecology of different urban areas; (iv) how the ecology of urban
areas will alter as a consequence of climate change and other global change
pressures; (v) more comparative work between different towns and cities; and
(vi) more experimental studies.

Acknowledgements
I am grateful to Z. Davies for helpful comments.

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CHAPTER TWO

Urbanisation
KEVIN J. GASTON

Unsurprisingly, knowledge of the pattern and process of urbanisation is key

background to an understanding of urban ecology. However, of course, the
necessary material lies largely in other fields of study. Moreover, it is widely
scattered and rather poorly synthesised. This chapter therefore provides a
broad but selective overview of some of the most important issues of especial
relevance to urban ecology. In particular, it addresses what constitutes an
urban area, how urbanisation occurs, the history of urbanisation, the scale of
urbanisation (in terms of population and household numbers, land cover and
ecological footprints), the structure of urban areas (in terms of location, size,
form and composition) and, finally, the sustainability of cities. Whilst an
attempt has been made to draw on an extensive range of examples, there is
an inevitable and unfortunate bias towards the developed regions on which the
vast majority of research, and the associated literature, has been focused.

What is urban?
Urbanisation is the process by which a rural area becomes an urban one, or the
degree to which an area is urbanised (although some differentiate the former
process as urbanisation and the latter as urbanicity). But what is an urban area?
There is no simple answer to this question, although most of us would have an
intuitive sense of what constitutes such an area and what does not, probably
based on the numbers of people and the level of cover by buildings, transport
networks and other such infrastructure. More formally, urban areas have, in
practice, been distinguished in a wide variety of ways. These include using
administrative or geopolitical boundaries (e.g. towns, cities, metropolitan
districts), functional boundaries (based on flows of people or resources), human
population size or density (of varying accuracy, depending on census tech-
niques), housing numbers or density, land cover (commonly derived from
satellite imagery) and/or other indicators of human activity (e.g. artificial

Urban Ecology, ed. Kevin J. Gaston. Published by Cambridge University Press.

# British Ecological Society 2010.
 URBANISATION 11

night time lights; Theobald 2002; Millennium Ecosystem Assessment 2005;

Small et al. 2005; Kasanko et al. 2006; UN-HABITAT 2006; Decker et al. 2007).
Almost invariably, the cut-offs between what is and is not regarded as urban are
essentially arbitrary, and arise from applied rather than theoretical consider-
ations. They variously depend, for example, on precisely which administrative
boundary scheme is used (and how through time it is amended), what levels of
flows are employed to identify functional boundaries, what levels of population
or housing are regarded as urban, and how urban land cover is distinguished
from others (including how small any associated green spaces and water bodies
need to be if they are to be treated as urban cover). These thresholds may, in turn,
be influenced by variations in the perceptions of users as to what constitutes an
urban area. Those living in regions in which human population densities and
building coverage are generally lower often regard lower thresholds of such
variables as being more appropriate cut-offs for defining urban areas.
Obviously, the definitions employed to distinguish urban areas can make
a substantial difference to their measured extent. Different authorities use different
definitions, and these may also alter through time (Cohen 2004; Montgomery
2008). For instance, the administrative definition of an urban area in New Zealand
is a settlement of more than 1000 people, with no reference to the population
density (Statistics New Zealand 2006). In contrast, in Canada it is defined as land
with more than 400 people km 2 (Statistics Canada 2001), and in Japan as areas
with a population density greater than 4000 people km 2 that contain at least 5000
people (Japanese Statistics Bureau 2005). As a consequence of such differences,
estimates of the level of urbanisation of a particular region or country may differ
markedly between different sources, and often may not be strictly comparable.
There seems to be little apparent concerted effort to resolve this formidable prob-
lem, although this may perhaps change in response to research analyses requiring
more standardised measures of urbanisation. This variation in approaches to
measuring urbanisation needs to be borne in mind when considering much of
what follows in this and other chapters in this volume. Indeed, in this spirit, I shall
subsequently treat urban areas rather broadly to include towns, cities and associa-
ted infrastructure. Arguably, even quite substantial variation in estimates of levels
of urbanisation would not markedly alter many of the generalisations reported.

How does urbanisation occur?

The urbanisation of a region can take place through three distinctly different
routes, and combinations thereof. First, it may result from the migration of
people from rural areas (or urban areas elsewhere) to existing urban areas,
leading to their intensification and/or expansion. This currently accounts for a
substantial proportion of urban population growth. Second, differences
in birth and death rates may result in greater natural population growth in
existing urban than rural areas, again leading to the intensification and/or
12 K. J. GASTON

expansion of the former. At the present time, although fertility rates tend to be
lower, health is generally better and longevity greater in urban areas than
rural ones (Dye 2008; Mace 2008). Third, natural or planned (e.g. through the
location of specific new developments or initiatives) population growth in
rural areas may result in their becoming urban, either through the expansion
of existing urban areas into their periphery or the development of independent
urban areas. These different routes to urbanisation can individually, or in
combination, lead to three distinct urban growth types: (i) infilling, in which
unurbanised land that has been surrounded by previously urban land itself
becomes urbanised; (ii) edge-expansion, in which existing urban areas spatially
expand; and (iii) spontaneous growth, in which new urban areas are formed
with no direct spatial connection to existing urban areas (Xu et al. 2007).
Models of urbanisation, both informal and formal, have a long history
(Berling-Wolff & Wu 2004a). Traditionally, urban areas have been pictured as
comprising a series of concentric circles of different land-use, radiating from
the central business district, that over time progressively expand outwards
(e.g. Burgess 1925). More recent models acknowledge that this is unduly
simplistic, and particularly highlight (i) leap-frog development, in which fresh
housing in particular is established beyond the pre-existing urban fringe;
(ii) coalescence, in which large cities emerge as a consequence of development
between pre-existing towns; and (iii) polycentric developments, in which urban
regions emerge as collections of historically distinct and independent cities
(e.g. Manrubia & Zanette 1998; Meijers 2005; Decker et al. 2007; Batty 2008).
In addition, more complex approaches to the processes involved in urbanisa-
tion have been incorporated, including, for example, reaction-diffusion, based
on simple assumptions about human reproduction and migration, and/or
fractal structures in which cities show self-similarity or hierarchical clustering
in patterns of land coverage at multiple spatial scales.
One of the most frequently used approaches to modelling urban systems spatially
employs cellular automata, typically comprising a lattice of cells, each of which
takes a particular state of land-use or land cover which may change through time
following a set of transition rules that reflect the influence of the states of neigh-
bouring cells. Complex spatial patterns can result even when the transition rules
are rather simple. Such approaches have been used both to study the general
interaction between particular sets of processes and resultant urban forms, and
to simulate the growth of specific cities (e.g. Li & Yeh 2000; Barredo et al. 2003;
Berling-Wolff & Wu 2004b; Divigalpitiya et al. 2007; He et al. 2008). They have also
been used to predict the future growth of particular urban areas (e.g. He et al. 2008).

A brief history of urbanisation

Relative to the time since the human species first appeared, urbanisation is a
rather recent phenomenon. Urban societies are thought initially to have
developed about 5000 to 6000 years ago, emerging as a consequence of humans
 URBANISATION 13

organising themselves into larger groups, establishing sedentary settlements in

which tools and facilities could be increased and agriculture pursued, and
developing cultural mechanisms for assembling and redistributing knowledge
(Redman 1999). Mesopotomia is argued to have been the first region of the world
to become urbanised (Leick 2001). The environmental impacts of some early
urban societies were severe. Typically resulting from regional degradation as a
consequence of the overexploitation of resources associated with rising popula-
tion levels, these led to the abandonment of entire cities and the collapse of some
civilisations (Redman 1999; Ehrlich & Ehrlich 2004; Diamond 2006).
Even by 1700, only a small proportion (<10%) of the world’s population
inhabited urban areas, with these individuals distributed amongst a relatively
limited number of small urban centres (Figure 2.1; Berry 1990). Parts of western
Europe in the eighteenth century were the first to achieve more sizeable
urban populations, with capital cities in particular accounting for much of
this growth. Although throughout the history of urbanisation cities have risen
and fallen in size, both in absolute and relative terms (Lanaspa et al. 2003; Batty
2006), only one city (Beijing) had a population of more than one million in
1800 (there were 16 in 1900, and more than 400 by 2000; Chandler 1987;
United Nations Centre for Human Settlements 2002).
Up until the early twentieth century, at least in Europe, urban dwellers
experienced higher rates of infant mortality than those living in rural environ-
ments, with cities apparently being maintained in large part through immi-
gration (Woods 2003; Birchenall 2007). In the first half of the century,
urbanisation was predominantly confined to countries with the highest levels
of per capita income (Cohen 2004). Indeed, rapid urbanisation began in what
are today the more developed regions of the world, with around 30% of their
population being urban in 1920, and more than a half by 1950 (United Nations
2008). In 2007, more than 80% of the populations of Australia, New Zealand
and North America were urbanised, and 72% of the population of Europe
(which, perhaps surprisingly, is the least urbanised major region of the
developed world). The level of urbanisation of a region, as reflected in the
proportion of the population living in urban areas, has thus historically tended
to increase with levels of wealth, although there may be no simple causal link
(Bloom et al. 2008). The entire abandonment or collapse of cities has been
relatively infrequent over the past two centuries, even as a consequence of
conflict and natural disasters.
The urban transition that is currently taking place across much of the world
is rather different from those that have gone before, being characterised as
(i) unprecedented in spatial scale; (ii) rapid (albeit not unprecedented) in pace;
(iii) occurring more rapidly in countries that have relatively middle to low
levels of per capita income (indeed, in some countries the growth of cities has
not had concomitant growth in economic activity); (iv) heavily dependent on
the global economy; (v) accompanied by a convergence in rural and urban
14 K. J. GASTON

Figure 2.1 Estimates of the distribution of the human population across the world
between 1700 and 2000. Reproduced with permission from Klein Goldewijk (2005).

lifestyles, particularly as a consequence of ease of transport and communica-

tion; and (vi) occurring under a different set of demographic regimes, particu-
larly abrupt declines in fertility and mortality (Cohen 2004).

The scale of urbanisation

The present and predicted scales of urbanisation are, by multiple metrics,
extraordinary. Indeed, to many minds, this is one key reason why rapid con-
tinued development of the research field of urban ecology is essential.

Population and household numbers

The percentage of the population that is urban varies markedly from one
country to another. Estimates for 2007 give a mean of 58%, ranging between
10 and 100% (Figure 2.2; United Nations 2008). However, in 2008, for the first
time in history, the global urban population was estimated to have equalled
the rural population, and has subsequently exceeded it (Figure 2.3; United
Nations 2008). That makes this the urban century, and a key point of transition
in human history, at least in terms of simple descriptive statistics. Whilst this
may conjure visions of modern cities and suburbs, it is important to recognise
that 2007 saw the number of slum dwellers – those living in urban areas in
inadequate housing with few or no basic services – exceed one billion, equating
to one in every three urban dwellers (UN-HABITAT 2006). This number is also
predicted to continue to grow.
 URBANISATION 15

Figure 2.2 The percentage of the population of each of the countries of the world that
resides in urban areas. From data in United Nations (2008).

Figure 2.3 The percentage of the population of the world (filled circles), more
developed regions (open squares) and less developed regions (open circles) estimated
(1950–2005) and projected (2010–2050) to reside in urban areas. From data in United
Nations (2008).
16 K. J. GASTON

Annual growth (%) 7

6
5
4
3
2
1
0
Vienna

Munich

Dresden

Prague

Copenhagen

Lyon

Milan

Tallinn

Dublin

Brussels

Bilbao

Porto

Helsinki

Bratislava

Palermo
Figure 2.4 Annual percentage growth of built-up areas of 15 European cities from the
mid 1950s to the late 1990s. Diamonds, 1950s–1960s; squares, 1960s–1980s; and
triangles, 1980s–1990s. Modified from Kasanko et al. (2006).

The growth rates of even well-established cities can be high (Figures 2.4–2.6).
Cairo (Egypt) grew from 6.7 million people in 1976 to more than 10 million in
2002 (49% increase; Stewart et al. 2004), Las Vegas (USA) from 557 000 people in
1985 to nearing 1.7 million in 2004 (205% increase; UNEP 2007), Shanghai
(China) from 10.8 million people in 1975 to 13.4 million in 2003 (24% increase)
and from 159.1 km2 in 1975 to 1179.3 km2 in 2005 (641% increase; Zhao et al.
2006), and Nanjing metropolitan area (China) from 128 km2 in 1979 to 461 km2
in 2003 (260% increase; Xu et al. 2007). Significant differences exist in the
growth patterns of urban areas, variously as a consequence of available
resources, juxtaposition to trade and transportation routes, investment, innov-
ation and cultural factors.
The global urban population is predicted to increase from 3.29 billion in
2007 to 4.58 billion in 2025, and 6.40 billion in 2050. This is equivalent to
almost doubling by 2050 to approximately the same size as the entire popula-
tion (rural and urban) in 2004 (United Nations 2008). Indeed, urban areas
in less developed regions will account for most of the world’s population
growth. However, whilst the majority of people have lived in urban areas in
more developed regions since at least 1950, they will not be in the majority
in less developed regions until around 2019.
The rate of growth in numbers of households or dwellings is currently
greater than that of the populations in much of the developed world, even
increasing in some regions in which population size is declining (Liu et al. 2003;
Lepczyk et al. 2007). Foremost, this has arisen as a consequence of social
changes, including increases in longevity, single occupancy and divorce rates,
as well as in some regions a growing number of second homes.
 URBANISATION 17

3
2

5
4
6
11

7
10

8
9

Kilometres
0 5 10 20 30 40

Figure 2.5 Urban expansion of Beijing. Light grey indicates urban area in 1991; dark
grey, urban area in 1991–2004. 1, Yanqing; 2, Huairou; 3, Miyun; 4, Pinggu;
5, Changping; 6, Shunyi; 7, Mengtougou; 8, Fangshan; 9, Daxing; 10, Tongzhou;
11, Capital airport. Modified from He et al. (2008).

Land cover
Various attempts have been made to determine the global coverage of urban
or built-up areas. Estimates include 260 092 km2 (0.2%; Hansen et al. 2000),
260 117 km2 (0.2%; Loveland et al. 2000), 3 million km2 (2.3%; Wackernagel
et al. 2002), 3 673 155 km2 (2.8%; Millennium Ecosystem Assessment 2005),
2 213 496 km2 (1.7%; Salvatore et al. 2005), 3 485 596 km2 (2.7%; Salvatore et al.
2005) and 500 000 km2 (0.38%; Sterling & Ducharne 2008; all percentages
calculated using a global land extent, excluding permanent ice cover, of
c.130 million km2). Much of the marked variation in these figures results from
differences in the spatial resolution and approach of analyses, particularly in
18 K. J. GASTON

(a)

28

24
Number of cities

20

16

12

0
–2 –1 0 1 2 3 4 5 6 7
Population
(b)
40

35

30
Number of cities

25

20

15

10

0
0 2 4 6 8 10 12 14 16
Area

Figure 2.6 Annual percentage change in (a) the population and (b) the built-up area
(km2) of 90 cities of over 100 000 individuals, calculated for various combinations
of years between 1984 and 2002. From data in Angel et al. (2005).

the extent to which smaller settlements and infrastructure (e.g. roads) are
included. Nonetheless, whichever figures are used, the percentage coverages
are low in comparison with croplands covering c.18 million km2 and grazing
lands comprising 36 million km2 (Sterling & Ducharne 2008). However, the
consequences of this urban coverage, particularly in terms of resource
demands, are far reaching (see below).
 URBANISATION 19

35

30
Number of countries

25

20

15

10

0
–4.0 –3.5 –3.0 –2.5 –2.0 –1.5 –1.0 –0.5 0.0 0.5 1.0 1.5 2.0
Log10 Percentage urban

Figure 2.7 Percentage of land area in 2000 that is urban and built-up for 163
countries for which this percentage is greater than zero. From data in World
Resources Institute (2007).

Angel et al. (2005) estimated that in 2000, cities with populations in excess of
100 000 covered c. 200 000 km2 both in developing countries and in industrial-
ised countries. These numbers are expected to increase to 600 000 km2 and
500 000 km2, respectively, by 2030.
The percentage of the areal extent of individual countries that is covered
by urban areas is hugely variable, ranging from close to zero to about 32%
(Figure 2.7). Unsurprisingly, the coverage of urban areas generally increases
with the size of the associated human population. However, the intercept and
slope of this relationship vary between regions. For example, the intercept is
greater and the slope is shallower for large cities in Europe than for large
cities in the USA (Figure 2.8). Thus, whilst the former house more people for a
given urban area, they accrue people at a slower rate for a given increase in
that area. The first of these outcomes is likely to be a consequence of the
suburban sprawl that is particularly marked around many cities in the USA
(Theobald 2005).
Not only are population size and urban area positively correlated for particu-
lar points in time, they also tend to scale positively through time for any
given urban area. For much of the world this relationship is commonly such
that the area occupied increases faster than the population size, with new
residents effectively occupying more land per capita than do existing residents
(Marshall 2007). This follows from the previously mentioned tendency for
the rate of growth in numbers of households to outstrip that in population
20 K. J. GASTON

4.5

4.0

3.5
Log10 City area

3.0

2.5

2.0

1.5

1.0

0.5
4.8 5.2 5.6 6.0 6.4 6.8 7.2
Log10 Human population

Figure 2.8 The relationship between city area (km2) and the number of human
inhabitants, for cities exceeding 100 000 inhabitants in Europe (filled circles) and the
USA (open circles). From unpublished data kindly provided by R. A. Fuller.

size, and is exacerbated by rising living standards (with demands for more
space per person) and changing land-use policies (e.g. attitudes to urban sprawl;
Kasanko et al. 2006).

Ecological footprint
Industrialised cities have energy fluxes of between 100 000 and 3 000 000 kcal
m 2 yr 1, in contrast to natural ecosystems which flux between 1000 and
40 000 kcal m 2 yr 1 (Odum 1997). This is one reflection of the influence of
urban areas reaching far beyond their geographic limits, often drawing on
resources and ecosystem services from distant lands. The ecological footprint of
a city is the total area of biologically productive land and water required to
generate the renewable resources that it consumes and assimilate the waste it
produces, using prevailing technology (Rees 1992, 2001; Wackernagel et al.
2006). Although the details of the accounting vary between studies, and the
approach has its limitations (Chapter 12), cities commonly have ecological
footprints one or two orders of magnitude larger than the urban areas them-
selves (Rees 1992, 1999; Wackernagel et al. 2006). For example, one estimation
of the ecological footprint of Greater London in 2000 was 293 times its
geographical area, which is twice the size of the UK, and roughly the same size
as Spain (Best Foot Forward 2002). Likewise, in 1996, metropolitan Toronto had
an ecological footprint that was 290 times its geographical area, with each
citizen having a footprint of 7.7 ha (Rees 2001). It is important to remember,
 URBANISATION 21

however, that in many cases per capita footprints may be smaller for urban
than rural areas, because of the efficiencies of urban living.
At a global scale the ecological footprint of humanity is estimated already
to exceed the available capacity, which if sustained will lead to the loss of
ecological capital (Wackernagel et al. 2006; Kitzes et al. 2008). This demand
arises disproportionately from the urban segment of the population, particu-
larly that in developed nations.

The structure of urban areas

Location
The human population is not randomly distributed across the Earth’s terres-
trial surface (Figure 2.1). Although the precise figures are dependent on the
spatial resolution of analysis, on a grid of 2.5 arc-minute quadrilaterals, half
of the potentially habitable area contains less than 2% of the population,
whilst 50% of the population occurs in less than 3% of the potentially
habitable area (Small & Cohen 2004). The density of people is greatest at
sea level and c. 2300 m elevation (the densely populated volcanic highlands
of Mexico), increases at high levels of variability in precipitation, peaks at
tropical and temperate mean temperatures and towards arid (but not the
driest) climates, and declines with distance from coastlines and major rivers
(Small & Cohen 2004). The physiographic variables appear to have a much
stronger influence than do the climatic ones, although one consequence of
the combination of influences is that, at reasonably coarse spatial reso-
lutions, in many regions of the world human density tends to increase with
levels of net primary productivity (or at least correlates thereof; Chown et al.
2003; Evans & Gaston 2005; Gaston 2005). Many of these variables tend also
to be correlated with native and endemic species richness, resulting in
important patterns of covariation with human density at these low reso-
lutions (Chapter 5).
Looking at different broad ecosystem types, the highest proportion of the
human population is urbanised within coastal and inland water zones, with
these also having the highest urban population densities, and the highest propor-
tions of the areas of coastal and cultivated lands being urbanised (Table 2.1).
Conversely, the lowest proportion of the human population is urbanised within
mountain and forest lands, which also have the lowest proportions of their
areas urbanised (Table 2.1). The proportion of the population that is urbanised
is quite similar across Latin America, North America, Europe and Oceania
(c. 70–80%, depending on source), and much lower in Africa and Asia (c. 35–40%;
Millennium Ecosystem Assessment 2005).
The probability that an area becomes urbanised and the intensity of urban-
isation have been shown to be a function of a large number of variables,
including topography, soil type, water availability, population density,
Table 2.1 Estimated population size, population density and land area of different broad ecosystem types, divided into urban and rural. Note
that population numbers for ecosystems do not sum to totals, as systems are not mutually exclusive.

Population Population density Land area

Total Urban Rural Urban share Overall Urban Rural Total Urban Rural Urban share

(million) (%) (person per square km) (square km) (%)

Coastal zone 1 147 744 403 64.9 175 1 119 69 6 538 097 664 816 5 873 281 10.2
Cultivated 4 233 1 914 2 309 45.3 119 793 70 35 475 983 2 412 618 33 063 350 6.8
Dryland 2 149 963 1 185 44.8 36 749 20 59 990 129 1 286 421 58 703 698 2.1
Forest 1 126 401 725 35.6 27 478 18 42 092 529 839 094 41 253 435 2.0
Inland water 1 505 780 726 51.8 51 826 25 29 439 286 943 518 28 495 767 3.2
Mountain 1 154 349 805 30.3 36 636 26 32 083 873 548 559 31 535 242 1.7

World 6 052 2 828 3 224 46.7 46 770 25 130 669 507 3673 155 126 996 316 2.8

Source: Modified from Millennium Ecosystem Assessment (2005)

 URBANISATION 23

proximity to population centres, markets and transportation networks, and

land-use policies (e.g. Kline et al. 2001; Angel et al. 2005; Aguayo et al. 2007;
Conway & Hackworth 2007; Divigalpitiya et al. 2007; Xu et al. 2007; Davies et al.
2008). The particular weighting given to different factors varies between
regions and through time (particularly as the dependence of urban populations
on local food production declines, and the role of transportation of goods and
services, and thus the importance of good transportation pathways, increases).
However, within reasonably homogeneous regions empirical models can be
built to predict which areas have the greatest probability of being developed in
the near future.

Size
The size of a city or urban area is dependent on a complex set of factors,
including its history, regional topography and environment, and socioeco-
nomics (including production patterns, income distribution and economic
growth). There are many more small urban areas than large ones. Indeed, the
frequency distribution of the sizes of cities or urban areas more generally, as
reflected in the numbers of people, is strongly right-skewed.
As has been the case with species-abundance distributions (the frequency
of species of different abundances) in community ecology, there has been
much vigorous debate as to the precise form taken by the frequency distri-
bution of settlement or city sizes. It has principally been argued to follow
either a power function or a lognormal distribution (Zipf 1949; Manrubia &
Zanette 1998; Eeckhout 2004; Decker et al. 2007; Batty 2008). Zipf’s law
describes the distribution of city sizes in terms of a power function in
which, on logarithmic scales, the population size of a city is linearly and
inversely related to the rank of that population size, with an exponent that
approximately equals unity (Zipf 1949). However, this seems to apply best to
larger cities, with the full distribution being better described as lognormal
(Eeckhout 2004; Decker et al. 2007). An approximate lognormal would accord
with Gibrat’s law (Gibrat 1931), that such distributions emerge when the
growth rates of entities, in this case cities, are random (or at least independ-
ent of entity size) and so is their variance. The extent to which this is
actually true for cities is, however, contentious (Batty 2008; Garmestani
et al. 2008). Although there is some empirical support, at least for some
periods and regions of the world (Eeckhout 2004; Resende 2004), there is
also contradictory evidence for declines in growth rates of cities at larger
sizes (i.e. density dependence; e.g. Garmestani et al. 2007).
Whatever its detailed form, the frequency distribution of settlement
sizes is such that most people live in smaller towns and cities (Figure 2.9).
In 2007, more than half of the world’s urban population lived in cities or
towns with fewer than half a million inhabitants (United Nations 2008).
24 K. J. GASTON

1800 60

1600
50
1400
Population (millions)

1200 40

Percentage
1000
30
800

600 20

400
10
200

0 0
0 to 0.5 0.5 to 1 1 to 5 5 to 10 >10

Size class (millions)

Figure 2.9 The population of the world residing in each of five size classes of urban
settlement in 2005 (filled circles; left axis), and the percentage of total urban population
in each class (open circles; right axis). Size classes of settlement are as follows: fewer
than 500 000 individuals; 500 000 to 1 million; 1 to 5 million; 5 to 10 million; 10 million
or more. From data in United Nations (2008).

These are likely to absorb much of the urban growth that is projected to
occur in the next few decades, in part because they are destinations for
many rural migrants.
The largest cities (or urban agglomerations) are extremely large. Tokyo
(35.3 million) is at present the only so-called hypercity or metacity of more
than 20 million inhabitants, although a number of others are projected to
reach this level in the near future (e.g. Mumbai, São Paulo, Ciudad de México
(Mexico City), Delhi, New York–Newark; United Nations 2008). Megacities of
more than 10 million inhabitants currently comprise Ciudad de México,
New York–Newark, São Paulo, Mumbai, Delhi, Shanghai, Kolkata (Calcutta),
Dhaka, Buenos Aires, Los Angeles–Long Beach–Santa Ana, Karachi, Cairo, Rio de
Janeiro, Osaka–Kobe, Manila, Beijing and Moskva (Moscow) (United Nations
2008). However, these largest cities account for only a relatively small propor-
tion of the total urban population (<9%; United Nations 2008). Many important
demographic, socioeconomic and behavioural characteristics, such as housing,
employment, energy and water consumption, gross domestic product, wages,
bank deposits, numbers of patents and numbers of crimes, are scaling func-
tions of city size (Bettencourt et al. 2007).
 URBANISATION 25

350
Jakarta
300
Population density

1998
(persons ha–1)

250
1990
200
1980
150
1970
100
50
0
0 5 10 15 20 25
Distance (km)

350
Bangkok
300
Population density

1998
(persons ha–1)

250
1990
200
1980
150
1970
100
50
0
0 5 10 15 20 25
Distance (km)

500
Metro Manila
Population density

400 1995
(persons ha–1)

300 1990
1980
200
1970
100

0
0 5 10 15 20 25
Distance (km)

Figure 2.10 Population density as a function of distance from the centre of the cities of
Jakarta (Indonesia), Bangkok (Thailand) and Metro Manila (Philippines). Reproduced
with permisson from Murakami et al. (2005).

Form
Urban areas vary greatly in structure or form (the patterning of land cover and
transport networks). Particular focus has fallen on the sprawl that epitomises the
fringes of many metropolitan areas in the USA, and on the notion of compact
cities, with a high density of buildings and limited interstitial space, which is more
typical of parts of Europe (Breheny 1997; Radeloff et al. 2005; Tsai 2005; Irwin &
Bockstael 2007). However, these reflect only some of the major differences.
26 K. J. GASTON

(a) (b)

(c) (d)

Figure 2.11 Variation in the urban form of the city of Leicester, UK, as reflected in eight
1 km  1 km areas. Images # Bluesky International and # Infoterra 2006.

Differences in urban form result from the complex variation in, and inter-
plays between, early settlement, industrialisation, land ownership, planning,
regulation and infrastructure development (Iverson & Cook 2000; Huang et al.
2007). Nonetheless, some patterns do exist. The structure of many cities has
traditionally been characterised in terms of a curvilinear decline in human
density with distance from the central business district (a distance-decay func-
tion, commonly represented as a negative exponential), albeit with the precise
form of this gradient varying greatly from city to city (Clark 1951; Edmonston
et al. 1985). However, the explanatory power of this pattern has clearly been
declining with the suburbanisation (decentralisation) of cities particularly in
western countries, including both the loss of population in the more built-up
 URBANISATION 27

(e) (f)

(g) (h)

Figure 2.11 (cont.)

areas and growth in low-density peripheral developments (Zheng 1991;

Bunting et al. 2002; Stewart et al. 2004). In terms of these density gradients,
available data suggest that the Asian megacities, for example, are at different
stages of urbanisation, with Jakarta undergoing suburbanisation, Metro Manila
at an earlier stage and Bangkok at an intermediate stage (Figure 2.10).
Cities of developed regions tend to be more complex (i.e. irregular in
shape), less compact and more porous (greater ratio of open space to
urban area), and have less dense human populations than those in developing
regions (Huang et al. 2007). Much of this difference is associated with variation
in levels of mobility, and particularly car transport, which facilitates low-
density living.
28 K. J. GASTON

Composition
Urban areas are heterogeneous at multiple spatial resolutions (Figure 2.11).
A wide variety of schemes have been employed to characterise the land cover
and land-use of urban areas (e.g. Iverson & Cook 2000; Pauleit & Duhme 2000;
Cadenasso et al. 2007; Gill et al. 2007). From the perspective of urban ecology, the
first key distinction is between impervious surfaces (buildings, roads etc.), ‘green
spaces’ (parks, gardens/backyards, allotments, road verges, school playing fields,
sports fields, green landscaping) and ‘blue spaces’ (streams, rivers, ponds, lakes).
The coverage of these different kinds of areas varies enormously between cities.
Published estimates of the extent of green space in different urban areas
are rather scarce. However, a survey of 386 cities across Europe using remote-
sensed land cover data revealed that whilst averaging 18.6%, green coverage
ranged from 1.9% (Reggio di Calabria, Italy) to 46% (Ferrol, Spain; Fuller &
Gaston 2009). Per capita green space provision varied by two orders of magni-
tude, from 3 to 4 m2 per person in Cádiz, Fuenlabrada and Almerı́a (Spain) and
Reggio di Calabria (Italy) to more than 300 m2 in Liège (Belgium), Oulu (Fin-
land) and Valenciennes (France). Coverage was lowest in the south and east,
increasing to the north and northwest of Europe.
Owing to their much larger number, small parcels of green space contribute
disproportionately to overall coverage of urban areas (Fuller et al. 2010), so the
spatial grain at which green space is measured will have a large impact on the
resulting figures. This is an asymmetric effect, such that if one starts with areas
that at a coarse resolution are scored as urban/built-up, increasing the reso-
lution of the mapping will result in more green spaces being found. The
converse would occur if mapping a rural area, but such areas are of course
excluded from urban studies by definition.
Each of the different categories of surface (impervious, green, blue) and
each of its components may exhibit complex patterns of spatial variation
(Figure 2.12). Surprisingly, however, these have been characterised in rather
few studies.
Of course, patterns of land cover are not static. For example, in the UK
available evidence suggests that the extent of green space within urban areas
has declined substantially in recent years (Pauleit et al. 2005). This has resulted
chiefly from a combination of the loss of brownfield sites (including aban-
doned industrial areas) and municipal green spaces (e.g. school playing fields)
to development (brownfield sites are commonly treated as green spaces in this
region), and the increasing density of housing within new developments and
within existing housing areas through infilling and backland development
(Royal Commission on Environmental Pollution 2007). This has been further
exacerbated by the extension of existing buildings, the paving over of front
gardens to provide off-road parking, road widening schemes and the general
progressive erosion of smaller green spaces.
 URBANISATION 29

Figure 2.12 Environmental surfaces for Sheffield at 250 m  250 m cell resolution:
(a) extent of green space (m2); (b) vegetation cover Normalised Difference Vegetation Index
(NDVI); (c) tree cover (m2); (d) elevation (m); (e) degree angle of slope; (f) buildings area (m2);
(g) total road length (m); (h) road nodes (junctions); (i) density of households in blocks of
flats; (j) terraced housing density; (k) semi-detached housing density; (l) detached housing
density; and (m) total housing density. All housing densities were measured as
households per hectare. Reproduced with permission from Davies et al. (2008).
30 K. J. GASTON

Sustainable cities
Urbanisation has profound consequences for the environment. These include
changes in:

• habitat cover (Blair 1996; Zhao et al. 2006);

• habitat fragmentation (Radeloff et al. 2005; Irwin & Bockstael 2007; Di
Giulio et al. 2009; Evans et al. 2009);
• nutrient flows (Chapter 3);
• light levels (Cinzano et al. 2001; Small et al. 2005);
• noise levels and other acoustics (Bolund & Hunhammar 1999; Warren et al.
2006);
• atmospheric and aquatic chemical composition (Chapter 3);
• carbon stocks and flows (Chapter 3);
• water cycles (Chapter 3);
• ground and air temperatures (Chapter 3);
• disturbance regimes (Evans et al. 2009);
• species phenology (Chapter 4);
• species life history (Chapter 4);
• species dispersal and migration (Chapter 4);
• species composition and abundance (Chapter 5); and
• biotic interactions (including parasites and diseases; Chapter 4).

Subsequent chapters in this book address many of these differences.

Concerns about such consequences have contributed to the notion that
cities need to be made much more sustainable than is the case at present.
A variety of approaches are available for so doing (Chapter 12). Critical to the
notion of sustainable cities is the issue of the spatial extent over which sustain-
ability is considered. With their high concentrations of people, cities necessar-
ily draw large amounts of resources from elsewhere and export waste
materials. This will always be so, and as such cities will never be sustainable.
Whilst the notion that the inflows and outflows to and from cities should
not exceed the capacity of their immediate hinterlands is an attractive one,
this is obviously difficult or impossible in the context of regional, continental
and global markets, and indeed this has long been the case. What is important
is that the efficiencies inherent in bringing large numbers of people into
close proximity are maximised, and the impacts of their resource use
minimised.

Acknowledgements
Z. G. Davies, F. Eigenbrod, K. L. Evans, R. A. Fuller, S. Gaston and
P. Johnson are kindly thanked variously for their advice, assistance and
comments.
 URBANISATION 31

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CHAPTER THREE

Urban environments
and ecosystem functions
KEVIN J. GASTON, ZOE G. DAVIES
AND JILL L. EDMONDSON

Urbanisation profoundly changes both the abiotic and biotic properties of

ecosystems. It does so not just within urban areas, but also in surrounding
landscapes and often much further afield. Traditionally, the foremost focus for
research has been on the negative impacts of these changes, particularly for
human health and wellbeing, and how these can most effectively be mitigated.
This is readily understandable given that, for much of their history, urban
environments have often been associated with high rates of infant mortality,
disease outbreaks and a generally poor quality of life, and that this still
remains true for many of those living in cities today (Woods 2003; UN-HABITAT
2006; Birchenall 2007). From a broader ecological perspective, urban areas have
also long been considered depauperate in comparison to their rural counter-
parts in terms of flora and fauna, with the exception of a few notable species
that were widely categorised as pests.
More recently, research into urban environments has increasingly shifted
towards examining the positive contributions that such areas can make both to
the human population and to other species. Of course, viewing urban environ-
ments in terms of the benefits they can provide or the costs they can exact are
essentially two sides of the same coin. However, closer consideration of the
potential advantages has served to broaden the range of environmental issues
that have received emphasis; rather than focusing almost exclusively on the
undoubtedly vital human health concerns resulting from poor air quality,
unclean water and inadequate sanitation, there is now a growing appreciation
of the benefits that greener and more ecologically diverse urban areas have
on the mental and physical status of residents, on economic markets and for
biological conservation (Chapters 7–11; Fuller et al. 2010; Gaston & Evans 2010;
Irvine et al. 2010).
This focus on the positive contributions that urban areas can make to human
health and wellbeing is closely associated with the concept of ecosystem service

Urban Ecology, ed. Kevin J. Gaston. Published by Cambridge University Press.

# British Ecological Society 2010.
36 K. J. GASTON, Z. G. DAVIES AND J. L. EDMONDSON

provision, which can be broadly defined as the benefits people obtain from
ecosystems (e.g. Bolund & Hunhammar 1999; Pataki et al. 2006; Tratalos et al.
2007). Such services can be divided into four main categories (Millennium
Ecosystem Assessment 2005): (i) supporting services (e.g. soil formation,
photosynthesis, primary production, nutrient cycling, water cycling); (ii) regu-
lating services (e.g. air quality regulation, climate regulation, water regulation,
erosion regulation, water purification and waste treatment, disease regula-
tion, pest regulation, pollination, natural hazard regulation); (iii) provisioning
services (e.g. food, fibre, genetic resources, biochemicals, natural medicines,
pharmaceuticals, fresh water); and (iv) cultural services (e.g. cultural diversity,
spiritual and religious values, knowledge systems, educational values, inspir-
ation, aesthetic values, social relations, sense of place, cultural heritage values,
recreation and ecotourism). Plainly, urban ecosystems supply services from
each of these different categories, although some are obviously more pertinent
than others. In certain cases this is foremost a matter of provision to local
human populations (e.g. pollination). Alternatively, for services such as carbon
sequestration, although the immediate benefits to the local population may
be less significant, urban centres can contribute substantially to provision at
a regional scale.
The transition towards appreciating the benefits that can be obtained from
urban areas, rather than simply mitigating their negative influences, may also
increasingly be driven by the recognition that the long-held belief that rural
landscapes are better for native biota is breaking down in some parts of the
world. This is particularly true in regions where agriculture has become very
intensive and has restricted natural or semi-natural vegetation to small and
highly fragmented areas, resulting in some components of urban environments
(especially suburban locations with extensive coverage by green space and high
habitat heterogeneity) acting as havens for wildlife (Beebee 1997; Gregory &
Baillie 1998; Gehrt & Chelsvig 2004; Peach et al. 2004; Gaston & Evans 2010).
This chapter provides a broad and selective overview of the changes that
urbanisation can have on ecosystem functions and services. Issues specifically
relating to biodiversity and cultural services are addressed in other chapters
(Chapters 5 and 8).

Temperature
Urban environments across a wide range of latitudes and climatic regimes
commonly experience ‘heat island’ effects, in which air temperatures are
elevated compared with surrounding landscapes (e.g. Karl et al. 1988; Taha
et al. 1999; Pickett et al. 2001; Baker et al. 2002; Arnfield 2003). For larger cities
the maximum difference between urban and rural ambient temperatures is
6–12  C (Kaye et al. 2006), and air temperatures are often 2–10  C higher in
urban areas than neighbouring ones (Shepherd 2005). Indeed, in general, mean
 URBAN ENVIRONMENTS AND ECOSYSTEM FUNCTIONS 37

(a) Mesoscale

Urban ‘plume’

Mixing layer
PBL
UBL

(b) Rural BL
Surface layer

Rural Urban Rural

(b) Local scale (c) Microscale

Inertial
Surface sublayer
layer
Roughness
sublayer
Roughness (c)
sublayer UCL
UCL

Figure 3.1 Urban environments at different spatial scales, with an emphasis on

atmospheric components: (a) mesoscale; (b) local scale; (c) microscale. PBL, planetary
boundary layer; UBL, urban boundary layer; UCL, urban canopy layer; BL, boundary
layer. Reproduced with permission from Shepherd (2005), modified after Oke (1987).

and minimum temperatures are often increased, and daily periods of cool
temperatures, cool seasons and frost days are all reduced. These effects are
exhibited at ground level, in the urban canopy layer (which extends from
ground to roof level, and within which are urban street-canyon flows) and in
the urban boundary layer (above roof level, and developing downwind of the
leading edge of a city; Figure 3.1), although the patterns may not be entirely
congruent (Arnfield 2003) and may be spatially complex (Gaffin et al. 2008).
Ordinarily, the heat island phenomenon is greatest within large cities, during
the summer and at night, and lessens with rising wind speed and more
extensive cloud cover.
Urban heat island effects are exacerbated by a number of different factors,
but principally tend to increase with proportional coverage by impervious
surfaces (e.g. buildings and roads, which often have albedos and heat capacities
that result in the storage of incoming solar radiation as sensible heat during
the day and its release as long wave radiation at night). Conversely, they will
decrease relative to the proportion of unsealed surface, and more particularly
green space (Henry & Dicks 1987; Lo et al. 1997; Chen & Wong 2006; Chen et al.
2006; Jenerette et al. 2007; Weng et al. 2007; but see Gaffin et al. 2008).
The benefits accrued from planting urban vegetation tend to be non-linear,
38 K. J. GASTON, Z. G. DAVIES AND J. L. EDMONDSON

such that the addition of even small amounts of green material where previ-
ously there was none makes a much greater difference to ambient tempera-
tures than small increases in areas where the coverage was already extensive.
This vegetation can be effective in a wide diversity of forms, including green
roofs and green walls (Alexandri & Jones 2008).
In regions where urban temperatures become uncomfortably warm, the
demand for air-conditioning in buildings is likely to rise. The planting of
vegetation can therefore also contribute to reductions in fossil fuel consump-
tion and resultant carbon dioxide (CO2) emissions (Akbari et al. 1997; Taha et al.
1999). In particular, levels of tree cover can have a marked influence on energy
use in buildings, as well as outside air temperatures, by (i) reducing the
heat gain by lowering ambient temperatures through evapotranspiration
(the transfer of water to the atmosphere by plants); (ii) increasing the latent
air-conditioning load by adding moisture to the air through evapotranspira-
tion; (iii) reducing the outside air infiltration rate by lowering ambient wind
speeds; (iv) reducing solar heat gain through windows, walls and roofs by
shading them; and (v) reducing radiant heat gain from the surroundings by
shading (Akbari 2002). Trees can also improve the thermal comfort of urban
dwellers more generally (Stone & Rodgers 2001; Georgi & Zafiriadis 2006), with
the positive impacts tending substantially to outweigh any negative influences
on temperatures during the winter (e.g. through shading buildings). Given that
the determinants of tree cover can be both biophysical (e.g. soil type, drainage,
herbivores) and socioeconomic (e.g. wealth, cultural norms), this provides an
example of how in urban areas most ecosystem functions routinely change in
response to these two sets of factors, and the limited applicability of models
derived from more natural environments.
The influence of urbanisation on air temperatures appears to be much
smaller than the effects of global climate change, although not insignificant
when combined with other land cover changes ( Jones et al. 1990; Kalnay & Cai
2003). Nonetheless, the temperature increases associated with urbanisation can
have a variety of ecological consequences, some of which extend beyond the
urban area itself, including adjustments relative to rural areas in the timing of
germination, leaf flush, leaf drop, flowering and length of growing season of
plants, and in the breeding of animals (Chapter 4; White et al. 2002; Partecke
et al. 2004; Zhang et al. 2004a, 2004b; Neil & Wu 2006). The potential knock-on
impacts that these phenological differences may have on species survival,
assemblage structures and community dynamics are clearly diverse.

Water quality
Urbanisation is commonly, and perhaps almost universally, associated with
declines in the quality of the water carried in streams and rivers (e.g. Paul &
Meyer 2001; Walsh et al. 2005; Zhao et al. 2006). Indeed, it has been argued that
 URBAN ENVIRONMENTS AND ECOSYSTEM FUNCTIONS 39

urbanisation is second only to agriculture in its negative impacts on water

quality (Paul & Meyer 2001). This quality is typically measured by some com-
bination of (i) temperature, which may be altered in urban areas as a conse-
quence of the loss of riparian vegetation, decreased groundwater discharge,
industrial outputs and the heat island effect, and affects oxygen solubility and
the rate of biochemical processes; (ii) pH, which influences the dissolving of
ions; (iii) dissolved oxygen, which is important for aquatic fauna and the taste
of water; (iv) biochemical oxygen demand; (v) faecal coliform bacteria, indica-
tive of inadequate wastewater treatment or sewage plant overflow; (vi) phos-
phates and nitrates, which may be at heightened concentrations over hundreds
of kilometres from the point of origin and are agents of eutrophication; and
(vii) turbidity and total suspended solids, both of which have an effect on
aquatic fauna and the treatment cost of drinking water (Paul & Meyer 2001;
Duh et al. 2008). Water pollutants arise from a variety of sources, including
wastewater, fertilizers and sewage discharges. Stormwater runoff washes pol-
lutants from impervious surfaces, providing a significant non-point source of
pollution in urban areas (Characklis & Wiesner 1997; Bibby & Webster-Brown
2005). Nonetheless, there is no evidence to suggest that, in general, the level of
water pollutants increases in relation to the size of an individual city (Duh et al.
2008). There is potential within urban areas to improve water quality by
construction of, or improved management of existing, wetland ecosystems.

Water flows
Cities almost invariably draw the bulk of their water supplies from outside
their boundaries, as they typically have only a small area of standing water,
directing large quantities in and out through often heavily engineered infra-
structure. In most cases, these quantities are considerably greater than would
otherwise have passed through such environments, although occasionally
they may be reduced. Urbanisation thus changes the hydrology of an area
by altering the water supply, drainage and outflow. Indeed, in simple terms
of mass, water is by far the largest component of the metabolism of an urban
centre, vastly exceeding the amounts of food and fuel used by the human
population (Kennedy et al. 2007). It is, therefore, perhaps unsurprising that
although the natural density of water channels tends to be much reduced in
urban ecosystems, with many streams being filled in, paved over or cul-
verted, the overall density may actually be higher owing to the number of
artificial ones that are created. In addition, the development and structure
of such channels tends to differ between natural and urban settings (Paul &
Meyer 2001).
To date, much of the research into urban water flows has focused on the
relationship between impervious surface coverage and water regulation
(Arnold & Gibbons 1996). The construction of such sealed areas, which cover
40 K. J. GASTON, Z. G. DAVIES AND J. L. EDMONDSON

Figure 3.2 Relationships between impervious surface cover and surface runoff,
infiltration and evapotranspiration. Reproduced from Federal Interagency Stream
Restoration Working Group (1998).

9% of Europe alone (Scalenghe & Marsan 2009), results in marked increases in
surface runoff and decreases in infiltration (Figure 3.2). The outcome of this,
especially when associated with infrastructure to drain paved surfaces as
quickly and efficiently as possible (e.g. piped stormwater drainage systems), is
a substantially faster flow of water through systems and higher peak discharges
during storm events. In turn, this can lead to a greater frequency and severity
of flooding. At the same time, base flow rates may be influenced in a variety of
ways, sometimes becoming much diminished because of the lower infiltration
rates of water (reducing groundwater recharge). However, this may be offset by
irrigation, leakage from water supply or sewerage infrastructure, and waste-
water treatment plant outputs (Paul & Meyer 2001; Walsh et al. 2005).
Urbanisation can also influence precipitation both within and downwind
of cities, although usually to a smaller and less consistent extent than is the
case with temperature (Shepherd 2005; Collier 2006; Hand & Shepherd 2009).
 URBAN ENVIRONMENTS AND ECOSYSTEM FUNCTIONS 41

Deep, moist convection within urban environments is primarily enhanced by

the heat island effect (which destabilises the urban boundary layer; Figure 3.1),
the surface roughness of the area caused by buildings, and potentially by
vegetation. Conversely, precipitation may be suppressed by atmospheric pollu-
tion, particularly aerosols which change the microstructure of clouds by pro-
viding sources of condensation nuclei. The magnitude of some of these effects
may be related to the size of a city, and it has been suggested that the expan-
sion of urban areas may be having non-trivial influences on global patterns of
precipitation (Shepherd 2005).
The distribution of green space is crucial in explaining variation in water
regulation across the urban landscape. Increasing the extent of vegetated areas
enhances water regulation and improves water loss from the ground through
evapotranspiration. Generalised models of the impacts of various land covers
on runoff tend to rely on simple coefficients for rainwater infiltration of
different surfaces, usually obtained from the literature and not tailored to
particular urban areas (e.g. Pauleit & Duhme 2000; Whitford et al. 2001; Pauleit
et al. 2005; Tratalos et al. 2007). Nonetheless they frequently reveal non-linear
relationships with the extent of green space; initially, runoff rapidly increases
as green space declines, but at low levels of vegetative cover, changes in the
remaining green space have little effect (Tratalos et al. 2007). This is a conse-
quence of the relatively low infiltration rates of even many of the non-sealed
surfaces, due to factors such as extensive compaction of soil surfaces
(Scalenghe & Marsan 2009), when building densities become high.

Carbon cycles
Urbanisation influences local carbon cycles in two main ways. First, it changes
land cover into a mosaic of green spaces and impervious surfaces. Second, it
generates CO2 emissions as a consequence of the importation and subsequent
combustion of fossil fuels (especially oil, gas and coal) for residential and
industrial energy production, as well as for transportation.
The two major biological carbon pools in urban green spaces are vegetation
and soils. Most attention given to vegetation tends to fall on trees, which have
been shown to provide significant levels of carbon storage and sequestration in
urban environments, albeit substantially less than equivalent areas of forest
stands (Nowak & Crane 2002). Estimates of carbon sequestration by urban trees
tend to be generated by using simple functions of tree cover, which is logical
since this is both likely to have a dominant influence and is extremely variable
amongst different urban areas (Rowntree & Nowak 1991; Whitford et al. 2001;
Tratalos et al. 2007). However, in practice, the relationship will depend on a
variety of factors, including growth rate, size at maturity, demographic struc-
ture and species composition (Nowak et al. 2002). These in turn will be influ-
enced by land-use, planning policies, management and more immediate
42 K. J. GASTON, Z. G. DAVIES AND J. L. EDMONDSON

environmental conditions, such as soil quality, water potential and coverage by

impervious surfaces (Quigley 2004). In particular, on public lands, trees may be
removed or subject to surgery in response to concerns over issues of subsidence
or human safety. For instance, although over a 5-year period there was no
overall change in the number of trees in a study conducted in London, UK,
there was nonetheless a rapid turnover of individual trees and a disproportion-
ate loss of mature native trees (London Assembly Environment Committee
2007). Despite the potential to use these removed trees and waste wood prod-
ucts as an alternative energy source to fossil fuels, such bioenergy generation
still produces CO2 emissions (MacFarlane 2009). The degree to which urban
trees provide net carbon sequestration will also depend on the intensity of
management which they receive using carbon-emitting maintenance equip-
ment (e.g. chainsaws, chippers, motor vehicles), and the extent to which
decomposition is limited by long-term timber storage (e.g. wood products,
landfills; Nowak et al. 2002; MacFarlane 2009). Similar concerns have been
previously expressed about the environmental costs of management of other
urban land covers (e.g. grasslands; Falk 1976; Golubiewski 2006).
Soil carbon pools naturally comprise both labile organic carbon (with
relatively quick turnover rates) and black carbon, formed from incomplete
combustion of organic matter or fossil fuel (which is highly recalcitrant with
a long soil residence time). The black carbon constituent within urban soils is
often enhanced by vehicle and industry emissions (Lorenz et al. 2006; Rawlins
et al. 2008). Below-ground carbon storage in urban environments has vari-
ously been projected to be equal to, or considerably bigger than, that associ-
ated with aboveground vegetation, depending partially on the nature of this
vegetation (Figure 3.3; Jo & McPherson 1995; Pataki et al. 2006; Pouyat et al.
2006). On a per unit area basis, there is also evidence that the carbon pool in
urban soils is often high compared with other types of ecosystem, and may
be greater than that in equivalent rural soils because of the profound differ-
ences in management (Pouyat et al. 2002, 2006, 2009; Golubiewski 2006;
Pataki et al. 2006; Lorenz & Lal 2009). Indeed, it seems probable that urban
soils will often be markedly richer in organic carbon than those in rural
landscapes with a significant history of intensive agriculture (this is less
likely to be true in forest-dominated regions). Unfortunately, understanding
of urban soil carbon pools and fluxes is limited, primarily because the soils
are often spatially very heterogeneous, are poorly mapped and characterised,
in substantial part lie beneath impervious surfaces, and are often within
mosaics of different aged urban green spaces (Effland & Pouyat 1997; Lorenz &
Lal 2009). Nonetheless, it is clear that the patterns and rates of decomposition
and nutrient cycling may be profoundly altered in urban areas compared with
rural ones, owing to differences in leaf litter quality, because of altered biotic,
chemical and physical environments (often including high levels of heavy
 URBAN ENVIRONMENTS AND ECOSYSTEM FUNCTIONS 43

(a)
14
Belowground carbon density (kg m–2)

12

10

0
0 1 2 3 4 5 6
Aboveground carbon density (kg m–2)

(b)
160
Belowground carbon storage (million tonnes)

120

80

40

0
0 10 20 30 40
Aboveground carbon storage (million tonnes)

Figure 3.3 Relationships between estimated belowground and aboveground (a) urban
carbon density and (b) urban carbon storage estimated for different states in the
USA. From data in Pouyat et al. (2006).
metals), and owing to management practices and levels of disturbance
(McDonnell et al. 1997; Pouyat et al. 1997, 2009; Carreiro et al. 1999; Pickett
et al. 2001; Shen et al. 2008).
Given the relatively limited coverage by urban areas worldwide (see Chapter 2),
the immediate influence of urbanisation on vegetation and soils has arguably
also been reasonably small globally. Hence this is also true at a regional scale
(e.g. Erb 2004), although of course less so for those regions that have become
44 K. J. GASTON, Z. G. DAVIES AND J. L. EDMONDSON

more significantly developed. However, in many regions the expansion of urban

areas has disproportionately been on some of the best soils at the expense of
agricultural and forested land, and thus the ecosystem goods and services that
they provided (Nizeyimana et al. 2001; Alig et al. 2004; Zhao et al. 2006; Zhang et al.
2008). There are, therefore, significant concerns that this has become a seriously
problematic addition to the other pressures on such high-value land covers,
especially where agricultural land can be very productive. For example, in the
USA urbanisation is disproportionately taking place on the most fertile lands, with
a substantial overall negative effect on net primary production estimated to be
equivalent to the caloric requirement of 16.5 million people or c. 6% of the human
population (Imhoff et al. 2004).
Cities are major contributors to CO2 emissions, being responsible for
c. 75–90% of global anthropogenic CO2. Clearly, the levels of these emissions
are such that they cannot be offset by local natural carbon sequestration.
Nonetheless, the per unit area and the gross sizes of biological urban carbon
pools can be substantial (Nowak & Crane 2002; Pouyat et al. 2002; Kaye et al.
2005; Lorenz et al. 2006). Indeed, at least in some areas, carbon sequestration
within urban environments may not be an entirely trivial consideration (Jo &
McPherson 1995; Johnson & Gerhold 2003; Golubiewski 2006; Pataki et al.
2006), and there has even been discussion of the potential for these pools to
be cost effective in carbon credit markets (McHale et al. 2007).

Atmospheric chemistry
The atmospheres of urban areas tend to have heightened concentrations of
CO2, nitrous oxides (NOx), sulphur dioxide (SO2), ozone (O3), aerosols, metals
and suspended particulates, with profound implications for nutrient flows in
particular. Nitrous and sulphurous oxides are mainly generated through fossil
fuel combustion, and can be transformed in the atmosphere into acidic pre-
cipitation. Ozone is principally derived from photochemical reactions in the
atmosphere that involve nitrous oxides and hydrocarbons, and tends to occur
in particularly high concentrations downwind of urban centres. Suspended
particulates include larger particles (c. 2.5–100 microns diameter), usually
consisting of smoke and dust caused by industrial processes, agriculture,
construction and road traffic, and smaller particles (<2.5 microns) mainly from
the burning of fossil fuels (World Resources Institute, the United Nations
Environment Programme, the United Nations Development Programme and
The World Bank 1998).
The temporal dynamics of these pollutants are often complex, depending
on patterns in the history, growth and composition of an individual city, the
balance between increasing demand and output in power generation, indus-
try and transport, and the effectiveness of any active attempts to reduce the
 URBAN ENVIRONMENTS AND ECOSYSTEM FUNCTIONS 45

levels of pollutants (including through new technologies). Thus, for example,

in Shanghai, China, concentrations of SO2, total suspended particulates and
acid precipitation all fell from 1983 to 2004, mainly as a consequence of
reductions in the usage of coal, but levels of NOx have risen, primarily with
growing numbers of motor vehicles (Zhao et al. 2006). However, SO2 and acid
rain have tended to increase in the suburbs of Shanghai, in response to a
policy to move factories from urban centres to suburban areas, and NOx has
decreased in more urbanised areas as a consequence of policies towards
cleaner vehicle emissions.
Generally, atmospheric pollution in the urban areas of industrialised nations
has declined over the past few decades whereas, conversely, there have been
increases in pollutants within the developing world (Mayer 1999). However,
even where the levels of atmospheric pollutants have fallen in urban environ-
ments in recent years, they commonly still remain above desirable (albeit
variable) regional standards. The changes made to atmospheric chemistry by
urban areas can have influences on multiple scales. Positively, they can affect
nutrient cycling and plant production, particularly through fertilisation
effects, both locally and regionally (Trusilova & Churkina 2008). More nega-
tively, air pollution from urban centres is a major contributor to global climate
change and acid deposition, and many studies have shown that such pollution
and its biogeochemical consequences influence both terrestrial and aquatic
organisms differentially, leading to shifts in assemblage structure (Lovett et al.
2009). Minimisation of atmospheric pollutants within urban areas is also of
utmost importance for human health (Brunekreef & Holgate 2002). For
example, in Beijing, China, increases in atmospheric air pollutants were asso-
ciated with a rise in emergency room admissions of patients with cardiovascu-
lar disease (Guo et al. 2009). Indeed, the World Health Organization has
identified particulate pollution as one of the most important contributors to
poor human health, with high concentrations being responsible for triggering
a wide range of respiratory diseases, exacerbating heart disease and aggravat-
ing a variety of other conditions. In many cities across the world, the concen-
tration of suspended particulate matter greatly exceeds the recommended air
quality standard of less than 90 micrograms m 3 (Figure 3.4).
Urban vegetation may help to improve air quality through the uptake of
gaseous, aerosol and particulate pollutants (Freer-Smith et al. 1997), depending
on the type and the location of the planting. In urban ecosystems, the presence
of trees increases the surface area and roughness onto which pollutants can be
deposited (Beckett et al. 1998), meaning that heavily forested cities reap consid-
erable benefits (Bolund & Hunhammar 1999). Likewise, in Guangzhuo, China,
areas of the city with the highest tree cover were responsible for the largest
proportion of pollutant removal (Jim & Chen 2008).
46 K. J. GASTON, Z. G. DAVIES AND J. L. EDMONDSON

Helsinki
Moscow

Paris
Toronto Kiev Beijing Seoul
Rome Tokyo
Montreal Lanzhou
New York
Delhi Osaka
Taiwan
Mexico City Mumbai
Manila

Jakarta

3 São Paolo
SPM (micrograms per m ) Rio de Janeiro
<99 Sydney

100–199
200–299
>300

Figure 3.4 Levels of particulate air pollution in 1995 within the world’s largest cities.
SPM, Suspended Particulate Matter. From data in Soubbotina (2004).

Species interactions
The effective functioning of any ecosystem depends on the interactions that
occur between species. These take many forms, including competition, preda-
tion, parasitism and mutualism. It is clear that urbanisation can have profound
effects on such relationships, although there is a lack of information regarding
how alterations in species abundances across urban landscapes will affect the
interactions between them (Faeth et al. 2005). For example, the urbanisation of
Phoenix, Arizona, has led to a dramatic increase in available water, a limiting
resource in the surrounding landscape, which has resulted in a shift in trophic
dynamics including a greater top-down influence of predators on trophic
interactions (Faeth et al. 2005).
In much of the world, an important element in maintaining an urban green
infrastructure (e.g. parks, gardens) is an adequate level of insect pollination,
without which many plant species cannot produce fertile seeds and the yields of
others are substantially reduced. Unfortunately, several studies have documented
declines in the species richness and abundance of key groups of insect pollinators
in response to urbanisation (e.g. McIntyre & Hostetler 2001; Zanette et al. 2005;
Matteson et al. 2008). However, others have actually found elevated richness and
abundance in urban areas, which has been attributed to heat island effects,
reduced exposure to agricultural chemicals and the wide variety of microhabitats
present in urban landscapes (Eremeeva & Sushchev 2005; Fetridge et al. 2008).

In conclusion
Key to an understanding of urban ecology is the recognition that ecosystem
function in urban areas has been profoundly shaped by human activities, such
that there are often only weak parallels with natural ecosystems. This has two
 URBAN ENVIRONMENTS AND ECOSYSTEM FUNCTIONS 47

major consequences. First, it requires the continued development of a specific-

ally urban approach to studying ecosystem function, recognising that the
opportunities to generalise from what is known about other ecosystems may
be very limited. Second, it provides an opportunity, as yet largely unexploited,
to test broad ideas about ecosystem function in an arena in which the forces
are very different.

Acknowledgements
K. L. Evans, S. Gaston and P. Johnson are kindly thanked variously for their
advice, assistance and comments. Z.G.D. and J.L.E. were supported by EPSRC
grant EP/F007604/1 to the 4M consortium: Measurement, Modelling, Mapping
and Management: an Evidence Based Methodology for Understanding and
Shrinking the Urban Carbon Footprint. The consortium has five UK partners:
Loughborough University, De Montfort University, Newcastle University, the
University of Sheffield and the University of Leeds.

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CHAPTER FOUR

Individual species and urbanisation

KARL L. EVANS

Intraspecific variation in ecological and life history traits has interested ecolo-
gists since at least the nineteenth century (Bergmann 1847; Allen 1878; Gloger
1883; Jordan 1891). Such variation is still the focus of intensive research, but
somewhat surprisingly the environmental factors driving large-scale patterns
have rarely been firmly established and are intensely debated (Gaston et al.
2008). Turning to finer spatial scales, local variation in environmental condi-
tions can promote marked trait divergence between adjacent populations. This
raises important evolutionary questions regarding the role of phenotypic plas-
ticity in generating such divergence, and the interactions between genetic
adaptation and gene flow (Postma & van Noordwijk 2005; Senar et al. 2006;
Ghalambor et al. 2007; McCormack & Smith 2008). The patterns and processes
associated with trait divergence have, with the exception of a few classic
studies concerning the effects of pollution, traditionally been assessed in
relatively natural rural systems. Recently, however, there has been an explo-
sion of research focusing on how the marked ecological differences between
rural and urban areas influence the traits of conspecific populations. So far this
work has largely emerged as a series of isolated case studies of trait differences,
with the few attempts to synthesise the available information being confined
to a subset of taxa or traits (Marzluff 2001; Bradley & Altizer 2007; Chamberlain
et al. 2009). Assessments of intraspecific variation among rural and urban
populations have also rarely been placed in the broader context of trait vari-
ation in more natural systems.
Against the background of the profound influence of urbanisation on eco-
system form and function (Chapters 2 and 3), this chapter provides a detailed
synthetic overview of trait differentiation between conspecific rural and urban
populations, and its consequences. Studies assessing which traits are associated
with interspecific variation in species responses to urbanisation (e.g. Croci et al.
2008; Thompson & McCarthy 2008; Evans et al. in press, a) are not covered here

Urban Ecology, ed. Kevin J. Gaston. Published by Cambridge University Press.

# British Ecological Society 2010.
54 K. L. EVANS

(a)
60

50
Number of studies

40

30

20

10

0
–40 –20 0 20 40 60
Latitude

(b)
40
Number of studies

30

20

10

0
s

ts

ns

es

sh
ird

al

te
an

til
ia

Fi
m

ra
B

ib

ep
Pl

am

eb
ph

R
rt
M

ve
A

In

Figure 4.1 Assessment of the ecological and life history traits in urban populations is
biased towards (a) northern temperate regions and (b) certain taxonomic groups,
particularly birds. Data concern those studies cited in this chapter; multiple papers
reporting divergence in more than one trait for the same population are considered as a
single study. Positive and negative latitudes indicate northern and southern
hemispheres, respectively.

but are discussed in Chapter 5. I consider a number of broad groupings of

ecological traits and, taking each trait in turn, summarise intraspecific pat-
terns of divergence between rural and urban populations. Where possible such
variation is assessed in relation to the mechanisms proposed to drive spatial
patterns in intraspecific trait variation in more natural areas. The aim is to
illustrate the predominant patterns that have been documented to date and
their causal mechanisms. Examples are drawn from as large a taxonomic and
 INDIVIDUAL SPECIES AND URBANISATION 55

geographical scope as possible, although the dominance of research conducted

in northern temperate regions and on birds creates inevitable biases (Figure 4.1).
Finally, I assess the implications of trait divergence between urban and rural
populations, and highlight a number of issues that have received insufficient
attention.

Species traits
Abundance
Population density is an important trait in its own right, and its response to
urbanisation may influence the nature of other patterns of trait divergence
between urban and rural populations if these traits are density dependent
effects. In addition, the effect of urbanisation on demographic and other traits
must be viewed in the context of the suitability of urban environments for the
focal species, which can be assessed by the density of urban populations,
especially in comparison to rural densities (Evans et al. in press, a). Urbanisation
typically influences population density in one of three ways. Abundance may
be lowest in urban areas, peak in areas of intermediate development or be
greatest in highly urbanised sites (Blair 1996; Tratalos et al. 2007; Grimm et al.
2008; Figure 4.2). Species exhibiting these patterns have been termed respect-
ively urban avoiders, exploiters and adapters (Blair 1996).
The relative proportion of species exhibiting each of the three main ways
in which population density responds to urbanisation varies taxonomically.
Amongst plants, urban adapters appear particularly frequent, but this pattern
is largely driven by the positive responses to urbanisation of many introduced
species (Grimm et al. 2008). Amongst birds, it appears that a larger proportion
of species, compared with most other taxonomic groups, have peak densities
at intermediate levels of urbanisation; this is perhaps in response to
increased habitat diversity in such areas (Marzluff 2005; Tratalos et al. 2007).
Insufficient studies have been conducted in other taxonomic groups to formu-
late generalisations, but as relatively few species occur in highly urbanised
areas (Chapter 5) it seems highly likely that in most taxonomic groups the
majority of species will occur at higher densities in rural environments than
highly urbanised ones.
That urbanisation influences species densities is unsurprising. What is
more interesting is that the nature of a species’ response to urbanisation
can vary spatially, the most dramatic examples of which concern those
species which are absent from urban areas in parts of their range despite
occurring at high densities in urban areas elsewhere. One such example is
the dunnock Prunella modularis, which has high population densities in
British cities but is largely confined to rural areas elsewhere in its European
range (Vogel & Tuomenpuro 1997). Similarly, wild boar Sus scrofa are urban-
ised in parts of Europe, such as Germany, but are absent from some urban
56 K. L. EVANS

(a)

Carrion crow density (km–2)

30

20

10

0
0

00
00

00

00

0
5.

.0

.0

.0

.0

.0

25
5.

0.

0.
0–

40

50

00

50

00
–1

10

50

>
1–

–2

–5

–7

25
01

1–

–1
.0

01

01

01

1–
5.

.0

01
15

0.

0.

0.

.0
40

0.
10

25

50

00
75

15
Housing density (km–2)

(b) 60
Robin density (km–2)

40

20

0
0

00
00

00

00

0
5.

.0

.0

.0

.0

.0

25
5.

0.

0.
0–

40

50

00

50

00
–1

10

50

>
1–

–2

–5

–7

25
01

1–

–1
.0

01

01

01

1–
5.

.0

01
15

0.

0.

0.

.0
40

0.
10

25

50

00
75

15

Housing density (km–2)

(c)
Willow warbler density (km–2)

18
15
12
9
6
3
0
0

00
00

00

00

0
5.

.0

.0

.0

.0

.0

25
5.

0.

0.
0–

40

50

00

50

00
–1

10

50

>
1–

–2

–5

–7

25
01

1–

–1
.0

01

01

01

1–
5.

.0

01
15

0.

0.

0.

.0
40

0.
10

25

50

00
75

15

Housing density (km–2)

Figure 4.2 Urbanisation influences abundance in one of three ways, (a) density may
peak in highly urbanised areas, (b) density may peak in moderately urbanised areas, and
(c) density may peak in rural areas. These patterns are illustrated respectively by the
changes in carrion crow Corvus corone, European robin Erithacus rubecula and willow
warbler Phylloscopus trochilus densities in 1-kilometre squares in the UK across a gradient in
housing density. Error bars represent standard errors. Modified from Tratalos et al. (2007).
 INDIVIDUAL SPECIES AND URBANISATION 57

areas in the Baltic states despite occurring in the nearby countryside

(Zorenko & Leontyeva 2003). Even species that are frequently highly urban-
ised may be absent from towns and cities in part of their range. The feral
pigeon Columba livia is amongst one of the most urbanised birds in the
world, but in central Asia the species is lacking from some towns despite
the occurrence of nearby rural populations (Obukhova 2001).
The mechanisms driving spatial variation in intraspecific responses to urban
environments have seldom been identified, but four main mechanisms are
likely to generate such patterns (Evans et al. in press, b). Differences in the
history of urban development will affect the opportunity for urban populations
to arise by influencing both the time available for colonisation and the number
of sites that can be colonised. The occurrence of urban populations will also be
linked to biotic factors, such as the availability of potential colonists, which
may vary spatially according to rural population size and the pressure on such
individuals to disperse. The quality of urban habitats will also change with
location owing to differences in factors such as human attitudes to urban
wildlife, soil type, pollution levels, vegetation structure and composition, and
the occurrence of competitors, pathogens, predators and pollinators. Finally,
rural populations may differ in their ability to acquire specific traits that
increase fitness in urban environments owing to variation in either their
genetic composition or their ability to exhibit sufficient phenotypic plasticity.
There is certainly increasing evidence that the contrast in the selection pres-
sures of rural and urban environments is large enough to generate marked
intraspecific trait variation between urban and rural populations. The rest of
this chapter assesses the nature and consequences of such divergence.

Body size
It is relatively straightforward, in comparison with many other types of traits,
to assess the effect of urbanisation on body size, and a number of studies have
thus done so. Reduced body size has been reported in urban populations from a
wide range of taxonomic groups including invertebrates (Weller & Ganzhorn
2004), snakes (Luiselli et al. 2002) and birds (Richner 1989; Ruiz et al. 2002;
Rasner et al. 2004; Liker et al. 2008; Rodewald & Shustack 2008). In contrast, a
smaller number of studies have reported larger body size in urban populations
(amphibians: Ovaska 1991; snakes: Savidge 1991; mammals: Liro 1985). More-
over, the impacts of urbanisation on body size may be population-specific
(Evans et al. 2009a) or gender-specific (Auman et al. 2008), and urbanisation
certainly does not have a universal impact on body size.
Most studies of body size divergence in rural and urban populations have
focused on describing patterns, rather than assessing the driving mechanisms.
A large literature has, however, arisen concerning factors that may drive
spatial patterns in body size across larger spatial scales. Three broad groups
58 K. L. EVANS

of mechanisms have been proposed, and it is useful to assess their potential

role in driving the impacts of urbanisation on body size. First, thermodynamic
hypotheses that relate body size to heat conservation and dissipation predict
reduced body size in warmer environments (Bergmann 1847; Brown & Lee
1969), and thus that urbanisation will generate smaller bodied individuals
because of the urban heat island effect. Second, amongst ectotherms, the
effects of temperature on cell division and replication, or on growth and
differentiation, may generate larger cells and therefore larger individuals in
cold environments (Partridge et al. 1994; Walters & Hassall 2006). This mechan-
ism thus also predicts reduced body size in urban populations. Finally, a suite
of alternative hypotheses states that the optimum body size is determined by
trade-offs between rates of reproduction and mortality that are mediated by
factors related to food availability, such as primary productivity, starvation
resistance and seasonality (Roff 1980; Lindstedt & Boyce 1985; Zeveloff & Boyce
1988; Kozłowski et al. 2004). These trade-off mechanisms thus suggest that rural
and urban populations will diverge in body size in a manner dependent on
numerous factors, but with relative resource availability in rural and urban
areas playing a major role.
The thermodynamic and physiological hypotheses may drive changes in
body size in some urban populations, but they cannot provide a general
mechanism as they fail to explain the increased body size of some urban
populations. The trade-off mechanisms predict that temporal increases in food
supply in urban areas as a consequence of increased human activity is likely to
generate increased body size, at least in commensal species, and such patterns
have been documented (Yom-Tov 2003). The trade-off mechanisms also predict
that body size will exhibit contrasting responses to urbanisation in closely
related species that experience different levels of food availability in urban
environments. Such a pattern is exhibited by two sympatric cobra species.
Urbanisation reduces food availability for Naja melanoleuca, and urban popula-
tions of this species are smaller bodied than rural ones, whilst N. nigricollis
experiences equally high food availability in urban and rural areas, and its
body size does not vary with habitat type (Luiselli et al. 2002). Whilst the effects
of urbanisation on body size will often arise through changes in food availabil-
ity, this is probably not a universal mechanism. Indeed, the smaller body size of
urban house sparrows Passer domesticus, relative to rural ones, is maintained
when individuals are raised under identical conditions with an unconstrained
food supply (Liker et al. 2008).
Additional mechanisms, other than those proposed to explain large-scale
patterns in body size, probably also contribute to the impacts of urbanisation
on body size. First, increased predation pressure may favour lighter individ-
uals, at least in birds, as this facilitates predator avoidance (Witter & Cuthill
1993). Urban populations frequently coexist with higher densities of some
 INDIVIDUAL SPECIES AND URBANISATION 59

predators (Sims et al. 2008); predator density may not always determine preda-
tion risk, but it strongly influences the perceived risk of predation, which is
often the most important factor in determining species’ responses to predators
(Cresswell 2008). Second, body size often has high heritability (Glazier 2002),
and therefore in isolated urban populations changes in body size, or any other
genetically inherited trait, could simply be a stochastic consequence of the
characteristics of founding individuals. Finally, body size is also often a correl-
ate of individual quality, so intraspecific competition may result in smaller
individuals occupying less preferred habitats, which may include urban ones.
The Acadian flycatcher Empidonax virescens is a possible example of this mech-
anism; urban populations occur at lower densities and have less breeding
success than rural ones, suggesting that urban habitats are of poorer quality,
and urban birds are also smaller than rural ones (Rodewald & Shustack 2008).
Urban and rural populations frequently differ in their body size, but insuffi-
cient studies have been conducted for general trends to emerge. Additional
studies are clearly required, and these should focus on testing driving mechan-
isms, rather than just describing patterns, and the adaptive value of any
documented changes. Ideally, such studies should assess patterns of body size
divergence in multiple paired rural and urban populations, and do so in
species for which urban environments provide both high and low quality
habitats.

Communication
A number of theoretical and empirical studies have documented how popula-
tions in rural areas have adapted acoustic signals to the divergent sound
transmission properties of contrasting habitat types in order to reduce attenu-
ation (loss of volume), degradation (change in acoustic structure) and masking
by background noise (Morton 1975; Nottebohm 1975; Hunter & Krebs 1979;
Ryan et al. 1990). Anthropogenic noise has been demonstrated to reduce the
effectiveness of acoustic communication in many taxa, including fish
(McCauley et al. 2003; Popper et al. 2003), frogs (Bee & Swanson 2007), birds
(Leader et al. 2005; Habib et al. 2007) and mammals (Nowacek et al. 2007;
Weilgart 2007). As urban environments are typically noisier than rural ones
(Butler 2004), individuals occupying such habitats are likely to experience
selection pressures that will promote divergence of their vocalisations from
those of rural conspecifics.
The Lombard effect predicts that in urban areas the masking effects of
background noise can be reduced by increasing the amplitude (i.e. volume) of
vocalisations (Lombard 1911). It is thus predicted that urban birds will sing
louder on weekdays, when traffic noise is greater, than at weekends; this
pattern has been documented in urban nightingales Luscinia megarhynchos
(Brumm & Todt 2002; Brumm 2004), but not urban great tits Parus major
60 K. L. EVANS

(B. Hawkins & A. G. Gosler, unpublished data). This contrast may arise because
the focal populations may differ in their ability to pay the energetic cost of
louder vocalisations.
Californian ground squirrels Spermophilus beecheyi in areas subject to noise
from wind-turbines had higher-pitched vocalisations than those in quieter areas
(Rabin et al. 2003); such changes are typically considered to be adaptive because
of the low pitch of most anthropogenic noise which may otherwise mask signals.
Increased traffic noise is also associated with increased pitch of the mating calls
of some frogs (e.g. Litoria ewingii, Parris et al. 2009), but other anurans do not
exhibit such responses (e.g. Hyla arborea, Lengagne 2008). The increased pitch of
vocalisations of urban populations has been documented most frequently in
birds (great tit: Slabbekoorn & den Boer-Visser 2006; Slabbekoorn & Ripmeester
2008; Mockford & Marshall 2009; house finch Carpodacus mexicanus: Fernández-
Juricic et al. 2005; Bermúdez-Cuamatzin et al. 2009; song sparrow Melospiza
melodia: Wood & Yezerinac 2006; blackbird Turdus merula: Nemeth & Brumm
2009). Male great tits appear to exhibit reduced territorial responses when
hearing song from a territory where background noise differed from their
own, which at the very least suggests that these changes in vocalisations influ-
ence their effectiveness as a signal (Mockford & Marshall 2009).
Despite the frequent interpretation of the increased pitch of urban popula-
tions’ vocalisations as being adaptive, there are alternative perspectives.
Changes in pitch in response to urbanisation may simply be a consequence of
release from the constraints of sound transmission in densely vegetated habi-
tats, which readily absorb high-pitched sounds (Morton 1975), or may be an
unintended consequence of other differences between urban and rural popula-
tions (Nemeth & Brumm 2009). The higher population densities of urban
populations (see above) may result in more intraspecific competition and thus
greater arousal status, which can result in higher-frequency avian vocalisations
(Dabelsteen & Pedersen 1985). Moreover, some urban bird populations are
characterised by lower testosterone levels than their rural counterparts
(e.g. blackbird; Partecke 2005), and these hormonal changes can increase the
pitch of vocalisations (Cynx et al. 2005). Similarly, the pitch of anuran vocalisa-
tions increases with temperature (Gerhardt & Mudry 1980), and thus changes
in vocalisations of urban populations may be a consequence of the urban heat
island effect. Switches to higher-frequency vocalisations in urban environ-
ments may also be maladaptive because such signals degrade more rapidly
(Leader et al. 2005). Experimental studies are thus required to confirm that
changes in the pitch of urban populations’ vocalisations are adaptive.
Increasing the effectiveness of acoustic communication in noisy environ-
ments may also be achieved by switching the timing of vocalisation to quieter
periods. A number of frog species reduce calling rates following exposure to
anthropogenic noise (Sun & Narins 2005; Lengagne 2008), and many bird
 INDIVIDUAL SPECIES AND URBANISATION 61

species commence singing earlier in the day in urban areas than in forest
(Bergen & Abs 1997; Nemeth & Brumm 2009); the extent to which such changes
are adaptive mechanisms for avoiding peaks in ambient noise is, however,
debatable. More convincingly, European robins Erithacus rubecula breeding in
noisier urban sites increase the amount of singing during quiet periods (i.e. at
night), and this effect is not attributable to increased ambient light pollution
in such territories (Fuller et al. 2007). Moreover, urban great tits and house
finches tend to produce shorter songs than conspecifics in more rural areas
(Fernández-Juricic et al. 2005; Slabbekoorn & den Boer-Visser 2006). This pattern
may arise because shorter songs are more likely to transmit fully during brief
and unpredictable lulls in anthropogenic noise, but the driving mechanisms
are uncertain and not all urban bird populations have reduced their song
length (e.g. blackbirds: Nemeth & Brumm 2009).
It would be useful to extend studies of urbanisation impacts on avian song
to other vocalisation types, such as begging calls, the structure of which can
be markedly influenced by the acoustic environment (Leonard & Horn 2008).
The influence of urbanisation on non-acoustic communication systems has
also received very little attention. As an example, the amount of white in the
tail of dark-eyed juncos Junco hyemalis is a very important signalling trait in
rural environments, but although urban and rural populations differ in the
amount of white in the tail, the causes and consequences of this variation are
unknown (Yeh 2004; Price et al. 2008). More generally, high rates of back-
ground movement can alter the efficiency of visual displays (Peters 2008).
Therefore, in areas where wind frequently moves vegetation, lizards with
rapid displays signal territory boundaries more effectively than conspecifics
with slower displays, leading to habitat-specific divergence of display charac-
teristics (Ord et al. 2007). Relatively rapidly moving objects, such as traffic and
people, may be more frequent in urban areas than rural ones. This could
generate equivalent divergent selective pressures on the rapidity of visual
displays in the two habitats, but this has not yet been assessed. Similarly,
pollution of aquatic systems through eutrophication and increased turbidity
reduces the effectiveness of visual signals, thus reducing the intensity of
sexual selection (Seehausen et al. 1997; Järvenpää & Lindström 2004; Wong
et al. 2007) and promoting the use of non-visual signals (Heuschele & Candolin
2007). Water pollution in urban areas may generate equivalent effects, but
again this remains untested.
Research conducted to date has tended to assume that changes in the
vocalisations of urban individuals are adaptive, but this may not always be
the case. Future research should thus determine the consequences of alter-
ations in species’ vocalisations associated with urbanisation, and this should
include further assessment of how rural individuals respond to the altered
vocalisations of urban conspecifics. As case studies accumulate, it is also
62 K. L. EVANS

becoming apparent that all species do not alter their vocalisations in the same
manner when occupying urban localities, and the species’ traits and environ-
mental factors driving this variation need to be determined. Much more work
on species other than birds is also needed. Despite these requirements for
further research, it is apparent that the communication signals of many
urbanised populations have changed in the direction predicted by theory
(Patricelli & Blickley 2006; Warren et al. 2006), providing a clear example of
how urban ecology interlinks with that in more natural areas.

Physiology
Physiological traits can determine spatial patterns in biodiversity and its
response to environmental degradation (Chown & Gaston 2008). Adjustment
of physiological traits may thus make it easier to exploit urban environments,
and whilst such divergence between rural and urban populations has seldom
been explored, a number of very different suites of physiological traits have
received initial investigation.
The markedly altered climatic regimes of urban areas may generate diver-
gence in the physiological tolerances of rural and urban conspecific popula-
tions. As an example, urban populations of Aedes mosquitoes have greater
tolerance to desiccation than rural conspecifics, presumably because the
former tend to breed in more temporary waterbodies (Mogi et al. 1996). Simi-
larly, urban leafcutter ants Atta sexdens have much greater tolerance to high
temperatures than their rural conspecifics (Angilletta et al. 2007). Despite the
general nature of the urban heat island, it is unclear whether most urban
populations will exhibit increased thermal tolerances, and such divergence is
perhaps particularly unlikely to occur in cooler temperate regions.
Stress physiology can differ in rural and urban populations, at least in
birds. Urban blackbirds have reduced concentrations of the stress hormone
corticosterone than rural individuals; this appears to be genetically con-
trolled, and may be an adaptive response that enables urban individuals to
tolerate closer approach by humans (Partecke et al. 2006). Reduced fear of
humans is certainly a common trait of urban populations (reptiles: Burger
2001; birds: Cooke 1980; Mller 2008; mammals: Shargo 1988; Gliwicz et al.
1994; Smith & Engeman 2002), suggesting that reduced levels of stress hor-
mones in urban populations may be a general pattern (Partecke et al. 2006).
Other factors will, however, also influence corticosterone concentrations in
urban populations; they may be lowered by increased food availability in
urban areas arising from anthropogenic supplementary feeding (Schoech
et al. 2007), or increased in urban adapters because the higher population
densities elevate sexual competition (Bonier et al. 2007). Furthermore, the
rural and urban populations of some bird species have very similar baseline
corticosterone levels (Fokidis et al. 2009). General trends in the stress
 INDIVIDUAL SPECIES AND URBANISATION 63

physiology of urban populations and their driving factors are thus highly
uncertain, although they illustrate the potential for rural and urban popula-
tions to exhibit divergent physiologies.
The frequently divergent diets of rural and urban populations (e.g. Harris
1984; Kiat et al. 2008) may have physiological consequences. In particular,
urban populations tend to ingest more food from anthropogenic sources
than do rural ones. Such shifts have been documented in a wide range of
birds (e.g. rufous-collared sparrow Zonotrichia capensis: Ruiz et al. 2002;
Eurasian starling Sturnus vulgaris: Mennechez & Clergeau 2006; Eurasian
kestrel Falco tinnunculus: Kubler et al. 2005) and mammals (e.g. long-nosed
bandicoot Perameles nasuta: Scott et al. 1999; raccoon Procyon lotor: Zeveloff
2002; red fox Vulpes vulpes: Contesse et al. 2004; coyote Canis latrans: Morey
et al. 2007). The biochemical implications of these changes in diet have
rarely been explored, but increased blood cholesterol and glucose levels
have been reported, and are likely to have adverse physiological effects that
compromise fitness (Ruiz et al. 2002; Ishigame et al. 2006).
Urbanisation, mainly via its high levels of anthropogenic pollution, increases
oxidative stress in both avian and mammalian urban populations, and thus the
demand for antioxidants (C. Isaksson, unpublished meta-analysis). Despite
these challenges to the immune system, it appears that levels of oxidative
damage in urban populations are similar to those in conspecific rural popula-
tions (Isaksson et al. 2009; Isaksson, unpublished meta-analysis). This is perhaps
because urban individuals tend to invest more in immune defence than rural
ones. Such patterns have certainly been documented in urban birds (Isaksson
et al. 2007), and there is some evidence for this in urban Rana frog spp.
(Romanova & Egorikhina 2006). In birds, this increased investment is mainly
due to internal up-regulation, since many urban populations experience
reduced food quality such as lower availability of dietary carotenoids (Hõrak
et al. 2001, but see Isaksson & Andersson 2007), which are extremely important
immuno-stimulants and may play additional roles in avian immune systems
(Costantini & Mller 2008, 2009). The population-level consequences of these
patterns are not well understood. Indeed, little research has addressed the
impacts of urbanisation on the immune physiology of (non-human) animals.
Many more studies are required that investigate how the environmental char-
acteristics of urban environments, such as alterations in nutrition and disease
risk (see below), interact with each other to determine how investment in the
immune system diverges between urban and rural populations.
Finally, and turning to plants, traits related to photosynthesis can be
markedly altered by urbanisation. In urban areas, the greater concentrations
of photochemical pollutants, such as ozone, nitrous oxides and sulphur
dioxide, and of heavy metals frequently reduce photosynthetic rates
(NEGTAP 2001; Baycu et al. 2006). In less polluted urban sites experiments
64 K. L. EVANS

suggest that photosynthetic rates may be greater than in rural areas,

perhaps partly in response to increased carbon dioxide concentrations
(Gregg et al. 2003). Moreover, urban plant populations can exhibit adapta-
tions to pollution that minimise its impacts on photosynthesis; examples of
such trait differentiation include genetic adaptations to heavy metals
(Wilson & Bell 1985; Shaw 1999), increased chlorophyll concentrations (Gra-
tani et al. 2000; but see Joshi & Swami 2009) and higher stomatal densities
(Alves et al. 2008). The increase in stomatal densities has occurred despite
higher carbon dioxide concentrations in urban areas which, over longer
timescales, are associated with a reduction in stomatal densities (Kouwen-
berg et al. 2003). However, the extent to which the impact of urbanisation on
stomatal densities depends upon water availability rather than carbon diox-
ide concentrations remains untested. Indeed, much further work is required
to ascertain how the numerous changes in photosynthetic traits induced by
urbanisation interact to determine relative photosynthetic rates in urban
and rural plant populations.
The above studies clearly demonstrate that conspecific rural and urban
populations can differ in a diverse range of physiological traits. The variation
documented so far is, however, likely to be a small proportion of the physio-
logical traits that are affected by urbanisation, and insufficient studies have
been conducted for general trends to emerge.

Disease risk
There is increasing evidence that urbanisation can dramatically alter para-
site loads and disease risk (Bradley & Altizer 2007). One example concerns
urban blackbirds, which have lower abundances of Ixodes ticks, ectoparasites
that impose direct fitness costs and transmit pathogens, and also often have
reduced prevalence of avian malaria than rural birds (Figure 4.3; Grégoire
et al. 2002; Evans et al. 2009b). Such reductions in disease risk seem likely
to arise largely because the abundance of parasites and pathogens is
adversely affected by urban environments; this may happen through three
mechanisms. First, pollution may kill pathogens and parasites. This has
frequently been documented in freshwater systems (Marcogliese & Cone
1997; King et al. 2007), and plant pathogens also often respond negatively
to air pollution (Saunders 1966; Jarrauld 2000). Second, alterations in urban
habitats or climatic conditions may directly reduce parasite abundance by
increasing mortality (Lafferty 1997). Finally, parasite and pathogen abun-
dance may be lowered indirectly through disruption of their life cycle,
because urban environments contain fewer intermediate hosts or vectors
(Reperant et al. 2007; Page et al. 2008).
Conversely, a number of studies have reported increased disease risk in
urban populations (Bradley & Altizer 2007). Four mechanisms seem likely to
 INDIVIDUAL SPECIES AND URBANISATION 65

(a)
18
16
Mean tick abundance

14
12
10
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(b)
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Malaria prevalence

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Figure 4.3 Urbanisation can markedly influence exposure to parasites and pathogens,
but the nature of this influence may vary across cities and with the identity of the
parasite/pathogen. For example, (a) urbanisation has consistent impacts on exposure to
Ixodes ticks (no ticks were detected in urban or rural blackbird populations in Sheffield,
Tunis or Valencia), but (b) the impact of urbanisation on avian malaria prevalence in
paired European urban and rural blackbird Turdus merula populations is very variable.
In both panels, black represents urban. Error bars represent standard errors. Modified
from Evans et al. (2009b).

generate such patterns. First, and counter to the effects described above,
susceptibility to pathogens and their vectors can be increased by the
immunosuppressive effects of pollutants (Dohmen et al. 1984; deSwart et al.
1996; Galloway & Depledge 2001; Kiesecker 2002), which tend to be com-
moner in urban areas. This may be one factor driving observations of
increased virulence and density of ranaviruses and trematode parasites in
urban amphibian populations ( Johnson et al. 1999; Carey et al. 2003). Second,
pathogen encounter rates may increase in urban areas owing to increased
introductions of exotic species through the horticultural trade and other
66 K. L. EVANS

human activities (Brasier 2008; Hamer & McDonnell 2008). Third, urbanisa-
tion may alter assemblage diversity and composition in a manner which
increases the transmission of some vector-borne diseases owing to reduced
abundance of less competent reservoir hosts, which would otherwise dilute
pathogen transmission between vectors and more competent hosts. This is
the converse of the dilution effect (Ostfeld & Keesing 2000), and may contri-
bute to the greater prevalence of West Nile virus in North American urban
bird populations compared with rural ones (Bradley et al. 2008). Finally, the
higher densities of many urban populations, relative to rural ones, seem
likely to increase disease transmission rates.
Urbanisation can clearly alter biotic interactions through a number of path-
ways that can generate both increases and decreases in disease prevalence in
urban populations. No trends have yet emerged regarding whether urban
populations typically experience reduced or increased disease risk (Bradley &
Altizer 2007). The occurrence of spatial variation in such patterns (Evans et al.
2009b) and variation with the taxonomy of both hosts (Fokidis et al. 2008) and
parasites (Fenoglio et al. 2009) does, however, generate opportunities to assess
the relative importance of the alternative causal mechanisms.

Demographic traits
Timing of reproduction
The timing of reproduction can vary markedly between urban and rural
populations. Many aspects of vegetation phenology, such as flower emergence
and bud burst, are frequently advanced in urban environments as a conse-
quence of the urban heat island effect (Roetzer et al. 2000; Zhang et al. 2004; Lu
et al. 2006; Neil & Wu 2006). However, this pattern is not universal; it is
particularly unusual in the tropics, where vegetation phenology is often
dependent upon humidity, which is less uniformly influenced by urbanisation
than is temperature (Gazal et al. 2008).
Turning to animals, the relative timing of reproduction in urban popula-
tions has most frequently been studied in birds, which typically breed
earlier in towns and cities (Chamberlain et al. 2009). The mechanisms driving
this pattern warrant further investigation, but improved body condition of
urban individuals due to anthropogenic provision of additional food is
considered to be a major factor (Schoech & Bowman 2003; Chamberlain
et al. 2009). Although empirical data are lacking, it also seems likely
that the urban heat island effect will result in earlier availability of inverte-
brate food sources in towns and cities, at least in temperate regions where
humidity is typically unimportant, further contributing to earlier reproduc-
tion. Advancement of avian reproduction in urban environments is
not, however, a universal pattern (Rodewald & Shustack 2008). Indeed, for
 INDIVIDUAL SPECIES AND URBANISATION 67

many species urban environments are less suitable than rural ones, which
may delay the acquisition of sufficient resources for breeding. Furthermore,
in those species for which urban environments provide poorer-quality
habitat than rural ones, intraspecific competition may result in urban
populations comprising a greater proportion of low-quality individuals
which may thus breed later in the year.

Reproductive success
Urbanisation can also exert strong influences on reproductive output.
Pollination, and thus seed production, may be disrupted in towns and cities
because of reduced abundances of animal pollinators (Cheptou & Avendaño
2006; R. A. Fuller et al., unpublished data), or the fragmented nature of urban
habitats (Andrieu et al. 2009). An increase in alternative food sources can also
disrupt pollinator behaviour, leading to reduced pollination even when pollin-
ator abundance is unaltered. The ornamental and exotic plants that typically
dominate urban floras can have this effect (Kandori et al. 2009; Morales &
Traveset 2009), and in the New World, the provision of hummingbird feeders
can have similar consequences (Arizmendi et al. 2007). Alternative pollinators
can, however, sometimes compensate for reduced availability of core pollin-
ators, resulting in altered biotic interactions but no reduction in reproductive
success (Arizmendi et al. 2007; Roberts et al. 2007). Effects of urbanisation on
wind pollination have not yet been studied. It seems possible though that, at
least in highly developed areas, buildings and other infrastructure will inter-
cept pollen, and may reduce pollination rates and thus the connectivity of
fragmented populations. Lower rates of successful cross-pollination in urban
plant populations can also be associated with increased self-compatibility
(Cheptou & Avendaño 2006).
The impacts of urbanisation on reproductive success have been most rigor-
ously studied in birds. Clutch size, nestling weight and productivity per nesting
attempt are frequently, but not universally, reduced in urban populations
(Chamberlain et al. 2009). These patterns appear to be driven mainly by reduced
abundance of natural food, although the role of density dependence in urban
adapters has not been adequately explored. Nest predation risk is often con-
sidered to be influenced by urbanisation, but there is little evidence for a
general trend (Chamberlain et al. 2009). Some studies provide support for the
‘safe nesting zone’ hypothesis (i.e. reduced nest predation rates in urban areas;
Gering & Blair 1999; Kosinski 2001; Newhouse et al. 2008); the mechanisms
generating such reductions in predation are poorly understood, although urban
noise can mask the acoustic signals used by predators to locate nests (Francis
et al. 2009). In contrast, other studies suggest that in towns and cities the
increased abundance of key nest predators, such as cats and many corvid species
(Sims et al. 2008) increases nest predation rates (Thorington & Bowman 2003;
68 K. L. EVANS

Jokimäki et al. 2005; Beck & Heinshon 2006; Marzluff et al. 2007). Many other
factors, including habitat structure, the responses of predators to human
disturbance, and human attitudes, are also likely to influence the relative rates
of nest predation in urban and rural landscapes. Data from other taxa are
much rarer and provide conflicting results, with urbanisation being associated
with increased (amphibians: Severtsova et al. 2002), decreased (reptiles: Rubin
et al. 2004) or little significant variation (reptiles: Endriss et al. 2007; mammals:
Scott et al. 1999; McCleery 2009) in reproductive success and recruitment
relative to rural populations. It is thus not yet possible to ascertain if the trends
documented for urban bird populations generalise to other groups.

Survival
It is remarkable how few studies have assessed the effect of urbanisation on
survival. It is clear that increased predation or pollution can reduce survival
markedly in some urban populations, and such reductions can even result in
local extinction (e.g. Scott et al. 1999; Hamer & McDonnell 2008). Other studies
have, however, found that survival rates differ little in rural and urban popula-
tions (McCleery et al. 2008; Rodewald & Shustack 2008), or are higher in urban
populations (Liro 1985; Endriss et al. 2007). Indeed, supplementary feeding
seems likely to increase survival rates of many urban bird populations,
although conclusive data are not yet available (Brittingham & Temple 1998;
Robb et al. 2008). The nature of differences in rural and urban survival rates
may also be dependent on spatial and temporal variation in mortality agents,
such as predation rates and disease outbreaks (Gosselink et al. 2007).
Many more case studies are required across a wide range of taxonomic
groups, and across species that exhibit both positive and negative responses
to urbanisation, before broad generalisations can be made concerning the
effects of urbanisation on demographic traits and the mechanisms driving
these patterns. Survival rates in urban populations, relative to rural conspeci-
fics, are especially poorly known. Most species that occur in urban environ-
ments do not reach markedly higher densities in towns and cities than in rural
areas, which increases the probability that most urban populations will not
have higher survival rates or reproductive output. This prediction receives
further support from evidence that, even in species that are abundant in towns
and cities, urbanisation can adversely affect demographic traits (Mennechez &
Clergeau 2006), suggesting that some urban populations may be sinks main-
tained by immigration from surrounding rural areas.

Population genetic structure

Urban populations may arise through development of a site that converts an
existing rural population into an urban one. In such cases the genetic struc-
ture of the urban population will partly reflect underlying patterns that were
 INDIVIDUAL SPECIES AND URBANISATION 69

present before urbanisation commenced (Swei et al. 2003). In addition,

however, habitat alteration is likely to alter dispersal rates between the
recently created urban population and surrounding rural ones, resulting in
changes to genetic structure that are similar to those created by habitat
fragmentation (i.e. reduced genetic diversity and increased genetic drift and
divergence; Keyghobadi 2007).
Alternatively, urban populations may arise through colonisation (Wandeler
et al. 2003; Marzluff 2005; Abs & Bergen 2008; Evans et al. in press, b). These
cases provide rarely documented examples of internal range expansion, the
colonisation of unoccupied habitat types within a species’ original geographic
range, rather than the more frequently studied cases of colonisation of novel
areas external to a range (Evans et al. in press, b). During colonisation, founder
effects are likely to promote reduced genetic diversity in novel populations,
and genetic drift and divergence from ancestral populations. The magnitude of
such changes will be influenced by whether colonisation occurs from a sur-
rounding rural population, or in a leap-frog manner in which urban adapted,
or imprinted, individuals colonise other towns and cities (Evans et al. in press, b).
In the latter situation the source individuals will be derived from an urban
population that has already experienced at least one founder event, thus
increasing the loss of genetic diversity and divergence from rural populations
(Evans et al. 2009c).
The effects of urbanisation on population genetic structure have been
assessed in a wide range of species, but these still represent a very small
proportion of those that are urbanised. The plethora of metrics used to mea-
sure genetic structure hinders direct comparisons between studies, but it is
clear that although urbanisation typically reduces genetic diversity there is
marked variation in the magnitude of such reductions (Table 4.1). Three factors
are likely to have a strong influence on the genetic diversity of urban popula-
tions. First, and perhaps most obviously, the magnitude of bottlenecks arising
through founder effects will exert a large effect on genetic diversity, at least in
recently established urban populations. Second, loss of genetic diversity is
likely to be greater in native species with limited dispersal capacity (Desender
et al. 2005; Aruda & Morielle-Versuta 2008). Gene flow between horticultural
specimens and native plants can, however, result in increased genetic diversity
of urban populations (Roberts et al. 2007), and similar effects may arise if urban
populations are founded from multiple rural ones (Aruda & Morielle-Versuta
2008). Finally, reductions in genetic diversity are likely to be marked in species
that require genetic adaptation to urban environments, as not all genotypes
will contain the necessary genetic adaptations (Keane et al. 2005).
Most urban populations exhibit genetic divergence from rural ones,
albeit again with considerable variation in the magnitude of differentiation
(Table 4.1). In part, this is simply a consequence of reduced genetic diversity,
Table 4.1 Studies comparing the population genetic structure of rural and urban populations. Differences that are not statistically significant
are indicated by ns. Genetic diversity is measured by a – number of alleles, aR – allelic richness, GD – Nei’s gene diversity, GI – mean number
of genotypes per individual, h0 – Shannon’s diversity index, Ha – number of haplotypes, HaD – haplotype diversity, ho – observed
heterozygosity, PB – percentage of polymorphic bands, and PL – percentage of polymorphic loci. Genetic divergence is measured by
dS – Nei’s genetic distance, dNL – Nei and Li’s genetic distance, DIS – Dice’s similarity value, fST and its derivative ΦPT (see the cited sources
for how these metrics were calculated).

Change in genetic
diversity of urban Divergence of urban Divergence within
Species Region populations and rural populations urban populations Source

Plants
[moss] Leptodon smithii Italy Ha 36% FST 0.370 Pop A FST 0.242 Spagnuolo et al.
PL 20% Pop B FST ns (2007)
McGillivray Grevillea Australia A þ95% Roberts et al.
macleayana Ho þ27% (2007)
Common dandelion USA GI 41% Keane et al.
Taraxacum officinale (2005)
Broad-leaved helleborine Scotland A 10% FST 0.328 Hollingsworth &
Epipactis helleborine Ho 29% Dickson (1997)
Rue-leaved saxifrage Germany GD H0 and PB ns ΦPT 0.30 Reisch (2007)
Saxifraga tridactlites
Downy yellow violet USA H0 and PL ns DNL 0.453 Culley et al.
Viola pubescens (2007)
Oxlip Primula elatior Belgium FST 0.054 van Rossum
(2008)
Insects
Small white butterfly Japan PL ns Takami et al.
Pieris rapae (2004)
[butterfly] P. melete Japan PL ns Takami et al.
(2004)
[ground beetle] UK AR and Ho ns Desender et al.
Pterostichus madidus (2005)
[ground beetle] Belgium AR and Ho ns Desender et al.
P. madidus (2005)
[ground beetle] UK AR 15% Desender et al.
Abax ater Ho 88% (2005)
[ground beetle] Belgium AR and Ho ns Desender et al.
Abax ater (2005)
Amphibians
[frog] Leptodactylus Brazil GD 46% DS 0.21 Aruda &
ocellatus PB 32% Morielle-
Versuta (2008)
[frog] L. fuscus Brazil GD 10% DS 0.04 Aruda &
PB 8% Morielle-
Versuta (2008)
[frog] L. podicipinus Brazil GD þ50% DS 0.04 Aruda &
PB þ37% Morielle-
Versuta (2008)
Table 4.1 (cont.)

Change in genetic
diversity of urban Divergence of urban Divergence within
Species Region populations and rural populations urban populations Source

[frog] L. labyrinthicus Brazil GD þ11% DS 0.06 Aruda &

PB þ6% Morielle-
Versuta (2008)
Common toad Bufo bufo UK A 22% FST 0.230 (cf. rural Hitchings &
Ho 15% FST 0.066) Beebee (1998)
Common frog Rana UK A 10% FST 0.35 (cf. rural Hitchings &
temporaria Ho 25% FST 0.11) Beebee (1997)
Red-backed salamander Canada A 29% FST 0.189 FST 0.064 (cf. rural Noël et al. (2007)
Plethodon cinerus AR 39% FST  0)
Ho34%
Reptiles
Caissaca viper Bothrops Brazil PL 28% FST 0.19 Dutra et al.
moojeni (2008)
Blanding’s turtle USA PB 18% DIS ns Rubin et al.
Emydoidea blandingii (2001)
Birds
Dark-eyed junco Junco USA AR 41% FST 0.079 Rasner et al.
hyemalis Ho 21% (2004)
Blackbirda Turdus merula Europe AR 10% FST 0.037 Evans et al.
Ho 2% (2009c)
Eurasian kestrel Falco Poland AR 10% FST 0.05 Rutkowski et al.
tinnunculus Ho 8% (2006)
Mammals
Red fox Vulpes vulpes Switzerland A 14% FST 0.04 FST 0.068 (cf. rural Wandeler et al.
Ho 17% FST 0.009) (2003)
Large Japanese Japan Ha 77% FST 0.49 FST 0.77 (cf. rural Hirota et al.
fieldmouse Apodemus HaD 71% FST 0.20) (2004)
speciosus
Eurasian badger Meles UK AR 15% Huck et al.
meles (2008)

Note: aData for the blackbird are means from 12 paired urban and rural populations.
74 K. L. EVANS

with urban populations that have lost substantial diversity exhibiting greater
divergence from rural populations. Limited dispersal of urban individuals will
further reduce gene flow and increase divergence. It is thus noteworthy that
urban populations of some bird species appear to be more sedentary than those
of rural conspecifics (Eurasian kestrel: Plesnik 1990; blackbird: Partecke &
Gwinner 2007; K. L. Evans et al. unpublished data), and that urbanisation
promotes reduced dispersal in some plant species (Crepis sancta: Cheptou et al.
2008). Divergence between rural and urban populations will also be increased
by biased dispersal patterns arising from individuals imprinting on their natal
habitat type. This mechanism, often termed natal habitat preference induc-
tion, has been documented in some species (Davis & Stamps 2004; Mabry &
Stamps 2008), but has not yet been investigated in an urban context. Genetic
divergence is also likely to be greater when differential selection pressures act
on rural and urban populations, as strong selection pressure can overcome the
homogenising influence of gene flow (Garant et al. 2005; Senar et al. 2006).
Evidence for habitat-specific selection pressure is provided by studies indicat-
ing that trait divergence between urban and rural populations is a conse-
quence of genetic adaptation. Such traits include heavy metal tolerance in
plants (Keane et al. 2005), reduced investment in extra-floral nectaries that are
associated with ant attraction in the plant Chamaecrista fasciculate (Rios et al.
2008), and changes in avian stress physiology and migratory behaviour
(Partecke et al. 2006; Partecke & Gwinner 2007). The impact of time since
colonisation on the genetic structure of urban populations has rarely been
established, but there is evidence for both decreasing (Wandeler et al. 2003) and
increasing divergence (Evans et al. 2009c) over time, with the latter occurring
when selection pressures appear to differ in rural and urban environments.
The final impact of urbanisation on genetic structure concerns genetic
differentiation within urban populations. Although data are very limited, this
appears to be greater than equivalent differentiation within rural populations
(Table 4.1). Genetic structure within urban populations can arise if urban areas
are colonised by genetically differentiated rural populations (Hollingsworth &
Dickson 1997), but probably arises most frequently from intensive fragmentation
of urban habitats reducing gene flow (Vandergast et al. 2007). There is a time-lag,
however, between fragmentation and impacts on population genetics, such that
long-lived species tend to show little genetic structure within urban environ-
ments (Rubin et al. 2001; van Rossum 2008). Human activities, such as movement
of soil between habitat patches, and thus associated faunas and seed banks, can
also promote gene flow between isolated populations to a sufficient extent to
mitigate the effects of fragmentation (Field et al. 2007).
The trends for urban populations to contain less genetic diversity than rural
populations, and for the two to be genetically divergent from each other, have
important consequences. Theory and empirical data indicate that reductions in
 INDIVIDUAL SPECIES AND URBANISATION 75

genetic diversity will limit the ability of populations undergoing range expan-
sion to adapt to novel environments (Hewitt 2000; Pujol & Pannell 2008).
Therefore, reduced evolutionary potential of urban populations may hinder
their adaptation to urban environments, possibly contributing to their typic-
ally lower species richness relative to rural areas (Chapter 5; Grimm et al. 2008).
Conversely, genetic differentiation between urban and rural areas is indicative
of reduced gene flow, which may facilitate adaptation to towns and cities, as it
may buffer locally adapted genotypes from being swamped by genes from rural
populations (Hoffmann & Blows 1994; Bridle & Vines 2007). The relative impor-
tance of reduced genetic diversity and gene flow in determining evolutionary
potential is highly debatable, but urban studies clearly have great potential to
contribute to this debate.

Synthesis and directions for future research

Owing to a recent explosion of interest, divergence between conspecific urban
and rural populations has been demonstrated in a remarkably wide range of
traits. Despite this rapid progress, a number of issues require further attention.
First, the studies conducted to date almost certainly concern just a fraction of
the traits that are affected by urbanisation, and a broader range of studies is
required. Second, studies are currently markedly biased towards northern
temperate regions and birds; it is particularly surprising that relatively few
studies have looked at intraspecific trait divergence in rural and urban plant
populations. These geographic and taxonomic biases must be addressed before
general patterns in trait divergence can be ascertained fully. Third, most
studies have focused on a single urban population, and simultaneous assess-
ments of divergence across multiple sites and species are required to assess
fully the extent to which patterns generalise (but see Figure 4.3). Fourth, the
impacts of urbanisation on multiple traits have been assessed in very few
species (but see Figure 4.4); more studies of this kind are required to enhance
understanding of the interactions and trade-offs that seem likely to influence
trait divergence. Fifth, much more attention should be paid to assessing which
characteristics of the urban environment drive trait divergence and to assess-
ing the fitness consequences of such divergence in order to establish its adap-
tive value. Sixth, few studies assess whether divergence arises through
phenotypic plasticity or genetic adaptation (but see Partecke et al. 2004, 2006;
Partecke & Gwinner 2007; Liker et al. 2008; Rios et al. 2008), and even fewer
adequately distinguish between genetic adaptation and epigenetic effects. Sev-
enth, there is very little understanding of the rate at which trait divergence
between urban and rural populations has arisen. This is unfortunate, as discus-
sion of evolutionary rates in human-altered environments is currently based on
studies of a limited range of selection pressures that do not include the impacts
of urbanisation (Hendry et al. 2008; Smith & Bernatchez 2008). It would thus be
76 K. L. EVANS

Figure 4.4 The blackbird Turdus merula is emerging as a model species with which to
assess trait divergence in urban and rural populations. It provides a very scarce example
of a species in which such divergence has been assessed for multiple traits; these include
demography, disease risk, migratory behaviour, morphology, population genetic
structure, stress physiology and vocalisations (Grégoire et al. 2002; Partecke et al. 2004,
2006; Partecke & Gwinner 2007; Evans et al. 2009a, 2009b, 2009c and unpublished data;
Nemeth & Brumm 2009). For some traits the relative contributions of phenotypic
plasticity and genetic changes to differentiation have been assessed, and divergence in
other traits has been assessed in multiple urban populations. Photograph supplied by
Z. G. Davies.

useful to assess the magnitude of trait divergence in populations for which the
timing of urbanisation is known. Despite these requirements for additional
work, it is clear that selection pressures differ enough in many urban and rural
areas to generate trait divergence between conspecific populations occupying
these contrasting environments. Moreover, the nature of this trait divergence
can often be related to theories derived from investigations of the more natural
systems on which ecology has traditionally focused. Urban ecology should thus
not be viewed as an isolated discipline; rather, assessing how populations adapt
to towns and cities may shed light on many fundamental unresolved ecological
and evolutionary issues.

Acknowledgements
This work was funded by the Natural Environment Research Council and the
Leverhulme Trust. D. Chamberlain, C. Isaksson and an anonymous reviewer
provided assistance.
 INDIVIDUAL SPECIES AND URBANISATION 77

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CHAPTER FIVE

Species diversity and urbanisation:

patterns, drivers and implications
GARY W. LUCK AND LISA T. SMALLBONE

Spatiotemporal patterns in species diversity have intrigued ecologists for many

decades (MacArthur 1965; Brown 1981; Gaston 2000). These patterns occur at a
variety of scales, for example the latitudinal gradient of increasing species
richness from the poles to the Equator (Willig et al. 2003; Hillebrand 2004)
and altitudinal gradients across regions (Sanders 2002; McCain 2004). Their
appropriate identification and explanation have generated enormous interest
and debate (Rahbek & Graves 2001; Willig et al. 2003; Field et al. 2009). It is only
logical to extend this interest to urban systems and examine species richness
patterns, and drivers of these patterns, across gradients of urbanisation at
multiple spatial and temporal scales.
What sets urban studies apart is the implicit or explicit recognition of the
major influence of human landscape modification on species distribution.
Indeed, the effects of anthropogenic activity are virtually impossible to ignore
in urban systems. Moreover, integrating social, cultural, economic, human
demographic and ecological data, and exploring the role of human activities
in moderating patterns in species diversity, are exciting developments that are
forging new ground in interdisciplinary research.
In this chapter, we explore patterns in species diversity in urban systems
across space and time, and briefly review some of the key drivers of these
patterns and their implications. The chapter’s focus is on patterns in species
richness, particularly native species, as variation in species abundance and
density are covered in Chapter 4. Much more attention is given to spatial
patterns in richness, and this reflects the bias in the literature. We group
patterns under the headings ‘broad-scale’, ‘comparative’, ‘gradient’ and
‘urban-centric’. Broad-scale refers to patterns that occur over large spatial
extents (e.g. national, continental or global) with a large grain size (the size
of the sampling unit used; e.g. 1
grid cells) where researchers have examined
covariation in species richness and some measure of urbanisation (e.g. human

Urban Ecology, ed. Kevin J. Gaston. Published by Cambridge University Press.

# British Ecological Society 2010.
 SPECIES DIVERSITY AND URBANISATION 89

population density). Comparative studies are those that compare species rich-
ness in native habitats or rural landscapes with urban areas. A particular type
of comparative study is the rural–urban or native–urban gradient, and we treat
these studies separately since they are prominent in the literature. In compara-
tive and gradient studies, spatial extent is usually much less than broad-scale
studies, and grain size is almost always much smaller. Urban-centric refers to
patterns that occur when sampling locations are primarily nested within a
single urban centre or across centres. Researchers generally compare species
assemblages across different levels of urbanisation, but may occasionally
include direct comparisons with native or rural landscapes.
We use the terms species diversity and species richness interchangeably to
describe the number of species in an area, as this is consistent with their use in
the literature. However, we note that, technically, ‘diversity’ is a function of both
species richness and the abundance of each species and is often encapsulated in
diversity indices such as the Shannon index. Also, ‘urban’ is a term that is poorly
defined across studies and is occasionally used qualitatively rather than
quantitatively. Pickett and Cadenasso (2006) suggested that definitions of urban
will necessarily be flexible and case-specific, but nevertheless the term needs to
be defined in each study. This could be achieved by quantifying key measures
such as housing density, road density or percentage cover of impervious surfaces
(Chapter 2). We try to avoid confusion by referring to patterns in diversity with
increasing urbanisation or between urban and non-urban areas (as indicated
by authors). Hence, urban is used in a relative rather than absolute sense.
We make no attempt to compare levels of urbanisation across studies. While
this might be highly desirable, suitable data often do not exist in the literature.

Patterns in space
Broad-scale
At very broad scales (national to continental), human population density
(HPD) is a useful surrogate for the level of urbanisation, particularly in
developed countries. A growing number of studies have examined correlations
between HPD and species richness for a range of taxonomic groups across
various spatial extents and using a diversity of grain sizes (e.g. Balmford et al.
2001; Araújo 2003; Gaston & Evans 2004; Luck et al. 2004; Fjeldså & Burgess
2008). A somewhat surprising result from these studies is the consistent
reporting of positive correlations between HPD and species richness; surprising
because of the undoubted negative impacts of human landscape modification
on the persistence of many species.
Luck (2007a) conducted a major review of the relationships between HPD and
biodiversity, including a meta-analysis of studies correlating HPD with species
richness. He found strong, positive population effect sizes (a population effect
size is a single value combining correlation coefficients across studies using
90 G. W. LUCK AND L. T. SMALLBONE

1.0

0.8
Population effect size

0.6

0.4

0.2

0.0
Birds Mammals Plants Threatened Restricted Introduced
species species species

Figure 5.1 Population effect sizes (combining correlation coefficients across studies)
between human population density and the species richness of birds, mammals and
plants, and threatened, restricted and introduced species. Error bars are 95% confidence
intervals. Data from Luck (2007a).

meta-analytical techniques) between HPD and the species richness of plants,

birds and mammals (Figure 5.1). However, there was substantial variability
across studies owing to, among other things, different spatial extents, grain
sizes, sample sizes, habitats and approaches.
A key variable in these studies is grain size, which ranged from 100 m2 plots up
to countries in the research reviewed by Luck (2007a). As grain size increases,
so does the strength of the positive correlation between HPD and species
richness (also see Pautasso 2007), although this can be complicated by the fact
that higher population densities may be characteristic of smaller sampling areas.
At small grain sizes the correlation can be, not unexpectedly, negative rather than
positive (e.g. Drake & Pereira 2002; Urquiza-Haas et al. 2009). Moreover, broad-scale
studies using alternative measures of level of urbanisation may yield results that
differ from those relying on HPD. Clergeau et al. (2001) conducted a biogeograph-
ical meta-analysis of 18 studies from towns in temperate and boreal climatic
regions where degree of urbanisation was measured as peri-urban, suburban
and town centre. They found that bird species richness was negatively correlated
with latitude and increasing urbanisation. Lepczyk et al. (2008) found that,
across the mid-western United States, native bird richness was highest where
anthropogenic land cover was lowest and housing density was intermediate.
The general conclusion from the above studies is that people co-occur with a
diversity of species at the regional level (2500 km2), but only a few species
persist with humans at the local level, particularly in highly urbanised loca-
tions. ‘Regions’ are broad enough to encompass human settlements and rela-
tively undisturbed areas, and co-occurrence across regions seems to be driven
by people and other species responding similarly to energy and/or productivity
gradients moderated by human land-use (see ‘Drivers’ below, and Figure 5.7).
 SPECIES DIVERSITY AND URBANISATION 91

Lower species diversity at the local level results from substantial habitat loss in
highly urbanised locations. This realisation helps to reconcile the apparently
contradictory findings among broad-scale and local-scale studies.
Across regions, Luck (2007a) found that HPD was also positively correlated
with the number of threatened species, geographically restricted species
(endemic to a localised region) and introduced species (especially for plants;
Figure 5.1), although the last of these has been poorly explored at broad scales
(Chapter 6). While the correlation with threatened species infers that increas-
ing urbanisation adversely affects species persistence, such spatial correlations
are weak causal explanations. Stronger evidence of the negative impacts of
urbanisation on species persistence comes from studies that track persistence
over time (e.g. Thompson & Jones 1999; Parks & Harcourt 2002; Tait et al. 2005).
Yet there are only a handful of these studies in the literature and there is a
desperate need for long-term monitoring of species persistence in urban areas
as development proceeds.
Geographically restricted species are those with small geographic ranges.
The positive correlation between HPD and these species suggests their ranges
occur, more often than not, close to human settlements. This presents a conser-
vation challenge since there may be few locations within their range that are
distant from human settlements and associated negative impacts. It also under-
scores the dire need to implement effective conservation strategies in urban
areas (Schwartz et al. 2006; Lawson et al. 2008).
Patterns of HPD and species richness in Australia match those from other
countries and continents across a range of taxonomic groups. This is true even
if the effects of biased sampling effort and spatial autocorrelation are con-
trolled for (Luck et al., in press; Figure 5.2). However, reptiles represent an
important exception to the general pattern, as there is a slightly negative
correlation between HPD and reptile species richness across Australia at a grain
size of 1
(although strong positive correlations between HPD and richness are
recorded elsewhere, e.g. North America; Luck et al. 2004). This is because diver-
sity patterns in reptiles, which are ectothermic, are probably more closely
related to spatiotemporal variation in solar radiation or temperature than
other Australian species (including humans) which are responding to gradients
in rainfall and productivity. This results in many reptiles occurring in sparsely
populated arid and semi-arid regions.
The broad-scale trends described above mask more complex patterns
between species richness and urbanisation occurring at local levels. Neverthe-
less, these studies yield important information that may guide regional man-
agement strategies. They also encourage a hierarchical approach to examining
species–people relationships and demonstrate that different patterns and
drivers manifest themselves at different spatial scales. A multi-scaled approach
to ecological studies has gained substantial support in recent decades
92 G. W. LUCK AND L. T. SMALLBONE

Bird species richness (1 degree)

10–26
27–36
37–44
45–52
53–61
62–72
73–93

Human population density (0.05 degree)

1–3190
3191–15 690
15 691–27 190
27 191–48 460
48 461–78 440

0 500 1,000 2,000 Kilometers

Figure 5.2 The distribution of bird species richness and human population density
across mainland Australia. Bird data are at a resolution of 1
grid cells with equal survey
effort per cell (10 random surveys). Blank (white) cells represent insufficient survey
effort (fewer than 10 surveys). Human population density is number of people per 0.05

grid cell. Reproduced with permission from Luck et al. (in press).
 SPECIES DIVERSITY AND URBANISATION 93

(Lindenmayer & Franklin 2002), and urban ecology should continue to promote
this approach.

Comparative
Comparisons of species richness between urban areas and those dominated by
agriculture or native vegetation generally find that richness is lower in urban
areas, while total species abundance is higher with a handful of species contrib-
uting the majority of individuals. There are important exceptions to this
general pattern identified from studies across different taxonomic groups
and subsets of species from a particular group (e.g. cavity-nesting birds). It is
also important to note that some researchers confine their sampling entirely to
patches of native vegetation (e.g. surrounded by agriculture or urbanised
areas), while others sample the full range of available habitats. Taxonomic bias
inevitably exists in these studies, with much work conducted on birds and to a
lesser extent plants, and fewer studies on other taxonomic groups. This reflects
a combination of ease of sampling and the fact that a range of bird and plant
species have adapted relatively well to urbanisation.
Lower species diversity with increasing urbanisation has been found for birds
(Cam et al. 2000; Sandström et al. 2006), bats (Kurta & Teramino 1992;
De Cornulier & Clergeau 2001), terrestrial mammals (Tait et al. 2005) and
amphibians (Gagné & Fahrig 2007; see Hamer & McDonnell 2008 for a review
of urbanisation effects on amphibians), but the occasional study does not fit this
trend. For example, Palomino and Carrascal (2007) found that bird species
richness in native vegetation patches was not adversely affected with decreasing
distance to small cities (<15 000 people) in Central Spain, although roads had
generally negative impacts. Fewer studies have been conducted on invertebrates
(see McIntyre 2000 for a review), but Rickman and Connor (2003) found that the
species richness and total abundance of leaf-mining Lepidoptera in habitat
remnants around San Francisco Bay was not influenced by the extent of urban-
isation occurring in a 500-m radius around each remnant. However, increasing
urban density in Melbourne, Australia had substantial negative impacts on
macro-invertebrate communities of small streams, whereby metropolitan
communities were dominated by a few abundant species (Walsh et al. 2001).
Variation in the results among studies can be explained in part by the varied
responses of particular species groups. A number of studies have examined
changes in the diversity of subsets of bird species with increasing urbanisation,
and Chace and Walsh (2006) conducted a comprehensive review of the effects
of urbanisation on avifaunal assemblages. Results across studies are not always
consistent, but some general trends can be identified (Table 5.1). For example,
housing density was negatively correlated with the richness of territorial
species, forest interior species and neotropical migrants in some regions of
the USA (Mills et al. 1989; Friesen et al. 1995; Green & Baker 2003), while
94 G. W. LUCK AND L. T. SMALLBONE

Table 5.1 Generalised trends in species richness and abundance of

subsets of avian species in response to increasing urbanisation.

Exotic species ↑ Sedentary species ↑?

Habitat generalists ↑ Short-distance migrants #?
Habitat specialists # Widely distributed species ↑
Ground-nesters # Narrowly distributed species #
Cavity-nesters ? Omnivores ↑
Shrub-nesters #? Insectivores #?
Tree-nesters ↑? Granivores ↑?

Note: A question mark indicates a degree of uncertainty in likely responses, while

strong conflicting trends have been reported for cavity-nesters.

omnivores and exotic species have been shown to increase in urban areas in the
USA and Europe (Jokimäki & Suhonen 1998; Allen & O’Connor 2000). Pidgeon
et al. (2007) found that forest-dependent birds were mostly negatively affected by
increasing house density across the USA, although results varied for different
species groups and ecoregions. Cavity-nesters, short-distance migrants and nar-
rowly distributed species were some of the groups to show negative responses.
For bird communities in Singapore, insectivores, carnivores, shrub-nesters
and primary cavity excavators were adversely affected by increasing urbanisa-
tion, whereas frugivores prospered in areas of low-density housing (Lim &
Sodhi 2004). Other studies support the trend of insectivore decline with
increasing urbanisation, while also showing that granivores may adapt well
to urban environments (e.g. Allen & O’Connor 2000; Lindsay et al. 2002). In
Australia, urbanisation may favour behaviourally aggressive and medium- to
large-bodied bird species (Garden et al. 2006). Many of these species are respond-
ing to resource availability in the urban landscape. For example, large honey-
eaters are attracted to streetscapes with a high density of flowering native trees
and shrubs.
In contrast to the results of Pidgeon et al. (2007) and Sandström et al. (2006),
Chace and Walsh (2006) suggested that cavity-nesters are favoured by urbanisa-
tion (also see Miller et al. 2003). Such apparently contradictory findings high-
light the importance of acknowledging contextual and ecological differences
across studies. For example, Australia has many cavity-[hollow-]nesting birds,
but no species that can excavate cavities, which are formed from insect and
fungal attack. Hollow-bearing trees are rare in Australia’s urban environments,
since most are cleared during development, and hollows take many decades to
form in native trees so are mostly absent from plantings post development.
Hence, urban areas in Australia do not support a high diversity of hollow-
nesting species (Garden et al. 2006). It is also important to acknowledge that
few studies that examine how species groups with particular traits vary in
 SPECIES DIVERSITY AND URBANISATION 95

richness with urbanisation simultaneously assess multiple traits or address

species phylogeny.
A number of studies classify bird species into ‘urban-adapted’ or ‘urban-
sensitive’ (e.g. Blair 1996; Lim & Sodhi 2004). Classifications based purely on
variation in population abundance across gradients of urbanisation are not
particularly informative without exploration of the ecological traits that allow
species to adapt to urban environments. Thankfully, a handful of studies have
begun to explore this issue (e.g. Kark et al. 2007; Croci et al. 2008). Splitting
species into urban adapters or urban-sensitive can result in different patterns
in richness. For example, Tratalos et al. (2007) found that, across Britain, total
bird species richness and that of urban-adapted species increased from low to
moderate housing density then declined at higher density (following the
hump-shaped pattern often reported in rural–urban gradient studies; see
below), while the richness of urban-sensitive species consistently declined with
increasing density. Total species abundance and the abundance of urban-
adapted species increased with housing density, declining only at the highest
densities.
Richness patterns for plants often differ from those of faunal species. Urban
areas have been shown to harbour more plant species than surrounding non-
urban areas (Kühn et al. 2004), and species richness is positively correlated with
city size (Pyšek 1998), although this may reflect a simple species–area relation-
ship. Positive correlations still exist even when plants are split between native
and introduced species. For example, Pyšek (1998) found that, across 54 cities in
central Europe, the richness of native plants, aliens, neophytes (introduced to
central Europe after ad 1500) and archaeophytes (introduced before ad 1500)
increased with city area, number of inhabitants and HPD. The increase was
strongest for neophytes, the most recent introductions. An important result
from this study was that the proportional representation of aliens was much
higher in cities than in the regional species pool (Figure 5.3). This is indicative
of urban areas being highly modified landscapes, although still supporting a
rich diversity of plant species.
Kühn et al. (2004) demonstrated that native and naturalised alien plant
species richness were significantly higher in city grid cells than in non-city
grid cells in Germany. Native plant species richness was largely explained by
the number of geological types per grid cell (130 km2), and cities were mostly
settled in areas of high geological diversity. Hence, human settlers were
attracted to inherently diverse locations. Similar to Pyšek (1998), Kühn et al.
(2004) found that the proportional representation of alien (neophytes and
archaeophytes) versus native species increased in city (26%) compared with
non-city (rural; 19%) grid cells. Of course, contrary patterns can generally be
found. For example, Roy et al. (1999) showed that total plant species richness
and the richness of native plants did not increase with urbanisation in Britain,
96 G. W. LUCK AND L. T. SMALLBONE

120

100

Native species
Species richness

80

60

40
Introduced species
20

0
Urbanisation

Figure 5.3 A stylized representation of broad trends in plant species richness with
increasing urbanisation. Introduced species generally increase in richness and in their
proportional contribution to total species richness. Native species richness may either
increase slightly or decline with increasing urbanisation (dashed line). The relationships
are asymptotic, and total species richness may decline at very high levels of
urbanisation.

although alien species richness did. Importantly, they used a much smaller
grain size (2 km2) than the above studies.
In support of the results found for bird species, work on plants has also
determined that species with particular traits are more likely to persist in
urban environments. For example, Thompson and McCarthy (2008) demon-
strated that in Sheffield and Birmingham in England, urbanisation favoured
plant species that were larger (based on plant height) and that preferred base-
rich habitats and, to some extent, dry, unshaded and moderately fertile habi-
tats. Their results were largely consistent for both native and introduced
species. Plant height also dictated the likelihood of extinction for plants in
Middlesex, England, whereby short plants (mostly natives) were more likely to
suffer extinction (Preston 2000; a similar result was recorded in Auckland,
New Zealand by Duncan and Young 2000).
In an important follow-up to the study by Kühn et al. (2004), Knapp et al.
(2008) showed that plant species richness in German cities was not related to
phylogenetic diversity. That is, high species richness mostly resulted from a
greater number of more closely related, functionally similar species adapted to
urban areas. This suggests that functional diversity is reduced in urban versus
non-urban areas. Such a conclusion is well supported in the growing literature
on biotic homogenisation, whereby increasing urbanisation reduces species
diversity across sites and leads to greater community similarity among
urban areas than among non-urban areas (Rahel 2000; Olden & Poff 2003;
McKinney 2006; Olden et al. 2006; Devictor et al. 2007). Homogenisation of
 SPECIES DIVERSITY AND URBANISATION 97

species assemblages in urban areas appears to be driven by invasions or intro-

ductions of generalist, urban-adapted species, local extinctions of specialist
species (McKinney & Lockwood 1999; Olden et al. 2004) and the promulgation
of similar habitat components across urban localities. The process of homogen-
isation raises substantial challenges for protecting diverse species assemblages
within human settlements.

Gradient studies
One way to look at patterns of urbanisation is to sample along a gradient of
increasing settlement intensity from native habitats or rural areas to city
centres. Sampling can occur at non-contiguous points in different land-use/
urbanisation categories along a transect running from city centre to rural/
native zone, or at points located randomly throughout the entire study area.
Studies of this type have been undertaken across a range of settlement sizes
from major cities (population >1 000 000) to regional towns (population
<250 000) (e.g. Sewell & Catterall 1998; Melles et al. 2003; Caula et al. 2008).
Grain sizes varied from <10 ha (Sewell & Catterall 1998; Smith & Wachob 2006;
Caula et al. 2008) to much larger plots (>2000 ha; Weng 2007; Gaublomme et al.
2008). Native–urban or rural–urban gradients are generally defined by some
index of urbanisation using physical (e.g. building density, land-use types,
vegetation cover, density of roads or distance to the central business district;
Melles et al. 2003; Clergeau et al. 2006; Pillsbury & Miller 2008), demographic
(e.g. population density or demographic indices; Rubbo & Kiesecker 2005;
Hahs & McDonnell 2006) and/or landscape metrics (e.g. patch fragmentation,
patch size, edge or land-use heterogeneity; Luck & Wu 2002; Weng 2007).
Studies across gradients often report species diversity peaking at moderate
levels of development with reduced species richness occurring at high levels
(urban) and low levels (rural and native) of development (Sewell & Catterall 1998;
Blair 1999, 2004; Smith & Wachob 2006). Rural–urban gradients in particular
often display a hump-shaped pattern in species richness (Figure 5.4). This is
consistent with the intermediate disturbance hypothesis, which predicts that
species richness will be highest at intermediate levels of disturbance (Connell
1978). Many studies have observed this relationship for a variety of taxonomic
groups including birds (Sewell & Catterall 1998; Blair 2004; Chace & Walsh
2006), bats (Gehrt & Chelsvig 2003; Duchamp et al. 2004) and lizards (Germaine &
Wakeling 2001). This response may be the result of increased resources at
moderate levels of urbanisation, with gardens, parks, reserves and other land-
uses characterising these areas and providing habitat for particular groups of
species (French et al. 2005; Gaston et al. 2007; Young et al. 2007).
McKinney (2008) conducted a comprehensive review of studies on
plants, invertebrates and non-avian vertebrates (i.e. mammals, reptiles and
amphibians), comparing species richness along a gradient from low (rural) to
98 G. W. LUCK AND L. T. SMALLBONE

Figure 5.4 Stylised trend of species richness (synthesising results of gradient studies)
along a rural–urban gradient defined by increasing dwelling density with
corresponding aerial photos of landscape types. Each figure represents
an area of 600 by 600 m.

moderate (suburban) to high (urban core) levels of urbanisation. He found that

11 of 17 plant studies recorded an increase in species richness from low to
moderate urbanisation, while studies of invertebrates and non-avian vertebrates
were more likely to record a decrease in richness (30 of 47 studies and 14 of 17
studies, respectively; although 14 of the invertebrate studies documented an
increase in species richness from low to moderate levels of urbanisation). Con-
versely, in the transition from moderate to high urbanisation, no study on plants
or vertebrates and only one study on invertebrates recorded an increase in species
richness. Most studies recorded a decrease in richness for vertebrates and inver-
tebrates, while the number of plant studies recording a decrease in richness or no
change was evenly split. McKinney (2008) concluded that plant species richness
tended to peak at moderate levels of urbanisation, while the richness of inverte-
brates and non-avian vertebrates peaked at low levels of urbanisation.
Species display different tolerances to urban disturbance and have been
grouped as ‘exploiters’, ‘adapters’ or ‘avoiders’ of urbanisation (Blair 1996;
Kark et al. 2007; Croci et al. 2008). Species classified as adapted to moderate
levels of disturbance will tend to be dominant at the rural–urban interface
where landscapes are the most heterogenous (Sewell & Catterall 1998; Melles
et al. 2003; Caula et al. 2008). This is a distinct group of species able to use
resources typical in this landscape, but the assemblage is very different from
the species that occurred in the area prior to development (see Tait et al. 2005;
McKinney 2006).
 SPECIES DIVERSITY AND URBANISATION 99

Another recurring pattern along rural–urban gradients is the greater

abundance of species at the urban centre, with a few urban exploiters contrib-
uting the majority of individuals. This has been noted for birds and micro-bats
(Kurta & Teramino 1992; Lesinski et al. 2000; Blair 2004; Palomino & Carrascal
2007). Other taxonomic groups show a different pattern even when a few
species are dominant. For example, abundance has been observed to decrease
at the urban centre for amphibians (Rubbo & Kiesecker 2005; Pillsbury & Miller
2008), lizards (Germaine & Wakeling 2001), butterflies (Blair 1999) and ground
beetles (Gaublomme et al. 2008).
The difference in responses of certain taxa may be related to their ability to
disperse through the landscape, their habitat preferences and the location of
habitat along the rural–urban gradient (Niemelä et al. 2000; Atauri & De Lucio
2001). For example, frogs are highly sensitive to habitat fragmentation and
road density (Rubbo & Kiesecker 2005; Parris 2006; Pillsbury & Miller 2008),
which is highest at the city centre, and they often rely on ephemeral ponds for
breeding, which are more likely to be found in the peri-urban and rural zone
(Hamer & McDonnell 2008). The level of urban disturbance a species is able to
tolerate depends on its ecological traits, and these will determine a species’
capacity to occupy niches along the gradient (see ‘Urban-centric’ below).
Some studies have observed seasonal variation in species richness patterns
along rural–urban gradients, whereby richness generally declines with increas-
ing building intensity during favourable seasons when resources are abundant
(e.g. spring), whereas in the harshest seasons with greater resource restrictions
(winter/summer) more species are recorded using urban environments of inter-
mediate disturbance. For example, in Brisbane, Australia, Sewell and Catterall
(1998) found more bird species in moderately disturbed suburban environ-
ments than large vegetated remnants (>100 ha) during summer compared
with winter (summer is a harsher season for this location). In Montpellier,
Southern France, Caula et al. (2008) found more bird species using suburban
environments compared with croplands and woodlands during winter than in
spring. Hence, some species may be exploiting resources within moderately
disturbed urban environments when harsh climatic conditions are affecting
resource availability in the surrounding landscape. This suggests that urban-
isation may limit seasonal variation in important resources (e.g. water), and
this is a crucial question for future research. Understanding the patterns of
species distribution along rural–urban gradients can help to focus conserva-
tion and restoration efforts where they will be the most effective.

Urban-centric
Comparative studies generally show that species richness declines with increas-
ing urbanisation or that richness peaks at some intermediate level of develop-
ment especially across rural–urban gradients. However, patterns vary across
100 G. W. LUCK AND L. T. SMALLBONE

taxonomic groups and for species with specific ecological traits, and some
species can readily adapt to or exploit urban habitats. Simple comparisons of
native versus urban landscapes sometimes treat ‘urban’ as a homogeneous
unit, ignoring the substantial heterogeneity that can occur within an urban
environment. This heterogeneity leads to changes in species diversity and
composition across neighbourhoods within towns and cities.
The species richness of fauna across neighbourhoods is often positively
correlated with the cover of native and/or exotic vegetation. Studies on birds
consistently report on the importance of retaining native tree canopy cover,
vegetation structure and streetscape vegetation to support a higher number of
species (e.g. Mills et al. 1989; Fernández-Juricic 2000; Hennings & Edge 2003;
White et al. 2005; MacGregor-Fors 2008). White et al. (2005) found that parks and
streetscapes with native vegetation supported more bird species than recently
developed streetscapes or those with exotic vegetation in Melbourne suburbs.
Hennings and Edge (2003) found that urban canopy cover promoted native bird
richness in Portland, Oregon, and Green (1984), also working in Melbourne,
reported that total native vegetation cover was positively correlated with native
bird richness, but negatively correlated with exotic bird richness. Studies also
report the importance of native remnants and riparian zones within urban
environments as potential conservation reserves (Bush et al. 2003; Hennings &
Edge 2003; Hodgson et al. 2006; Pennington et al. 2008).
Surveys confined entirely within green space surrounded by urbanisation
(e.g. remnant vegetation or recreation parks) generally show that species rich-
ness is highest in the areas that most closely resemble the previous land cover
(e.g. forest reserves), and the size of remaining vegetation patches is positively
correlated with species richness, consistent with well-established species–area
relationships (e.g. Mörtberg 1998; Koh & Sodhi 2004; Palmer et al. 2008).
Vegetation area seems to be particularly important for those species that
cannot readily use the surrounding urban landscape, although Antos et al.
(2006) found that vegetation patch size was also positively correlated with
introduced bird species richness in urban areas in Melbourne. Moreover, area
might not be the most critical factor driving patterns in species richness in
some taxonomic groups. Garden et al. (2007) found that habitat structural
elements were the most important determinants of reptile and mammal
assemblages in urban vegetation patches in Brisbane.
In sum, these results show, unsurprisingly, that native faunal species
respond positively to the retention of native vegetation and that richness
increases with increasing vegetation cover. Hence, they demonstrate the
importance of retaining vegetation in urban landscapes. Not only can this
promote species conservation, but vegetation offers human residents a range
of additional benefits including microclimate regulation (Harlan et al. 2006;
Jenerette et al. 2007), control of air and water pollution (Randolph 2004),
 SPECIES DIVERSITY AND URBANISATION 101

carbon storage (Nowak 1994), recreational opportunities (Miller 2006), and

cultural and health benefits (Fuller et al. 2007; Tzoulas et al. 2007).
Some of the more interesting urban-centric work links household or neigh-
bourhood socioeconomic characteristics with vegetation cover and species
diversity. This begins to tease apart some of the driving forces of neighbour-
hood variation in urban ecosystems and also highlights demographic groups
that may be disadvantaged through lack of direct access to the benefits of
nature. For example, Hope et al. (2003) found that plant species richness was
positively related to, and primarily a factor of, family income and housing age
across neighbourhoods in Phoenix, Arizona. Income is a key variable in studies
of vegetation cover (e.g. Grove & Burch 1997), and this suggests that people
with greater financial security are either attracted to neighbourhoods with a
higher level of ‘naturalness’ or promote vegetation cover and diversity through
their activities. The flipside of this trend is that residents with a ‘lower’ socioeco-
nomic status often live in neighbourhoods with few species and little vegeta-
tion cover (Iverson & Cook 2000; Pauleit et al. 2005).
Urban-centric studies are now beginning to focus on species richness pat-
terns in household gardens and how householder behaviour may influence
these patterns. For example, Thompson et al. (2003) found that a third of all
plant species in gardens in Sheffield, England, were native and, collectively,
garden quadrats contained twice as many species as semi-natural habitats
(although mean species richness per quadrat was lower in gardens). Moreover,
plant species richness increased with garden area, with the relationship being
potentially stronger for natives than exotics (Smith et al. 2006). Householder
activities such as watering and fertilising can play an important role in main-
taining small plant populations and increasing the diversity of gardens, and
some potential exists for coordinating householder activities to achieve
broader conservation objectives in line with incentive schemes currently used
in rural landscapes.
The structure and composition of household gardens may also have a
stronger influence on bird assemblages than broader landscape factors (Daniels
& Kirkpatrick 2006). Garden area has been shown to be positively related to the
species richness of garden birds, and native birds may be particularly attracted
to native plants (Thompson et al. 1993; Chamberlain et al. 2004). For example,
French et al. (2005) found that native nectarivores, a substantial component of
Australian urban bird communities, preferred to feed on native Banksia spp.
and Grevillea spp. than on introduced Camellia spp. and Hibiscus spp. in Sydney,
Australia.
Provision of supplementary food will generally increase bird species richness
and abundance in gardens (Savard et al. 2000; Daniels & Kirkpatrick 2006;
Parsons et al. 2006). Fuller et al. (2008) documented evidence that bird assem-
blages had important links with bird feeding stations and with the proportion
102 G. W. LUCK AND L. T. SMALLBONE

Socioeconomic status
Low High
Species richness

Figure 5.5 Suburban streetscapes in southeastern Australia along a gradient of

socioeconomic ‘status’ based on income and the proportion of residents with a tertiary
education. Bird and plant species richness tend to be highest in neighbourhoods with
higher status (i.e. higher income and education levels; G. W. Luck and L. T. Smallbone,
unpublished data). Photographs used with permission of L. T. Smallbone.

of households that provide supplementary food in the neighbourhoods of

Sheffield. Interestingly, they also found that species richness was higher in
middle- and high-income neighbourhoods, mirroring the results of Hope et al.
(2003) for plants in Arizona. We have found similar results in towns (17 000 –
80 000 people) across southeastern Australia showing that vegetation cover,
bird species richness and bat activity are all higher in high-income suburbs
where a greater proportion of residents have completed tertiary education
(Luck et al. 2009, and unpublished data) (Figure 5.5).
While there is much variation in the results of urban-centric studies, some
general patterns are still evident, such as the decline in species diversity from
the edge to the interior of metropolitan areas (Green 1984; Clergeau et al. 1998;
Cam et al. 2000; Melles et al. 2003), although it is unclear how such a pattern
might be influenced by town size. The drivers of these patterns reflect complex
interactions among environmental gradients, neighbourhood socioeconomic
characteristics, householder behaviour and species ecological traits. Potential
drivers of species richness patterns in urban landscapes across scales are
further explored below.

Patterns in time
Although there are many studies on spatial patterns in diversity across urban
areas, there are relatively few on patterns over time. This is not unusual in
ecology owing to the rarity of accurate, long-term databases. The most compre-
hensive work relates to temporal changes in vegetation cover in urban areas,
which is relatively easy to map with satellite images or aerial photographs
(e.g. Morawitz et al. 2006; DiBari 2007). Moreover, some studies have tracked
changes in urban land cover or housing development in urban and peri-urban
areas and matched this with loss of vegetation (e.g. Hammer et al. 2004; Tian
et al. 2005; Gonzalez-Abraham et al. 2007). Hence, we have reasonably good
 SPECIES DIVERSITY AND URBANISATION 103

information on temporal patterns in urban development and changes in

vegetation cover, but few data exist on changes in species diversity over time
(and there are even fewer studies on changes in population size, although see,
for example, Brichetti et al. 2008).
Nevertheless, a key study in this area is that of Tait et al. (2005). These authors
tracked changes in the species richness of native and introduced plants,
mammals, birds, reptiles and amphibians in Adelaide, Australia, from 1836
to 2002 (Adelaide is currently a city with over one million people). Some results
were predictable, while others were surprising. Native species richness
declined in most groups, but only marginally for reptiles (3.6% of 56 species)
and not at all for amphibians. Native mammals suffered the most, losing 50%
of species (n ¼ 40; a similar result was reported by van der Ree and McCarthy
(2005) for mammal species loss in Melbourne since European settlement).
Interestingly, total species richness remained practically the same for birds
and reptiles. That is, while 21 native bird species were lost, 20 introduced
species established themselves in the city. Similarly, two native reptiles were
replaced by two introduced species. The same was not true for mammals (a loss
of 20 species, but a gain of only 9), while there was an overall increase of 46% in
the total number of plant species.
Other than for mammals, these patterns do not represent the devastating
loss of species that is normally associated with urban development. However, it
is important to note that Tait et al. (2005) included the entire metropolitan area
in their study, and this incorporates patches of native and other vegetation.
Moreover, species richness is not necessarily the best measure of human impact
on nature since many populations can be lost, with dire implications for
species persistence and ecosystem functioning, even though a species may
remain extant (Hughes et al. 1997; Luck et al. 2003).
Puth and Burns (2009) also reviewed studies that examined changes in
species richness over time for the metropolitan region of New York. They found
that of the 26 studies documenting temporal change, 65% recorded declines in
species richness (increasing to 77% when only trends in native species were
considered). Only six studies found that species richness increased over time,
and most of these examined recolonisation after disturbance. Decreasing
species richness was recorded across a variety of taxonomic groups including
plants, fish, amphibians, reptiles and birds. Other studies of temporal trends in
New York flora concur with these findings, showing that the area has lost over
40% of its native plant species in the past century or so (Robinson et al. 1994;
DeCandido et al. 2004). These latter studies also found that exotic plant species
richness increased over the same time period.
A decline in native plants and corresponding increase in introduced species
over time, sometimes leading to a net gain in plant species richness, is
consistent with studies conducted over broader areas (e.g. McKinney 2002;
104 G. W. LUCK AND L. T. SMALLBONE

Pyšek et al. 2005), although site-specific results may vary from this trend
(e.g. Chocholouskova & Pyšek 2003). Parody et al. (2001) reported no change in
bird species richness in moderately populated areas over a 50-year period in
Michigan, while Jones and Wieneke (2000) recorded the same result for a single
suburban bird community studied over 16 years in Townsville, Australia. Yet
some studies report a substantial decline in bird species richness over time for
parks surrounded by urban development (e.g. Diamond et al. 1987; Corlett
1988; Recher & Serventy 1991), although it is unclear how much this species
loss is a factor of park size and isolation (or habitat changes) and how much can
be attributed to increasing urbanisation.
Lane et al. (2006) estimated that, since 1819, between 33% and 72% of bat
species have been lost from Singapore, an island that has been substantially
developed (50% ‘built-up’). The upper bound is based on a species assemblage
inferred from a neighbouring mainland area and may be an overestimate. The
authors make the important point that many species are probably lost during
initial habitat clearance prior to extensive urban development and prior to the
advent of comprehensive and systematic species monitoring. Therefore, it is
likely that the species richness of original assemblages used as baselines for
comparing historical changes is almost always underestimated, and this is
problematic for detailing changes over time with increasing urbanisation.
Matching spatial variation in settlement or neighbourhood age with species
richness can also allude to likely temporal changes. For example, broad-scale
studies by McKinney (2001, 2002) showed positive correlations between intro-
duced plant richness or net gain in the number of plant species (i.e. introduc-
tions minus local extinctions) and HPD and time since settlement in the United
States. The same was not true for fish species analysed in the same studies.
Other research found that bird species richness was higher in older suburbs,
probably reflecting the development of urban vegetation over time (e.g. Jones
1981; Munyenyembe et al. 1989). This probably occurs when vegetation is
cleared at the time of settlement or settlement develops on previously cleared
land (e.g. rural areas) and the establishment of domestic gardens increases
vegetation cover and species diversity.
In sum, changes over time appear to follow these general trends (Figure 5.6).
For plants, early human settlers clear native vegetation, but introduce many
new species (for agricultural or domestic purposes) such that the number of
introductions outpaces the number of local extinctions leading to a net gain in
richness over time. The number of species introductions is likely to decline
over time and the same is probably true for local extinctions as most urban-
sensitive species have already been lost from long-established settlements. This
should lead to a plateau in total species richness (although the number of
exotic species will probably continue to increase slowly over time, especially
for plants). A similar plateau would be reached for animal (i.e. vertebrate)
 SPECIES DIVERSITY AND URBANISATION 105

10 000
Mammals Plants
Birds Introduced species
Species richness (log scale)

1000

100

10
Increasing urbanisation over time

Figure 5.6 General patterns in total species richness and the richness of native
and introduced species with increasing urbanisation over time. From data in
Tait et al. (2005).

species richness, but possibly through different processes. These could be

a replacement of native species with a similar number of introduced species
(e.g. birds), no change in native species (e.g. reptiles) or a substantial decline in
natives resulting in a new, and much lower asymptote in total species richness
(e.g. mammals).
The trends in Figure 5.6 are, of course, simplifications and ignore the fact
that rapid gains and losses in species richness can occur over short periods
(Tait et al. 2005). More importantly, native species richness almost always
declines with increasing urbanisation and highly developed locations have
substantially different species assemblages from those that existed prior to
development. Spatiotemporal patterns in species diversity in urban landscapes
result from a myriad of complex, interacting processes occurring over various
spatial and temporal scales (Kent et al. 1999).

Drivers
The drivers of patterns in urban diversity vary across spatial scales (Figure 5.7).
What drives the broad-scale congruence between species richness and HPD
across regions is subject to some speculation, but little comprehensive assess-
ment. People and other species may co-occur either because both are respond-
ing to underlying abiotic or biotic conditions (e.g. climate) or human landscape
modification leads to increased species richness near human settlements (e.g.
through creation of more heterogeneous landscapes). Moreover, humans are
106 G. W. LUCK AND L. T. SMALLBONE

(a) (b) (c)

0 25 50 Kilometres 0 5 10 Kilometres 0 1 2 Kilometres

Key to maps a & b Key to map c

Main roads Dwelling density/ha
Metropolitan area 0–11
Census districts 12–19
Tree cover 20–28
Tree cover

Figure 5.7 Summary of drivers across scales. (a) Across broad extents, using large grain
sizes (e.g. 1
), human population density and species richness positively covary, probably
owing to common drivers such as available energy and primary productivity. This
relationship is moderated by human land-use policy (e.g. area of conservation land near
human settlements). (b) Within a region, rural–urban gradients often show peaks in
species richness in fringe suburbs of metropolitan areas (intermediate disturbance)
owing to greater vegetation cover and landscape heterogeneity in these locations
compared with rural areas and city centres. (c) Within urban centres, the socioeconomic
characteristics of neighbourhoods are linked to vegetation cover and diversity, and this in
turn affects faunal assemblages. Neighbourhoods characterised by relatively low housing
density, high-income and high education levels support a greater diversity of species.

often responsible for deliberately introducing or promoting colonisation of

exotic species in urban environments. This is especially true for plants, but
these introductions do not explain the positive correlation between HPD
and species richness across regions, because this correlation still exists when
examining only native species (Luck 2007a).
A useful starting point is to assess whether well-established drivers of
biogeographic gradients in species richness also explain patterns in human
distribution. Drivers in common may include potential energy and/or primary
productivity, habitat heterogeneity and evolutionary time/long-term climatic
stability (Rahbek & Graves 2001; Willig et al. 2003). Potential energy and
primary productivity are positively correlated with HPD and species richness
in some locations and offer possible explanations for their co-occurrence across
regions (Evans & Gaston 2005; Luck 2007b). Fjeldså and Rahbek (1998) sug-
gested that historical climatic stability may have led to the spatial congruence
between people and species richness.
 SPECIES DIVERSITY AND URBANISATION 107

Alternatively, human land-use policy may be solely responsible for the

regional co-occurrence of human settlements and species richness through
anthropogenic landscape heterogeneity (Fairbanks 2004), increased productiv-
ity (Hugo & van Rensburg 2008) or conservation policy (Gaston 2005). The first
two explanations suggest that humans increase heterogeneity (e.g. through
establishing a diversity of land-uses) or productivity (e.g. through irrigated
agriculture) near human settlements, leading to greater species richness. The
third predicts that HPD and species richness are correlated because both
positively covary with conservation activities, or that conservation policy
mediates the relationship between people and richness. The former is highly
unlikely given reports of strong negative correlations between HPD and
the size of conservation reserves. This relationship is likely to increase the
probability of species extinction inside reserves (Brashares et al. 2001; Harcourt
et al. 2001; Luck 2007b).
Hugo and van Rensburg (2008) tested each of the human land-use hypotheses
for HPD and bird species correlations across South Africa at a quarter degree
resolution. They found support for the anthropogenic heterogeneity and prod-
uctivity hypotheses, and concluded that the co-occurrence of humans and
other species is strongest in regions with greater cover of conservation land--
use. Extending this work, we tested the explanatory power of a range of
hypotheses for biotic drivers (primary productivity and inherent habitat
heterogeneity) and for human land-use drivers (anthropogenic landscape
heterogeneity, increased productivity and conservation policy) of HPD and bird
species richness co-occurrence across Australia at a 1
resolution (Luck et al. in
press). We found that potential energy (annual rainfall) and net primary
productivity appeared to be the key drivers of the people–richness correlation
after controlling for spatial autocorrelation and biased sampling effort. Inher-
ent habitat heterogeneity was more strongly, positively related to species
richness in low-energy than high-energy regions, contrary to previous findings
(Kerr & Packer 1997). Similar to Hugo and van Rensburg (2008), we found that
conservation policy appears to mediate the relationship between humans and
other species, whereby the HPD–diversity correlation was strongest in areas
with a greater percentage cover of conservation land-use and declined steadily
with a reduction in conservation land-use. This means that in regions where
conservation land-use is low, species are restricted to a few small conservation
reserves that are mostly distant from human settlements. In contrast to Hugo
and van Rensburg (2008), we found little support for the anthropogenic or
increased productivity hypotheses.
Another potential driver of the HPD–species richness correlation that has not
received adequate attention is biased sampling effort. This is particularly
problematic when species lists are derived from atlas data collected by volun-
teer observers. Sampling effort has been shown to be greatest in regions of high
108 G. W. LUCK AND L. T. SMALLBONE

HPD using such data (Luck et al. 2004), and this may lead to more species being
recorded in these regions, confounding relationships with other potential
drivers. However, recent studies have now demonstrated that the HPD–richness
correlation still exists after controlling for biased sampling effort (e.g. Evans
et al. 2007; Pautasso & McKinney 2007).
At smaller scales, the correlation between species richness and urbanisation
is generally negative. Therefore, the examination of potential drivers is usually
focused on the question of why more species occur in some urban areas
than others. Landscape variables change considerably when moving from
highly urbanised centres to peri-urban and rural areas. Vegetation patch size
and density, land-use category and fragmentation show varying patterns
across degrees of urbanisation (Luck & Wu 2002; Hahs & McDonnell 2006;
Weng 2007). As urbanisation intensifies, patch size generally becomes smaller
and fragmentation increases (Luck & Wu 2002; Smith & Wachob 2006;
Weng 2007).
Across rural–urban gradients, faunal species richness may peak at intermedi-
ate locations (e.g. outer suburbs) owing to greater vegetation cover and the
diversity of land-uses in these locations compared with rural areas (which are
largely cleared) and urban centres (Luck & Wu 2002; Weng 2007). Moreover,
intermediate locations are areas where species that respond positively to
urbanisation may coexist with those more dependent on native vegetation.
Landscape/habitat heterogeneity (i.e. the variety of landscape elements) plays
an important role in driving faunal species richness patterns (e.g. Kerr & Packer
1997; Atauri & de Lucio 2001; Tratalos et al. 2007), although promotion of
heterogeneity per se is unwarranted without some understanding of the key
landscape elements required by particular taxonomic groups.
At various scales, bird species have been shown to respond positively to
increasing vegetation cover, composition and structure (Mills et al. 1989;
Munyenyembe et al. 1989; Pidgeon et al. 2007; Bino et al. 2008). Vegetation cover
is often negatively correlated with housing density and the cover of impervious
surfaces (e.g. roads and footpaths). The availability of anthropogenic food may
also have a strong influence on bird assemblages in urban environments
(e.g. Jokimäki & Suhonen 1998; Fuller et al. 2008), although this may increase
the abundance of a few readily adaptable species rather than species richness
per se. At the scale of individual streets and gardens, plant species composition
has an important influence on bird assemblages. For example, Young et al.
(2007) found that the presence of flowering native trees was more important
in driving nectarivore assemblages in urban streets in Adelaide than the
area surrounding the tree.
In sum, faunal species will respond to the availability of critical resources
(e.g. for feeding and breeding) across urban landscapes, and use of these
resources will be dictated by their capacity to move around the landscape,
 SPECIES DIVERSITY AND URBANISATION 109

interspecific interactions (e.g. competition) and species’ ecological traits.

Arguably, a more interesting question then is what dictates the distribution
of resources. Vegetation is an important resource for many faunal species and
has been shown to vary greatly across urban areas. In contrast to fauna, plant
species richness (natives and exotics) tends to be higher in urban than rural
areas and increases over time with increasing urbanisation (although the
relationship is likely to be asymptotic). This largely reflects the introduction
of new plant species for agricultural or domestic purposes (Pyšek 1998).
Moreover, humans are able to maintain very small plant populations (e.g. one
to two individuals) and ensure the persistence of non-endemic species through
watering, fertilisation and other maintenance activities, and this leads to an
‘artificially’ high diversity (i.e. many of these populations or species would
become ‘extinct’ without human intervention). Also, in some regions,
home gardeners have access to a huge species pool through local nurseries
(Thompson et al. 2003).
While underlying heterogeneity (e.g. topographic or geological) may have an
influence on plant species diversity in urban areas (Pyšek 1998; Kühn et al.
2004), an increasing number of studies demonstrate a relationship between the
socioeconomic profile of neighbourhood residents or householder behaviour
and vegetation cover and plant species diversity (Hope et al. 2003; Grove et al.
2006). These studies often show that income or some other measure of social
‘status’ (e.g. education) is positively correlated with species richness or vegeta-
tion cover. Teasing apart the causal direction of this relationship will take some
detailed interdisciplinary work. For example, do people with high-incomes
prefer to live in more vegetated suburbs (and have the financial capacity to
do so) or do they increase vegetation cover through their behaviour? Some of
our own work has shown that level of education may be a better predictor of
vegetation cover than income (although the two are often correlated), whereby
cover is positively correlated with the proportion of residents with a tertiary
education (Luck et al. 2009).
If resident behaviour is important in determining levels of vegetation cover,
then improving social status is a potential mechanism for increasing
vegetation and subsequently plant and fauna richness in neighbourhoods. Of
course, not everyone can be rich or highly educated (although income and
education levels can always be improved), and local government planning is
still key to improving the nature of urban environments. Moreover, more easily
managed factors like housing density also have strong relationships with
vegetation cover and species richness (e.g. Hope et al. 2003; Tratalos et al.
2007; Luck et al. 2009), and a lot more research is required to determine how
interactions between landscape planning, socioeconomic status, resident
behaviour and species’ ecological traits influence flora and fauna in human
settlements.
110 G. W. LUCK AND L. T. SMALLBONE

Implications and future research

There are two primary implications of the broad-scale congruence between
people and species richness. The first is the substantial conservation challenges
raised by this congruence given the negative impacts of human activities on
nature. Alleviating spatial conflict between people and biodiversity has been
explored in a number of studies through conservation planning strategies
(e.g. Balmford et al. 2001; Araújo et al. 2002; Luck et al. 2004; O’Dea et al. 2006).
These studies suggest that opportunities still exist for promoting conservation
in the midst of human development, but these opportunities are more
restricted when conservation objectives become more demanding (e.g. protect-
ing 20% of a species range rather than 10%).
The second implication is that there is substantial opportunity for people to
experience species-rich locations near where they live if conservation close to
human settlements can be improved (Schwartz et al. 2006; Lawson et al. 2008).
Maintaining high diversity can be important for the production of various
ecosystem services, many of which are used locally (Dı́az et al. 2006). Future
research should continue to focus on key drivers of spatial congruence and
the development of intra- and inter-regional management strategies that
ensure species persistence through appropriate settlement planning. More
work is needed to assess the ecological, social and environmental (e.g. energy
use) trade-offs between concentrated, high-density living in increasingly large
cities versus more dispersed, smaller regional centres. We know little, for
example, of how rural–urban gradients vary with city size or if neighbour-
hoods in small towns are characterised by the same spatial variation in species
richness as those in large cities. At a smaller scale, clustered development,
although promoted as a tool to reduce the impacts of human settlement on
nature, offered no greater conservation benefits than dispersed housing in a
study in Boulder County, Colorado (Lenth et al. 2006).
Within human settlements, people living on the rural–urban fringe, in low-
density housing or ‘wealthier’ neighbourhoods have the greatest opportunities
to conserve and interact with a diverse nature. However, the stark reality for
most people, particularly in large cities, is that their neighbourhood is likely to
be species-poor and dominated by a handful of abundant urban exploiters
(Turner et al. 2004). A reduction in the opportunities for people to interact with
nature can impact negatively on human health and psychological wellbeing
(Harlan et al. 2006; Tzoulas et al. 2007), and there is some evidence to suggest
that species diversity per se is important in promoting the latter (Fuller et al.
2007). For an increasing number of people, their entire experience with nature
may be dominated by interactions in urban environments. Interactions with
nature can influence people’s perception of and desire to protect nature, and
may link directly with environmental activism (Dunn et al. 2006). Since most of
 SPECIES DIVERSITY AND URBANISATION 111

the Earth’s population now live in urban areas (United Nations 2008), and these
people will have a substantial influence on conservation policy through voter
numbers, improving human–nature interactions in human settlements is not
only important for meeting local conservation objectives, but could have major
implications for global conservation (Dunn et al. 2006).
In the future, researchers must build on the ground-breaking interdisciplin-
ary studies that link ecological, socioeconomic and environmental data
(e.g. Hope et al. 2003). Only through integrated research can we begin to
understand the patterns of diversity that occur in urban landscapes and tease
apart the complex, interconnected drivers of these patterns.

Acknowledgements
This work was funded by an Australian Research Council Discovery Grant
(DP0770261) to G.L. Thanks to S. McDonald (Spatial Data Analysis Network,
Charles Sturt University) for assistance with remote sensing and Geographical
Information System analyses. K. J. Gaston and two anonymous reviewers
provided thoughtful comments on a draft manuscript.

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CHAPTER SIX

Urbanisation and alien invasion

S T E F A N K L O T Z A N D I N G O L F K Ü H N

Introduction
Urbanisation is a rapidly developing process of global change. The world’s
human population in 1900 was around 1.6 billion, of which 13% lived in cities
(UN-HABITAT 2003, 2006). Within 100 years the world population increased to
6.5 billion and the urban population increased to 50% (UN-HABITAT 2006).
This increase will continue rapidly. In Europe and other highly industrialised
regions the percentage of urban population is already much higher (>70%).
This urban population growth is strongly connected with an increase in the
size and intensity of urban land-use. As a consequence of land-use change, more
and more plant and animal habitats have been lost. Urbanisation is a major
driver of plant and animal extinction on a regional scale (Fuller & Gaston
2009). The species most affected have been those of wet and very pristine
habitats, as well as those of extensive agricultural lands like meadow and
pasture, all of which have become rarer in heavily used landscapes such as
those of Germany (Klotz 1989).
The concentration of the human population has resulted in an increase in
traffic and transport of food, raw materials etc., and growing needs for open
space recreation activities such as gardening and walking. Additionally, land-
scaping and gardening, as well as vegetable and ornamental plant production,
are concentrated in urbanised regions. Therefore we have two of the general
prerequisites for species introductions into cities. First, there is the concen-
tration of intentional introductions of plant and animal species. Second,
resulting from the increasing traffic and transportation of different materials
(Hulme et al. 2008), urban areas are hotspots of unintentional introductions.
Cities are hence not only rich in native plant species (Kühn et al. 2004a) but
even more so in alien plant species (Pyšek 1998; Kühn et al. 2004a; for a
general overview of species richness in urban areas see Chapter 5). This can
easily be seen, for example, in Germany (Figure 6.1). Hotspots of neophyte

Urban Ecology, ed. Kevin J. Gaston. Published by Cambridge University Press.

# British Ecological Society 2010.
 URBANISATION AND ALIEN INVASION 121

3
1

6
4 7
Figure 6.1 Hotspots of neophyte
species (alien plant species introduced
after the discovery of the Americas)
5
in Germany (percentage of total
plant species richness). 1: Berlin;
2: Hamburg; 3: Bremen and the
< Weser valley; 4: Ruhr area;
= 2.5%
>2.5–5% 5: Rhine–Main (greater Frankfurt)
>5–9%
>9–12% area; 6: Leipzig–Halle; 7: Saxonian
>12%
industrial region.

distribution (i.e. alien plant species introduced after the discovery of the
Americas; Pyšek et al. 2004) are the major urban areas of Berlin, Hamburg,
the Ruhr and Rhine–Main areas, Leipzig–Halle and the Saxonian industrial
region. Another striking feature is the richness in neophytes of the Weser
river valley. This river is characterised by high salinity due to potash mining
in the upper reaches.
The urban environment has a small grain size and is very patchy, being
made up of very different land-use structures and habitats such as residential
areas with gardens, lawns and shrubby vegetation, as well as parks and
cemeteries and areas of industrial production and traffic (roads, railways,
ports, airports etc.). Former mining lands as well as solid waste dumps and
landfills may also be parts of urban conglomerations. Additionally, remnants
of pristine habitats as well as forests, woodlands and agricultural habitats
remain in urban areas (see Gilbert 1989). However, owing to the small spatial
sizes of the different habitats, all biological communities are influenced by
urbanisation in general. Totally new urban habitats as well as the degraded
pristine remnants are target habitats for new alien species arriving in cities
(Wania et al. 2006).
Thus far, most studies of alien species in urban areas have been carried out in
Europe. Far fewer are from other parts of the world, and these have often
mainly been descriptive or have covered single species. Most studies have
122 S. KLOTZ AND I. KÜHN

analysed plant species as focal organisms of urbanisation. General studies

using animal groups are much rarer and often restricted to selected habitats
within the city. Birds are the best-studied group of animals (e.g. Fuller et al.
2009; Loss et al. 2009). Knapp et al. (2008a) compared different species groups in
selected protected habitats within a city and in the surrounding agricultural
landscape of Halle, Germany. They found that the total species number for
carabid beetles and butterflies was higher in the rural protected areas than in
the urban protected areas. Large-scale and multi-species comparisons of alien
animals between habitats of different degrees of urbanisation are, however,
mostly lacking. In this chapter we will focus on plant species in particular.

Ecological characteristics of urbanisation

and their importance for invasion
Urbanised areas are different from other landscapes in many environmental
characteristics (Chapter 3; Sukopp 1998). First, micro-climates and even
regional climates are clearly distinct from those of surrounding landscapes.
Urban climate studies in cities of the temperate zone have shown that global
radiation is decreased by air pollution, and sunshine duration decreased
because of increased development of clouds. The greenhouse effect leads to
an increase in temperatures (Landsberg 1981; Oke 1982). Frost events are rarer
than or not as strong as in the surrounding landscapes. At higher latitudes,
cities can thus be seen as outposts of warmer and milder climates. Wind speed
and storm events are diminished in cities with consequences for biotic pro-
cesses such as species dispersal and pollination. The high coverage of non-
vegetated surfaces (including areas paved by asphalt and concrete) increases
the water runoff after rainfall and decreases evapotranspiration. The main
consequence is a reduction in air humidity. Precipitation is increased in cities
owing to the heat island effect and the higher cloud density. Snow fall is
reduced by the generally higher temperatures.
Besides climatic changes, soils are heavily affected by urban land-use, includ-
ing by construction activities, pollution and sedimentation of dusts, as well as
by frequent disturbances. Typical processes in urban soils are the destruction of
the native soil layer structure by soil intermixing, covering of surface soil
layers, and the addition of natural (sand, gravel etc.) and/or artificial substrates
(asphalt, demolished bricks etc.). The covering by impermeable surfaces results
in fossilisation of soils over a longer period. Gardening activities lead to
nutrient-rich hortisols. Pollution increases the nutrient content and the pH
level of soils in general. The use of de-icing salt in winter leads to higher salt
concentrations locally along roads and on dumps.
The hydrology of urban ecosystems mainly depends on the canalisation of
the original waters as well as on the water treatment systems. Generally, wet
sites are ameliorated and wet habitats become rarer. Conversely, modern
 URBANISATION AND ALIEN INVASION 123

gardens often include artificial waters such as small ornamental ponds with
swamp and mire plants. Vegetation strips along waterbodies are often heavily
disturbed and open enough for the immigration of new species.

Typical urban species: natives and aliens

Despite the heterogeneity of urban areas, specific urban species can be found.
Wittig et al. (1985), using grid mapping techniques, identified species which
are more frequent or less frequent in urban areas. They thus distinguished
urbanophilic, urbanoneutral and urbanphobic plants. Wittig (2002) presented
a list of most of the urbanophilic species in Germany. Most of the species on
that list are alien. That means that in the native species pool, plants which are
well adapted to urban conditions are rare. Urbanophilic aliens are concen-
trated in the inner part of the cities where the climatic differences and the
general environmental changes from non-urban regions are greatest. Using
gradient analyses, Kunick (1974) found a clear zonation in the relative
numbers of alien species, decreasing from the centre of urban agglomerations
to the margin of Berlin, Germany. This pattern highlights that inner urban
habitats are targets for alien species (mostly thermophilous species and those
adapted to a high frequency of disturbances). These thermophilous alien
species are mostly restricted to urban areas, and it is reported that they
invade the countryside only at lower latitudes. One such example is the
Tree of Heaven Ailanthus altissima (Mill.) Swingle in several European cities
(e.g. London, Paris, Berlin, Halle, Leipzig). In central and northern German
cities, the species is restricted to the urban heat islands. In southern Germany
and in the neighbouring southern countries, the tree can invade the open
countryside (Gutte et al. 1987).
An analysis of the levels of invasion of urban and non-urban habitats across
Europe (Pyšek et al. 2009) shows that intentionally and unintentionally intro-
duced species occur in highest numbers in urbanised habitats (Table 6.1).
The process of acclimatisation and further spread starts here. Urban areas
can thus be the sources of further immigration of plants and animals.
Former intentionally introduced and widely used plants colonise new habi-
tats and new landscapes. Common chicory Cichorium intybus, formerly widely
sown in Central Europe, was used as a surrogate for coffee, but lost import-
ance because of the increase in coffee production and the decrease in coffee
prices. Now, this species is only rarely planted but is present to some degree
in many city meadows and lawns. A similar example in subtropical and
tropical cities worldwide is tree tobacco Nicotiana glauca, currently one of the
most important invaders. It was formerly used as a surrogate for tobacco.
Another example is Canada golden-rod Solidago canadensis. This species was
mainly introduced to Central Europe as an ornamental plant and widely
grown in gardens. It is well adapted to grow on fallow land, urban
124 S. KLOTZ AND I. KÜHN

Table 6.1 Levels of invasion of European EUNIS (European Nature Information

System) habitats by alien plant species in Europe and those to Europe
(after Pyšek et al. 2009).

No. of aliens in No. of aliens to

Category Europe Europe

Number of species classified 2122 1059

% classified of the total 56.6 57.7
A. Marine habitats 12 7
D. Mires, bogs & fens 220 118
B. Coastal habitats 343 170
C. Inland surface waters 444 260
F. Heathland & scrub 462 206
H. Inland sparsely vegetated
habitats 497 211
G. Woodland & forests 668 310
E. Grasslands 793 276
I. Arable land, gardens & parks 1240 533
J. Industrial habitats 1360 658

Note: Aliens in Europe are species which are native to some regions in Europe and alien to
others; aliens to Europe are species with the area of origin outside Europe. The letters A to J
are those used in the European Nature Information System (EUNIS) classification.

brownfields and along railways. In late summer and autumn the species
flowers intensively. By escaping and flowering in ruderal habitats the
species lost its former importance as an ornamental.
In analyses of the flora of the city of Halle in central Germany and the rural
surroundings, Wania et al. (2006) identified on a small scale (grid cells with 250-m
resolution) those species which are typical for the city and for the rural region.
Of 86 species identified as being significantly more frequent in cities than the
surroundings, 27 (31.4%) were alien and 16 (18.6%) were neophytes. Of seven
species characteristic of the more rural surroundings, only one (14.3%) was
alien and none were neophytic.
To survive, grow and reproduce in particular urban habitats, plant and
animal species need specific adaptations to these environments. Urban plant
species in Germany, for example, more often are wind-pollinated, have sclero-
morphic leaves and are dispersed by animals, and less often are insect-pollinated,
have hygromorphic leaves and are dispersed by wind (Knapp et al. 2008c). In the
Czech Republic, Lososová et al. (2006) found that, compared with weed vegeta-
tion of arable land, in the ruderal vegetation of settlements species were
often stronger competitors, wind-pollinated, flowered in mid-summer, repro-
duced both by seeds and vegetatively, dispersed by wind or humans, and had
 URBANISATION AND ALIEN INVASION 125

high demands for light and nutrients and more continental distribution
ranges. Hence they are often adapted to grow in warm, dry and nutrient-rich
habitats. Species which have several reproductive strategies and are adapted to
human dispersal therefore have an advantage in urban regions. Interestingly,
although in general the phylogenetic diversity of species in urban areas is
reduced (Knapp et al. 2008b), species sharing traits which are better adapted
to urban environments showed a higher phylogenetic diversity than those
better adapted to rural environments. Many rare species (alien as well as
native), especially those preferring cool or acidic habitats, might already have
disappeared from urban areas (Knapp et al. 2009).
Evolutionary adaptability might play some role in urban areas, but is not
well studied. New ‘anthropogenic species’, such as hybrids between native and
alien species, or between alien species from different biogeographical regions
and escaped cultivars of ornamental plants, could play an important role.
A first nationwide list of such plant species is available within the BIOLFLOR
database of biological and ecological traits of the flora of Germany (Kühn &
Klotz 2002; Kühn et al. 2004b).

Immigration pathways and urbanisation

There are several different pathways by which alien species arrive in new
regions which might eventually become their non-native range. Understand-
ing the processes involved is crucial for successful management but also for
improved projections of future invasions. Cities here are just a special, albeit
probably a most important, system. As stated previously, new species may be
introduced intentionally or unintentionally. This, however, is no clear dichot-
omy but rather represents the two extremes in a gradual change. Hulme et al.
(2008) present a simplified framework to categorise six major pathways of
initial introduction, namely release, escape, contaminant, stowaway, corridor
and unaided. Releases are intentional introductions of a commodity, such as
biocontrol agents, game animals or plants for erosion control. Escapes are
also intentionally introduced as a commodity but escaped unintentionally,
such as feral crops and livestock, pets, garden plants and live baits.
Contaminants are unintentionally introduced with a specific (intentionally
introduced) commodity, such as parasites, pests and commensals of traded
plants and animals. Stowaways are unintentionally introduced while being
attached to or within a transport vector. Typical examples are hull fouling,
and ballast water/soil/sediment organisms. Corridors are defined as human
infrastructures linking previously unconnected regions. This can result in
unintentional introductions of, for example, marine organisms moving
through the Suez Canal or Ponto-Caspian organisms migrating along rivers
and newly established canals to the Baltic Sea. Unaided introduction is of
course also unintentional through natural dispersal of species from a region
126 S. KLOTZ AND I. KÜHN

Figure 6.2 Relative contributions of pathways of introduction shown for naturalised

aliens to Europe, i.e. plant species with the area of origin outside Europe (modified from
Pyšek et al. 2009; based on 1983 naturalised aliens; from data in Lambdon et al. 2008).

where they are already alien across political borders. Most of the European
alien plant species were intentionally introduced into this region and were
released deliberately or escaped unintentionally (Figure 6.2; Pyšek et al. 2009).
Of these, the majority were introduced for ornamental, agricultural, horticul-
tural or forestry reasons or as an amenity. Contaminants also play a consider-
able role but fewer species arrived unintentionally attached to or within a
transport vector.
The relative importance of invasion pathways specifically to cities is not as
well analysed. Still, one can assume that the proportion of species introduced
for ornamental or horticultural reasons is even higher than for the complete
European species set and makes up a high proportion of those species intro-
duced into Europe. Several of the ornamental species are typical for urban
parks, gardens or even historic garden cultures (Kowarik 2003). Nonetheless,
agricultural seeds contributed considerably to seed rain along road tunnels in
Berlin (von der Lippe & Kowarik 2007a, 2007b).

Cities as sources of invasions

There is considerable concern that cities, being so rich in alien species, can act
as foci for the spread of alien species into the wider environment (e.g. Sullivan
et al. 2005; Houlahan et al. 2006). Road verges are known to be rich in non-
native plant species (Parendes & Jones 2000; Gelbard & Belnap 2003). Alien
species such as the South African ragwort Senecio inaequidens DC. in Europe
 URBANISATION AND ALIEN INVASION 127

(Ernst 1998; Heger & Böhmer 2005) or common milkweed Asclepias syriaca L.
and purple loosestrife Lythrum salicaria L. in North America (Wilcox 1989;
Wyatt 1996) are reported to spread along roadsides. Therefore, several studies
have explored to what extent, for example, traffic and traffic infrastructure in
cities might be able to facilitate biological invasions from cities into the
surroundings. Von der Lippe and Kowarik (2007b) attempted to describe the
spatial and quantitative effectiveness of traffic as a dispersal vector by address-
ing the role of vehicles in long-distance dispersal and by quantifying seed
deposition at roadsides in Berlin. They analysed seed rain on the roadsides of
five different lanes within three tunnels along a single urban motorway. They
found that half of the plant species were alien, with 33.8% of all species in
their seed traps being neophytes. Non-native species were therefore slightly
overrepresented, compared with the local species pool. Approximately one-
third of the species found in the tunnel traps were not present in any of the
tunnel surroundings, indicating long-distance dispersal, with alien species
(on average 38.5%) showing significantly higher proportions of such dispersal
than native species (4.1%). In a second analysis, von der Lippe and Kowarik
(2008) differentiated between seed deposition along the inbound and out-
bound tunnel lanes. Native and non-native species richness of the tunnel
samples was significantly higher along lanes leading out of the city compared
to the inbound lanes. The same pattern was true for seed deposition, but one
order of magnitude higher. The export of seeds from non-native species out
of the city by vehicles was not only higher in numbers than the import, it
was also higher in relation to their overall proportion in the flora. Cities
can therefore act as foci for the spread of alien species into the wider
environment.
Brunzel et al. (2009) found that in a region north of Frankfurt, Germany,
many species of neophytes increased their abundance between a first
survey (1974–81) and a second one (2003). Furthermore, neophytes were
more abundant in urbanised settlements closer to Frankfurt than those
further away. The occurrence of neophytes was better predicted by envi-
ronmental variables, using a multiple regression framework, than was
that of native plant species. Distance to Frankfurt (negative), number of
inhabitants (positive) and connectivity to Frankfurt (assessed as the quality
of the transport infrastructure; positive) were the three most important
significant predictors explaining neophyte plant species richness. In another
analysis of the same dataset, dispersal kernels based on patterns of human
movement between settlements led to a better match with the observed
distribution pattern than a null model simulating pure distance-dependent
dispersal for all species (Niggemann et al. 2009). Interestingly, alien species
seemed to benefit more from human dispersal than native species. Nonethe-
less, not only vehicles and transport infrastructure facilitate species
128 S. KLOTZ AND I. KÜHN

dispersal. Even walkers can contribute considerably to the long-distance

dispersal of plant species by carrying seeds underneath and on their shoes
(Wichmann et al. 2009).
Different patterns have been found for the United Kingdom (Botham et al.
2009). Comparing two mapping schemes (1987–88 versus 2003–04), the authors
reported a strong association of neophytes with urban land cover but no
apparent spread into the wider countryside. Neophytes as a group showed
some reduction in the strength of their association with urban land cover over
time. This was true even for the species for which strong positive urban
associations were identified. However, very few species changed in their indi-
vidual urban association over time. There are several factors which could
potentially mask the evidence of species spreading into the wider countryside:
(i) urban sprawl may outweigh species dispersal; (ii) the study is based on
presence/absence only and not on abundances; and (iii) the time period of
16 years may not be long enough. Alternatively, contrasting patterns may
genuinely exist, perhaps as a consequence of the situation in Britain being
different in several respects from the one in Central Europe. Unlike many other
studies (e.g. Araújo 2003; Hope et al. 2003; Kühn et al. 2004a; Wania et al. 2006),
Britain did not show an increase in species richness in urban areas compared
with rural ones (Roy et al. 1999).

Alien species and biotic homogenisation in urban regions

The spread of alien species into the wider countryside as well as the introduc-
tion of the same species into cities around the globe may lead to a phenom-
enon called biotic homogenisation (McKinney & Lockwood 1999; Lockwood
et al. 2000). This is often interpreted as formerly distinct biota becoming more
similar by the replacement of rare native species with widely spread alien
species. There are, however, two mechanisms involved, the introduction of
species and the extinction of species, which can lead to increased, decreased
or unchanged similarity among biota (Olden & Poff 2003). Increasing similarity
is generally called homogenisation, decreasing similarity called differenti-
ation. This is not restricted to species but can also occur at a phylogenetic or
functional level (Olden & Poff 2003; Winter et al. 2008).
The process of species introduction can, however, have different effects on
different scales. Especially on small scales and at short distances, differenti-
ation can occur (e.g. Qian et al. 2008; Winter et al. 2008, 2010), whereas on larger
scales and across larger distances homogenisation is frequently found
(e.g. McKinney 2005; Qian & Ricklefs 2006; Winter et al. 2010).
Considering cities, McKinney (2006) discusses the special case of urbanisation
and homogenisation. The expansion of towns and cities promotes the loss of
native species and their replacement by alien species. And not only can the
same plants be found in cities all over the globe (plants: Tree of Heaven,
 URBANISATION AND ALIEN INVASION 129

goldenrain tree Koelreuteria paniculata, common fig Ficus carica; animals: house
sparrow Passer domesticus) but urban habitats also tend to be very similar. In this
paper, McKinney showed that plant communities of state parks and national
wildlands are less similar than those of urban habitats.
For California, Schwartz et al. (2006) found that in densely populated coun-
ties there is a low similarity of plant species that have been extirpated but a
high similarity of noxious weeds. This will lead to homogenisation. For
Germany, the pattern is slightly more complex. Kühn and Klotz (2006) com-
pared 60 urbanised grid cells of the German floristic mapping scheme with
more rural cells, and found that urban areas are more similar to each other in
plant composition than rural areas (i.e. homogenised). Disentangling the con-
tributions of the different groups of species, assemblages of native species and
archaeophytes (i.e. alien plant species introduced before the discovery of the
Americas; Pyšek et al. 2004) are more homogenous in urban areas compared
with rural ones while assemblages of neophytes are more differentiated. Hence,
urbanisation can lead to homogenisation, but this is not caused by neophytes.
Similarly, La Sorte et al. (2008) found for 22 cities across Europe that archae-
ophytes were associated with a higher compositional similarity while neo-
phytes were associated with a lower compositional similarity. The former can
be interpreted as leading to homogenisation, the latter to differentiation.
A comparable pattern was also found in a cross-continental analysis of eight
US cities and seven European cities (La Sorte et al. 2007).
Interestingly, while all plant species (natives and aliens) were more species-
rich in urbanised areas in Germany than in rural areas, neophytes were
disproportionately more frequent (Kühn & Klotz 2006). While high species
richness is usually considered beneficial for biodiversity conservation, this
can often be associated with homogenisation. Of course, the identity of species
is important and whether they are generalists or specialists. Still, this leads to a
conservation challenge: cities are species-rich despite the effect of homogenisa-
tion (McKinney 2006). Appropriate management strategies therefore need to be
developed to maintain a high native diversity but counteract the homogenis-
ing processes.

Summary
Because of their specific environmental conditions as well as a high hetero-
geneity of habitats, urbanised areas support very many plant species. Some
are native species but alien (i.e. non-native) species are especially prevalent.
Typical alien plant species in Central Europe are, for example, Tree of Heaven
or Canadian golden-rod. The former is especially adapted to the warmer
climates of cities, which may anticipate more general climate warming.
Typical urban plant species are often better adapted to dry or nutrient-rich
soils and are more often wind-pollinated than insect-pollinated. Most alien
130 S. KLOTZ AND I. KÜHN

plant species were intentionally introduced, often as ornamentals or for

horticultural reasons; fewer were unintentionally introduced, often as con-
taminants of commodities (seeds, wool, minerals etc.), especially as a result of
transport infrastructures to and within cities. But cities act also as sources of
biological invasions into the wider environment. Here, traffic infrastructure
especially facilitates spread into surrounding landscapes. This may potentially
result in a homogenisation process (i.e. formerly more distinct biotas become
more similar). However, currently urbanisation has mostly been observed to
result in homogenisation (among urban floras) of native species, whereas
alien species show differentiation (i.e. becoming more dissimilar). Therefore,
specific management strategies need to be developed to meet the specific
demands of conserving a high biodiversity with many specialised or rare
species and concurrently preventing homogenisation to preserve distinct
regional species compositions.

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CHAPTER SEVEN

Interactions between people

and nature in urban environments
RICHARD A. FULLER AND KATHERINE N. IRVINE

Since the dawn of human civilisation, people have interacted with nature,
most notably to harness the resources that have fuelled the human enterprise
(Vitousek et al. 1997). The sheer rate and scale of human appropriation of
natural resources has precipitated a biodiversity crisis currently being mani-
fested in rapid rates of species extinctions, extensive transformation of the
structure and function of ecosystems, and rapid alterations to the Earth’s
climate (Vitousek et al. 1986; Pimm & Raven 2000). The biodiversity crisis is a
result of human activity, so the solutions to it will depend largely on human
actions, on understanding and enhancing the way that we all interact with
nature (Collins et al. 2000; Ehrlich 2002). Because most people on the planet live
in towns and cities, the majority of our daily interactions with nature take
place in urban environments, and this has led to a recent upsurge of interest in
the dynamics of these relationships (Bradshaw & Bekoff 2000; Miller & Hobbs
2002; Pyle 2003; Saunders et al. 2006). Despite the manifest impoverishment of
the natural environment in urban areas, or perhaps because of it, many urban
dwellers seek out interaction with nature in some form, for example by visiting
a local green space, or feeding backyard birds. Yet our understanding of these
interactions is nascent, principally because their study requires work across
several disciplinary boundaries (Alberti et al. 2003; Braun 2005). For example,
ecologists often lack the interest or the tools to study people (Collins et al. 2000).
Interactions between people and urban nature are widespread across those
societies in which they have been studied. Small-scale urban agriculture for the
sale of produce in local markets is prevalent across much of the developing
world (Silk 1986), and gardening of private plots for subsistence is common
across many African cities (Obudho & Foeken 1999). Fifty-nine per cent of
respondents to a survey in Guangzhou, China, reported visiting urban parks
often or very often (Jim & Chen 2006a), and 92% of participants in a recent UK
survey reported visiting urban green spaces (GreenSpace 2007). In the USA,

Urban Ecology, ed. Kevin J. Gaston. Published by Cambridge University Press.

# British Ecological Society 2010.
 PEOPLE–NATURE INTERACTIONS IN URBAN ENVIRONMENTS 135

about 68 million people engage in wildlife-watching activity in or within a mile

of their home, spending about US $33 billion in the process (US Department of
the Interior, Fish and Wildlife Service 2006). In parts of Europe, North America
and Australia, between one-fifth and one-third of households provide supple-
mentary food for wild birds (Clergeau et al. 1997; Rollinson et al. 2003; Lepczyk
et al. 2004). Total annual expenditure on outdoor feeding of birds in the UK has
recently been estimated at £200 million (British Trust for Ornithology 2006),
while in the USA $23.2 billion is spent annually on wildlife-watching equip-
ment (US Department of the Interior, Fish and Wildlife Service 2006).
Despite this, there is concern that urbanisation is progressively isolating
people from the natural environment, and that the problem is getting worse
as cities continue to grow in size and density (Turner et al. 2004; Miller 2005;
Louv 2008). Even the term ‘outside’ suggests a degree of separation from nature
(Irvine & Warber 2002a). Modern humans spend most of their lives inside, and
urbanisation exacerbates this problem by adding substantial geographic dis-
tance from natural, or even sometimes any vegetated, landscapes. About one-
third of Hispanic children in the USA are routinely kept indoors because their
neighbourhoods are perceived as dangerous (Dye & Johnson 2007). In short, we
are seeing an ‘extinction of experience’ (Pyle 1978). This is of profound concern,
because interactions between people and nature lead to a variety of measurable
benefits, at both individual and societal levels (Irvine & Warber 2002b; Bird
2004, 2007; Brown & Grant 2005). It is also concerning from a conservation
viewpoint because demand for venues that allow interaction with nature imply
provision of a green space network that will also benefit biodiversity, and
access to nature in turn predicts active engagement (Schultz 2000; Williams &
Cary 2002; Lohr & Pearson-Mims 2005). A simple example illustrates this.
Surveys in the USA have discovered that people living in large cities are less
likely to be birdwatchers than residents of small cities and rural areas
(La Rouche 2003; Sali et al. 2008). The explanation for this is no doubt partly a
practical one – suitable birdwatching areas might be a considerable distance
away in a large city – but there is also the possibility that levels of daily
interaction with nature partly determine engagement in birdwatching in a
kind of positive feedback loop. It is thus perhaps ironic that much of the
philosophy of conservation is built around the idea of separating biodiversity
from people and the threats associated with them, a thought to which we shall
return in closing.
However, the story is more complicated than this. Urbanisation embodies a
shift from an economy based on directly harvesting nature to one based on
industrial and service sectors, a shift correlated with the growth of protective
environmental values at least in the post-war USA (Mertig et al. 2002). This
is manifested in a change from direct utilitarian interactions with nature, to
a mutualistic relationship with ‘fabricated or disrupted’ remnants of nature
136 R. A. FULLER AND K. N. IRVINE

(Bell 1976). While there are obvious benefits of concentrating people into cities,
it is critical that we identify the potential costs to human quality of life, and
how we can manage them.
While urbanisation can be viewed as a manifestation of economic growth
and developmental progress, from a biological viewpoint it is profoundly
disrupting, causing (i) the loss and degradation of natural habitats; (ii) reduc-
tions in ecosystem services such as climate and water regulation; (iii) increased
levels of pollution and disturbance; and (iv) major changes to the structure of
biological assemblages including invasion by non-native species (Chapters 3–6;
for reviews see McKinney 2002, 2005; Chace & Walsh 2006). Although urban
areas are clearly biologically disrupted systems, they can support significant
levels of biodiversity (Fuller et al. 2009) and have some regional conservation
value (Mason 2000; Bland et al. 2004; Vähä-Piikkiö et al. 2004). This is particu-
larly so in developed regions such as Europe and North America, where inten-
sive agriculture has led to substantial population declines across the wider
landscape, and increased the relative importance of urban areas to sustaining
species’ overall abundances (Gregory & Baillie 1998; Mason 2000). For example,
the density of birds in urban Sheffield, UK, is estimated at more than six times
the national average, although most of the abundant species in the city’s
avifauna are also those common across the UK at large (Fuller et al. 2009).
Resource fluxes, geochemical cycles and land cover composition are under
continuous control by humans in urban environments, and thus the pattern
of daily human activity will shape the biodiversity value of towns and cities.
Large-scale patterns in human activity result ultimately from a collection
of individual decisions by urban citizens. For example, the establishment of
a new pond in just 10% of gardens within urban Sheffield (a city in central
UK with a population of c. 500 000) would result in the formation of 17 500
new habitat patches (Gaston et al. 2007a). Widespread engagement in citizen
science can reveal much about the dynamics of urban biodiversity (Green-
wood 2007), and provide alerts to changes in populations, for example of
garden birds (Cannon et al. 2005).
There is enormous variation in the extent of urban green space networks
across 386 cities in Europe (Fuller & Gaston 2009), which leads not only to
variation in the biological quality of those urban environments, but also
to disparities across human society in access to experiences of nature (Barbosa
et al. 2007). Likewise, reaping the human benefits of such interactions with
nature depends on the presence, but also the qualities, of accessible nature in
the local environment. These reciprocal effects lead to a complex interplay
between human activity and biodiversity. Consequently, management for
urban biodiversity should not, and probably cannot, be separated from
programmes to improve human quality of life in urban environments. Provi-
sion of green spaces within urban areas historically focused almost exclusively
 PEOPLE–NATURE INTERACTIONS IN URBAN ENVIRONMENTS 137

on the area provided for each inhabitant (Turner 2006), although attention has
recently switched to an emphasis on green space quality (CABE Space 2004,
2005a). Indeed, emerging evidence of a positive relationship between biodiver-
sity value and environmental benefits to human wellbeing (e.g. Fuller et al.
2007) suggests that management to enhance biodiversity value could also
benefit the human population in a win–win scenario (Irvine et al. 2010).

What are people–nature interactions?

Dictionary definitions of the term ‘interaction’ emphasise its mutual or recip-
rocal quality; two elements engaging and each influencing the other. As such,
we consider interactions between people and nature to be those actions that
result in measurable changes both to people and to nature; in other words,
interactions are two-way relationships (Irvine et al. 2010). This is distinct from
studying, for example, the effects of exposure to nature on people, or human
impacts on ecological systems in isolation. Examples of interactions in urban
environments include visiting urban public green spaces, maintaining a pri-
vate garden and feeding wildlife. In each of these cases, human activities result
in measurable changes to the natural environment (e.g. providing habitat for
plants and animals, enhancing survival of garden birds, disturbing wildlife).
Simultaneously, being in the natural environment, or the very act of engaging
with nature, has measurable impacts on various aspects of human quality of
life (e.g. physical health, psychological wellbeing, economic gain). In all cases,
the outcomes of interactions can be positive or negative. Research on positive
experiences has predominated in the literature thus far, although negative
experiences are doubtless important (Bixler & Floyd 1997).
Interactions occur via the full range of human senses, from the visual
appreciation of a brightly coloured bird to the smell of fragrant flowers, the
taste of wild blackberries or garden-grown tomatoes, the sound of birdsong,
wind in the trees, or intermittent silence, the feel of the wind on one’s skin or
kicking through autumn leaves (Irvine & Warber 2002a). There is growing
interest in the ‘sensescape’ of urban environments that challenges the domin-
ance of visual senses in the research agenda thus far (Rodaway 1994).
Human interactions with nature may usefully be thought of as falling along
a continuum from passive to active (Irvine & Warber 2002b; Brown & Grant
2005). Merely knowing that a garden or urban green space is nearby represents
one of the most passive forms of interaction (Kaplan et al. 1996; Bell et al. 2004),
increasing through watching wildlife documentaries on television, having
pictures of nature on the wall or having a view through a window. More active
forms of interaction progressively include, for example, sitting in or walking
through a local green space, tending a garden, regularly feeding garden birds,
campaigning for local environmental change, volunteering for local conser-
vation management groups or designing a garden around its suitability for
138 R. A. FULLER AND K. N. IRVINE

biodiversity. These interactions can take place in private, in public or within

institutions (e.g. workplaces, prisons, hospitals, schools).
People–nature interactions are many and varied, and each will have particu-
lar dynamics. However, most are poorly studied, and it remains unclear
whether they stem from a single phenomenon, such as an underlying need
for human interaction with the natural world. One such unifying explanation
is biophilia, the idea advanced by Wilson (1984) that humans have a funda-
mental connection with the natural world, and an innate need to be exposed to
its complexity of form and function. Rapid global urbanisation has lent
urgency to discovering the importance of people–nature interactions, because
opportunities for them are declining in both quality and quantity (Pyle 2003;
Miller 2005). Research in this field is particularly critical because (i) much of
the focus thus far has been on interactions in the countryside or wilderness,
and (ii) there is evidence for substantial cultural variation in people–nature
interactions (Teel et al. 2007).
In response to cultural bias in research into people–nature interactions, the
Wildlife Values Globally project is documenting variation in wildlife values
across cultures in the USA, China, Thailand, Estonia, the Netherlands, Mongolia,
Malaysia, Uganda and Kenya (Teel et al. 2007). The results of the project thus far
support the possibility of a consistent shift in wildlife value orientations
towards a mutualistic rather than consumptive or utilitarian view of nature
across all of these diverse cultures. This said, there is ample evidence for
enormous cultural variation in values, attitudes and forms of people–nature
interactions. History suggests that cultural change can proceed rapidly, and so
a clearer understanding of culturally specific attitudes toward nature conser-
vation will help in devising ways to shape environmental attitudes in the
future (Ehrlich 2002). For example, in the USA traditional utilitarian values
of wildlife long held by those consumptively using nature (hunting, fishing)
have given way to a dominance by mutualistic and protective values (wildlife
appreciation), and the degree of this shift is positively correlated with urban-
isation (Manfredo et al. 2003). However, the situation in China appears to be
very different. Perhaps partially driven by lower income levels, materialistic
dimensions of valuing wildlife emerged as dominant in a small interview-based
survey of urban and rural residents (Zinn & Shen 2007).
Other aspects of people–nature interactions are less culturally variable. For
example, evidence for environmental altruism (a negative relationship
between value of self-enhancement and pro-environmental behaviour) was
consistent across respondents in Mexico, Nicaragua, Peru, Spain and the USA
(Schultz & Zelezny 1998). On balance, though, the evidence thus far suggests
that many interactions are culturally specific, and much remains to be done to
document global variation in the motivations for, and the form, prevalence
and outcomes of, interactions between people and urban nature.
 PEOPLE–NATURE INTERACTIONS IN URBAN ENVIRONMENTS 139

Examining three people–nature interactions that are reasonably well under-

stood, namely visiting urban public green spaces, maintaining a private
garden and feeding urban wildlife, we will consider how people–nature
interactions manifest themselves, their prevalence across human society,
motivations for engaging in them and their consequences for both people
and ecosystems. We conclude by drawing out common themes, and suggest-
ing ways in which the research agenda can be moved forward in this
emerging field.

Visiting urban green spaces

Urban green spaces have been the arena for much of the work on the
relationship between people and the natural world (Speirs 2003; Balram &
Dragicevic 2005; Pincetl & Gearin 2005; Jim & Chen 2006b). More than half of
the respondents to a survey in Guangzhou used urban green spaces more than
once per week (Jim & Chen 2006a), and the equivalent figure was 46% in an
English survey (Dunnett et al. 2002), equating to about 33 million people
making over 2.5 billion visits annually to urban green spaces. A visit to an
urban green space might involve no more than indirect exposure to its
natural, semi-natural or managed vegetation while engaging in another activ-
ity such as sport or walking the dog. More intimate interactions might involve
sitting and relaxing, or actively watching and engaging with nature by
birdwatching or feeding the wildlife. Whatever form the interaction might
take, access to urban green spaces is an important aspect of quality of life for
urban dwellers, given that it leads to a wide range of personal and societal
benefits including those to physical and psychological health, urban regener-
ation, economic gain, social cohesion, crime reduction, sense of community
and environmental awareness (Chapter 9; Bird 2004, 2007; Department of
Transport, Local Government and the Regions 2006; Maller et al. 2006). As a
result of this, access to green space is emphasised in numerous policy docu-
ments across the world, and is a key dimension of the sustainable city
(Bengston et al. 2004; Irvine et al. 2010).
While there is more or less complete consensus that green spaces are
beneficial to human quality of life, rather less is known about how the
presence of nature delivers these benefits (Stilgoe 2001). To answer this kind
of question, ecological datasets must be brought directly alongside those from
different disciplines, including, but not limited to, psychology, sociology and
geography. This approach was recently taken in a study of urban green space
users in Sheffield, where the degree of psychological benefit gained by visitors
to 15 urban parks was correlated positively with plant species richness in the
sites, suggesting that the biological complexity of the environment is import-
ant in determining the outcome of interactions between people and nature
(Figure 7.1; Fuller et al. 2007).
140 R. A. FULLER AND K. N. IRVINE

(a)
4.5

4
Reflection

3.5

3
1.8 2 2.2 2.4 2.6 2.8 3
log10 Plant species richness

(b)
4.5

4
Distinct Identity

3.5

2.5
1.8 2 2.2 2.4 2.6 2.8 3
log10 Plant species richness

(c)
4.5

4
Reflection

3.5

3
1 2 3 4 5 6 7
Number of habitats

Figure 7.1 Two measures of psychological wellbeing among visitors to 15 urban

green spaces in Sheffield, UK, increased with the richness of plant species in those
spaces. The two measures are Reflection (the ability to think and gain perspective) and
Distinct Identity (the degree of feeling unique or different through association with a
particular place). Reflection also increased with the number of habitats present in the
green space. Reproduced with permission from Fuller et al. (2007).
 PEOPLE–NATURE INTERACTIONS IN URBAN ENVIRONMENTS 141

This said, the proportion of respondents who mention direct contact with
nature as a reason for visiting green spaces varies markedly, with reported
values ranging from 22% (GreenSpace 2007) to 54% (Chiesura 2004) and 55%
(Irvine et al. 2010). A study in Sheffield that asked green space visitors to give
specific reasons for visiting discovered that explicit mentions of flora and
fauna were rare, but that broader constructions of nature such as fresh air,
being outside, peace and quiet, open space and topography were commonly
mentioned (Irvine et al. 2010). This suggests that specific elements of biodiver-
sity value (e.g. species richness, habitat heterogeneity) may not be directly
perceived as important, yet the combination of these components into a
natural scene is part of the reason why green spaces are used. Visitors to the
Vondelpark in Amsterdam, the Netherlands, indicated that relaxation was
their primary motivation for visiting (Chiesura 2004), yet without further
analysis it is hard to ascertain how much of this relates to the importance of
the natural environment. Thus, while contact with nature might not be cited
explicitly by park visitors, frequent contact with nature via visits to green
spaces features in the lives of many urban dwellers.
Preferences for particular features within urban green spaces often reveal
ambivalence. This is typified by residents of Guangzhou, who give low explicit
value to the presence of nature and wildlife, but state a preference on the one
hand for large, well-vegetated green spaces with lots of mature trees, and on
the other hand for those with a wide range of recreational facilities (Jim &
Chen 2006a). This paradox is also evident in Sheffield, where people avoided
biodiverse sectors within urban green spaces, spending most time in areas of
low-density vegetation and on paved surfaces, yet highly endorsed the presence
of trees when asked about their preferences for park design (Irvine et al. 2010).
In addition, park visitors in the city were able accurately to assess relative
biodiversity levels within urban green spaces, suggesting that urban residents
can be highly attuned to the presence of natural features within urban green
spaces (Fuller et al. 2007). Hearing mechanical sounds is consistently rated
less pleasant than natural sounds both within green spaces and in public
squares (Yang & Kang 2005). Irvine et al. (2009) reported that the sounds most
frequently mentioned by visitors to green spaces in Sheffield were those of
natural origin (e.g. bird song, dogs barking, wind in the trees), and vegetation
buffers are widely used to reduce the prevalence of mechanical sounds (for
review see Bucur 2006).
Human interactions with nature in urban green spaces influence biodiver-
sity both directly and indirectly. First, although many urban green spaces are
provided chiefly for human benefit, they directly enhance the ability of an
urban landscape to support biodiversity. Green spaces provide habitat net-
works for plants and animals, including those of regional and national
conservation priority (Vähä-Piikkiö et al. 2004; Angold et al. 2006; Bryant 2006;
142 R. A. FULLER AND K. N. IRVINE

Mason et al. 2007). Indeed, nature conservation was explicitly considered in UK

green space policy from 1983 onwards, latterly emphasising ecological quality
and connectivity in the green space network (Turner 1992, 2006), concepts
directly applied from landscape ecology. The ways in which green spaces are
managed will also directly affect their biological quality and ability to deliver
ecosystem services such as temperature regulation and flood alleviation
(Cilliers & Bredenkamp 2000; Cornelis & Hermy 2004; Jim & Chen 2006b). For
example, clearing of dead wood and removal of dense vegetation can reduce
habitat quality for birds (Sandström et al. 2006); availability of certain exotic
plants can benefit urban butterflies (Shapiro 2002); and extensive mowing of
grasses reduces invertebrate populations and the richness of native plants
(Helden & Leather 2004; Kirkpatrick 2004).
Second, because interacting with nature in urban green spaces has been
empirically linked to positive attitudes toward nature (Lohr 2007), it is possible
that facilitating such interactions will encourage interest in conservation
issues beyond the arena of the interaction itself (Miller 2005). The intriguing
possibility of this indirect feedback has received much recent attention and has
been mooted as a possible long-term outcome of urban environmental educa-
tion programmes (Hale 1993).
The history of urban green space provision, and thus the social and geo-
graphical distribution of opportunities to interact with nature, varies mark-
edly across the world. The availability of public green space in European cities
was primarily shaped during the period following the Industrial Revolution
when publicly funded parks were provided for general use. These areas were
conceived as places to facilitate both physical and social wellbeing among the
working class by being a resource for fresh air, a place of beauty, a space for
relatively passive recreation such as picnicking, carriage riding or walking, and
an area for social interaction (Conway 2000). These parks were characterised by
formal ornamental horticulture (e.g. ornamental flowerbeds, rose gardens;
Kendle & Forbes 1997). Since the Victorian era, many European urban parks
have predominantly been in the form of recreational grounds consisting of
mown grass areas for a range of more active recreation (Kendle & Forbes 1997),
although chronic underinvestment in the UK’s system of 27 000 urban parks
resulted in the loss and degradation of much infrastructure during recent
decades (CABE Space 2005b).
In the USA, urban green spaces were a response to the development of
large cities, prompted by concerns about access to nature and clean air
(Pincetl & Gearin 2005). In the twentieth century, public park design empha-
sised function over experiences of nature, based on the idea that social
reform depended on the availability of areas for structured play (Draper
1996), and only recently were more naturalistic open spaces developed
alongside urban parks, most notably in suburban areas (Rome 2001). Most
 PEOPLE–NATURE INTERACTIONS IN URBAN ENVIRONMENTS 143

recently, green space has been seen as a tool for social regeneration, particu-
larly in cities where provision is very low (Wolch et al. 2002). For example,
parks cover only about 5% of Los Angeles, USA, falling well short of national
standards (Wolch et al. 2002).

Access to urban green spaces

Visiting green spaces is a popular activity, but access varies markedly among
the human population. Many cities in the developing world are characterised
by low levels of green space provision (Jenks & Burgess 2000). For example, open
space provision totals about six square metres per urban inhabitant of Kuala
Lumpur (Dali 2004), and only one square metre per inhabitant of Bangkok,
Thailand (Fraser 2002). In 2001, urban green space accounted for about one-
third of the surface area of Guangzhou, a city with a population of 3.22 million
people in Guangdong, China; this is one of the highest proportions in China
(Jim & Chen 2006b). There is a strong drive to create a comprehensive urban
green space network in South Africa (South African Cities Network 2006), yet
there are no clear standards for green space provision, nor the policy frame-
work to implement them (Sutton 2008).
Levels of green space provision among the white population of Los Angeles
exceed that of the African-American population by a factor of 20 (Wolch et al.
2002), and recent efforts by the Californian government are thus focused
jointly on raising the quantity of access to natural green spaces, but also on
ensuring equity across society (Garcia & White 2007). A negative relationship
between green space availability and human socioeconomic deprivation has
been found in New Delhi, India (Sliuzas & Kuffer 2008), although the opposite
result was found in a study of Sheffield, with public green space provision
being biased toward less affluent sectors of society, at least in terms of its
quantity (Barbosa et al. 2007). In part these differences might reflect the differ-
ent histories of green space provision among countries. However these patterns
arise, correspondence between the distribution of human social diversity and
green space provision is one component of a wider relationship between
human socioeconomics and patterns of biodiversity across urban landscapes
(Jim & Liu 2001; Pickett et al. 2001; Kinzig et al. 2005).
Although simply accessing green space itself is a form of interaction with
nature, the quality of that interaction is critical. This will depend on both
proximity of the green space to potential users, and also its recreational value.
The role of proximity in determining green space visitation seems to vary. More
than 90% of respondents to a survey in Guangzhou indicated that proximity
was important or very important in their decision to visit an urban green space
(Jim & Chen 2006a), and studies in both the UK and USA have found that people
generally desire a green space within 3–5 minutes’ walk of their home or
workplace (Godbey et al. 1992; Comedia & Demos 1995). Despite this, 36% of
144 R. A. FULLER AND K. N. IRVINE

respondents to a UK study reported that their nearest urban green space is not
the one they visit most frequently (GreenSpace 2007), indicating that decisions
are not based on distance criteria alone. Reflecting this, the quantitative targets
for green space provision that dominated UK urban planning policy from 1925
to 1976 were replaced by a requirement to provide a hierarchy of parks based
on accessibility to a range of different amenities provided across the green
space network (Turner 1992, 2006). A series of government-commissioned
reports in the UK has resulted in a recent emphasis there on the quality and
not just the quantity of urban green spaces (Urban Task Force 1999; CABE Space
2004, 2005a, 2005b, 2007; GreenSpace 2007).

Outcomes of interactions with nature in urban green spaces

The human need for green spaces close to where we live has arguably been
instrumental in moderating the impacts of urbanisation on nature, resulting
in the retention of large parks, well-wooded remnants and encapsulated
patches of undeveloped land within many city boundaries. Although small
patches of green spaces typically make up a high proportion of overall vegeta-
tion cover across a city (Fuller et al. 2010), larger patches promote the integrity
of ecological processes, and the pattern of green space provision will directly
affect the ability of species to persist in urban environments. For example,
larger green spaces in Birmingham, UK, contained a higher diversity of plants,
and had a higher probability of occupancy by individual plant species (Bastin &
Thomas 1999). Inadequate retention of green spaces as an area becomes urban-
ised results in local extinction of species that are habitat specialists, require
large habitat patches, utilise the interiors rather than the edges of patches or
are associated with complex vegetation structures (Pickett et al. 2001; McKinney
2002; Chace & Walsh 2006).
Although green spaces can clearly moderate some of the negative effects of
urbanisation on biological assemblages, the way in which they are used and
managed is critical to their biodiversity value, and this is where human activity
and biodiversity must be simultaneously considered. An extreme example
might be that a repeatedly mown and trampled football field will not support
as many plant species as an unmown long grassy margin; the value for recre-
ation trumps biodiversity needs in this instance. However, there are many
more subtle cases. For example, the species richness of woodpeckers, forest
birds and hole-nesting birds in urban parks in Örebro, Sweden, was positively
correlated with the number of trees, and an increase in the proportion of
survey plots with a shrub layer was related to an increase in bird species
richness (Sandström et al. 2006). Species richness of birds in urban green spaces
in Greater London was positively related to the amount of unmown rough
grassy habitat (Chamberlain et al. 2007). Each of these features of green spaces
(tree cover, the presence of a shrub layer and mowing regimes) is subject to
 PEOPLE–NATURE INTERACTIONS IN URBAN ENVIRONMENTS 145

management decisions that must simultaneously consider human visitor

needs and biological value (CABE Space 2005a).
Safety of person and property is one of the main issues in urban green space
management, and there is some evidence that a high density of vegetation in
urban parks is associated with negative perceptions of safety (Schroeder &
Anderson 1984; Lyytimäki et al. 2008). Vegetation influences perceptions of
safety by affecting the ability to see long distances, or the ease of locomotion
(e.g. Nasar et al. 1993, Nasar & Jones 1997). This has led to the clearance of
vegetation and opening up of lines of sight being a dominant paradigm in the
management of many urban green spaces (Nasar & Fisher 1993; Kuo & Sullivan
2001), no doubt with negative impacts on the ability of a green space to support
biodiversity.
However, this paradigm is contradicted by research on landscape preference,
in that vegetation structure and structural designs that obscure the view,
thereby providing ‘mystery’ and interest, are part of the reason that some
landscapes are well liked (Kaplan & Kaplan 1989; Herzog & Miller 1998). There
is a need for a more refined understanding of the perceptions of safety in urban
green space, and several strands of research have begun to explore the relation-
ships among safety, preference, visitor use and naturalistic designs that pro-
mote biodiversity and ecosystem service provision in greater depth. For
example, Bjerke et al. (2006) found a greater preference for more densely
vegetated park landscapes among middle-aged, educated individuals as well
as those with an interest in wildlife and a more pro-environmental attitude.
Jim and Chen (2006a) found a similar preference for more naturalistic design
in China, while Özgüner and Kendle (2006) concluded that both naturalistic
and more manicured designs are equally appreciated. In contrast, signs of
neglect may also influence perceptions of safety. Respondents to a survey in
Sheffield mentioned signs of neglect just as much as vegetation-related factors
in contributing to places feeling unsafe (Irvine et al. 2010). These related to the
types of people and their activities (e.g. antisocial behaviour), the impact of
poor maintenance (e.g. cracked hardscape under children’s play area, dilapi-
dated buildings), and also the potential impact of litter (e.g. broken glass).
Given the interconnectedness of these various elements, future research should
continue to study these issues in combination, rather than isolation, and
investigation of cultural variation is required.
The heavy use of urban green spaces creates a paradox in the sense that
amenity use can reduce biodiversity value, but without the amenity value to
humans many of these green areas would cease to exist. Disturbance is intim-
ately tied up with the concept of open access to green spaces, and it is not clear
how the two can be separated. Disturbance effects include the simple presence
of large numbers of people, startling noises, trampling, disturbance by pets,
dumping of garden waste, litter, campsites and firewood gathering (Matlack
146 R. A. FULLER AND K. N. IRVINE

1993; Ditchkoff et al. 2006). Such disturbances extend well into vegetation
patches in urban green spaces, and markedly exceed natural edge effects in
penetration and severity (Matlack 1993). Disturbance data can be used to
formulate park management guidelines particularly where sensitive species
are known to be present in a green space. For example, flushing distances of
birds have been used to suggest design parameters for urban parks in Madrid,
Spain (Fernández-Juricic et al. 2001).
Naturalistic or more ecologically appropriate management of urban green
spaces necessitates a change in expectations of visitors and a willingness to
accept some ‘messiness’ in the landscape, to let go of a certain degree of
control, and to accept the surprises and vagaries of natural processes. For
example, the inclusion of culturally relevant ‘cues to care’ into more naturalis-
tic design can result in higher preference for and acceptance of such land-
scapes (Nassauer 1997), and Sheffield park users were willing to accept
management for biodiversity where it was compatible with human uses (Irvine
et al. 2010). Green space management that includes both human and biological
factors is already enshrined in law in several developed nations, although it is
rare in the developing world (Pierce et al. 2005). Plans for urban green space
management that fail to include citizens’ perceptions will not be widely
accepted or understood (Balram & Dragicevic 2005).
Public green space and private gardens might, to some extent, provide
alternatives for contact with nature in urban settings. Indeed, in Sheffield
there is a negative correlation between the extent of public green space and
private garden space across the city (Barbosa et al. 2007). However, the degree to
which these radically different kinds of green space can substitute for one
another is unclear as the two very frequently play different roles (Kellett
1982). For example, public green space can promote community integration
while social interactions in gardens are focused around a private social net-
work (Bernardini & Irvine 2007). Research that explicitly compares measures of
perceptions, activities, biodiversity contact and human wellbeing in public
versus private green spaces will be required to assess the consequences of
changes in the proportion of urban green space that is provided via the two
different routes.

Maintaining a private garden

Despite their generally small size in comparison with publicly available urban
green spaces, private domestic gardens (backyards) are probably the most easily
accessed and close at hand elements of nature for many urban dwellers.
Frequent contact with plants and animals is possible, and the garden forms
the arena for many childhood experiences of nature (Bernardini & Irvine 2007;
Louv 2008). Individual gardens are a dense mosaic of many microhabitat types,
with UK gardens averaging six, but reaching up to 12 land cover types despite
 PEOPLE–NATURE INTERACTIONS IN URBAN ENVIRONMENTS 147

median garden size being only 100–200 m2 (Loram et al. 2007, 2008b). Although
urban gardens are a habitat type with no real natural analogue (Gaston et al.
2007a), they can be surprisingly biodiverse (Owen 1991). A sample of 267
private gardens in the UK together contained more than 1000 plant species,
many non-native or planted (Loram et al. 2008a). As such, gardens can provide
people with exposure to a concentrated and intensified spectrum of natural
processes through seasonal and life cycles, and thus contribute substantively to
quality of life (Martin & Stabler 2004).
In the USA, at least 78% of households participate in some form of gardening
activity (Clayton 2007), and 87% of UK households have access to a garden (Davies
et al. 2009). Forty per cent of African urban dwellers engage in urban agriculture
in private gardens (Mougeot 1994), and this proportion has continued to
increase since the 1990s, in line with decreasing food security and ongoing
urbanisation of the African population (Foeken et al. 2002). In 1998, about 30%
of the population of 239 000 people in Nakuru, Kenya was engaged in some form
of urban agriculture, producing 8000 tonnes of crops per annum, enough to
satisfy about 30% of the city’s food requirements (Foeken et al. 2002). There is
substantial variation within cities, with 27% of households in inner city loca-
tions in five UK cities reporting no access to any type of outside space, while the
corresponding figures for outer suburbs and areas situated between the suburbs
and the inner city were 3.1% and 4.5% respectively (Gaston et al. 2007b).
According to Francis (1988), gardens are a venue to exert and maintain
control in contrast to the world beyond the garden fence, although Catanzaro
and Ekanem (2004) found no evidence that control of nature is important to
gardeners. Whether or not control is an important driving factor, vegetation
managed by citizens in their private gardens forms a significant component of
all urban plant assemblages. Gardens cover about one-quarter of a typical UK
city, and form up to a half of all urban green space coverage (Loram et al. 2007),
as well as affecting patterns of global environmental change (Niinemets &
Peñuelas 2009). Enhancing the quality of garden habitat emerged as more
likely to improve ecological connectivity than the specific establishment of
green corridors across urban landscapes in British Columbia, Canada (Rudd
et al. 2002). Plant species composition, richness, evenness and density are
continually influenced by human intervention in private gardens (Faeth et al.
2005), and floras are typically dominated by non-native species (Loram et al.
2008a). Heavy subsidies of nutrients and control of competition by gardeners
leads to the persistence of species at lower densities than could occur in
unmanaged populations (Smith et al. 2006b; Loram et al. 2008a). Human control
over urban plant assemblages therefore appears to overwhelm geographic,
historical and climatic variation among cities, with species richness, diversity
and composition of garden plant assemblages being highly conserved across
five disparate UK cities (Loram et al. 2008a).
148 R. A. FULLER AND K. N. IRVINE

In contrast to the findings from Europe, urban gardeners in Boa Vista, Brazil,
concentrate on only a few species of exotic trees, primarily because of their
superior fruit production (Semedo & Barbosa 2007). Likewise, urban gardens in
Limete, Democratic Republic of Congo, show low plant species richness, with
only 18 tree species in 201 plots (Makumbelo et al. 2002). A recent assertion that
urban gardens in China support few species of birds, bees and butterflies was
accompanied with a call for research into how such places might be improved
for biodiversity conservation (Wang et al. 2007).
Cultural differences in how gardens are used relate in part to the historical
provision of gardens as housing stock was built in successive planning cycles,
and in part to the pervasive economic necessity associated with using vegetable
and livestock products from small garden plots. In developing countries, urban
gardening is often carried out for subsistence purposes, e.g. producing food
from urban gardens in Mozambique (Sheldon 2003), the Philippines (Miura
et al. 2003) and Peru (Works 1990). Garden maintenance in Africa for subsist-
ence or as a small-scale commercial enterprise occurs wherever small plots of
land are available, around the margins of the home, or in vacant plots or those
owned by a third party. As cities grow, former peri-urban areas become densely
settled, plots are subdivided and agriculture continues in the land available in
the vicinity of the home (Foeken et al. 2002).
An urban garden in Cape Town of only 30 m2 is capable of providing half the
vegetable subsistence requirements for a household (Eberhard 1989, cited in
Slater 2001), although potential productivity is higher than currently realised,
and there is much interest in improving yields and horticulture techniques
(Nugent 2000). Extensions of subsistence gardening include sale of produce at
local markets, as occurs in the closely settled peri-urban regions around Bobo-
Dioulasso, Burkina Faso (Freidberg 2001), and many African cities depend on
‘garden belts’ around their perimeter for a high proportion of their food
requirements (Guyer 1987; Linares 1996). Small-scale urban agriculture com-
petes with intense land-use pressures in rapidly growing cities across the
developing world. Although urban agriculture is potentially economically
viable in South Africa, intense competition for land in Cape Town, ironically
in part from nearby nature reserves, means that plans for its continued expan-
sion are uncertain there (Reuther & Dewar 2005).
While there are tremendous cultural differences in the way gardens are
managed, and in the motivation for cultivating particular plants, their eco-
logical ramifications are scarcely understood. Uses of private gardens varies
markedly in California, such that they are seen by white-collar suburbanites as
places of escape or retreat from the stresses of the day, but by poor, urban
African-Americans as a source of food, a tie to cultural practices passed down
from parents and grandparents, as well as a safe haven in an otherwise less
than safe neighbourhood (McNally 1987, 1990). Economic factors can drive
 PEOPLE–NATURE INTERACTIONS IN URBAN ENVIRONMENTS 149

biodiversity in private gardens in rather direct ways. For example, in urban

Barcelona higher-income households can afford more water-consuming plants,
and tend to have gardens dominated by Atlantic species, while lower-income
households tend to use species adapted to the drier local Mediterranean cli-
mate (Domene et al. 2005). Thus a major dichotomy in garden plant assem-
blages is driven by the affordability of irrigation.

Managing nature in private gardens

Gardening provides multiple opportunities for interacting with nature.
Decisions about garden management are made at the household level and
are strongly influenced by the personal characteristics of the householders
(Figure 7.2). Fieldwork in Sheffield revealed a wide range of garden manage-
ment regimes from minimal active planting or vegetation control, through
assiduous planting and management of vegetation for a range of purposes, to
completely paving over the space and continually treating it with herbicide to
ensure no vegetation grows within the garden (Irvine et al. 2010).
Perhaps most obviously, and whatever the management motive, gardeners
control the range of plant species and the abundance of individual species
present in their garden. Gardeners typically favour non-native plants, either
alone or in combination with natives (Zagorski et al. 2004; Head & Muir
2006), and this is reflected in the high proportion of non-native species
available at horticultural suppliers (Thompson et al. 2003), planted in private
gardens (Loram et al. 2008a) and escaping therefrom to form established
populations in the wild (Dehnen-Schmutz et al. 2007). While it seems
logical that native plants will provide food for coevolved native inverte-
brates, assertions that non-native garden plants can have a significant
negative impact on the natural environment (e.g. Steinberg 2006) require
more detailed investigation, and there is much debate about the relevance
of exotic status alone in predicting biological quality of garden plants
(Gaston et al. 2007a). In one of the few direct tests of this issue, the richness
and abundance of invertebrates in Sheffield gardens were not dependent
on the proportion of plants that were of native origin (Smith et al. 2006a,
2006c). In addition, the presence of structurally complex vegetation was
more important than its species composition in predicting occupancy of
urban sites by small mammals and terrestrial reptiles in Brisbane, Australia
(Garden et al. 2007).
Garden management decisions depend strongly on garden size. Across five
cities in the UK, many garden features were more likely to be present in larger
gardens (cultivated borders, lawns, paths, uncultivated land, compost heaps/
bins, ponds, vegetable patches, greenhouses, unmown grass, tall trees, and
linear features within the garden such as hedges and walls; Loram et al.
2008b). As well as affecting the biodiversity value of gardens, such features
150 R. A. FULLER AND K. N. IRVINE

Figure 7.2 Divergent approaches to garden management are evident in the contrast
between neighbouring gardens in suburban Sheffield, UK. Such variation in how
gardens are managed makes for a highly heterogeneous resource for wildlife when
summed across an urban landscape. Photograph used with permission of R. A. Fuller.

contribute to ecosystem service provision by gardens. For example, paving

of front gardens in the UK has led to significant concerns about elevated
stormwater runoff and has led to additional management costs for local
authorities (Perry & Nawaz 2008). About 12% of respondents to a Sheffield
survey indicated that they parked their car within their garden space (Table 7.1).
The likelihood of large trees being in a garden reduces significantly in smaller
gardens (Smith et al. 2005), so provision of carbon sequestration depends on
how garden management decisions interact with garden size. Urban areas
experience heat island effects, in which temperatures are elevated in compari-
son with surrounding landscapes, particularly at night and in colder weather
(Pickett et al. 2001). Decisions by individual householders can moderate these
effects insofar as cover by trees and impervious surfaces determines the magni-
tude of the effect (Chen & Wong 2006). Urban parcel sizes are already small in
comparison with those in peri-urban and rural areas (Luck & Wu 2002), and
ongoing reductions in garden sizes through increasing residential density will
no doubt influence future land cover change.
 PEOPLE–NATURE INTERACTIONS IN URBAN ENVIRONMENTS 151

Table 7.1 Activities carried out at least weekly in

Sheffield, UK, gardens in summer and winter.

Activity Summer Winter

Relax 76 4
Gardening 70 10
Observe nature 52 40
Think about things 51 10
Socialise 42 2
Play sports 23 2
Park car 13 12

Note: Figures are percentages of 671 respondents to

a postal questionnaire sent to 2016 households with
garden access in Sheffield.

Concern has recently been raised about the consequences of garden manage-
ment decisions for areas outside the garden itself. For example, the use of peat
fertilisers in horticulture is widespread despite the fact that large-scale peat
extraction causes loss of biodiversity and ecosystem services (Chapman et al.
2003). The likelihood of plant species escaping horticultural use and becoming
established in the wild in the UK is positively related to their frequency of
occurrence in the nineteenth-century horticultural market, indicating that
social trends in gardening can influence the rate of establishment of non-
native species (Dehnen-Schmutz et al. 2007).
The degree to which decisions about the management of different gardens
are independent is uncertain, but critical if we are to understand how
garden management decisions influence biodiversity and determine the eco-
logical quality of urban landscapes. Gardens form an interesting gradient
between the public arena of the street and the private interior of the home.
The garden itself is in many ways a semi-public, semi-private space; many
urban gardens are overlooked by neighbours in the back or people passing by
in the front, yet are frequently enclosed by a hedge or fence. Because of the
semi-public nature of gardens, perceptions of social norms are one source of
dependency that could lead to homogenisation of garden management within
neighbourhoods, but starker differences among neighbourhoods. We are not
aware of any work on this issue, although intriguing work in Tasmania has
shown that front and back gardens show consistently different floristic com-
position and vegetation structure (Daniels & Kirkpatrick 2006). While there was
no difference in plant species richness between front and back gardens, the
former were dominated by trees and shrubs, and lacked features indicating
production and function. Relegation of production and function to backyards
is also evident in the USA (Dorney et al. 1984; Richards et al. 1984), and may
152 R. A. FULLER AND K. N. IRVINE

relate to security concerns (Daniels & Kirkpatrick 2006). Interestingly, there

was no evidence that Tasmanian front gardens were more showy (of complex
ornamental design rather than productive or utilitarian) than back gardens;
rather it appeared that the degree of showiness reflected householder attitudes
to, and interest in, gardening, and was thus manifest in all their garden space.
However, the proportion of gardens that were showy in the front and not
showy in the back was negatively related to suburb age.
Rented and shared accommodation are major exceptions to the rule that
domestic gardens are privately owned. In Tasmania, the proportion of gardens
in a suburb with non-showy front and back components increased with the
proportion of households that were rented (Daniels & Kirkpatrick 2006). This
could either represent a difference in the characteristics of the people occupy-
ing owner versus rented accommodation, or a difference in motivation when
the property is not personally owned.

Motivations for garden management

While urban agriculture is motivated primarily by the need for food, it can also
provide employment, food security, income (direct or fungible), and impetus
for urban sustainability and development in general (Smit et al. 1996). About
one-third of crops are self-consumed in Nakuru, Kenya (Foeken et al. 2002),
and garden crops can add 20% to household income in Bangkapi, Bangkok
(Fraser 2002). As well as utilitarian benefits, there are substantial additional
advantages such as empowerment of women via control over household
food consumption, the formation of social networks and community develop-
ment (Slater 2001). For example, small-scale urban gardening in Brazil pro-
motes social interaction between urban and rural communities as well as
providing subsistence (Winklerprins 2003). Entrepreneurial urban agriculture
is a growing phenomenon in the USA, and provides a focus for community
activity as well as enhancing local food security (Kaufman & Bailkey 2000;
Saldivar-Tanaka & Krasny 2004).
Where it is not directly focused on growing food for subsistence, gardening is
perceived as both work and recreation. Fifty-six per cent of householders in
Perth, Australia, indicated gardening to be both work and recreation, but over
70% regarded it as a valuable way to spend time (ARCWIS 2002). We found
comparable results in Sheffield, and the distribution of responses was surpris-
ingly similar across the two cities (Figure 7.3). Studying customers recruited
from a garden centre in Ohio, USA, Clayton (2007) identified a range of
motivations for gardening. In order of importance, these were spending time
outdoors, observing nature or natural processes, relaxation, controlling
appearance of the garden, working with the hands, novelty, producing food
or herbs, demonstrating effort and demonstrating expertise. Spending time
outdoors, observing nature and relaxation were significantly more important
 PEOPLE–NATURE INTERACTIONS IN URBAN ENVIRONMENTS 153

60

50
Percentage of respondents

40

30

20

10

0
Totally Work Both work Recreation Totally
work & recreation recreation

Figure 7.3 Attitudes to gardening are remarkably similar in studies across two cities.
Responses were to the same stem question, ‘Do you consider gardening to be work
or a recreation activity?’ Open bars are responses by 615 householders in Perth,
Australia (data from ARCWIS 2002), and shaded bars are responses by 663
householders in Sheffield, UK.

than all of the others. This closely echoes our finding from Sheffield that
observing nature was the only important use of gardens in winter, when many
other garden activities ceased (Table 7.1). Reasons for specific garden manage-
ment decisions in Clayton’s (2007) Ohio study included, in order of importance,
achieving colourful appearance, safety, reducing or eliminating weeds, main-
taining a healthy ecosystem, enhancing property value, ease of maintenance,
demonstrating care, minimising resource use, low cost, using native plants and
meeting neighbourhood standards. Perhaps surprisingly, individuals were neu-
tral as to whether the practices used in their yard affected the local environ-
ment. Practical concerns as drivers for gardening practices were negatively
correlated with use of the garden for interacting with nature, suggesting that
the two forms of motivation for garden management are somewhat opposed
(Clayton 2007).
There is marked variation across society in the use of ‘wildlife-friendly’
garden features and the propensity to engage with nature, but rather few
investigations of the motivations for particular management decisions. From
a policy perspective, both the conservation of urban biodiversity and the
enhancement of public health depend on a better understanding of this issue.
154 R. A. FULLER AND K. N. IRVINE

Benefits of gardening
Gardening is generally perceived as being important to overall wellbeing.
Responses to a similar question, though ranked on different scales, about
the importance of spending time in the garden to overall wellbeing were
remarkably similar for householders in Perth and Sheffield, with the majority
of respondents indicating that it was quite important or very important
(Figure 7.4).
Gardening produces measurable changes to psychological processes, enhan-
cing self-esteem, self-efficacy and personal identity (Bernardini & Irvine 2007).
Texan gardeners reported higher satisfaction with life and better self-reported
health than non-gardeners, and had higher physical activity levels (Waliczek
et al. 2005). Among gardeners the act of gardening itself brings satisfaction and
a pleasant break from household chores or routine (Kaplan 1973; ARCWIS
2002), and the availability of places to garden increases neighbourhood satis-
faction (Frey 1981). Gardens are full of cues to connect people with their
personal history. For example, daily or seasonal cycles, visits by particular bird
species or flowering of a favourite plant bring reminders of people, places and
events (Francis 1988; Bernardini & Irvine 2007). Twenty-eight of forty interview-
ees in Leicester, UK, expressed positive place identity associated with their
garden (Bernardini & Irvine 2007). Other benefits include the joy of anticipa-
tion and sense of accomplishment that come through the processes of garden
planning, planting and harvesting, sustained engagement with the natural
environment and reduction in stress levels (Kaplan 1973; Kaplan & Kaplan
1989; Catanzaro & Ekanem 2004). Kaplan and Kaplan (1989, p. 170) suggest that
‘an important source of satisfaction derived from gardening is that it holds
one’s attention in a multitude of ways, even when the garden lies dormant’.
A range of health measures in Scotland, UK, are related to household
tenure, such that rented housing in the public sector is associated with
particularly high levels of health problems. However, these effects are par-
tially offset by the availability of gardens, with 56.4% of respondents without
access to a garden describing their health as fair/poor, compared with 37.3%
among those with access to a garden (Macintyre et al. 2003). Having access to
and visiting one’s garden frequently were both associated with lower self-
reported sensitivity to stress among people across nine Swedish cities
(Stigsdotter & Grahn 2004).
Benefits of gardens need not involve direct interaction. For example, having
a ‘green’ view increased cognitive functioning of children living in single-
family homes in the USA (Wells 2000). Similarly, having access to a view over
gardens is positively associated with residents’ sense of community (Kaplan
1985), and satisfaction with the neighbourhood, as well as an individual’s
ability to function effectively (e.g. plan), among individuals living in multi-
storey, multi-family buildings (Kaplan 2001).
 PEOPLE–NATURE INTERACTIONS IN URBAN ENVIRONMENTS 155

(a)
50

40
Percentage of respondents

30

20

10

0
Most Very Somewhat Not
important important

(b)
40
Percentage of respondents

30

20

10

0
Very Quite Somewhat A little Not at all

Figure 7.4 The importance of spending time in the garden to overall wellbeing in
studies across two cities for (a) 616 householders in Perth, Australia (data from ARCWIS
2002), and (b) 667 householders in Sheffield, UK.

Feeding urban wildlife

In one of the most direct people–nature interactions, many people provide food
for urban wildlife. Food is provided for birds at an estimated 12.6 million of the
26 million homes in the UK (Davies et al. 2009), 43–50% of the households in the
USA and 25–57% of the households in Australia (Jones & Reynolds 2008). Such
156 R. A. FULLER AND K. N. IRVINE

activity is not limited to private gardens; 36% of visitors to urban parks in

Sheffield indicated that they feed wildlife, most notably ducks and squirrels
(Irvine et al. 2010). Although birds are the most frequent recipients, mammals
are also given supplementary food, with 11% of respondents to a British survey
of urban residents indicating they fed mammals (Gaston et al. 2007b), and 35%
of Americans who feed wildlife around their home explicitly targeting
mammals (United States Fish and Wildlife Service 2001). Three of 134 respond-
ents to a garden wildlife feeding survey in Brisbane, Australia, reported specif-
ically feeding blue-tongued skinks Tiliqua scincoides (Rollinson et al. 2003).
Observing and feeding wildlife predicts neighbourhood satisfaction (Frey
1981), but we are aware of no information on the motivations for wildlife
feeding. Because it is one of the most direct forms of people–nature inter-
actions, an appreciation of the perceptions and motivations surrounding the
activity could throw light on why people seek out interactions with nature
close to where they live. Rather more is known about the socioeconomic
correlates of wildlife feeding. Householders engaging in bird feeding in south-
east Michigan tended to be older, were more likely to be women and had
achieved higher educational qualifications than those not participating
(Lepczyk et al. 2004), although bird feeding activity was not related to the
number of people in the household, their occupation or the size of the dwelling.
What are the costs and benefits of wildlife feeding? On the one hand,
increasing public participation in wildlife feeding activity is an explicit policy
target in the UK, based on the idea that enhancing this interaction with nature
will simultaneously benefit both human wellbeing and biodiversity (DEFRA
2002). On the other hand, wildlife feeding is highly controversial in some
countries. For example, it is strongly discouraged or illegal in Australian
protected areas, particularly where dangerous wildlife are present (Mallick &
Driessen 2003; Petrie et al. 2003). It has been suggested that the strongly
seasonal climate of northern latitudes makes supplementary feeding a logical
activity to ‘help’ wildlife through the cold winter months when food is scarce,
but that this paradigm does not translate well to less seasonal environments
such as those in Australia, and as a consequence bird feeding is not beneficial
(Parsons 2007). However, there is no good evidence that wildlife feeding has
fundamentally different effects on Australian faunas compared with those of
Europe or North America. In fact, the evidence base surrounding the impacts of
supplementary wildlife feeding in general, both positive and negative, is so
weak that it is hard to draw any definitive conclusions at this stage. Studies
that can clearly disentangle feeding effects from urbanisation, habitat and
individual quality effects are urgently required (Jones & Reynolds 2008).
Despite the controversy surrounding wildlife feeding in Australia, many
visits to national parks are motivated by wildlife feeding, and participation
rates among the Australian public remain high (Woodall 1995; Rollinson et al.
 PEOPLE–NATURE INTERACTIONS IN URBAN ENVIRONMENTS 157

2003; O’Leary & Jones 2006). In response to this, best practice guidelines have
been published, alongside arguments that habitat enhancement rather than
feeding is the best way to encourage wildlife to visit a domestic garden (Parsons
2007; Plant 2008). The evidence base to support the details within such best
practice guidelines is acknowledged as limited, and work on this issue is
urgently needed (Jones 2008; Jones & Reynolds 2008). For example, although
the spread of mycoplasmal conjunctivitis in house finches Carpodacus mexicanus
in the USA has been linked to bird feeders (Fischer et al. 1997), and tubular
feeders in particular (Hartup et al. 1998), none of 25 samples taken from 25 bird
feeding stations in Sheffield contained Salmonella spp. (Cannon 2005).
Such widespread resource provision is likely to have significant impacts on
urban wildlife populations (Cannon 1999; Shochat 2004; Faeth et al. 2005).
A study in Sheffield found a positive relationship between levels of bird feeding
and the abundance of species known regularly to take supplementary food
from gardens (Fuller et al. 2008). There was no association between bird feeding
and the abundance of birds that did not regularly take supplementary food,
and there was also no effect on avian species richness. Experimental work to
determine the extent to which direct feeding of wildlife changes assemblage
structure at a landscape scale would be valuable. Such work should incorporate
differences in the frequency of feeding, given that 46% of households feeding
birds across five UK cities did so less than once per week (Gaston et al. 2007b).
Feeding wildlife in private gardens also shapes non-urban ecosystems. The
seeds of around 30 species of plant are regularly used in the feeding of birds in
the UK, but seed mixes often contain small numbers of many other species.
More than 400 plant species are believed to have been imported in this way
with bird seed into Britain from various countries around the world, and
subsequently have been found growing in the wild (Hanson & Mason 1985).
Moreover, given that they are grown precisely because of their attractiveness to
birds, cash crops aimed at the bird seed market, such as sunflower Helianthus
species, are at risk of in situ damage by wild birds (Blackwell et al. 2003). Avicides
are used to control such damage, but this is a highly controversial practice
(Fenwick 2001), and ironic given that part of the crop is destined for bird
feeders in domestic gardens.
Wildlife feeding could potentially be harnessed explicitly on a large scale to
influence the conservation status of a particular species occurring within
urban areas. Urban environments support nationally important populations
of some species (Bland et al. 2004; Cannon et al. 2005; Chamberlain et al. 2005),
yet the degree to which this people–nature interaction could be used for
conservation remains unknown, and seems an exciting avenue for further
work. Such interactions might also contribute to targets relating to urban
liveability, yet we know almost nothing about the benefits to people of
engaging in wildlife feeding.
158 R. A. FULLER AND K. N. IRVINE

People–nature interactions: common threads

People–nature interactions shape urban environments
It is fast becoming apparent that the environmental outcomes of people–nature
interactions in urban areas are significant. Most obviously, many vegetation
communities in urban areas are deliberately planted, or regularly managed, be
they in public green spaces or in private gardens. Thus the composition of urban
plant assemblages, and the distribution of vegetation types and habitats across
the urban landscape, are continually shaped by human management decisions.
Such management is often motivated by the need for public accessibility, safety
and amenity, and the presence of large numbers of people leads to disturbance
effects of various kinds. The dynamics of decisions about garden management
have a large potential impact, especially if we consider how they sum to generate
landscape level effects (Niinemets & Peñuelas 2009). Gardens are places for self-
expression, and thus they are places where the human psyche is linked intim-
ately with the distribution and abundance of species in the environment.
Thus to a first approximation, it is human decisions that structure most of
Bell’s (1976) ‘fabricated or disrupted’ urban nature. Fingerprints of societal
changes over time are reflected in components of urban nature such as
changing preferences for planting certain species, the age, density and matur-
ity of street trees, and evolving paradigms of urban park management
(Richards 1983; Zipperer & Zipperer 1992). Despite this, there remains rela-
tively little direct integration of human processes into the study of urban
ecology in the 10 years since Hostetler’s (1999) call for action.

Interacting with nature benefits people

It is clear that people derive a range of benefits from interacting with nature.
Benefits to mental and physical health and wellbeing are relatively well docu-
mented (for reviews see Chapter 9; Irvine & Warber 2002b; Brown & Grant
2005), but the implications for quality of life more broadly have scarcely been
studied. Strategies to improve quality of life among urban dwellers urgently
require this knowledge. Other aspects of human quality of life that also depend
to varying extents on access to urban nature include economic sustainability,
social cohesion, the success of urban regeneration programmes, crime reduc-
tion, sense of community and environmental awareness (Department of Trans-
port, Local Government and the Regions 2006; Maller et al. 2006).
Given the many documented benefits of local interactions with nature, low
levels of ambient biodiversity close to where people live can be interpreted as
an axis of deprivation. In Sheffield there is a negative relationship between bird
species richness and the level of deprivation among neighbourhoods, as meas-
ured by the Index of Multiple Deprivation (Irvine et al. 2010). Less privileged
sectors of society have lower levels of ambient biodiversity around the places
where they live, suggesting that lack of access to biodiversity can compound
 PEOPLE–NATURE INTERACTIONS IN URBAN ENVIRONMENTS 159

0.3
Proportion feeding

0.2

0.1

0
0 20 40 60 80
Index of Multiple Deprivation

Figure 7.5 Socioeconomic deprivation predicts engagement with nature. The

proportion of households at which birds are fed declines among 35 neighbourhood
types in Sheffield, UK, with increasing deprivation. Reproduced with permission from
Fuller et al. (2008).

more traditional measures of social and economic deprivation. This contrasts,

however, with the finding that access to local green space was better in more
deprived neighbourhoods in the same city (Barbosa et al. 2007). Socioeconomic
deprivation is also linked to the likelihood of interacting with nature. For
example, people in more deprived neighbourhoods of Sheffield were also less
likely to feed garden birds, perhaps reflecting either constraints to the activity
among those neighbourhoods, or a response to few bird species being present
in those areas (Figure 7.5; Fuller et al. 2008). Thus, there is some evidence that
interactions with nature might decline in less affluent sectors of human
society, potentially compounding other components of deprivation.
Clearly, there is also potential for negative outcomes of some people–nature
interactions, such as those with household and garden pests, encounters with
dangerous species, or negative perceptions of safety in densely vegetated areas.
Management of such conflicts between humans and wildlife in urban areas has
been amply reviewed elsewhere (Conover 2001).

People and urban nature are interdependent

The increasingly well-documented associations between human and biological
organisation suggest that there is a close interplay between the configurations
of ecosystems and human communities in urban environments (Hostetler
1999; Hope et al. 2003). Evidence of such mutual feedback mechanisms in
people–nature interactions suggests that individual components of the inter-
actions cannot properly be studied independently. For example, escape of non-
native species in urban environments can create problems that are expensive to
deal with, but resolving the issue depends on a good understanding of the
160 R. A. FULLER AND K. N. IRVINE

decision-making process surrounding species selection (Ehrenfeld 2008).

Expending time and effort to attract wildlife into a domestic garden is perhaps
more likely to continue if the activity is rewarded by the presence of certain
target species (Rollinson et al. 2003).
Perhaps one of the most important questions in this regard is how perceptions
resulting from experiences gained in the urban environment translate into
wider attitudes to nature and biodiversity conservation (Lohr & Pearson-Mims
2005; Cheesman & Key 2007; also see Clergeau et al. 2001). Are people more likely
to conserve nature when they have positive and direct experiences of biodiver-
sity on a regular basis? Childhood interactions with nature are cited as import-
ant formative experiences by those active in conservation later in life (Tanner
1980; Chawla 1999), and so the ‘extinction of experience’ resulting from low
levels of contact between today’s children and biodiversity has been cited as one
plausible reason for the lack of a coherent global strategy for biological conser-
vation (Pyle 1978; Miller 2005). Perhaps what is needed is a re-learning of how to
interact with the natural environment. Not only is it important to understand
factual information about the landscape, such as the names of birds visiting
one’s back garden, but also to have procedural or skill-based knowledge (e.g. how
to set up a compost pile) that may facilitate more active engagement. Research
on this issue is timely, because there has been a recent upsurge of interest in how
local living relates to global conservation, most notably through contributions
to climate change. If such links can successfully be made, local experiences of
biodiversity could well prove to be important in determining the future of the
global response to the biodiversity crisis (Ehrlich 2002).

People–nature interactions are diverse

There are many more interactions between people and nature than we have
been able to cover specifically here, including knowing that nature is nearby,
watching wildlife on television, hanging pictures of nature on the wall,
viewing nature through a window, visiting urban zoos and botanic gardens,
campaigning for environmental change, joining local conservation manage-
ment groups and engaging in citizen science. Many activities are hard to
delineate given that humans are almost perpetually in contact with some form
of nature in one way or another. A clear typology of people–nature interactions
is lacking, but would be useful as research in this field continues to develop.
People–nature interactions vary with socioeconomic factors and the cultural
background of participants. The literature is strongly biased toward Western
cultures in terms of overall quantity, but there is also cultural variation in
emphasis (Teel et al. 2007). Studies in Western societies tend to focus on human
quality of life and wellbeing, and maintaining the biological quality of the
urban landscape, while those in non-Western societies frequently emphasise
subsistence, economic gain and developmental potential of interactions
 PEOPLE–NATURE INTERACTIONS IN URBAN ENVIRONMENTS 161

between people and nature. Perhaps most notably, there is a large literature on
urban agriculture in developing countries (see, for example, the 526 references
in Obudho & Foeken 1999). Because of its inherently interdisciplinary nature,
work on people–nature interactions is scattered through the natural science
and social science literatures (for example, the citations in this chapter arise
from journals in 25 major subject areas). Our impression is that there is far
more work by social scientists considering the importance of nature for
humans than by biologists considering the importance of human decisions in
structuring biological communities.

Concluding remarks
Experiences of nature have declined in quantity and quality over the past
few decades, and concerns are mounting that this process will accelerate as
the world’s population becomes increasingly urbanised. Anthropogenic simpli-
fication of ecosystems through harvesting and agriculture, the setting aside
of protected areas for biodiversity conservation, and urbanisation of the
world’s population have all contributed to a reduction in contact between
people and biodiversity, and a growing separation between people and nature
that shows no sign of slowing. This separation appears to have two major
consequences, (i) a reduction in the contribution of experiences of nature to
human quality of life, and (ii) a lack of broad-based support for biodiversity
conservation (Miller 2005).
Biodiversity conservation is a response to anthropogenic impacts on ecosys-
tems, and as such depends on a good understanding of the motivations and
drivers of human behaviours that lead to such impacts. Given that most of the
world’s population lives in urban areas, this is where we must look to study
human relationships with biodiversity, because the outcomes of interactions with
nature appear to be important in shaping our values and attitudes toward nature
conservation. Implementing solutions to the biodiversity crisis will depend on
interdisciplinary research efforts as well as systems of implementation that can
actively trade off ecological value and benefits to human wellbeing (Polasky et al.
2008). Much remains to be done, and we encourage increased collaboration
among researchers whose disciplines intersect on these issues.

Acknowledgements
R.F. is supported by the Applied Environmental Decision Analysis research hub,
funded through the Commonwealth Environment Research Facilities pro-
gramme, Australia, and additionally by a grant from the University of Queens-
land. We thank O. Barbosa, Z. Davies, P. Devine-Wright, C. Fuller, K. Gaston,
S. Herbert, S. Kark, J. Rhodes, J. Tratalos, S. Warber and P. Warren for helpful
discussions on people–nature interactions, and useful comments were pro-
vided by two reviewers.
162 R. A. FULLER AND K. N. IRVINE

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CHAPTER EIGHT

Urban ecology and human social

organisation
PAIGE S. WARREN, SHARON L. HARLAN,
CHRISTOPHER BOONE, SUSANNAH B. LERMAN,
EYAL SHOCHAT AND ANN P. KINZIG

Introduction
Consider a tree, growing in a forest. Its fate, in terms of growth rate, longevity
and reproduction, is regulated by a suite of biotic and abiotic factors, such as
soil moisture, competition with other plants and interactions with natural
enemies. Its branches, leaves and roots contribute to nutrient and water cycles.
The tree provides habitat for animals – insects, birds, mammals. Understanding
these interactions falls squarely in the domain of classical ecology. Now,
imagine that the tree and its surrounding forest is part of a city, the trees
interspersed with houses, streets, lawns and gardens. Here, the fate of the tree
becomes strongly intertwined with the lives and decisions of humans. Its
longevity, reproductive rate and contributions to the larger ecosystem are
mediated by a complex suite of anthropogenic processes. In this setting, the
tree’s life cycle may differ from that of all of its ancestors: its ‘birth’ perhaps in
a commercial nursery, its dispersal through an economic system of marketing
and transportation, its growth determined by local application of fertiliser by a
resident or a lawn care firm, its ultimate removal (death/decay) perhaps medi-
ated by city policies on tree hazards, its reproduction quelled entirely. Yet, the
tree and its urban forest still contribute to ecosystem processes. It provides
habitat for animals, and its leaves, branches and roots still influence the flow
of water through the urban ecosystem. Humans, therefore, both consciously
and unconsciously sculpt biodiversity and ecosystem functioning in cities
(and elsewhere).
We use this story of a tree in an urban forest, not to focus attention on trees,
but to illustrate how human social organisation drives the characteristics of a
city’s green infrastructure (e.g. vegetation, animals, soil and water). Urban
ecology has emerged in recent years as an interdisciplinary or trans-disciplinary

Urban Ecology, ed. Kevin J. Gaston. Published by Cambridge University Press.

# British Ecological Society 2010.
 URBAN ECOLOGY AND HUMAN SOCIAL ORGANISATION 173

endeavour that aims to integrate understanding of human social organisation

with traditional ecological approaches to the study of ecosystems.
Over the past century and more, social scientists in virtually all disciplines,
including sociology, geography, anthropology, economics, environmental
psychology, landscape architecture, urban planning and design, have gener-
ated rich and sometimes conflicting bodies of knowledge and theory about
human life and the environment in cities. Ecologists have come to study the
city far later in the game, and frequently find challenges far different from
those in their traditional arenas of study. How should an ecologist proceed in
such an environment, where understanding ecological processes necessitates
crossing the boundary into social sciences? Here we provide a necessarily brief,
but ecologist-friendly, survey of key theories from the social sciences and a
review of findings from recent integrated empirical research. As a point of
entry, we focus on the spatial pattern and temporal dynamics of urban ecosys-
tems, these being fundamental components of disciplinary-based ecology.

The social organisation of cities

Historical context
Most scholars agree that cities, defined as dense settlements incorporating great
physical and human social diversity, originated in Mesopotamia five to six
thousand years ago. A necessary precursor to cities was the invention of agricul-
ture, which generated a surplus of food that could be traded for other types of
food, goods or services. From the earliest city-states to modern metropolitan
areas, the agglomeration economies of urban settlements allow for increasing
divisions of labour and productivity, which are key drivers of urbanisation
(Batty 2008). Until the Industrial Revolution, relatively few people could afford
to live in cities, dependent on others to grow most of their food. Significant
advances in the productivity of agriculture and manufacturing, made possible
through steam power and fossil fuels, pushed surplus labour from farms and
drew workers to cities in larger numbers. By 1850, England became the first
urban nation in the world, with more than half its population living in towns
and cities (Bédarida 1979). By 1920, the United States passed this urban thresh-
old, and in 2008, half of humanity was housed in urban areas (UNFPA 2007).
Distinct hierarchies and social inequalities are as much characteristics of
modern-day Lagos as ancient Babylon. Yet despite very visible and persistent
poverty, urban areas continue to offer opportunity for rural migrants or
shantytown dwellers to find work, education, health care and entertainment.
Efforts to reduce rural to urban migration have largely failed, and have often
worsened economic conditions for the very poor. Inadequate planning and
mismanagement of resources have amplified poor living standards and envir-
onmental impacts that can accompany urban migration, especially in the
developing world. Those on the lowest rungs of the socioeconomic ladder
174 P. S. WARREN, S. L. HARLAN, C. BOONE ET AL.

Box 8.1 Divisions in cities

Marcuse and van Kempen (2002) illustrate the complexity of urban forms,
drawing on the following definitions of function, status and culture. Pre-industrial
cities were divided by economic function, according to the kind of market
activity taking place in particular locations. Public commercial spaces were
identifiable and demarcated from residences. Neighbourhoods were often
organised around residents engaged in similar kinds of craft occupations. More
complex divisions of land by types of industry, residences, open spaces,
highways and so forth are characteristic of industrial cities, which formalise
restrictions on the functional use of space with legal restrictions (zoning).
(See also the section on ‘Temporal dynamics’ for further discussion of historical
changes in division by land-use.)
Hierarchical status divisions in cities embody the power of higher-status
residents spatially to separate themselves from subordinate status groups and
less desirable functional activities. Socioeconomic status (a set of indicators
measured by household income, occupation and education) taps an elusive
underlying dimension of social status. Cultural divisions are not themselves
hierarchical, though they complicate urban spatial patterns because they may or
may not overlap with status hierarchies and economic function. Cultural divisions
are most readily identified by language, ethnicity and visible symbols of
associated beliefs and lifestyles.
Most urban ecological studies to date have sought to relate ecological
outcomes, such as patterns of species diversity, to socioeconomic indicators
(see ‘Social stratification and ecological outcomes’). Fewer studies have
addressed cultural divisions (see ‘Other social dynamics’).

suffer most from environmental consequences of urbanisation, including pol-

luted air, water and soil (UNFPA 2007). It is increasingly recognised that quality
of life in urban areas depends in part on an adequate supply of ecosystem
services (Martine 2008). However, inclusion of the benefits of ecosystem services
into urban planning is rare, and addressing social and environmental inequi-
ties and vulnerabilities remains a challenging and pressing issue.

Spatial organisation
Evidence of partitioned urban space exists in ancient cities and extends with
remarkable consistency through time and across virtually every historical
urban tradition (Marcuse & Van Kempen 2002). Urban archaeological records
identify ‘zones’ that correspond to administrative and economic functions and
demarcate residential areas with different population characteristics (Keith
2003; Cowgill 2007; Smith 2008). In a sweeping analysis of world cities, past
and present, Marcuse and Van Kempen (2002) concluded that three social
divisions in cities – function, status and culture (see Box 8.1) – constitute the
 URBAN ECOLOGY AND HUMAN SOCIAL ORGANISATION 175

foundation for spatial boundaries. While these boundaries, which demarcate

power differences in segments of society, have been blurred at times in the past
and ‘may be merging again in today’s economies’ (Marcuse & Van Kempen
2002, p. 14), significant segregation remains in many modern cities ( Johnston
et al. 2007). These authors and others (e.g. Briggs 2004) acknowledge that cities,
both past and present, differ widely in the contours, permeability and charac-
teristics of their sociospatial boundaries. Social complexities introduced by
industrialisation and global capitalism have increased the variety and degree
of sociospatial differentiation within cities ( Johnston et al. 2007). For ecologists
interested in the environmental consequences of spatial partitioning in cities,
these are relevant points of agreement across historical cases and different
branches of urban theory:

• Spatial patterns are socially constructed by human actors;

• Spatial patterns often reflect social hierarchies;
• Particular spatial configurations of social activities fragment environments
within a city and increasingly beyond city boundaries, fragmenting envir-
onments in peripheral areas.

Social theories differ, however, in their explicit recognition of the institutional

forces through which fragmented urban space is intentionally constructed and
controlled ( Joseph 2008).
American sociological theory on cities and spatial differentiation originated
with the human ecologists associated with the University of Chicago in the
early twentieth century (Park 1915; Burgess 1928). They were the first to detail
a theoretical model of urban growth and development based on concepts
borrowed from ecologists, including competition for space, and the succession
and segregation of new population groups with city expansion and increas-
ingly diverse economic activities (Palen 2005). Among the contributions of the
Chicago School was conceptually linking:

urban spaces to distinctive social groups, creating a spatiality to the urban form and
to cultural difference that was previously undeveloped. Urban space came to be seen
as divided and organized by social boundaries that were connected to class, race,
ethnicity, and degree of assimilation . . . in a way that recognized the extent of
concentration and segregation, and linked cultural assimilation to spatial movement
from the inner city to the outer rings.
( Joseph 2008, p. 9)

Largely inspired by this tradition, subsequent sociologists have created a richly

contextualised body of literature that explores both the positive aspects of
residential segregation in the form of ethnic enclaves and the negative aspects
of concentrated poverty and racial discrimination in slums and ghettos
(e.g. Whyte 1943; Wirth 1956; Duneier 1999; Johnston et al. 2007). These studies
176 P. S. WARREN, S. L. HARLAN, C. BOONE ET AL.

Box 8.2 Defining terms

A central city is the core municipality of a large metropolitan region.

Growth around a central city was originally conceived as a spreading set of
concentric rings corresponding to functional and status divisions. This model
is no longer subscribed to by most urban geographers. Suburbs, largely
residential communities, historically arose on the margins of central cities through
abandonment of central cities by wealthy elites. Suburban sprawl or
suburbanisation refers to the largely post-World War II phenomenon of the
expansion of metropolitan regions through low-density development.
Post-industrial cities, such as Los Angeles, are more often polycentric, with
reduced dependencies of suburbs on a single central city such as London,
Paris or Chicago. These metropolitan regions have multiple nodes of
higher-density development dispersed throughout a mixed landscape of
low-density and remnant rural lands. Many quantitative definitions exist for terms
like urban and suburban, but authors generally agree on these qualitative
definitions (McIntyre et al. 2000).

of urban social environments serve as valuable precursors and counterparts to

studies of various environmental inequalities among city neighbourhoods.
The specific urban core–periphery spatial model proposed by the human
ecologists was modified and ultimately rejected by later social theorists who
contrasted pre-industrial cities with industrial cities (e.g. Sjoberg 1960) and
those who studied the emergence of Los Angeles, USA, and other world mega-
cities with distinctly different sociospatial patterns (e.g. Soja 2000). Thus, the
urban to rural gradient is an overly simplistic assumption in the social sciences
because urban spatial patterns vary according to the contours of multiple and
overlapping functional and cultural divisions with socioeconomic hierarchies
(Box 8.1). In polycentric cities, for instance (Box 8.2), fragmentation appears to
characterise urban space better than linear distances from a central city.
Second, the human ecologists’ assumption that spatial segregation is the result
of ‘natural’ selection as opposed to economic and political power exercised by
elites (Chaskin 1997) was also abandoned by later theorists who focused on how
the political economy of urban development is intentionally structured to
benefit elites and disadvantage society’s marginalised groups (Palen 2005).
Logan and Molotch (1987) argued in a widely acclaimed book that land in the
USA, as well as in the developing nations, is highly commodified, privatised and
controlled by development corporations with the acquiescence of local govern-
ments. In response to profit-making opportunism, municipal governments
respond to the ‘urban growth machine’, leaving local residential communities
without consideration or resources to obtain a good quality of urban life.
 URBAN ECOLOGY AND HUMAN SOCIAL ORGANISATION 177

Political economists differ on whether they attribute urban social and

environmental degradation primarily to investment capital decisions by local
governments (Harvey 1973; Logan & Molotch 1987) or to withdrawal of indus-
trial capital from the urban economy (Wilson 1997), but they speak with one
voice about the deliberate construction of urban sociospatial inequalities.
Access to existing environmental amenities and proximity to disamenities
(such as pollution) is stratified locally between privileged neighbourhoods
and neighbourhoods that are politically disenfranchised by legally enforced
restrictions on land, socially burdened population groups and lack of economic
resources (Boone 2008).
The production of sociospatially segregated cities is a multi-scale process,
influenced by underlying choices about urban growth policies, society’s provi-
sions for the education and employment of urban residents, and the political
economic realities that manipulate consumers’ desires and tastes. Household
decisions about where to live are experientially conditioned by all these factors
even before they are also limited by household income, knowledge and lifestyle
preferences (Bourdieu 1984; Gottdiener & Hutchinson 2001). Racism and leg-
acies of colonialist ideologies, reflected in real estate markets and loan prac-
tices, often play on the majority’s prejudice and fear to exacerbate the isolation
of minority communities (Krysan & Farley 2002; Quillian 2002).
The extent and salience of contemporary urban residential segregation by
social class and race/ethnicity has been amply documented in the USA (Massey &
Denton 1993; Lee et al. 2008), in other developed nations (Musterd 2005;
Johnston et al. 2007) and in some cities in developing countries (King 1976;
Monteiro 2008). Social scientists have focused attention on the ramifications of
sociospatial structuring of cities for vulnerabilities to problems such as air
pollution and toxic waste sites (Bolin et al. 2005; Grineski et al. 2007). Attention
to sociospatial patterns has followed more slowly in the ecological literature. In
later sections, we examine spatial associations between social and ecological
patterns, such as vegetation structure and composition of biotic communities.

Temporal dynamics
Cities grow, decline and change in response to long- and short-term cycles and
rhythms. Tall grass on lawns of abandoned homes and swimming pools filled
with algae are reminders of the cyclical nature of capitalist economies. Karl
Marx remarked on the cyclical patterns of economies 140 years ago and others
have since paid careful attention to how the ‘crisis of overaccumulation’, the
result of surplus capital unable to find profitable markets, affects processes and
patterns of urbanisation (Castells 1977; Smith 1996). For Harvey (1982, 2003),
the crisis of overaccumulation in cities culminates in large-scale infrastructure
development as a means, if only temporary, of averting an economic down-
turn. New infrastructure projects provide room for spatial expansion of
178 P. S. WARREN, S. L. HARLAN, C. BOONE ET AL.

capital as well as a temporal deferral of economic decline, what he terms a

spatiotemporal fix. Kuznets (1967) recognised the same infrastructure
investment as a 15- to 25-year long wave economic cycle. Other economists
have predicted economic cycles for technology (Kondratieff ’s 40–60 years),
fixed investment (Juglar’s 5–7 years) and inventory (Kitchin’s 3–5 years) (Hall
1988). What is important for our understanding of cities is that these waves
ripple through the built environment, expanding suburban limits and
renewing older neighbourhoods during up years, and leading to abandoned
homes on the periphery and disinvestment in the down years.
Similar to all landscapes, cities record generations of decisions and actions in
the built forms that remain. Older cities like London, UK, or Beijing, China,
show remnants of urbanisation that are hundreds or even thousands of years
old (Vance 1990). The morphology of cities reflects the dominant economic
function and prevailing technologies, especially for transportation, in eras of
building and growth. Indeed, most cities are about an hour across (Marchetti’s
Constant), corresponding to the average travel time people will tolerate on a
daily basis, regardless of the transportation mode (Newman & Jennings 2008).
Configuration of a city like Phoenix, USA, characterised by low-density suburbs
and segregated land-use, is largely a product of post-World War II growth in an
era of widespread automobile use, inexpensive energy and rising incomes. Pre-
industrial cities, by contrast, were constructed when movement of goods and
labour happened on foot or by beast of burden, resulting in high densities,
narrow roads and mixed land-use. While these pre-industrial morphologies
may now be ringed by low-density suburbs, they continue to have important
impacts on metropolitan function. Old historical cities like metropolitan
Boston, USA, or Paris, France, poorly accommodate the automobile, but are
well suited to mass transit, walking and mixed land-uses, elements of the
compact city that are now celebrated and promoted in smart growth policies
and the planning and design philosophy of New Urbanism (Boone & Modarres
2006). Similar to ecosystems, past urban practices continue to have legacy
effects on present-day city structure and function.
Urban areas are dynamic by their nature, but some components are persis-
tent. Road networks, for instance, can persist and define urban morphology for
hundreds or even thousands of years. Private residences, on the other hand,
may last only two or three generations before being replaced with new struc-
tures. Social organisation in cities can also display the same persistence and
dynamism. In the USA, white and later black middle-class abandonment of
central cities occurred swiftly after World War II, drawn to the suburbs by
federally guaranteed mortgages and subsidised automobile transportation, in
addition to the tactics of estate agents (realtors) looking to capitalise on
residents’ fear, racial prejudices and discrimination ( Jackson 1985). Earlier
research on neighbourhood succession, drawing on ecological theory, neg-
lected some of the institutional and political factors, such as redlining (the
 URBAN ECOLOGY AND HUMAN SOCIAL ORGANISATION 179

twentieth-century practice of delineating areas, usually black inner city neigh-

bourhoods, where banks would not invest) that accelerated the process of
neighbourhood change (Hillier 2003a, 2003b). While thresholds of poverty
and home ownership rates appear to be triggers of neighbourhood transform-
ation, much research remains to be done on the institutional and structural
drivers of change (Galster et al. 2000). Another promising avenue of research
from the environmental justice literature is the link between environmental
disamenities and neighbourhood dynamics over the long term (Boone 2008).
Urban growth on the periphery continues to accelerate throughout the
world, but central cities (see Box 8.2) are far from static. Because of the econo-
mies of agglomeration they provide, central business districts attract high-end
commercial, office and service sectors. Many older neighbourhoods in central
cities have enticed investors to purchase and renovate ageing houses.
Gentrification of inner city neighbourhoods is one example that has garnered
a good deal of attention, in part because the rejuvenation of historic houses can
drive up rents and drive out poorer residents. One theory on gentrification
argues that the return to the central city is a function of the rent-gap, or the
difference between actual and realised value of inner city property (Smith 1987).
Another argues that changing demographics, especially the rise of a new cre-
ative class, is the primary force behind infill housing, gentrification and
demand for inner city living (Ley 2003; Florida 2005). Net effects, however, are
small compared with the rapid expansion of suburbs. By 1970 the suburbs of the
USA contained more people than central cities or rural areas, and by 2000 more
people lived in the suburbs than urban and rural areas combined. Over the
course of 80 years, the USA moved from being an urban to a suburban nation.

Global context
Urban theorising in the social sciences since 1990 has demonstrated a broader
global perspective on cities, space and the physical environments as these
concepts relate to the world system of cities in the hierarchy of nations (Palen
2005). Political ecologists have assessed the impacts of global capitalism on the
economies of developing nations that are undergoing rapid urbanisation and
industrial transformation (O’Connor 1994). Political ecology ‘seeks to under-
stand the human processes leading to the destruction and creation of material
environments’ (Kirkby et al. 2001), and in the developing world, as in industrial-
ised nations, this amounts to the struggle between powerful agents over the
control of land (Bryant 1998).
Encroachment of human settlement beyond city boundaries is not a new
problem, but landscape fragmentation on the urban fringe is occurring now
on an unprecedented global scale (Alberti et al. 2003; Zhao et al. 2006). Suburban
sprawl in industrialised nations, which turned natural and agricultural land-
scapes into millions of houses, is one of the two great land-use changes of the late
twentieth century (Rudel 2009). The other is tropical deforestation by conversion
180 P. S. WARREN, S. L. HARLAN, C. BOONE ET AL.

Figure 8.1 The growth of shantytowns and squatter settlements, such as this one in
Mirpur, Bangladesh, exemplify some of the starkest inequities in environmental
conditions found in modern cities, particularly in the developing world. New urban
migrants in search of work struggle to gain accommodation in shantytowns and
squatter settlements. Photograph used with permission of A. Y. Hoque.

to agriculture, which according to Rudel’s (2009) analysis results from the same
political economic process of local elites forming ‘growth coalitions’ for profit.
Uncontrolled urban sprawl around cities in industrialising nations, however, is
the third-order spatial transition that further degrades landscapes and increases
the vulnerability of marginal populations and fragile ecosystems.
According to Massey (1996), the world has entered a new era of urban
extremes, in which rapid urbanisation combined with rising levels of income
inequality create ‘hypersegregation’, or an ‘ecology of inequality’, in world
cities. The economic, social, human health and, we argue, ecological conse-
quences of sociospatial stratification are profound. The consequences of these
changes for the physical environment can hardly be overstated. One billion
people, or almost one-third of the world’s urban population, live in slums,
defined as abject poverty where there is no fresh water, sanitation or public
infrastructure (UN-HABITAT 2003). This is projected to double in 30 years to
one-third of all humanity, with the vast majority of these people living in Asia,
Africa and South America (Whelan 2004). Many of the poor are housed in inner
city slums but even greater changes are taking place on the periphery of cities
as new urban migrants in search of work struggle to gain accommodation in
shantytowns and squatter settlements (Figure 8.1). ‘Squatter settlements are
 URBAN ECOLOGY AND HUMAN SOCIAL ORGANISATION 181

mainly uncontrolled low-income residential areas with an ambiguous legal

status regarding land occupation . . .’ (UNCHS (Habitat) 1981, p. 15). The
conundrum faced by environmental scientists and conservationists is that both
planned growth (urban growth coalitions) and unplanned growth (settlements
by desperate, marginalised people) seem to lead to increasing sociospatial
divisions within cities that can create social and environmental problems.

Linking social organisation and ecological outcomes

We now take a ‘one-way’ look at how the socioeconomic and temporal pro-
cesses outlined above lead to various ecological outcomes, from patterns of
nutrient loading to plant communities to animal communities inhabiting
urban spaces. We review spatial and temporal processes jointly because it is
clear that they are closely intertwined in the formation of the patterns detected
in recent studies. Work that links human social organisation to ecological
outcomes has generally fallen into three classes of study: (i) impacts of zoning
policies as exhibited by differences among land-use categories, (ii) correlates of
social stratification, such as associations between ecological variables and
income level, and (iii) environmentally oriented human actions at the house-
hold level, such as gardening, fertilising lawns or feeding birds. These three
roughly track a gradient in spatial scale of the effects with the household or
property parcel scale as the smallest zone of effect (Figure 8.2).
The patterns emerging from recent literature suggest a profound influence
of humans on the structure and functioning of urban ecosystems, and they
point to a new catalogue of environmental inequalities – of access to greenness
and biodiversity – playing out alongside inequities in exposure to toxic chem-
ical releases and other classic examples that led to the founding of the environ-
mental justice movements of the late twentieth century. But the survey we
present here also points to the many broad gaps in our knowledge.

Differentiation by land use

Historical economic and technological changes have differentiated the growth
forms of cities. For heuristic purposes, a simple classification of urban
morphologies yields two major types: pre-industrial cities with high densities,
narrow roads and mixed land-use, sometimes surrounded by modern expanses
of lower-density suburbs; and post-World War II cities dominated by low-density
suburbs and segregated land-use (see ‘Temporal dynamics’ above). Ecologists
have generally addressed these distinct morphologies by applying a land-use
classification, an urbanisation gradient approach or a combination of these
approaches (Rebele 1994; Pickett et al. 2001; Cadenasso et al. 2007). A broad
range of ecological outcomes are strongly correlated with land-use type, and
can usually be characterised as varying with increasing levels of urbanisation,
182 P. S. WARREN, S. L. HARLAN, C. BOONE ET AL.

KEY
Strength of ties
CITY/COUNTY
STRONG
MEDIUM
WEAK

NEIGHBOURHOOD
Social stratification
• Public rights-of-way

HOUSEHOLD
Lifestyle behaviour
• Private lands

INDIVIDUAL
Population
• Riparian Areas

Figure 8.2 An illustration of the connections between different levels of social

organisation in cities. Modified from Grove et al. (2006) with permission.

as measured by increasing density of human populations, built structure and

impervious surface cover (Blair 1996; Pickett et al. 2001; McKinney 2002).
Urbanisation drives local extinctions of many native plants while introdu-
cing other, non-native species (Kaye et al. 2006). Exotic species contribute to the
high species richness of urban communities (Chapter 6; Hope et al. 2003), which
may exceed diversity in wildlands. Land-use type also affects plant commu-
nities and diversity. In the Georgia Piedmont, USA, species richness of woody
plants decreased with impervious surface and landscape diversity. Alien species
richness was the highest in urban sites, whereas native stem densities were
lower in urban and agricultural lands, compared with unmanaged forests
(Burton & Samuelson 2008). Urban and suburban areas can also affect plant
species composition in adjacent forest patches, by serving as sources for the
spread of non-native species into forests. In Ottawa, Canada, forest fragments
surrounded by urban habitat had a 50% greater proportion of introduced
plant species than fragments surrounded by agricultural and forest habitats
(Duguay et al. 2007).
Whereas urban plant communities are normally more diverse than wildland
communities because of the introduction of many species, the opposite pattern
 URBAN ECOLOGY AND HUMAN SOCIAL ORGANISATION 183

has been observed for most animal taxa. Several studies on arthropods suggest
that urbanisation causes a reduction in diversity, although within the urban
core, land-use type may also influence diversity. In Phoenix, Arizona, Shochat
et al. (2004) found spider diversity in xeric residential yards (those with
drought-tolerant plants) to be as high as in the adjacent Sonoran desert,
whereas mesic residential yards (those with water-dependent plants) had spider
diversity as low as in agricultural fields. In Birmingham, UK, the richness and
diversity of carabid beetles were lower in urban and suburban habitats than in
rural zones (Sadler et al. 2006). As with many other taxa (McKinney 2002;
DeStefano & DeGraaf 2003; Fuller et al. 2007a), the decrease in arthropod
diversity in urban settings is normally associated with an increase in the total
density. Faeth et al. (2005) suggested, based on field experiments from Phoenix,
that this increase is the result of bottom-up control (the increase in plant
densities and overall productivity), despite the strong top-down control
(higher bird population densities in the urban habitat compared with the
Sonoran desert).
Bird diversity also tends to decrease with urbanisation (Chace & Walsh 2006),
although, along a land-use gradient, bird diversity may peak at moderate levels
of urbanisation. In Seattle, Washington, Marzluff (2005) found that bird diver-
sity peaked at intermediate levels of urbanisation, where the proportion of
forest cover in the landscape was still relatively high. Early successional birds
(species that are found in a variety of habitats around the area) contributed to
the high species richness in this part of the landscape. As the proportion of
the built environment in the landscape increased, loss of many of these species
and specialist forest species exceeded immigration rates of synanthropic
species, and bird diversity declined. Blair and Johnson (2008) also showed
similar patterns from Ohio, Minnesota and California, suggesting that subur-
ban land-uses, with their intermediate levels of development, serve as points of
extirpation for woodland birds as well as entry points for invasive species into
urban systems.
Patterns of association between land-use and nutrient cycles are more vari-
able, depending on ecological context. In Baltimore, Maryland, a pre-industrial
city in a predominantly forested region, nitrate concentration is lower in dense
urban areas than in either suburban or agricultural areas (Groffman et al.
2004), owing to differential inputs. The source of urban nitrogen is mostly
atmospheric, whereas suburban and agricultural areas include deposition but
also fertiliser (Pickett et al. 2008). In arid regions, such as Phoenix, Arizona
(another post-World War II city), concentrations of soil nutrients are less
strongly associated with building density, instead following differences in
landscaping designs. Concentrations of carbon, nitrogen and phosphorus in
the soil are higher in mesic residential and agricultural lands than in xeric
residential and desert habitats, although legacies of land-use influence these
184 P. S. WARREN, S. L. HARLAN, C. BOONE ET AL.

patterns (Kaye et al. 2006). These differences are driven not only by biophysical
variables, but also by sociocultural variables underlying differences in cover
of turfgrass, tree and impervious surface. Carbon accumulates in human-
dominated soils, although most of it has short residence times in mesic yard
and agricultural soils (Kaye et al. 2006).
In summary, many ecological outcomes can be tied to urbanisation gradients
and differences in land-use, and these urban morphologies in turn derive from
historical legacies of changing modes of transportation technology and
changing economies. Plant and animal communities shift in relatively predict-
able ways with increasing levels of urbanisation – increasing densities of exotic
species and synanthropic native species, declines in native species diversity,
with a possible peak in total biodiversity at intermediate levels of urbanisation
(reviewed in Chapter 5; McKinney 2002; Shochat et al. 2006; Pickett et al. 2008).
Water quality and aquatic food web structure degrades with increasing levels
of urbanisation (Paul & Meyer 2001; Groffman et al. 2003). Other ecological
outcomes, such as soil nutrient concentrations, show fewer clear generalisa-
tions across cities and land-use types (Kaye et al. 2006). Variations on land-use
themes, such as the differences between mesic and xeric landscaping designs in
cities of the southwestern United States, illustrate the limitations of solely
employing land-use classifications or gradient approaches.

Social stratification and ecological outcomes

As we search for ecological signatures of the sociospatial divisions in cities, the
first step has been to document correlations between patterns of social stratifi-
cation and various ecological factors. Income is a commonly used indicator of
social stratification, along with ethnicity and education level. Members of the
Chicago School suggested that environmental quality would follow wealth,
leaving impoverished zones of cities also in poor environmental condition
(see ‘Spatial organisation’ above). However, it took some time before ecologists
attended to this prediction.
A focus on vegetation emerged relatively early on in the urban ecological
literature, with a few early studies documenting correlations between
income level and properties of the vegetation in city neighbourhoods
(Whitney & Adams 1980; Talarchek 1990). In the 1990s, a rapid proliferation
of large-scale ecologically driven studies in cities generated large statistical
samples of vegetation structure and other ecological properties across metro-
politan regions. The vast majority of these studies have focused analysis on
relationships of vegetation to human population density or to characteristics
of built structures (e.g. along urbanisation gradients). This greater attention
to urbanisation gradients has also been true for studies of animal commu-
nities, soil properties, nutrient processes and water quality (Pickett et al.
2001; Shochat et al. 2006).
 URBAN ECOLOGY AND HUMAN SOCIAL ORGANISATION 185

Inference of underlying social processes from ecological studies taking an

urbanisation gradient approach is limited by the dynamic relationships over
time between social stratification and built structure (see ‘Temporal dynamics’
above). Although it might be expected that gradients in built structure are
associated with poverty and wealth, processes such as gentrification introduce
complexity to this relationship (Ley 2003). Also, as Talarchek (1990, p. 66)
points out, ‘sequential occupancy of various income and social groups could,
in time, result in no relationship between socioeconomic status and vegeta-
tion.’ Furthermore, urban or suburban neighbourhoods of similar population
density and forms of built structures can harbour distinct biotic communities,
with their characteristics driven by a combination of historical legacies and
contemporary management by residents (Whitney & Adams 1980; DeGraaf &
Wentworth 1986).
Direct tests of association between social stratification and local ecological
conditions increasingly show evidence for the predicted pattern of environ-
mental quality following wealth, though with some complexities emerging.
Documented associations with income level and related socioeconomic indica-
tors include: vegetation cover (Talarchek 1990; Iverson & Cook 2000; Smith et al.
2005; Grove et al. 2006; Jenerette et al. 2007; Tratalos et al. 2007), plant diversity
(Whitney & Adams 1980; Hope et al. 2003; Martin et al. 2004), bird diversity (Hope
et al. 2004; Kinzig et al. 2005; Melles 2005), densities of some bird species (Loss
et al. 2009), supplementary feeding of birds (Fuller et al. 2008; Lepczyk et al. in
press; but see Gaston et al. 2007), small mammal diversity (Nilon & Huckstep
1998), nitrogen application to lawns (Law et al. 2004), aquatic food web complex-
ity (Overmyer et al. 2005), ambient noise levels (Forkenbrock & Schweitzer 1999;
Warren et al. 2006) and temperature regime ( Jenerette et al. 2007). The levels of
confidence in these associations vary. For example, associations with nitrogen
application and aquatic food web complexity are represented by just one study
each, with low statistical replication (Law et al. 2004; Overmyer et al. 2005).
The amount of vegetation shows the strongest general patterns of association
with social stratification. Across a wide variety of studies in cities around the
world, wealthier neighbourhoods appear to harbour more greenery: greater
tree cover (Talarchek 1990; Iverson & Cook 2000; Smith et al. 2005; Grove et al.
2006; Kirkpatrick et al. 2007), more areas with trees and other vegetation
(Grove & Burch 1997) and greater vegetation density (Hope et al. 2003; Martin
et al. 2004; Jenerette et al. 2007). There are profound social and ecological
implications of this association. Greater vegetation cover, and tree cover in
particular, increases the delivery of many key ecosystem services, from regulat-
ing water flows and climate, to providing cleaner air (with some exceptions), to
harbouring greater biodiversity (e.g. Jenerette et al. 2007; Tratalos et al. 2007;
Wang et al. 2008). Trees and other vegetation may provide a variety of other
human benefits, including fostering increased physical activity (Frumkin
186 P. S. WARREN, S. L. HARLAN, C. BOONE ET AL.

Figure 8.3 Social stratification and housing age are strong predictors of vegetation
cover in Phoenix, Arizona, leading to effects on local climates and ecosystem services.
Reproduced with permission from Jenerette et al. (2007).

2003), lowering urban crime (Kuo et al. 1998) and increasing property values
(Morancho 2003; but see Troy & Grove 2008). In Phoenix, Arizona, higher-
income areas having higher vegetation cover are cooler (Figure 8.3; Jenerette
et al. 2007), leading to inequities in risk of heat exhaustion in this hot desert
city (Harlan et al. 2006). Jenerette et al. (2007, p. 362) point out that ‘along with
residing in warmer locations, lower-income inhabitants may be less able to
afford air-conditioning and may have fewer opportunities to avoid high
temperatures’.
A few studies have found that plant community structure may also be
associated with income level. In Phoenix, Arizona, two studies find that plant
diversity is strongly associated with income level at the census block group
level in parks and residential yards, and across a random selection of urban
sites (Hope et al. 2003, 2004; Martin et al. 2004). Higher-income neighbourhoods
are also more likely to have cacti or other desert-adapted plants (Martin et al.
2004). Particular forms of garden or landscaping are also associated with
income and other social stratification variables in two Australian studies
(Kirkpatrick et al. 2007; Luck et al. 2009). Gardens with trees are more common
in high-income suburbs, as are ‘simple native’ gardens and ‘shrub-bush-tree’
gardens (Kirkpatrick et al. 2007). All of these garden types are likely to support
native bird species in that region (Daniels & Kirkpatrick 2006). Low-income
areas, and those with high proportions of unemployed and renting residents,
are more likely to have ‘non-gardens’ and ‘no-input exotic gardens’ (Kirkpatrick
et al. 2007). In the classic work of Whitney and Adams (1980), two types of tree
communities are associated with higher-income areas in Akron, Ohio: ‘old oak’
and ‘mixed suburban’, and these two types contain the sites with highest tree
species diversity. The strong association of animals with particular plant
 URBAN ECOLOGY AND HUMAN SOCIAL ORGANISATION 187

assemblages would suggest that these differences are likely to have effects on
animal community structure and other ecological processes that are as pro-
found as the effects of total vegetation cover.
A small but increasing number of studies finds associations between social
stratification and animal communities: mammal species diversity in St Louis,
Missouri (Nilon & Huckstep 1998) and bird species diversity in Phoenix (Hope
et al. 2004; Kinzig et al. 2005), Vancouver, British Columbia (Melles 2005) and
Chicago, Illinois (Loss et al. 2009). In Phoenix and Vancouver, total bird species
richness is higher in parks as well as in residential neighbourhoods in census
units with higher income and education level and with lower proportions of
non-white residents, all classic indicators of social stratification (Kinzig et al.
2005; Melles 2005). Chicago, by contrast, seems to have higher native species
richness in lower-income areas, with higher exotic species richness in higher-
income areas (Loss et al. 2009). The differences in the findings may be due to
differences in study design. But they may also reflect the complex interactions
of social stratification with dynamic change over time in neighbourhood
occupancy. Phoenix is a post-World War II city, dominated by suburbs with
relatively invariant lot sizes (Gammage 1999). Most of the variation in bird
habitat in residential areas of Phoenix is likely to be due to vegetation charac-
teristics in yards, characteristics known to covary with income level in Phoenix
and elsewhere (Figure 8.4; Kinzig et al. 2005). Chicago, by contrast, is a pre-
industrial city with steeper gradients in built structure from one portion of the
city to another. Moreover, the study sites used by Loss et al. (2009) included a
variety of land-uses, such as residential areas, parklands and vacant lands,
while the Phoenix and Vancouver studies focused on residential areas. Since
urbanisation gradients are clearly a dominant factor in structuring bird com-
munities (Chapter 5; McKinney 2002), further studies may find a more accurate
signature of social stratification on bird communities by carefully controlling
for other factors like land-use and housing density (e.g. Mennis 2006).
Many of the spatial patterns described above are difficult to disentangle from
temporal ones. Time and space are intertwined in the development of socio-
spatial divisions. Processes like disinvestment and suburban flight can lead to
the formation of lower-income neighbourhoods in formerly moderate- to high-
income ones (Pickett et al. 2008). Vegetation structure appears to vary with
housing age in nonlinear ways (Grove et al. 2006), with tree cover increasing
over the early decades, but reaching an inflection point in Baltimore, Maryland,
at around 40 years old (Figure 8.5). For neighbourhoods of a fixed density,
vegetation can mature and become more complex, supporting more species,
over time. This process of maturation can lead to time-lags in the relationships
between social processes and vegetation patterns (Troy et al. 2007; Boone et al.
2009; Luck et al. 2009). Effects of time may also be dependent on ecological region.
New housing on old fields has a different impact on bird communities than new
housing on former forest land (DeGraaf & Wentworth 1986; Loss et al. 2009).
188 P. S. WARREN, S. L. HARLAN, C. BOONE ET AL.

Social, Indices:
economic & a Income, education, ethnicity
political Marketing data (e.g. Claritas)
processes

b
Distribution of
habitat,
resources c
and predators

Avian Key:
community causation
structure correlation

Figure 8.4 Diagram of hypothesised processes responsible for observed correlations

between income level and patterns of bird community structure. Social, economic
and political processes drive sociospatial divisions in cities (see also Box 8.1), leading to
variation in vegetation structure, resources and predators. These in turn drive variation
in bird community structure. Socioeconomic indicators such as income level serve as
indicators of sociospatial divisions (a); a growing body of literature finds that vegetation
structure is correlated with socioeconomic indicators (b); and recent studies have
found correlations between bird species diversity and income level at the
neighbourhood (census unit) scale (c) (Kinzig et al. 2005; Melles 2005; Loss et al. 2009),
suggesting that sociospatial divisions have implications for biotic structure at higher
trophic levels as well.

In arid environments, time may lead to a steady attrition of plants from

periodic drought, with wealthier residents capable of combating this attrition
with investment in water and re-planting (Martin et al. 2004). Water conser-
vation policies in arid cities may also lead to shifting emphases in plantings
toward native or drought-resistant species which generally support greater
diversity of native animals (McIntyre & Hostetler 2001; Daniels & Kirkpatrick
2006). Wealthier residents may be better able to accommodate these shifts; as
noted above, cacti (both native and drought resistant) are more common in
wealthier neighbourhoods in Phoenix (Martin et al. 2004). Historic shifts in the
patterns of housing and growth lead to apparent effects of age on plant and
animal communities, not due to differences in the maturation of vegetation.
For example, in cities like Chicago and Vancouver, age of neighbourhood is
associated with housing density (older neighbourhoods at higher density),
itself a potential factor driving patterns of diversity (Melles 2005; Loss et al.
2009). Thus, as cities age, processes of vegetation maturation overlay socio-
spatial patterns of investment and disinvestment, perhaps leading to the non-
linear relationships between age and vegetation cover found in Baltimore.
 URBAN ECOLOGY AND HUMAN SOCIAL ORGANISATION 189

0.25

0.2

0.15
P (Y = 1)

1 2

0.1

0.05

0
1 5 9 13 17 21 25 29 33 37 41 45 49 53 57 61 65 69 73 77 81 85
Median house age

Figure 8.5 Nonlinear changes in vegetation through time in Baltimore, Maryland, USA,
a pre-industrial city. Graph shows the probability that tree cover is equal to 100% of
private land as a function of median age of block group, based on quasi-likelihood logit
regression. Tree cover increases steadily for the first few decades, reaching an inflection
point in neighbourhoods of about 40 years old. Modified with permission from Grove
et al. (2006).

Other social dynamics

Alongside the socioeconomic divisions at the level of the city are finer-scaled
variations associated with the activities that people engage in around their
homes. For example, several authors have noted that front yards have become a
site for the ‘expression of individuality and ideology in a consumer society’
(Kirkpatrick et al. 2007, p. 314; see also Bhatti & Church 2004). Grove et al. (2006,
p. 580) argue that ‘a household’s land management decisions are influenced by
its desire to uphold the prestige of its community and outwardly express its
membership in a given lifestyle group’. Grove et al. go further to argue that the
social pressures exerted to adhere to group norms can have a stronger effect on
some environmental conditions than social stratification. They coin the phrase
‘ecology of prestige’ to describe this alternative process (Grove et al. 2006). In a
series of studies, they show that social stratification is a strong indicator of
‘plantable space’ or the amount of pervious cover in parcels (Troy et al. 2007). By
contrast, they find that the actual vegetation cover (trees versus grass versus
bare ground) is associated with finer social classifications according to lifestyle
(Grove et al. 2006; Troy et al. 2007).
Larsen and Harlan (2006) also refer to the ‘symbolic presentation of self ’
represented by front yards and gardens. They find differences in preferred
190 P. S. WARREN, S. L. HARLAN, C. BOONE ET AL.

Figure 8.6 Exemplars of landscape types used in a survey of landscape preferences

and behaviour in Phoenix, Arizona, USA. Both macro-level processes (legacies from
developers’ plantings), and micro-level processes (residents’ preferences) influenced
actual landscape types in residents’ front yards. Preferences for frontyard types
varied significantly with income level, with lower-income homeowners tending to
prefer lawn, middle-income homeowners preferring desert, and higher-income
homeowners evenly divided between desert and oasis. Reproduced with permission
from Larsen & Harlan (2006).

frontyard and backyard types associated with income level (Figure 8.6). They
caution, however, that particularly in newer portions of cities, developers’
choices function as a macro-level force constraining the degree to which, for
example, frontyard landscapes reflect the choices of individual residents (see
also Kirby et al. 2006). Larsen and Harlan (2006) found evidence of macro-level
forces shaping actual frontyard landscapes through developers’ planting
legacies.
Market analysis has been advanced by some authors as a method of quanti-
fying lifestyle and group identity and mapping these additional sociospatial
divisions in cities (Weiss 2000; Kinzig et al. 2005; Grove et al. 2006; Fuller et al.
2008). Marketing datasets incorporate records of purchasing decisions,
reasoning that people’s purchases reflect their group identities and ideologies
(Grove et al. 2006). These datasets have been criticised, however, for failing to
account for informal economic networks, thereby underestimating purchasing
power in lower-income communities (Pawasarat & Quinn 2001).
 URBAN ECOLOGY AND HUMAN SOCIAL ORGANISATION 191

Distinctions between the ‘ecology of inequality’ and the ‘ecology of prestige’

are their emphases on processes operating at different spatial scales and
different levels of social organisation (Figure 8.2). The ecology of prestige
emphasises the actions and activities engaged in by urban residents at local
scales that shape local environments, typically the household and neighbour-
hood (Figure 8.2; Grove et al. 2006). In contrast, theories of social stratification
emphasise the political and economic forces operating at larger scales to
constrain both real and perceived options available to urban residents, thereby
yielding strong inequities in local environmental conditions (Massey 1996; see
also ‘Spatial organisation’ above). Larsen and Harlan (2006) refer to these two
sets of processes as ‘micro-level’ and ‘macro-level’.
Ethnic and culturally mediated differences in preferences and values form a
related but distinct component of the sociospatial organisation of cities. As
outlined above, racial and ethnic divisions tend to be reinforced through
processes of social stratification. While this often leads to poorer economic
conditions for minority groups, such divisions also contribute to colourful
heterogeneities in both built and unbuilt portions of cities (Whyte 1943; Wirth
1956; Duneier 1999; Johnston et al. 2007). This human diversity is imprinted on
the architecture and landscape architecture of ethnic enclaves. The field of
environmental psychology also explores culturally mediated preferences for
different environments, such as the amount and configuration of trees versus
other vegetation (Kaplan & Talbot 1988). Much work remains to be done in this
area to understand whether and how these differences in environmental
preferences lead to varying ecological outcomes, although there are sugges-
tions that this is the case (Fraser & Kenney 2000).

Ecological outcomes from household decisions

Many small-scale decisions about land management occur at the household or
property parcel level through gardening and maintaining lawns, keeping
swimming pools, feeding birds and maintaining outdoor pets. These actions
can provide resources such as habitat and food for birds, bugs, mammals,
amphibians and other wildlife (Gaston et al. 2007), but they can also have
negative impacts on other ecosystem functions such as water quality (e.g.
runoff of pesticides and fertiliser), and animal populations (e.g. pesticides and
introduced predators, such as cats; Law et al. 2004; Lepczyk et al. 2004;
Overmyer et al. 2005). Because residential landscapes represent a large percentage
of urban land cover (Cannon 1999; Martin et al. 2003; Chamberlain et al. 2004),
private landowners collectively have a profound impact on urban ecosystems.
Landscaping is a multi-billion-dollar a year activity. In the UK, an estimated
60% of all households participate in gardening and spend £2.62 billion a year
on gardening products (Horticulture Trades Association Garden Industry
Monitor, unpublished data, cited in Gaston et al. 2005). The National Gardening
192 P. S. WARREN, S. L. HARLAN, C. BOONE ET AL.

Association (2007) estimated that 71% of Americans participate in gardening

activities, and that they spent $34 billion on lawn care and landscaping
products in 2007. The manner in which a particular garden is managed and
maintained influences urban biodiversity. Two studies in Australia found asso-
ciations between native plantings and the presence of native bird species
(French et al. 2005; Daniels & Kirkpatrick 2006). Burghardt et al. (2009) exam-
ined urban food webs in southern Pennsylvania, comparing butterfly larvae
and bird distribution in gardens landscaped with native plants to those in
gardens landscaped with exotic species. They also found that native landscap-
ing supported higher abundances of native birds and butterflies.
In cultivating lawns and gardens, humans generally remove weeds and pests,
as evidenced by the amount of pesticides and herbicides applied to residential
gardens. Unfortunately, these activities have negative implications for biodiver-
sity both directly (loss of invertebrate fauna) and through altered trophic
dynamics (Blackburn & Arthur 2001; Lepczyk et al. 2004). For example, pesticide
applications greatly decrease insect diversity and density, thus eliminating a
potential food source for insectivorous birds. Furthermore, as evidenced by
Carson’s (1962) exposure of DDT and the many toxicology studies since (e.g.
Bishop et al. 1998a, 1998b; Brickle et al. 2000), pesticides have devastating
impacts on the reproductive health, growth and survival of birds and other taxa.
Bird feeding is a popular activity with more than 52 million people feeding
wild birds in the USA (US Fish and Wildlife Service 2002) and about 12 million
UK households (48%) feeding garden birds (Davies et al. 2009). Like gardening,
feeding birds is a multi-billion-dollar endeavour. An estimated $30 billion per
annum is spent on seed and feeders according to a US Fish and Wildlife survey
in 2001 (US Fish and Wildlife Service 2002). Numerous studies have shown that
feeding birds benefits the people participating in the activity. Bird feeding
fosters a connection between people and the natural world (Fuller et al.
2007b). This exposure leads to an increased understanding of ecological issues
and thus allows people to incorporate sound ecological initiatives into public
policy (Rosenzweig 2003; Turner et al. 2004; Miller 2005). In a study across
Sheffield, UK, Fuller et al. (2008) found that densities of typical bird feeder
species were strongly correlated with the densities of bird feeders, although
species richness did not have a significant relationship. Lepczyk et al. (2004)
surveyed private landowners in southeast Michigan to determine their influ-
ence on bird populations. Two out of three respondents engaged in some level
of bird feeding, and fed birds on average nine months out of the year. One
elegant study of supplemental feeding replicated the behaviour of how indi-
vidual landowners feed birds ( Jansson et al. 1981). The authors concluded that
winter survival probabilities of willow tits Parus montanus and crested tits
Lophophanes cristatus increased because of the supplemental foods. Backyard bird
feeding most likely has contributed to the northern expansion of both the
 URBAN ECOLOGY AND HUMAN SOCIAL ORGANISATION 193

northern cardinal Cardinalis cardinalis and American goldfinch Caruelis tristis in

North America (Morneau et al. 1999).
A few studies have shown negative impacts of bird feeding (reviewed by
Jones & Reynolds 2008; Robb et al. 2008). The concentrated food source leads
to greater contact, with potential for increased transmission of disease. In
eastern North America, house finches Carpodacus mexicanus have encountered
an increase in mycoplasmal conjunctivitis (Dhondt et al. 2005). Bird feeders also
provide resources for avian predators (e.g. corvids), pest and exotic species
(Daniels & Kirkpatrick 2006; Marzluff & Neatherlin 2006). The higher densities
of these predators might lead to increased nest predation, although this has yet
to be tested. Effects of bird feeders on predation risk may vary according to life
stage. For adult birds, the aggregation of birds around feeders reduces the
risk per individual through dilution effects and increased levels of vigilance
(Dunn & Tessaglia 1994). Woods et al. (2003) actually found a negative relation-
ship between bird feeders and risk of cat predation.
The regular food source provided by bird feeders has been implicated in the
cessation of migration in some populations of some bird species ( Jokimäki et al.
1996). This leads to higher densities of sedentary species which possibly
increases competition with migratory species ( Jansson et al. 1981), potentially
contributing to reduced species diversity (Clergeau et al. 1998). Concerns voiced
by some bird enthusiasts regarding the dependence of birds on feeders have
not been supported. Studies in the UK (blue tits Cyanistes caeruleus), USA (Florida
scrub jay Aphelocoma coerulescens) and Australia (Australian magpie Cracticus
tibicen) demonstrated that the majority of food provisioning to nestlings con-
sisted of natural food items despite the prevalence of human-subsidised food
items (Cowie & Hinsley 1988; Fleischer et al. 2003; O’Leary & Jones 2006).
Poor nutritional content of food provided by humans might have detrimen-
tal effects on health and behaviour, though long-term damage is unknown.
When Australian magpies were provided with processed meats, this led to
elevated levels of plasma cholesterol (Ishigame et al. 2006). When Florida scrub
jays were given high-fat protein food supplements, they foraged 12% less than
birds in wildlands without supplementation (Schoech et al. 2004). Net effects of
supplemental feeding on avian populations and communities remains an
important area for further research.
Although bird feeding is by far the most popular wildlife gardening activity
(Gaston et al. 2007), a number of other features also provide resources for
wildlife. Lepczyk et al. (2004) found that 50% of those surveyed provided nest
boxes in their yards. The nest boxes could provide significant assistance for
cavity nesters in urban and suburban landscapes.
Collectively, garden features that enhance urban wildlife presence and sur-
vival could have a profound effect on urban biodiversity. Furthermore, land-
owners participating in at least one wildlife gardening activity are more prone
194 P. S. WARREN, S. L. HARLAN, C. BOONE ET AL.

to participate in additional activities (Lepczyk et al. 2004; Gaston et al. 2007),

thus increasing the benefits for a variety of taxa. Participation in these activ-
ities may provide a variety of benefits, directly and indirectly to the people
engaging in them (Miller 2005; Fuller et al. 2007b).

Synthesis and directions for future research

Our review of the social organisation of cities finds a rich literature in the
social sciences addressing potential processes underlying spatial structuring of
ecological outcomes in cities. They point to three social divisions in cities –
function, status and culture (Box 8.1). Signatures for all three social divisions
can be found in ecological outcomes. Much more attention has been devoted,
however, to ecological impacts of functional divisions (e.g. land-use effects)
than to status or cultural divisions. Interactions between these divisions and
temporal dynamics of technology and economy underlie variations in urban
forms. This differentiation in form among cities, along with ecoregional effects
and effects of city size on social organisation, may account for some of the
differences found among cities in patterns of biodiversity and ecosystem func-
tioning. Ecologists have not yet attempted to derive generalisations about the
ecological consequences of differences in urban morphology – e.g. pre-industrial
versus post-World War II cities. One finding of concern is that socioeconomic
inequities extend to ecological inequities, with lower-income portions of cities
typically having less vegetation and in some cases lower biodiversity than
wealthier portions. There are significant potential consequences of these
inequities for the delivery of ecosystem services, from climate regulation to
clean air and to aesthetic and recreational services.
Beneath these general findings, however, lie broad gaps in knowledge. Ecolo-
gists have yet fully to capitalise on the potential for integration with the social
sciences. Even basic description of the patterns of association between biotic
structure and socioeconomic divisions still has far to go before truly generalis-
able findings can be drawn. Studies should move beyond the well-trodden
ground of bird and plant community structure to address ecological processes
like trophic dynamics, extinction–colonisation processes and ecosystem func-
tions such as nutrient cycling. Finally, greater attention needs to be paid to the
developing countries, where the most rapid urban growth is occurring and
where disparities in income, health and environmental conditions on the
margins of cities are far more stark than those in industrialised countries.

Acknowledgements
We thank G. Cowgill, J. Novic, M. Smith, B. Stanley, B. Stark and A. York for
enlightening discussions and many references from our interdisciplinary
research project, ‘Urban Organization through the Ages’. We thank M. Nation
for her assistance with gathering material for this chapter; any omissions,
 URBAN ECOLOGY AND HUMAN SOCIAL ORGANISATION 195

however, are ours and not hers. This material is based upon work supported by
the National Science Foundation (NSF) under grant number DEB-0423704,
Central Arizona–Phoenix Long-Term Ecological Research (CAP LTER) and the
Baltimore Ecosystem Study (BES LTER), grant number DEB-0423476. Any opin-
ions, findings and conclusions or recommendation expressed in this material
are those of the authors and do not necessarily reflect the views of the National
Science Foundation. We thank K. Gaston and two anonymous reviewers for
many helpful comments.

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CHAPTER NINE

Urban ecology and human health

and wellbeing
JO BARTON AND JULES PRETTY

The importance of urban greening

Westernised societies are becoming more and more urbanised, and throughout
the twentieth and twenty-first centuries the number of people living in urban
settings has steadily increased. More than half of the world’s population
currently live in urban areas (UNFPA 2007) and this proportion is still set to
increase (Pretty 2007). Urban environments expose people to many stressors,
such as traffic noise and congestion, crowding and fear of crime, and are often
a source of continual demands prohibiting restoration from mental fatigue
(van den Berg et al. 2007). Everyday life revolves around complex information
processing activities requiring directed attention (Kaplan & Kaplan 1989). Our
capacity for this type of concentrated attention is finite, so it is regularly taxed
to its limit, leading to mental fatigue. This is a state characterised by inatten-
tiveness, indecisiveness and increased irritability, and we have fewer cognitive
resources available to manage everyday tasks, leading to increased stress
(Kaplan 1995). To restore our capacity for directed attention, we need to spend
time in settings that utilise involuntary attention requiring no cognitive effort.
Having contact with nature and green spaces promotes this type of attention
restoration, alleviates fatigue and reduces stress. Thus, with ongoing urban and
suburban sprawl, the importance of access to nearby nature is paramount,
especially for those regularly exposed to the pressures of urban life.
The type of green space close to where people live and work is important for
the quality of life of urban citizens and for the sustainability of towns and cities
(Chiesura 2004). Green spaces are defined as ‘open, undeveloped land with
natural vegetation’ (CDC 2009) and there are many types of urban green
space, ranging from larger parks and gardens (community, formal and private),
city farms and urban agriculture to smaller-scale communal squares, allot-
ments and green roofs. Other types include canals and riverbanks, tree-lined
streets, cemeteries, woods and grasslands, cycling routes, disused railways,

Urban Ecology, ed. Kevin J. Gaston. Published by Cambridge University Press.

# British Ecological Society 2010.
 URBAN ECOLOGY AND HUMAN WELLBEING 203

school playing fields and pitches, golf courses, informal recreation areas and
amenity green spaces. Some of these urban green spaces join to form continu-
ous green corridors or networks linking towns, cities and the countryside. The
rural–urban fringe consists of new and reinstated areas of woodland, wetland,
meadow, nature reserves, parks and many other diverse natural habitats. It can
also transport countryside biodiversity to the urban doorstep, thus increasing
opportunities for urban dwellers to encounter rarer flora and fauna as well as
having regular contact with nature (Countryside Agency and Groundwork
2004). All of these sources provide an important direct link to nature for many
people and often represent their sole exposure to nature.

The benefits of urban greening

The presence of urban nature is important for a number of reasons including
improving human health and wellbeing (Kaplan & Kaplan 1989; Frumkin 2001;
Irvine & Warber 2002; Health Council of the Netherlands 2004), improving
behaviour and cognitive functioning (Wells 2000; Taylor et al. 2001), facilita-
ting social networking (Kuo et al. 1998; Ward Thompson 2002) and exercise
(Giles-Corti & Donovan 2002; Giles-Corti et al. 2005), reducing levels of crime,
aggression and violence (Kuo & Sullivan 2001a, 2001b), providing an outdoor
classroom (Kaplan & Kaplan 1989; Kahn & Kellert 2002) and improving its
aesthetic value (Sheets & Manzer 1991).

Improving human health and wellbeing

The presence of green spaces and vegetation in the built environment can
influence human health and wellbeing (Judd et al. 2002; Frumkin 2003; Frumkin
et al. 2004). These terminologies are often used interchangeably, but the term
‘health’ incorporates physical health, mental or emotional health, social health,
spiritual health, lifestyle and functionality. The World Health Organization
(WHO) definition of health is still the most widely cited and states that ‘health
is a state of complete physical, mental and social wellbeing, and not merely the
absence of disease or infirmity’ (WHO 1948). A universal definition of ‘wellbeing’
is not available, as many sources interpret and define it differently. However,
wellbeing is generally considered in a broader context, and the UK Department
for Environment, Food and Rural Affairs has collaborated with other govern-
ment departments and stakeholders to develop a shared understanding of the
meaning of wellbeing within a policy context (Box 9.1; DEFRA 2007).
Attempts to establish a potential link between the urban environment and
an individual’s mental health have shown that the prevalence of psychiatric
morbidity is greater in urban areas and less common in rural domains, after
adjusting for confounding variables (Lewis & Booth 1994; White & Heerwagen
1998; Galea et al. 2005). For example, Lewis and Booth (1994) found that urban
residents’ prevalence of psychiatric morbidity (33.7%) was higher than that of
204 J. BARTON AND J. PRETTY

Box 9.1 Shared understanding of wellbeing

‘Wellbeing is a positive physical, social and mental state; it is not just the
absence of pain, discomfort and incapacity. It requires that basic needs are
met, that individuals have a sense of purpose, that they feel able to achieve
important personal goals and participate in society. It is enhanced by conditions
that include supportive personal relationships, strong and inclusive communities,
good health, financial and personal security, rewarding employment, and a
healthy and attractive environment. Government’s role is to enable people
to have a fair access now and in the future to the social, economic and
environmental resources needed to achieve wellbeing. An understanding
of the effect of policies on the way people experience their lives is important
for designing and prioritising them.’ Source: DEFRA (2007).

their rural counterparts (24.8%), after controlling for socioeconomic and other
extraneous variables. Income-related inequalities in health also depend on
exposure to green space. People who live in greener areas reported lower levels
of health inequality relating to income deprivation for both all-cause mortality
and mortality from circulatory diseases (Mitchell & Popham 2008).
A direct link between the amount of accessible local green space and health has
also been evidenced using large-scale epidemiological studies (Takano et al. 2002;
de Vries et al. 2003; Grahn & Stigsdotter 2003). In one, tree-lined streets, parks and
other green spaces were found to play a key role in longevity and decreased risk of
mental ill-health (Takano et al. 2002). This longitudinal study compared access to
local walkable green spaces and mortality rates in elderly residents of Tokyo,
Japan, over a period of five years. After controlling for demographic and socioeco-
nomic variables, they found that out of 3100 Tokyo citizens born between 1903
and 1918, 71% were still alive in 1992 and the probability of living for an
additional five years was linked to their ability to walk in a local park or tree-
lined street (Takano et al. 2002). However, although the study asked respondents to
assess the availability of walkable green spaces in their neighbourhood, they did
not establish how frequently these spaces were actually used for walking.
Self-reported health data from over 10000 Dutch respondents was correlated
with national environmental data characterising the type and quantity of blue
(e.g. rivers, lakes, canals) and green spaces present in their neighbourhood.
Socioeconomic and demographic characteristics were controlled for selection
effects and the study reported that people living in greener neighbourhoods
enjoyed better general health (de Vries et al. 2003). The type of green space
did not seem to alter effectiveness, but the total amount of green space in
the living environment seemed to be the most relevant predictor. A criticism
of the study is that the environmental characteristics were separated into
 URBAN ECOLOGY AND HUMAN WELLBEING 205

neighbourhoods and all individuals within that particular area were classed as
having equal access to green spaces. This crude measure does not acknowledge
that the exposure to green space may vary considerably between residents of
the same neighbourhood and that durations of exposure may also differ.
In a separate study, one in ten residents felt unhealthy when the majority of
the space surrounding their home was green (90%). In contrast, when only 10%
of the environment was green, 16% of the residents felt unhealthy (Maas et al.
2006). Groenewegen et al. (2006) have set up the ongoing ‘Vitamin G’ project,
aiming to build on previous research analysing the relationship between
the amount and type of green space and health and wellbeing. The project
has three different levels: (i) national Dutch data; (ii) green spaces in urban
environments; and (iii) allotment gardens. It combines land-use data and self-
reported health states. The findings are going to inform policy development,
aid urban planning and design, and raise awareness of the importance of local
green pockets.
Perceived neighbourhood greenness is also strongly associated with better
mental and physical health. Respondents who perceived their neighbourhood
as highly green were 1.37 and 1.60 times more likely to have better physical
and mental health respectively, in comparison with those who perceived it as
low in greenery (Sugiyama et al. 2008). The degree of species richness in urban
green spaces has also been positively associated with psychological wellbeing of
visitors (Fuller et al. 2007), emphasising the importance of locally managed
biodiversity for sense of place and reflection.
In terms of overall health, local park users reported fewer visits to a physician
for purposes other than routine check-ups in comparison with non-park users
(Godbey et al. 1998). This difference was apparent even when controlling for the
effects of age, income, education level, health status and other potential con-
founding variables. Frequently active park users also scored better on self-reported
health indices and perceived their health states to be better than passive users and
non-park users (Godbey et al. 1992). Thus, people engaging in leisure recreation in
local parks seem to be in disproportionately better health than non-users and are
also less likely to be obese than the general population (Ho et al. 2003).
Godbey and Blazey (1983) also investigated the leisure behaviour of adults
participating in light to moderate aerobic activity in urban parks and found
that over half reported better moods after visiting the park. More and Payne
(1978) also found that participants’ negative moods improved and that park
users reported lower levels of anxiety and depression. Often visitors started
their recreation experiences in a better mood and their positive moods
remained on leaving, implying that outdoor recreation and park use might
enhance positive moods, reduce negative ones and alleviate stress.
In a Swedish study, Grahn and Stigsdotter (2003) examined the relation-
ship between use of urban green spaces and health. They found that the level
206 J. BARTON AND J. PRETTY

of self-reported stress experienced showed significant relationships with the

proximity of urban green spaces, visiting frequency and duration of stay. The
findings implied that the more frequent the visits, the lower the incidence of
stress-related illnesses. Having access to a public or privately owned garden
adjacent to their place of residence was another principal factor, which has
implications for both policy and urban landscape planning.

Improving behaviour and cognitive functioning

Parents of children experiencing Attention Deficit Hyperactivity Disorder
(ADHD) reported that participating in activities such as fishing or camping in
green spaces improved behaviour in 85% of cases (Taylor et al. 2001). When
watching television or playing video games in indoor environments, behaviour
improved in only 43% of activities. In some indoor activities, behaviour actually
deteriorated (57%) and made the children less manageable. Following on from
the ideas of attention restoration described earlier, Wells (2000) conducted a
longitudinal study with children of low-income urban families and assessed
the effects of nature on their cognitive functioning. When the families were
relocated to houses with more nature in the window view, they had higher
levels of cognitive functioning and their ability to direct attention continued
for several months after moving. However, these findings should be treated
with caution because it could be argued that these types of families were able
to select these types of preferred homes. Therefore, cause and effect can be
difficult to disentangle and decipher.

Facilitating social networking

Green space in the form of parks, streets, squares and allotments can be
valuable in urban areas for facilitating social contact (Coley et al. 1997; Ward
Thompson 2002) and giving rise to stronger neighbourhood ties (Kuo et al.
1998). Green spaces can also foster social inclusion, community development,
citizenship and local pride by allowing local residents to assist in the design,
management and care of local spaces (DLTR 2002). Activities in green places
often occur in social groups, or indeed people undertake activities in order to
interact with others. Social capital is thus a component whereby relations of
trust and reciprocity tied together by social norms and institutions can help
people engage in activities, link to particular places and remain mentally and
physically healthy. Social capital is also closely tied to capacity for collective
environmental management (Pretty & Smith 2004).
Several key studies researching the link between nature and social contact
have all involved the same USA study population (Coley et al. 1997; Kuo et al.
1998; Kweon et al. 1998). The ‘Robert Taylor Homes’ (RTH) are located in public
social housing communities in an underprivileged area of Chicago. They con-
stitute a naturally occurring field experiment as residents are randomly
 URBAN ECOLOGY AND HUMAN WELLBEING 207

assigned to apartments, which are identical, with the exception of the quantity
of surrounding vegetation and greenness of common spaces. All other environ-
mental, cultural and social variables are held constant, and residents are socio-
economically homogeneous.
The presence of trees significantly increased the utilisation of public green
space by both adults and youths (Coley et al. 1997). The communal green spaces
provided opportunities for more face-to-face contacts and encouraged social
interaction. This led to stronger neighbourhood social ties, which were
assessed by the amount of socialising, contact with nearby neighbours and
local sense of community (Kuo et al. 1998). The greener areas promoted the
strongest neighbourhood social ties, although the reasons for visiting the
green spaces were not reported (e.g. requiring shade from trees in hot summer
months). In addition, the exposure time to communal green spaces was posi-
tively linked to social integration of elderly residents in the community (Kweon
et al. 1998). Outdoor spaces dominated by trees and grass had the greatest
effect, and active use of these spaces predicted the strength of neighbourhood
social ties and sense of community.

Facilitating green exercise activities

The term ‘green exercise’ stems from a programme of research which aims to
investigate the synergistic benefits of engaging in physical activities whilst
simultaneously being directly exposed to nature (Pretty et al. 2003; Barton
2009). Access to urban green spaces has been shown to promote healthy living
by encouraging participation in green exercise activities, such as walking,
jogging and cycling (Ross 2000; Berrigan & Troiano 2002; Craig et al. 2002;
Handy et al. 2002; Parks et al. 2003; Wendel-Vos et al. 2004; Bedimo-Rung et al.
2005; Godbey et al. 2005). However, the state and design of the surrounding
environment can either be conducive or restrictive to activity participation.
Socioeconomic variants in health outcomes are often determined by the imme-
diate environment and thus the individual’s behavioural preferences (Owen
et al. 2000). Typical physically active behaviours may include walking for
exercise or recreation, jogging or participating in a sporting activity, whereas
sedentary behaviours comprise sitting, socialising, spectating or dining. There-
fore, behaviour settings can potentially influence the level of activity experi-
enced and can either encourage or prohibit participation.
Large-scale Australian studies found that accessible public open spaces, such
as parks, were used more frequently for physical activity, although this effect
was significantly moderated by attractiveness and size (Giles-Corti & Donovan
2002; Giles-Corti et al. 2005). People with easy access to an attractive and large
public open space were 50% more likely to exceed physical activity recommen-
dations. Open spaces incorporating trees, water features and birdlife were
commonly used for walking or jogging (64%) and cycling (12%) activities.
208 J. BARTON AND J. PRETTY

Other studies have reported the physically active behaviour of visitors to

parks (Scott 1997; Godbey et al. 1998; Raymore & Scott 1998; Tinsley et al. 2002).
The average visitor spends about half of their time walking when visiting a
park, and seven out of ten park visitors engage in moderate to vigorous levels of
physical activity (e.g. brisk walking, cycling, jogging; Godbey et al. 1998). Par-
ticipating in vigorous physical exercise was reported to be the most important
outcome for many park users (Tinsley et al. 2002). Other studies estimate that
7% of urban park users in England visit parks to engage in sporting activities,
such as football, bowls, golf and cycling (CABE Space 2004), which represents
7.5 million visitors per annum (Woolley 2003). Walking increases if the local
area is attractive and scenic, and if there are safe footpaths and pavements, a
diversity of land-use, easy access to public transport, a friendly neighbourhood
and ease of accessibility (Humpel et al. 2002; Bird 2004; Owen et al. 2004).
Landowners and managing agencies therefore have a role to play in delivering
these requirements to increase visitor numbers and activity levels.
Parks et al. (2003) demonstrated a dose–response relationship between
the number of places available for exercise within a neighbourhood and
the probability of meeting physical activity recommendations. The majority
of people who exercise choose to do so in their local park, thus indicating
that exercise frequency and park use are both associated with park proximity
(Cohen et al. 2007). Bedimo-Rung et al. (2005) propose a conceptual model
to describe the relationships between park characteristics, such as the number
of visits and physical activity levels within the park, park use and overall
benefits, including physical and psychological health, social, economic and
environmental.
Residential environments with large amounts of green and minimal graffiti
and litter have been found to be associated with increased physical activity and
a reduced incidence of obesity (Ellaway et al. 2005). Residents of greener envir-
onments were 3.3 times as likely to participate in regular exercise compared
with those living in areas with nominal greenery. Although the study was cross-
sectional, the findings supported other research studies which also reported
improved physical health when good-quality, well-maintained public spaces
were easily accessible (CABE Space 2004). Thus, access to nature and green
spaces seem to play an important role in increasing physical activity levels in
urban communities.
Grahn and Stigsdotter (2003) suggest that urban citizens who live 50 m or
less from their nearest green space visit it three to four times per week. If the
distance is increased to 300 m, the number of visits reduces to an average of
2.7times per week. However, a distance of 1000 m reduces the number of visits
to only once a week. Residents of communities lacking greenery in their local
area do not compensate for this by visiting public parks or urban forests more
frequently, which highlights the importance of restoring and conserving
 URBAN ECOLOGY AND HUMAN WELLBEING 209

nearby local green space. There are also physical, social or cultural barriers
which may restrict usage even when green space is available. Perceptions of
personal safety and fear of crime may discourage visits, along with cultural
barriers where people feel that they are not allowed to use the spaces.

Reducing levels of crime, aggression and violence

Kuo and Sullivan (2001a, 2001b) have made compelling links between small
amounts of green in the urban environment of Chicago’s poorest public hous-
ing neighbourhood and crime, aggression and domestic violence. Vegetation
levels surrounding RTH homes were assessed by an independent panel using a
combination of photographs from a number of vantages. The assessment took
place in June when the grass was green and the tree canopy was in full leaf.
Residents living in greener surroundings reported lower levels of fear, fewer
incivilities and less generalised aggressive and violent behaviour. Buildings
with more vegetation also had 52% fewer property and violent crimes than
those with minimal vegetation. There is often a perception that well-vegetated
places offer more opportunities for criminals and drug-dealers to hide, so these
findings raise some interesting questions.
There was also a greater difference between buildings in non-green and
moderately green surroundings than between moderately and very green,
suggesting more of a benefit would accrue from a light greening of all urban
spaces rather than a dark greening of just a few. Indeed, well-maintained
vegetation may instigate new ways of thinking as local people start to care
for their environment and so are more vigilant (Kuo & Sullivan 2001a). Levels
of aggression and violence were also significantly higher for residents in
conditions lacking vegetation than for those who had access to nearby nature
(Kuo & Sullivan 2001b). Access to greener surroundings reduced aggression
by increasing concentration, which mediates the relationship between mental
fatigue and aggressive behaviour. Therefore, neighbourhood settings lacking
nearby nature and greenery had a significant impact on human social
functioning (Sullivan 2005).

Aesthetic value
Urban environments incorporating green spaces are perceived to be more
attractive than urban areas lacking vegetation (Sheets & Manzer 1991; Kuo
et al. 1998).
Visitors report a deep sense of personal satisfaction from experiencing the
aesthetic pleasures, such as ‘enjoying the changing seasons, feeling the sun,
the wind or the rain, being able to walk, run or just sit down and enjoy the
view’ (Burgess et al. 1988). Environmental aesthetics are also positively associ-
ated with walking for exercise. Compared with people who had access to a very
pleasing aesthetic environment, those reporting a moderately aesthetic
210 J. BARTON AND J. PRETTY

environment were 16% less likely to walk for exercise, and those who only had
access to a low-rated aesthetic environment were 41% less likely to walk for
exercise (Ball et al. 2001). Research figures report that 85% of people questioned
believe that the quality of public space and the built environment directly
affects their lives and the way they feel (CABE Space 2002).

Providing an ‘outdoor classroom’

Urban green spaces also provide ‘outdoor classrooms’ to facilitate learning and
enhance knowledge of the natural world and local environment. Outdoor
classrooms are important for children (Moore & Wong 1997; Kahn & Kellert
2002; Rickinson et al. 2004), with creative social play, concentration and motor
ability positively influenced by play in green space (Taylor et al. 2001). However,
this evidence has not yet been strong enough to change the design of schools,
though the emergence of the forest school movement is an indication that
green space is being seen as a contributor to positive cognitive outcomes
(Bishops Wood Centre 2005).

Levels of engagement with nature

We posit three levels of engagement with nature leading to potentially differ-
ent outcomes: viewing nature, functional engagement and active participation
(Figure 9.1).

3 Levels of Engagement with Nature

(1) Viewing nature – from the window or in a painting
(2) Functional engagement – incidental exposure to
nearby nature whilst engaging in some other activity
(passive or active)
(3) Active participation – positive decision to visit
nature and green spaces to participate in an activity
and interact with nature

1. Improves 2. Generates 3. Facilitates social

psychological physical health networking and
wellbeing benefits connectivity

Healthier communities and avoided

public health costs

Figure 9.1 Three levels of engagement with nature.

 URBAN ECOLOGY AND HUMAN WELLBEING 211

Viewing nature
Viewing nature through a window or in a photograph is an important way of
engaging with nature, when direct access is restricted. This type of passive
engagement allows the mind to digress and stimulates reflection and recovery,
aids recovery from illness, improves mood, reduces stress and improves mental
wellbeing.

From the home

Kaplan and Austin (2004) and Kaplan (2001) used black and white photographs
of views from the home to explore the importance of differing levels of vegeta-
tion for residential satisfaction. Participants were asked how closely the char-
acteristics in the view represented the view from their own home and rated
their preferences. The most preferred scenes were predominantly nature views,
and trees were strongly associated with feelings of relaxation. Views of gardens,
flowers and landscaped areas played an important role in participants’ residen-
tial satisfaction but characteristics that were most favoured were also rated as
least available. Woodlands enhanced residents’ satisfaction with their sur-
roundings and contributed to the community and sense of peacefulness
(Kaplan & Austin 2004). However, there may be an element of response bias
as the participants were aware of the aim of the study and were self-selecting so
results may have been skewed.
The value of the view from a window is also reflected in monetary terms as
various studies have demonstrated increased economic value for housing and
hotels (Peiser & Schwann 1993). The presence of green space affects room pricing
policy in hotels in Zurich, Switzerland (Lange & Schaeffer 2001), and increases
the value of homes with gardens overlooking lakes and paths in the Netherlands
by 25% (Luttick 2000). Street trees in Berlin, Germany, increase real-estate value
by 17% (Luther & Gruehn 2001), and the value of housing near to water is greater
in Merseyside, UK (Wood & Handley 1999; Lindsey et al. 2004).

From the workplace

Windows present in the workplace can buffer the stresses of work and reduce
the frequency of illness, headaches and frustration, and improve patience and
enthusiasm for work (Leather et al. 1998; Kaplan 2001). Workers with views of
trees and flowers have been shown to feel less stressed in their jobs and to
derive greater job satisfaction than workers overlooking built environments
(Kaplan 1993). A similar study also reported that natural views buffered the
negative impact of job stress on intention to resign (Leather et al. 1998). People
working in windowless workplaces were four times more likely to compensate
by displaying pictures of landscapes and outdoor natural scenes or indoor
plants (Heerwagen & Orians 1993).
212 J. BARTON AND J. PRETTY

From institutions
Two classic and widely cited studies from the 1980s (Moore 1982; Ulrich 1984)
suggest that prolonged exposure to window views of nature can have import-
ant health-related consequences. The first found that prisoners in Michigan,
USA, whose cells overlooked farmland and forests reported 24% fewer sick cell
visits compared with those in cells facing the prison yard. Cells were randomly
allocated so the findings implied a stress reduction effect. The second ‘classic’
study exploited the configuration of a hospital in Pennsylvania, USA, where
rooms in the surgical section overlooked deciduous trees or a brick wall. It
formed a 10-year comparative study of post-operative patients who had under-
gone identical surgical procedures. Patients were randomly assigned a room
which only differed in its view from the window. The hospital stay for those
patients with tree views was significantly shorter (7.96 days per patient com-
pared with 8.70); they also required fewer painkillers and took less strong or
moderate pain medication. Nursing staff also reported fewer negative com-
ments in the medical records for those with the tree views (1.13 per patient
compared with 3.96).

Whilst travelling
The view during the commute to work (e.g. the type and quality of roadside
verges) can also influence levels of stress. Participants in one study were
exposed to four different simulated types of roadside corridors: rural, urban,
golf course or mixed (lots of vegetation but visible commercial buildings;
Parsons et al. 1998). There was some evidence that those on the urban drive,
dominated by human artefacts, had higher levels of skin conductance, facial
muscle tension and blood pressure compared with other settings. Restoration
of standard heart rate readings was faster and more complete after viewing
scenes of golf courses. Viewing nature-prevalent drives of forests or golf courses
facilitated stress recovery quicker than urban settings. In addition, the nature
drives offered a protective effect against the negative consequences of future
stresses that might have arisen during the working day, implying that there
might be an immunisation effect (Parsons et al. 1998).

Using simulated scenes of being in nature

A number of studies have explored the effects of viewing simulated scenes of
being in nature by conducting experiments in laboratories. This setting allows
the limitation of potentially confounding variables and often participants have
been attentionally fatigued, using a variety of demanding mental tests, prior to
viewing a series of slides. Photographic simulations have predominantly com-
pared natural settings (e.g. green spaces, forests, woods, open countryside) with
contrasting urban scenes lacking nature. Stress reduction qualities of these
 URBAN ECOLOGY AND HUMAN WELLBEING 213

differential environments have been assessed using a combination of physio-

logical (blood pressure, heart rate, cortisol, muscle tension) and psychological
measures (e.g. self-reports of emotion, concentration or mood).
Nature slides incorporating water or vegetation were consistently preferred
to grey urban scenes lacking greenery and had a more positive effect on
emotional states (Ulrich 1981; Hartig et al. 1996). The natural settings sustained
attention more effectively, and for stressed or excessively aroused participants
they offered the greatest opportunity for stress reduction, with blood pressure,
muscle tension and skin conductance levels all reducing during recovery
(Ulrich et al. 1991). Participants’ heart rate recordings decreased whilst viewing
the nature video, whereas participants viewing the urban environments did
not report any change (Laumann et al. 2003). Therefore, this implies that
natural settings have a relaxing effect on autonomic functions. Recovery was
also faster when exposed to natural environments, which supports the theory
that nature exposures of short duration are important in urbanised societies.
When coloured slides depicting cityscapes with designated green spaces were
introduced, positive effect scores were significantly higher compared with no
greenery (Honeyman 1992).
Van den Berg et al. (2003) analysed the relationship between restorative
potential and environmental preference. Participants viewed a frightening
movie followed by either a video simulating a nature-based walk or a walk in
a built environment. Greater improvements in mood and concentration were
reported after viewing natural settings compared with viewing built environ-
ments, and the natural settings were perceived as more beautiful and restora-
tive. Higher levels of stress were associated with stronger preferences for
natural scenes and less liking for urban settings. Berto (2005) explored restora-
tive environments’ ability to facilitate recovery from mental fatigue by analys-
ing the relationship with increased attention performance. Mentally fatigued
subjects were exposed to either nature scenes (restorative) or urban streets
(non-restorative), and only subjects who viewed the natural scenes regained
sufficient attention capacity to perform well in a secondary task.
A study at the University of Essex, UK, tested the physiological and psycho-
logical health benefits of exercising on a treadmill whilst being exposed to a
series of rural or urban photographic scenes (Pretty et al. 2005). Each of these
was subdivided into pleasant and unpleasant categories in order to explore the
effect of rural scenes compromised with pollutants or other visual impedi-
ments (e.g. rubbish, abandoned cars, billboards or pipes carrying effluents)
and urban scenes enhanced by the presence of nearby nature in the form of
green space. A control group was included which involved exercising without
exposure to images. Only those subjects viewing rural pleasant scenes experi-
enced significant decreases in mean arterial blood pressure (Figure 9.2). The
urban pleasant pictures had no effect on mean arterial blood pressure, whilst
214 J. BARTON AND J. PRETTY

96 Rural pleasant
Rural unpleasant
Mean arterial blood pressure (mm Hg)
94 Urban pleasant
Urban unpleasant
92 Control

90

88

86

84

82
* Error bars   1 Standard error
80 *p < 0.001

Figure 9.2 Change in mean arterial blood pressure (MABP) after treadmill exercise
whilst viewing different scenes on a projector (change in MABP normalised to the
starting average for all five groups). See text for details.

the urban unpleasant slightly increased it. All participants viewing rural pleas-
ant pictures experienced a reduction in their mean arterial blood pressure
compared with only 60% of people when exposed to the other picture condi-
tions. As control subjects experienced a slight decrease in blood pressure, it is
clear that both pleasant and unpleasant urban scenes increased blood pressure
relative to the controls. The urban scenes therefore appear effectively to negate
the marginal, but potentially beneficial impact of exercise on blood pressure.
Self-esteem significantly improved in all five groups, but the control group
produced a greater improvement in self-esteem than the two unpleasant treat-
ments (rural and urban), implying that the latter have a depressive effect on
self-esteem relative to exercise alone. Both pleasant treatments, however, pro-
duced the greatest increases in self-esteem (Figure 9.3). When viewing both
urban and rural pleasant scenes, levels of self-esteem increased by 10% and 9%
respectively. This was in comparison with a 7.5% increase after viewing no
pictures and only a 4% increase after viewing unpleasant pictures.
For the six measures of mood, viewing rural pleasant scenes during exercise
produced consistent, though not always significant, improvements relative to
viewing other scenes. Viewing urban pleasant scenes also resulted in improve-
ments in all six mood measures. Unexpectedly, exercise whilst viewing urban
unpleasant scenes produced significant improvements for anger-hostility, con-
fusion-bewilderment and tension-anxiety. However, the rural unpleasant
scenes had the most differentiated effect on mood measures. There were
negative effects on three mood states, the most for any type of scene. This
suggests that views embodying threats to the countryside have a greater nega-
tive effect on mood than already urban unpleasant scenes.
 URBAN ECOLOGY AND HUMAN WELLBEING 215

Rural pleasant
Rural unpleasant
16 Urban pleasant
Urban unpleasant
Control
*** **
**
17
Index of self-esteem

**
*
18

19

Error bars   1 Standard error

20 *p < 0.05, **p < 0.01, ***p < 0.001

Figure 9.3 Change in self-esteem after treadmill exercise whilst viewing different
scenes on a projector (change in self-esteem normalised to the starting average
for all five groups). NB: High score ¼ low self-esteem. See text for details.

Functional engagement
The second category of engagement is more functional and often involves
incidentally being in the presence of nearby nature, whilst primarily engaging
in another activity. The nature setting acts as background scenery for
social activities such as walking the dog, cycling to work through an urban
park, reading on a garden seat or talking to friends in a park (Hayashi et al.
1999; Ulrich 1999). One of the first longitudinal studies conducted over a
decade targeted poorly functioning impoverished neighbourhoods undergoing
environmental improvements (Dalgard & Tambs 1997). The mental health
states of the residents were initially quite poor, but significant improvements
were reported post-intervention for those residents still present. Therefore,
selective migration was not the reason for this outcome, indicating that
health parameters could be positively influenced by changes in environmental
features.
A study at the University of Essex assessed changes in local people’s behavi-
our and health measures following ecological restoration of local green spaces
at three urban sites in the UK (urban park, canal and coastal path; Peacock et al.
2005, 2006). Prior to the restoration process all three sites were often unused,
but transforming these areas created new opportunities for outdoor recreation
and contact with nature and green space. Ecological restoration processes
involved re-landscaping grassland, creating a wetland environment, improving
biodiversity and improving canal towpaths and coastal paths. A higher propor-
tion of users visited the locations for all of the reasons listed post-restoration
216 J. BARTON AND J. PRETTY

Table 9.1 A comparison of the number and duration of visits before and after
restoration at three sites (see text for details).

Before After Increase (%)

No. of visitors (who completed question re visitation 133 150 12.8

patterns)
No. of visits per month 1535 2007 30.8
No. of visits per person per month 11.5 13.4 15.9

No. of visitors (who completed question re duration of visit) 129 144 11.6
Total time spent at site for all users during one visit (mins) 4580 6068 32.5
Average time spent at site per person (mins) 35.5 42.1 18.7
Total time spent at site per person per month (mins) 410 564 37.6

Note: A month refers to a 4-week period; the numbers of visitors do not match as calculations
do not include those who had never visited before the improvements or who had not completed
both sections of the questionnaire.

For exercise***

Walking the dog

Health–walking, fresh air etc.***

Reasons for visit

Scenery***
Before (%)
On the way to somewhere else After (%)
Wildlife***
Significance tested
Meet family or friends* with the McNemar test
(*p < 0.05; ***p < 0.001)
Other

Never visited before improvements

0 10 20 30 40 50 60 70
(%)

Figure 9.4 Primary reasons users visited sites before and after completion of ecological
restoration projects. See text for details.

(Figure 9.4). However, significant differences were found for exercise, meeting
family or friends, health, scenery and wildlife. The restored sites attracted new
visitors and many more users were now choosing to visit the sites to view the
scenery and wildlife and interact with the environment.
Although the environmental improvements encouraged more people to visit
the sites, they also visited more frequently and spent longer engaging with
nature on each visit (Table 9.1). During a 4-week period before restoration, a
total of 133 users visited the sites 1535 times, which equates to an average of
11.5 visits per person per month. However, following restoration of the sites, a
total of 150 people visited the sites 2007 times during an equivalent period of
 URBAN ECOLOGY AND HUMAN WELLBEING 217

5
Index of self-esteem

10
R 2 = 0.1104
15

20

25

30

35
0 10 20 30 40 50 60 70
Length of time exercising (mins)

Figure 9.5 Relationship between the length of time exercising at restored sites and
self-esteem scores. NB: High score ¼ low self-esteem. See text for details.

time. This equates to an average of 13.4 visits per person per month and
represents a significant increase ( p<0.0001).
On average, users were spending 35.5 minutes exploring the sites on each
visit prior to restoration. Following the environmental improvements, this
figure increased to 42.2 minutes and the total time spent at the sites per person
per month increased by 154.2 minutes, representing a significant increase of
37.6% ( p<0.0001).
A significant improvement in self-esteem was found when comparing
arrival scores with departing scores ( p<0.05). Thus, those individuals who
had been participating in physical activity for longer within the green sur-
roundings reported an improved self-esteem score. A positive correlation
between reported self-esteem scores and the length of time exercising within
the environment was found ( p<0.0001) (Figure 9.5). This implies that the
longer people spend exercising in urban green areas, the more their self-esteem
improves.

Active participation
The third category of engagement with nature is referred to as ‘active partici-
pation’. This differs from the second category as it implies a positive decision to
visit nature and green spaces and directly participate in an activity, such as
gardening, walking, mountaineering, running, cycling or water-based activ-
ities. Studies often use psychophysiological measures to compare walking in
urban nature parks to city streets. Findings have suggested that nature groups
report more positively toned emotional states, higher happiness scores, higher
ratings of positive effect, markedly better cognitive performance scores (Hartig
218 J. BARTON AND J. PRETTY

et al. 1991), a greater ability to reflect on problems (Mayer et al. 2006) and
improved directed attention abilities (Berman et al. 2008).
Other studies have included ambulatory blood pressure measurements
during walks in both nature reserves and urban areas with minimal landscap-
ing after sitting in a room with different views (Hartig et al. 2003). Sitting in a
room with tree views promoted more rapid diastolic blood pressure decline
than sitting in a windowless room. Walking in a nature reserve reduced blood
pressure more than walking along an urban, non-green street. After 20 minutes
of walking, the mean blood pressure values differed significantly between the
two settings, before the difference converged. This is probably because the
benefits of the exercise (the walk itself) started to surpass any unpleasantness
of the urban street. In both contexts, the green room and green walk, people
recovered more rapidly from attention-demanding tasks, regardless of antece-
dent condition. There were no significant differences in blood pressure read-
ings post environmental treatment, but there were visible effects on emotion.
The walk in the natural setting increased positive effect and reduced feelings of
anger/aggression, while the opposite occurred in the urban setting.
Research has also analysed runners’ cognitions and moods whilst exercising
in different settings (Bodin & Hartig 2003; Butryn & Furst 2003). A 1-hour run
through a nature reserve dominated by greenery, water and pleasant views was
compared with an urban route through sidewalks and streets with varying
traffic volumes and many buildings (Bodin & Hartig 2003). Running in a nature
reserve promoted emotional restoration more effectively than exercising in
the urban environment, although there were no significant differences in the
reduction in anxiety/depression and anger between the two conditions. The
low level of statistical power may have contributed to this, as there were only
12 runners. However, runners did state their preference for the nature reserve
as they perceived it to be more psychologically restorative. Butryn and Furst
(2003) also examined the effects of natural parks and urban settings on mood,
feeling states and cognitive strategies of non-elite female runners. Mood and
feeling states significantly increased following both runs, irrespective of set-
ting, but the natural vegetation park setting was overwhelmingly preferred
(93%) by the runners. Pre-run moods were elevated, perhaps in anticipation of
the run or from accumulated benefits from participation in regular runs.
Therefore, there was minimal room for improvement.
Mood change and stress during recreation in an urban park was also com-
pared to indoor leisure activities (Hull & Michael 1995). Findings suggested that
anxiety and energy levels decreased with time spent at the park but changes in
fatigue and calmness were minimal. A more recent study conducted by the
University of Essex assessed the role the environment plays in the effectiveness
of exercise for mental wellbeing with members of local Mind groups. Mind is a
mental health charity based in England and Wales and provides support for
 URBAN ECOLOGY AND HUMAN WELLBEING 219

17
*

18
Index of self-esteem

19
Indoor shopping-mall
walk
20

21
Green outdoor urban
22 park walk

23
Type of walk Error bars   1 Standard error
*p < 0.05
24

Figure 9.6 The change in self-esteem following participation in both walks (change
in self-esteem normalised to the starting average for both groups). NB: High score ¼ low
self-esteem. See text for details.

individuals experiencing mental distress. It provides numerous services to all

of its members through a network of local Mind associations. Local Mind
members participated in both a green outdoor urban walk (Belhus Woods
Country Park) and an indoor shopping-mall walk (Lakeside shopping centre),
both of which were of the same duration and intensity and led by the same
person, to ensure consistency of personality.
Improvements in self-esteem ( p<0.05) and overall mood ( p<0.01) were sig-
nificantly greater following the green outdoor urban walk than following
the equivalent indoor walk (Mind 2007; Peacock et al. 2007). Figure 9.6 com-
pares the significant changes in self-esteem after both walks, highlighting the
positive improvement after the green outdoor urban walk and the negative
effect after the indoor walk. Figure 9.7 illustrates the significant changes in the
subscale mood factors after both of the walks (anger, confusion, depression and
tension). Participants felt significantly less angry ( p<0.05), confused ( p<0.05),
depressed ( p<0.05) and tense ( p<0.01) after the outdoor green urban walk and
felt more confused and tense after the indoor walk.
The findings show that exercising outdoors in a green environment is a lot
more effective in enhancing mood and improving self-esteem than the equiva-
lent amount of exercise indoors. The enjoyment of engaging in green exercise
activities in groups was a valuable part of the experience, as well as the
opportunity to breathe in fresh air, admire the scenery and enjoy the wildlife.
The findings add significant value to the ever expanding green exercise
research programme as they focus on individuals experiencing mental health
issues and separate the elements that constitute the green exercise experience.
 (a) (c) Green outdoor urban
42
43 Green outdoor urban park walk
park walk
42

Index of depression
Index of anger 41
41

40 Indoor shopping-mall
40 Indoor shopping-mall walk
walk
39
39
38
*
37 Type of walk 38 Type of walk *

(b) (d)
42 41
41 40
Green outdoor urban
Green outdoor urban 39
40 park walk
Index of confusion

park walk 38

Index of tension
39
37 Indoor shopping-mall
38 36 walk
Indoor shopping-mall
37 walk 35

36 34
33
35
32 **
Type of walk *
34 Type of walk
31

Figure 9.7 The change in feelings following participation in walks. (a) anger; (b) confusion; (c) depression; (d) tension (change in self-esteem
normalised to the starting average for both groups). Error bars ¼  1 standard error; *p<0.05, ** p<0.01. See text for details.
 URBAN ECOLOGY AND HUMAN WELLBEING 221

Being part of a social group contributes to the green exercise experience, so it

is important to compare different group activities to ascertain the importance of
the exercise and the contact with nature. A study at the University of Essex
therefore evaluated the effectiveness of introducing a 6-week green exercise
programme (a series of short countryside and urban park walks) for individuals
experiencing a range of mental health problems. The walking group was com-
pared to two other Mind group programmes which were already in existence,
including a swimming group (indoor exercise) and a social club (met indoors,
but did not participate in any form of exercise). Key findings were that all groups
experienced a significant improvement in self-esteem, but the change in self-
esteem was significantly greater in the green exercise group than in the social
group. All three groups also experienced a significant improvement in overall
mood, and feelings of anger, confusion, depression, fatigue and tension all
decreased more in the green exercise and swimming groups compared with
the social club. These findings imply that participating in exercise is the primary
driver in positively enhancing mood, although engaging in sedentary social
activities can also still contribute to an improved mood, highlighting the import-
ance of the social contact. Thus, encouraging people to interact with green space
and be active outdoors has a potentially therapeutic role in positively influ-
encing emotional and physical wellbeing (Barton et al., unpublished data).

Concluding implications
The central themes emerging from the research are that regular contact with
urban green spaces improves human health and wellbeing. It improves phys-
ical health by providing opportunities for recreation and green exercise activ-
ities. Provision of green spaces encourages cycling and walking to move in and
between spaces, instead of relying predominantly on cars, which has numerous
health and environmental implications. It enhances psychological health by
creating a restorative environment which helps to reduce stress and encourage
relaxation. It affords spiritual connections which ensure people start to care
about their surrounding environment and influences behaviour by providing a
space people will choose to visit more frequently to engage in healthy activ-
ities. Urban green spaces also improve social cohesion by encouraging greater
social and cultural interaction, community empowerment and sense of place,
leading to better community spirit and neighbourhood social ties. Green areas
promote social inclusion, generate citizenship and local pride and contribute
towards reduced levels of crime and violence.
However, many of the daily settings to which people are regularly exposed
are highly dissimilar to the landscapes that shaped human evolution (Sullivan
2005). So, what happens to the health of residents in urban areas lacking access
to nature and green spaces? The United Kingdom, for example, is becoming an
increasingly urbanised society, and by definition urban areas enjoy less access
222 J. BARTON AND J. PRETTY

to nature and green space than rural environments. Some of this will be by
choice, as urban areas have more services and jobs concentrated together, with
better access to schools, hospitals, recreational facilities and other services.
However, disconnection from nature can impose new health costs by affecting
psychological wellbeing and reducing the opportunity for recovery from
mental stresses or physical tensions (Pretty et al. 2004).
The benefits of urban green spaces extend beyond ecosystem service provi-
sion and biodiversity conservation to have real impact on the physical and
psychological health and wellbeing of local residents. Yet the development of
‘urban sprawl’ is compromising public health (Frumkin et al. 2004) as housing
rapidly diffuses into green areas and competes for land. Expanding populations
as well as changing family demographics are also putting pressure on land for
housing developments within the urban envelope and on its fringes. Recent
research has found that provision of urban green space is more dependent on
city area than on the number of inhabitants (Fuller & Gaston 2009). This
implies that the green space provision per capita is very low in small cities of
high density, which affects residents’ quality of life.
Grahn and Stigsdotter (2003) found that urban citizens who live very close to
green space visit it three to four times per week. But at a distance of 1 km
the number of visits falls to only one per week. Natural England’s Accessible
Natural Greenspace Standards (ANGSt) model requires that no one should live
more than 300 m from their nearest green space (or 5 minutes’ walk) to ensure
they have the opportunity for exercise, relaxation and wellbeing (Handley et al.
2003), but what percentage of the population actually meet this recommenda-
tion? Harrison et al. (1995) recommend that local nature reserves should be
provided in every urban area, with a minimum of 1 hectare per 1000 people.
Local accessible green spaces should also be at least 2 hectares in size (Handley
et al. 2003), but how far are we from reaching this target?
The findings relating to accessibility to local green spaces across different
social groups vary. Some report inaccessibility issues for young people, low-
income groups, ethnic minorities and disabled people (CRN 2001; Natural
England et al. 2006), whose participation in outdoor recreation remains low.
Bedimo-Rung et al. (2005) also report that older adults, ethnic minorities,
women and lower-income families are more likely to visit parks infrequently
or be non-users. But other evidence suggests that the most deprived groups and
older people enjoy the greatest access (Barbosa et al. 2007). However, the
importance of locally accessible green spaces remains essential for all cohorts,
especially those enjoying the least access. In today’s modern society character-
ised by high stress, sedentary lifestyles, rising obesity problems, poor mental
health states and disconnection from nature, there is a new challenge: to find
ways to develop more local green spaces and improve accessibility to encourage
people to visit them on a more regular basis.
 URBAN ECOLOGY AND HUMAN WELLBEING 223

A recent report by CABE indicated that ‘91% of people believe that public parks
and open spaces improve their quality of life’. However, one in five people think
that it is ‘not worth investing money in the upkeep and maintenance of local
parks and public open spaces because they will just get vandalised’ (CABE Space
2005). A major concern is that neglected parks attract antisocial behaviour, yet
case studies show that often those marginalised and perceived as being a social
problem (e.g. disaffected young people, homeless) have become positively
involved in transforming space and managing its upkeep. Research suggests that
regular park use may result in long-term benefits for health and wellbeing, as
the combined benefits of engaging in physical activities and being exposed to the
green environment may have a cumulative effect on individuals. Therefore, the
approaches that researchers and practitioners use to increase local park and
recreation usage to achieve optimal health are becoming increasingly important.
Exposure to urban green spaces also enhances education by providing an
‘outdoor classroom’ for children to learn about nature and the culture and
heritage of communities. It provides them with the opportunity to learn about
the natural world in their local environment, engage in creative play and
improve their ecological consciousness. Research has indicated that childhood
exposure to nature and the frequency of visits to green places at a young
age correlate with adult patterns of behaviour. Infrequent woodland or green
space experiences as a child correlate with a lower frequency of visits during
adulthood. ‘Interestingly, not visiting as a child is more predictive of not visiting
as an adult than vice versa’ (Ward Thompson et al. 2008). The lack of outdoor
experiences during childhood may hinder desires to visit such places as adults to
engage in physical activity or benefit from its emotional restorative qualities.
Green spaces are often associated with an increased likelihood of exercise,
even without actively promoting the health benefits. The outdoor environment
therefore exerts a direct influence on the probability of taking leisure-based
physical activity and other specific sporting activities. The infrastructure of
vehicle-free routes and green open spaces allows more people, irrespective of
their social and economic circumstances, easily and safely to enjoy a countryside
experience close to their home. A more accessible and attractive rural–urban
fringe provides respite from the daily stresses of urban living and affords
opportunities for recreational activities such as walking, cycling, horse riding
or just relaxing. It also meets a demand for more adventurous sport and water-
based leisure activities on rivers and canals as well as making use of disused
gravel pits and quarries for fishing and angling. Encouraging more people to
engage in a range of recreational activities and interact with nature in the
rural–urban fringe will improve physical and mental health states, resulting in
reduced costs for both society and the economy. Activities in these areas can
serve the needs of both the rural and urban communities contributing towards
a more sustainable environment (Countryside Agency and Groundwork 2004).
224 J. BARTON AND J. PRETTY

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CHAPTER TEN

Bringing cities alive: the importance

of urban green spaces for people
and biodiversity
JON SADLER, ADAM BATES, JAMES HALE
AND PHILIP JAMES

Introduction
A plethora of papers exist that trumpet the value of urban green spaces as
providers of benefits to both people and wildlife ( James et al. 2009). This body
of work emphasises five means by which such spaces improve the urban
environment: (i) shaping the character of the city and its neighbourhoods
(Pauleit 2003); (ii) engendering a sense of place for city inhabitants (Frumkin
2003); (iii) providing a range of physical (Maas et al. 2006) and psychological
(Hartig 2008) health benefits to people; (iv) supporting rich assemblages of
wildlife, including many rare and endangered species (Gibson 1998; Mortberg &
Wallentinus 2000); and (v) possessing important environmental functions that
scale to provide a wide range of ecosystem services (Bolund & Hunhammar 1999;
Elmqvist et al. 2004).
It is estimated that the number of urban areas with over a million people
will grow by over 40% by 2015 (Crane & Kinzig 2005). To accommodate this
rapidly increasing population and to reduce the deleterious impact of global
sprawling cities (European Environment Agency 2006; Irwin & Bockstael 2007),
in many countries regulatory bodies have created a range of policies on urban
living, housing provision and city development that appear to be in conflict.
On the one hand, policies exist espousing the utilisation of as much open space
in cities as possible to meet construction targets for new-build housing (e.g.
ODPM 2002a), while on the other hand different policy documents highlight
the provision of green space for people and wildlife to enhance quality of life
(e.g. EEA 2009). The global move towards compact forms of cities (Dantzing &
Saaty 1973) has the potential to reduce urban sprawl, thereby preserving rural
food production and habitats. It may also contribute towards wider sustain-
ability goals (Cairns 2006) by reducing car use and fuel emissions, encouraging

Urban Ecology, ed. Kevin J. Gaston. Published by Cambridge University Press.

# British Ecological Society 2010.
 IMPORTANCE OF GREEN SPACES FOR PEOPLE & BIODIVERSITY 231

more walking, cycling and the use of public transport, and revitalising inner
city areas (Burton 2000). Pauleit et al. (2005) illustrated the flip side to this
process in Liverpool, UK, where green space has markedly reduced over the past
25 years. This loss was most acute within residential areas as a result of infill
development. This intensification of land-use threatens to compromise the
quality of the urban environment for both people and biodiversity (RCEP 2007).
Here we review the issues surrounding: (i) the provision and quantification of
green spaces in cities, (ii) the value of these spaces for biodiversity, (iii) their
importance for providing ecosystem services, (iv) their significance for people,
and (v) their design as multi-functional spaces. The chapter concludes by
identifying key challenges where further research is necessary. We focus par-
ticularly on the United Kingdom as a case study, although we also draw on
material from other regions.

Green spaces: classification and quantification

Policy spaces
We stress here that the policy frameworks surrounding the management and
provision of green spaces are heavily geographically contextualised (CABE
2009a), which means that generalisations that have widespread applicability
are not easily derived. In this section we use the situation in the UK as an
example of national initiatives to identify possible commonalities and issues of
importance. In the UK there has been significant interest and numerous policy
statements concerning the value of green spaces to citizens in cities. This
interest has been driven by concerns over health inequalities in cities in much
of the developed world (Smyth 2008), and the so-called ‘obesity epidemic’
(Joshu et al. 2008). Concerns over the quality and management of green space
in the UK have been driven by local people, and a wide range of reports by non-
governmental organisations and government departments which culminated
in the Urban Task Force Report (Urban Task Force 1999). This report strongly
mainstreamed the significance of urban green space for enhancing the quality
of city life into a national policy context. The government’s response was the
Urban White Paper (DETR 2000), which recommended the creation of the
Green Spaces Taskforce (GST), whose remit was to advise government on pro-
posals for the improvement of the quality of urban parks, play areas and other
green spaces. GST’s report Green Spaces, Better Places (DETR 2002) examined the
provision of green spaces and their loss, design and utility for people and
wildlife, as well as introducing the concept of green networks into planning
policy. The government’s response to the recommendations in this report
arrived the same year in Living Places – Cleaner, Safer, Greener (ODPM 2002b). This
acknowledged the need for a clearer management process and greater under-
standing of the provision, quality and access importance of green spaces in
cities, and led to the establishment of the Commission for Architecture and the
Built Environment (CABE), whose report, Open Space Strategies: Best Practice
232 J. SADLER, A. BATES, J. HALE AND P. JAMES

NATIONAL

Open space policies in key documents

REGIONAL

Regional Strategies

Green Infrastructure Strategies

SUB-REGIONAL

Green Infrastructure Strategies

Green Networks

LOCAL

Open Space Strategies

NEIGHBOURHOOD

Area Action Plans

Public Realm Strategies

SITE-BASED

Site Action Plans

Landscape Action Plans

Figure 10.1 Hierarchical policy framework for the management of green spaces
in the UK. Reproduced with permission from CABE (2009b).

Guidance (CABE 2009b), called for a hierarchical system (Figure 10.1) that linked
national strategies on green space management and provision to policy
guidance at local (neighbourhood) scales, a structural pattern that is common
to most countries in the world. These policy documents are useful in
 IMPORTANCE OF GREEN SPACES FOR PEOPLE & BIODIVERSITY 233

Table 10.1 UK planning policy guidance documents that relate to green spaces
and green infrastructure.

Spatial unit Planning policy guidance Other relevant planning policies

Green spaces PPS 9: Biodiversity and NI 197: Improved local biodiversity

geological conservation
PPG 17: Planning for open
space, sport and
recreation
Green PPS 1: Delivering NI 185 & 186: Reduction of CO2
infrastructure sustainable emissions per capita and via Local
development Authority operations
PPS 1 (Supplement): NI 188: Adapting to climate change
Planning and climate NI 189: Flood and coastal risk
change management
PPS 12: Local spatial NI 194: Level of air quality
planning NI 197: Improved local biodiversity
PPS 9: Biodiversity and geological
conservation
PPS 25: Planning and flood risk
PPS 22: Renewable energy

Note: Planning Policy Guidance (PPG) and Planning Policy Statements (PPS) are documents
created at a national level which local authorities are required to use as part of their planning
processes. National Indicators (NIs) are a sequence of measurable indicators that local
authorities have to report to government on an annual basis.

highlighting areas of need, and specifically they identify issues surrounding

green space definitions, typology, quantity, quality (including ecology) and the
need for standards for access and robust monitoring and management systems.
These are issues that are now wired into UK national policy documents and
government planning guidance, which is used by local and regional authorities
(Table 10.1).

Classification and typology

In the UK, ‘open space’ is defined in the Town and Country Planning
Act 1990 as ‘land laid out as a public garden, or used for the purposes of
public recreation, or land which is a disused burial ground’. This emphasises
land that is public or municipal space, but not the wider range of open
green space available in cities. It does, however, include areas of water such
as rivers, canals, lakes and reservoirs, which offer important opportunities
for sport and recreation and can also act as visual amenity. More recent
234 J. SADLER, A. BATES, J. HALE AND P. JAMES

definitions emphasise a broader portfolio of spaces and include references to

‘nearby nature’ or ‘natural green spaces’. Harrison et al. (1995, p. 4), for example,
define them as ‘Land, water and geological features which have been naturally
colonised by plants and animals and which are accessible on foot to large
numbers of residents’. In Canada a distinction is drawn between green space
and parkland, with the former relating to municipal natural open space
(reserves, floodplains etc.) and the latter relating to land set aside by the
municipality as part of an established public park (Evergreen 2004).
Green spaces form part of a linked social–economic–ecological system
(Yli-Pelkonen & Niemelä 2005) and when scaled up to a landscape or cityscape
they form what has been termed green infrastructure. Benedict and McMahon
(2002, p. 12) defined this as ‘an interconnected network of green space that
conserves natural ecosystem values and functions and provides associated
benefits to human populations’. The distinction between green spaces and
green infrastructure is subtle but worth stressing. Green infrastructure goes
beyond the site specific, and provides a means of characterising multi-
functional landscapes; it also considers private as well as public assets. This
chapter focuses on green spaces but stresses green infrastructure where the
issues being discussed demand a focus on a larger spatial scale (e.g. ecosystem
services and design and planning).
There is similarly wide variation in the typologies of what constitutes a green
space. Table 10.2 illustrates this by examining examples drawn from academic
and policy literatures from the UK. It is clear that there is an uneasy mix of
classifications that relate to use (e.g. recreational), type (e.g. public/civic space)
or habitats (e.g. woodland, wetlands) even within the same scheme. Some
typologies include private residential land such as domestic gardens, and most
highlight linear features (e.g. greenways, river, road and railway corridors and
green networks). This variability in classification has led to inconsistency in
adoption of schema in the literature. Handley et al. (2003), Tzoulas et al. (2007)
and Comber et al. (2008) used the typology of green spaces provided by the
Urban Green Spaces Task Force (DETR 2002), which has been formalised in the
UK by Planning Policy Guidance 17 (PPG17): Planning for Open Space, Sport
and Recreation (DCLG 2000; Table 10.2).
A recent international comparison of green spaces in 11 cities (CABE 2009a)
has highlighted that all had an element of typology in their policies, mostly
based around function and size, but that this varied very significantly across
the cities examined. For example, in Malmo, Sweden, and Tokyo, Japan, green
spaces were classified according to function, and this was used in the planning
system to create an even distribution of green space across the city. Some cities
had only broad categories, such as Paris, France, where all habitats other than
woodlands are classified as gardens, and Minneapolis, USA, where all green
space is called parkland. In some countries, such as Japan, the green space
 IMPORTANCE OF GREEN SPACES FOR PEOPLE & BIODIVERSITY 235

Table 10.2 Typologies of urban green spaces that exist in UK policy literatures.

Landscape
Institute Swanwick et al. 2003 DETR DCLG (PPG17)

Street trees, Recreation green Green roofs Parks and gardens

verges and space (not including
hedges private)
Green roofs and Incidental green Urban parks Natural and semi-
walls space natural spaces
Pocket parks Private green space Green corridors Green corridors
Private gardens Productive green Encapsulated Outdoor sports
space countryside facilities
Urban plazas Burial grounds Derelict land Amenity green
spaces
Town/village Institutional grounds Housing green Provision for children
greens and space and and young people
commons domestic gardens
Pedestrian and Wetland Churchyards, Allotments,
cycle routes cemeteries and community
school grounds gardens and urban
farms
Cemeteries and Woodland Open standing and Cemeteries, disused
churchyards running water churchyards and
other burial
grounds
Institutional open Other habitats Public/civic space
spaces (e.g. grassland, (e.g. plazas,
disturbed ground) pedestrianised
areas)
Ponds, streams Linear green space
and rivers
Woodlands
Nature reserves
Play areas
School grounds
Sports pitches
Swales, ditches
Allotments
Brownfields

Sources: Landscape Institute (2009); Swanwick et al. (2003); DETR (2002); DCLG (2000).
236 J. SADLER, A. BATES, J. HALE AND P. JAMES

typology is defined nationally as part of a policy to provide various kinds of

green space within walking distance of residential areas.
In this chapter we focus attention on a discussion of the public spaces in the
typology of Swanwick et al. (2003) and exclude a full discussion of the biodiver-
sity of gardens, even though they comprise a large amount of the green space
in many cities (e.g. Loram et al. 2007). The importance of urban residential
spaces, however, is touched on in our consideration of ecosystem services and
green space design.

Quantification
Green space in the UK accounts for 14% of urban space nationally, but this is
very variable, with some cities being greener than others. In Leicester, 25% of
urban land is green space (Comber et al. 2008), in Greater Manchester 32% (Gill
et al. 2008) and in Sheffield 45% (Fuller & Gaston 2009). Approximately 30% of
the total land surface of Greater London is occupied by open green spaces
(Wilby & Perry 2006). In Birmingham the proportion of green space is 33.7%
not including gardens and land associated with water bodies, roads and rail-
ways (unpublished data). Within Europe, available statistics show rather more
variability. In a recent and extensive analysis of 386 European cities, Fuller and
Gaston (2009) showed that green space coverage at a wider European level was
extremely variable, ranging from 1.9% (Reggio di Calabria, Italy) to 46% (Ferol,
Spain). Their analyses also illustrated that green space coverage was scaled to
city land area so that cities large in area had greater green space provision,
even though overall population densities were similar to cities smaller in area.
The association showed that green space area increased more than city area, so
that cities differing in area by an order of magnitude will have a 15-fold
difference in green space area. Perhaps worryingly for UK residents, there the
green space provision in 67 cities scales only at the same rate as city area. The
implications of the work are concerning, as Fuller and Gaston suggest that
access to green space may well decline as cities grow.
Unfortunately, there is a paucity of longitudinal (time series) data on change
in urban areas to assess the scale of green space loss, aside from snapshots of
individual cities such as Munich, Germany (Pauleit & Duhme 2000), and Stock-
holm, Sweden (Bolund & Hunhammar 1999), and a European-wide comparative
study carried out by the European Environment Agency (EEA 2002). The EEA
report suggested that the area of public green spaces in 25 cities had reduced as
a proportion of total city land areas between 1950 and 1990. There are no data
to show whether this is a result of the limited provision of green spaces in
sprawling city margins or whether it relates to real loss of city green spaces due
to landscape intensification within cities. Davies et al. (2008) showed that the
extent of green space is negatively correlated with total area of buildings and
the length of the road network in Sheffield, UK, suggesting that urbanisation
 IMPORTANCE OF GREEN SPACES FOR PEOPLE & BIODIVERSITY 237

causes progressive loss of green space via land-use intensification. In their study
of Mashad, Iran, Rafiee et al. (2009) also found a significant decrease in the extent
of urban green spaces had occurred between 1977 and 2006. There is, however, a
dearth of global data on this issue, and producing systematic and comparable
figures detailing the loss of green space in our cities is an urgent task.

Green space standards

The creation of standards for access to green space is an important part of any
policy agenda. In the 1970s, the United Nations suggested a standard of 6ha per
1000 head of population as a good target figure for urban environments (Wang
2009). Similar standards were subsequently adopted by many countries there-
after with figures ranging from 4 ha/1000 people in Japan, 4.05 ha (10acres)/
1000 people in the USA and Canada, and 14 ha/1000 people in Germany
(Sukopp et al. 1995). Within these broad categories there is considerable vari-
ation within individual nations. For example, a review of guidelines in Can-
adian municipalities showed that they ranged from 0.7 to 6 ha/1000 people
(Evergreen 2004). Although such area-based standards have the advantages of
being clear and measurable, it is obvious that they do not capture the complex-
ity of green space needs in cities. Other approaches are also evident, and they
include time spent travelling to the nearest green space, and the nature of
those spaces along with area-based guidelines.
The European Environment Agency recommends that accessible green
spaces should be within an average 15-minute walk from people’s homes.
Natural England, the UK’s statutory organisation that deals with nature conser-
vation and the countryside, created a suite of hierarchical recommendations
concerning accessible natural green space standards (ANGSt) (Harrison et al.
1995; Handley et al. 2003). They recommend at least 2 ha of accessible natural
green space per 1000 head of population, but also that:

• every home should be within 300 m (5 minutes’ walk) from an accessible

natural green space;
• there should be 1 ha of local nature reserves per 1000 people; and
• there should be an accessible 20-ha site, 100-ha site and 500-ha site within
2 km, 5 km and 10 km, respectively, of every home.

Pauleit et al. (2001) considered the uptake of the ANGSt model by 30 municipal
areas in the UK. Although many (80%) already had in place some kind of green
space standard, most (75%) of these related only to ‘amenity green space’ and
ignored other types (brownfields, nature reserves and the like). Some local
authorities, such as Birmingham, embraced the model, integrating it, albeit
in a modified form, into their Nature Conservation Strategy in 1997, while the
majority (90%) had not implemented it into planning policy. The situation has
improved substantially since this time with access standards figuring highly in
238 J. SADLER, A. BATES, J. HALE AND P. JAMES

many council and local authority planning documents, including nature

conservation strategies that emphasise the importance of green spaces and
green infrastructure for local wildlife.

Green spaces, biodiversity and ecosystem services

Green space and biodiversity correlates
Cities are characterised by highly modified and complex landscapes compris-
ing a rich mosaic of relict habitats and designed spaces juxtaposed in combin-
ations that rarely occur in nature (Angold et al. 2006). As a result they are home
to a wide diversity of organisms (Gilbert 1989), some of which are becoming
increasingly rare in the wider countryside (Gibson 1998; Woodward et al. 2003),
where ecological processes have been stymied by agricultural intensification
(Tratalos et al. 2007). There is now an extensive literature on the biodiversity of
urban areas (e.g. Chapter 5; McKinney 2006, 2008) examining changes in diver-
sity along gradients from the city margins to the city centre (McDonnell &
Pickett 1990; McDonnell & Hahs 2008), fragmentation effects (Sadler et al. 2006)
and studies of total city species pools (e.g. Puth & Burns 2009).
This literature illustrates that the type, structure and location of green
spaces are important correlates for the presence, abundance, species richness
and persistence of plant and animal species, yet there has not been a systematic
study of these effects. Here we draw on a wider literature search aimed at
identifying the green space correlates of animal biodiversity in cities. The
search identified just under 4000 articles which were subsequently examined
to extract information on green space correlates for the target groups
(Table 10.3). It should be stressed that the studies revealed a very large variety
of approaches to capturing information on both the green space type (target
study habitat) and wider urban landscape, so careful interpretation and cat-
egorisation were needed to isolate the potential correlates. We divided those
correlates into two groups representing landscape and local influences on
green space biodiversity. The most evident trend is that there are more local
than landscape correlates listed. This was anticipated because researchers
carefully monitored and measured the habitats of interest, often generating
many local variables, but far fewer landscape variables. Moreover, some
authors (e.g. Blair 1999) used multi-variate techniques to analyse their data,
and this retained a larger pool of variables than studies that used stepwise and
selective regression techniques. There is also considerable variability between
correlates, which makes establishing causation problematic. Notwithstanding
these complexities, several pervasive themes can be isolated relating to: (i) the
importance of local versus landscape influences, and the significance of
(ii) local, and (iii) landscape variables. We stress here that because of the scale
of the issue our discussion will be selective and focus on very evident and large-
scale patterns.
 IMPORTANCE OF GREEN SPACES FOR PEOPLE & BIODIVERSITY 239

Table 10.3 Local and landscape biodiversity/green space correlates of animals.

Taxa group Green space habitat Landscape correlates Local correlates

Vertebrates
Birds Urban land-use Housing density Green space area
Brownfield Building density Rough grassland
Wetland Urban land-use Supplementary feeding
Woodland Isolation Human disturbance
Public green space Adjacent gardens % Woody vegetation
Natural green space Connectivity Riparian vegetation
Gardens Habitat structure
Size
% Woodland cover
Bats Woodland reserve Distance to water Shrub density
Natural relict green Density of built land Tree diameter
space
Hedge lines Woodland Tree density
fragmentation
Connectivity Forest cover
Roads
Amphibians Ponds Woodland cover Pond depth
Pond isolation Woodland cover
Pond density Pond structure
Pond size
Fish presence
Pond vegetation
Roads
Reptiles Natural relict green % Urban land Habitat physiognomy
space
Gardens Distance to nearest Vegetation structure
similar patch
Housing density Vegetation type
Urban sprawl Grassland presence
Soil compaction
Introductions
(Vegetation)
Aspect
Geology
Invertebrates
Coleoptera Brownfield Site location Vegetation structure
Greenways % Urban land Disturbance
Gardens Amount of similar Site age
habitat locally
Woodland Corridors Urban land-use
Wetland Size
Trampling
240 J. SADLER, A. BATES, J. HALE AND P. JAMES

Table 10.3 (cont.)

Taxa group Green space habitat Landscape correlates Local correlates

Butterflies Grassland Isolation

Garden Habitat quality
Office park Amount of similar
habitat locally
Golf course % Urban land
Park
Reserve
Amenity space
Galls Gardens Edge effects
Woodland Plant richness
Natural green space
Road verge
Spiders Woodland Woodland Prey availability
fragmentation
Desert park Isolation Habitat productivity
Urban desert Size
remnants
Industrial Introduced species
Agricultural
Xeric- and mesic-
residential yards
Bees Gardens Roads Grazing density
Allotments % Urban land Plant diversity
Grassland % Suburban land Tree abundance
Woodland Fragmentation Shrub diversity
Ants Woodland % Urban land Vegetation cover
Gardens Tree density
Brownfields Nest resources
Orthoptera Reserve Road density Road density
Brownfield Landscape context Road age
Grassland Vegetation structure
Landscape context
Soil parameters
Site age
Hemiptera Brownfield Landscape context Vegetation structure
Traffic roundabouts Size
Soil parameters
Site age
Mowing
Floral evenness

Note: Data were derived from searches on the Web of Knowledge using a sequence of keyword
queries (e.g. URBANþBIRDS, URBAN*BATS and so on). The subsequent results were sifted
for duplication and then categorised into the groups in the table.
 IMPORTANCE OF GREEN SPACES FOR PEOPLE & BIODIVERSITY 241

Several researchers modelled variability and partitioned it to either

landscape or local factors or combinations of both. In these studies there is a
general tendency for local factors to be more important than landscape vari-
ables for biodiversity. The pattern is pervasive and cuts across taxonomic
groups. Parris (2006) showed that declining pond quality (measured as the
proportion of ponds with a surrounding wall) led to a 44–56% decrease in
amphibian species in ponds in urban Melbourne, Australia. Evans et al. (2009)
state that local correlates such as increased structural complexity, woody
vegetation diversity and supplementary feeding influence avian assemblages
more strongly than factors operating at larger spatial scales. Many invertebrate
taxa such as hemipterans and ground beetles exhibit similar responses, where
site-based variables such as age since last disturbance (Small et al. 2006) and
management regime (Helden & Leather 2004) account for a greater proportion
of the variability in the assemblage pools. These findings are important because
they suggest that management at the scale of individual green spaces may
provide significant benefits to local biodiversity. This is a strategy that is much
more easily operationalised than attempting to manage whole multi-functional
cityscapes, especially as planning tends to operate on a site-by-site basis.
Local-scale biodiversity–green space correlates tend to fall under the
umbrella of what is frequently termed ‘habitat quality’ and include variables
such as the size of the green space, its vegetation type, structural diversity,
disturbance regime, management and so on. Perhaps the most evident pattern
is the link between biodiversity metrics and site size which is found in a wide
range of studies of birds (Chamberlain et al. 2007), amphibians (Dickman 1987)
and insects (Small et al. 2003). This is not altogether surprising as many studies
have shown pervasive area effects in habitats (e.g. Buckton & Ormerod 2002), a
pattern often attributed to the Theory of Island Biogeography (MacArthur &
Wilson 1967). However, several researchers have argued that this is not directly
applicable to urban areas where green spaces are not fully isolated islands
(Niemelä 1999), rather the matrix habitats are permeable and enhance species
movements. Perhaps the best example of this is residential gardens (Davies et al.
2009), which are not only home to a wide range of organisms (e.g. Loram et al.
2008) but are permeable to a great many others. Indeed, the high turnover rate
exhibited by many taxonomic groups (Smith et al. 2006a, 2006b) suggests that
the presence of gardens may enhance species dispersal through the city. None-
theless, utilising the area effect relationship may provide a means of gaining
rough estimates of likely species richness loss in the face of increasing land
intensification in cities.
Other variables illustrate the importance of structural habitat influences
(Denoel & Lehmann 2006; Garden et al. 2007), especially vegetation structure
(Strauss & Biedermann 2006), vegetation cover (Loeb et al. 2009), vegetation type
(Palomino & Carrascal 2006), substrate type (Small et al. 2003), soil compaction
242 J. SADLER, A. BATES, J. HALE AND P. JAMES

(Grandchamp et al. 2000) and site age (Bolger et al. 2000). Several papers
illustrate the importance of biotic linkages (Miyashita et al. 1998; Bolger et al.
2008) and their likely influence on ecosystem services (Sandford et al. 2008).
Most of the physical factors are amenable to management at a local scale,
although their importance differs across taxa suggesting caution is needed
when integrating them into management plans.
Landscape variables used to contextualise green spaces in cityscapes are
usually correlates of urbanisation (e.g. the amount of built land, percentage of
concrete per unit area, building densities) and/or relate to fragmentation and
isolation (e.g. percentage of habitat per unit area, distance to nearest similar
green space). Many studies also highlight the importance of connectivity
and linkage between green spaces. Several taxa such as butterflies (Hardy &
Dennis 1999), beetles (Magura et al. 2004), bats (Gehrt & Chelsvig 2003, 2004),
birds (Evans et al. 2009), reptiles (Germaine & Wakeling 2001) and amphibians
(Pillsbury & Miller 2008) illustrate negative responses to increased urbanisation
pressure. Studies of fragmentation effects caused by urbanisation are also
widespread in the literature (Table 10.3), as are studies examining the impact
of roads in creating barriers to movement for poorly dispersing species, notably
amphibians (Forman & Deblinger 2000) and small mammals (Dickman &
Doncaster 1987). Recent research on taxa as diverse as bats (Gorrensen et al.
2005), bees (Ahrne et al. 2009) and beetles (Ishitani et al. 2003) has shown that
species responses to urbanisation and green space fragmentation are complex,
frequently species-specific and often best predicted by using ecological traits
(Cane et al. 2006). There are few data quantifying the likely loss of species from
urban green spaces as a result of fragmentation effects. For example, Parris
(2006) showed that the predicted amphibian species richness in the smallest
ponds in their Melbourne survey was 2.8–5.5 times lower than predicted in the
largest ones. She noted also that the most isolated pond was predicted to have
5–9 times fewer species than the least isolated pond. The potential loss of
species richness also has more pervasive functional implications. In a recent
paper, Flynn et al. (2009) used an extensive meta-analysis to show that
intensification in the agricultural landscape has led to widespread loss of
functional diversity (FD) in temperate and tropical New World mammal and
bird communities. Significantly, in over one-quarter of the bird and mammal
communities they analysed, declines in FD were steeper than predicted by
species number. It seems likely that the loss of urban green spaces may be
leading to similar losses of FD in our cities.
Connective features such as green networks and corridors have been influen-
tial in guiding planning policies in many areas of the world (Turner 2006; von
Haaren & Reich 2006), but are also subject to considerable debate and confu-
sion (Hess 1994; Hess & Fischer 2001). They have long been seen as providing
connectivity, linking green spaces and minimising the potential effects of
 IMPORTANCE OF GREEN SPACES FOR PEOPLE & BIODIVERSITY 243

fragmentation on wildlife ( Jongman et al. 2004), while providing important

recreational, leisure and nature experience possibilities for people (Gobster &
Westphal 2004). Only a few empirical studies have illustrated the successful
role of corridors as conduits for species movement, and most of these derive
from carefully controlled landscape-scale experiments (Beier & Noss 1998;
Haddad & Baum 1999; Haddad et al. 2003). This is a vastly different situation
from most urban areas where green corridors are multi-functional spaces, with
a lot of habitat variability, and in these instances they do not appear to provide
a strong functional conduit role (Angold et al. 2006). Nonetheless, some mobile
and generalist species do utilise them as habitat (Mason et al. 2007), and even
small-scale green features, such as street trees, are thought to enhance move-
ments between green spaces in cities by birds (Fernández-Juricic 2000) and bats
(Avila-Flores & Fenton 2005). Moreover, the role that green corridors may play
as key elements in defining the social-nature landscape in cities is undeniable
(Hartig et al. 1991; Evans 2007).

Ecosystem function and services

Constanza et al. (1997, p. 253) defined ecosystem services as ‘the benefits human
populations derive, either directly or indirectly, from ecosystem functions’.
Although widely studied in broader ecological research, an evaluation of the
quantities and flows of these services in cities is lacking (Elmqvist et al. 2004).
A fuller review of this burgeoning literature is the subject of another chapter in
this volume (Chapter 3), so we have limited our discussions to key areas that are
of significance to green space planning in cities. Green spaces provide a
wide range of locally generated ecosystem services in urban areas (Bolund &
Hunhammar 1999) as well as important habitat and refugia functions as
discussed above. These include a mix of regulatory and provisioning services
such as climate amelioration (Akbari et al. 2001; Nowak et al. 2001), carbon
sequestration and storage (Nowak & Crane 2002; Sahely et al. 2003, Pouyat et al.
2006), floodwater storage (Pauleit & Duhme 2000), water (Bolund & Hunhammar
1999) and air purification (McPherson et al. 1997; Beckett et al. 1998). It seems
likely that several of these services have the capacity to help future-proof cities
against global climate changes (Gill et al. 2007), but their quantification and
the science underlying their application in this manner require considerable
research.
The structure, composition, location and spatial configuration of green
spaces influence their ecological performance and hence the services they
provide (Pauleit & Duhme 2000; Whitford et al. 2001). Tratalos et al. (2007)
compared the performance of differing landscape units across five cities in
the UK and concluded that although ecosystem quality tends to decline con-
tinuously as urban density increases, there is variability evident in many of the
relationships between landscape type and service provision. This suggests that
244 J. SADLER, A. BATES, J. HALE AND P. JAMES

potential exists for maximising ecological services at a given landscape density

by careful consideration of the structure (i.e. proportion and configuration) of
landscape units. Currently planning systems still operate by focusing on indi-
vidual units or development schemes (Niemelä 1999) but to gain the full
benefit of ecosystem services, planning of the green space resource needs to
occur at larger spatial scales (Andersson 2006).

Green spaces and people

Use and access
The final report of the World Health Organization’s Commission on the Social
Determinants of Health (CSDH 2008) and the latest European Union report on
‘urban living’ (EEA 2009) both called for wide-ranging improvements in daily
living conditions. They both highlighted the need for improved access to green
space for all, especially for those elements of society that live in poverty and are
subject to deprivation. Green spaces are certainly well used in many regions of
the world (e.g. De Sousa 2003). Comparative data on green space usage are few,
but they are available from the UK, and they provide a useful benchmark. The
Urban Parks Forum (2001) estimated that there are somewhere in the region of
1.5 billion visits per year to historic parks, using visitor numbers from three-
quarters of local authorities. This is similar to an estimate of 2.6 billion visits
per annum, based on a detailed questionnaire survey in Sheffield and ten other
UK cities (Swanwick et al. 2003). There are also relatively few studies that
consider access, especially measured against any standards that have been
stipulated by governing bodies.
In a comparison of six major European cities, Stanners and Bourdeau (1995)
reported that all citizens have access to urban green space within a 15-minute
walk from their homes. Similarly, several large UK cities meet Natural Eng-
land’s basic ANGSt criteria of providing at least 2 ha of green space per 1000
head of population, with estimates ranging from 3.5 ha per 1000 in Leicester,
8.8 ha per 1000 in Manchester and 9.3ha per 1000 in Birmingham to 21.5ha
per 1000 in Greater London. This basic criterion, however, is the lowest base
level provision and a more rigorous application of the standard is certainly
needed, utilising the tighter, more stringent remaining rules, which are much
more discriminatory (Handley et al. 2003). The ANGSt rules have been more
fully tested recently by studies in both Sheffield and Leicester. Barbosa et al.
(2007) noted that green spaces are chronically underprovided for in Sheffield.
Only 36.5% of Sheffield’s households are located within 300m of their nearest
green space, which is the stipulated threshold figure recommended by Natural
England. In contrast, 95.6% meet the European guidance, living within a 15-min
walk from a green space. When considering the 85 larger municipal spaces (e.g.
parks and public gardens) only 18% and 58% of Sheffield households meet the
UK and European recommended access thresholds, respectively. A study of
 IMPORTANCE OF GREEN SPACES FOR PEOPLE & BIODIVERSITY 245

access in Leicester showed that only 10% of the population live within 300 m of
a local green space and 40% lack access to large (20-ha) sites within 2 km of
their homes (Comber et al. 2008). On the basis of the few available studies it
seems that, when measured against UK and EU standards, access to green space
is very limited.
The CSDH (2008) report highlights that access to green spaces should be for
all sectors of society. Another important question is who has access to green
spaces in cities. Barbosa et al. (2007) showed that there was great variability in
access by different socioeconomic groups in Sheffield, although deprived social
groupings and the aged fared better than the rest. Studies elsewhere have
found opposite income-related patterns. Heynen (2006) showed a strong posi-
tive correlation between access and median household income. Mitchell and
Popham (2008) studied the association between green space and socioeconomic
inequalities in England using health statistics data derived from the English
Index of Multiple Deprivation (EIMD). They noted strong covariation between
green space access and socioeconomic group where poorer groups had less
access.
Inequality of access is also a significant issue for people from ethnic groups.
Studies in the USA have shown that the use of, and preferences of citizens for,
green space vary in relation to ethnicity (Gobster 1998, 2002; Frumkin 2005).
This pattern is paralleled in the city of Leicester, UK, where Comber et al. (2008)
showed that certain ethnic groups (Indian, Hindu and Sikh) had limited access
to green space in the city. Although few comparative data and little research
exist, the findings are worrying as such patterns could be exacerbating other
health equality issues (Smyth 2008).

Health and wellbeing benefits of green space

Green spaces have figured large in studies examining the relationships of
‘place to health’ (e.g. Frumkin 2003). This work has a strong historical dimen-
sion that focuses on extraordinary places of healing (e.g. Lourdes, France)
(Smyth 2005) and on the sociopolitical context surrounding the rapid growth
and creation of green spaces during the nineteenth century, which aimed at
improving the lot of the working classes who were subjected to squalid and
overcrowded living conditions (Giles-Corti et al. 2005). There is now an influen-
tial and substantial body of evidence exploring the links between green
space provision in cities and human health (physical and psychological) and
wellbeing (Tzoulas et al. 2007). The evidence can be grouped into three classes:
(i) epidemiological studies linking health benefits to exposure to an improved
natural environment, (ii) epidemiological evidence linking green space to
behavioural changes leading to increased levels of physical exercise, and
(iii) improvements in psychological (mental) health engendered by exposure
to natural places and scenes. We examine each in turn.
246 J. SADLER, A. BATES, J. HALE AND P. JAMES

(i) Epidemiological studies of green space health links

De Vries et al. (2003) showed a strong positive association between self-reported
health and available neighbourhood green space. Their results suggest that a
10% increase in green space provision reduced symptoms of ill health in their
survey pool of people (n ¼ 13090) to a level comparable with a decrease in age
of 5 years (p. 1726). A recent evidence-based study of 250 782 people, which was
carefully constructed to avoid potential selection errors, highlighted a strong
positive association between their local provision of green space (i.e. within
1 and 3 km of their homes) and perceptions of their own health (Maas et al.
2006). Mitchell and Popham (2007) aimed to determine the association between
the percentage of green space in an area and the rate of self-reported ‘not good’
health, using health statistics and census data (n ¼ 32482). They report that,
generally, higher proportions of green space in an area were associated with
better health, but the association was mixed depending on the degree of
urbanisation and the level of income deprivation. Although these are all well-
controlled multi-level and cross-sectional studies which account for variable
socioeconomic status and age, their design does not help to identify or prove
causal relationships that underlie the associations.

(ii) Behaviour and physical activity

An increasingly overweight and obese population in many developed countries
is strongly correlated with reductions in the numbers of people taking part in
physical activity ( Joshu et al. 2008). Both issues are linked to major risk factors
for a wide range of ailments and potentially fatal diseases (e.g. diabetes,
cancers, heart disease, stroke). The many studies that have examined this
important health issue fall into two categories: those that use questionnaires
and self-reporting to examine levels of physical activity, and those that link
indicators of overall physical activity (e.g. data on body mass index, BMI) to
green space provision.
The association between green space provision and an increase in physical
activity levels has been illustrated in several studies, which controlled for age,
gender and educational level. Li et al. (2005) considered aspects of self-reported
walking activity in the aged (people over 65 years old; n ¼ 577) in Portland,
Oregon, USA. The results suggest that neighbourhood building density and
green infrastructure provision were important determinants of walking.
Takano et al. (2002) observed a positive effect of living in an area that has
walkable green spaces on the longevity of elderly people in Japan.
Frank et al. (2004) examined this issue in Atlanta, USA, by associating obesity
(i.e. BMI  30 kg m2) with other factors such as land-use mix, residential
density and car use in adults. The analyses indicated that an additional hour
spent in a car per day was associated with a 6% increase in the likelihood of
obesity, whereas an additional kilometre walked per day was associated with a
 IMPORTANCE OF GREEN SPACES FOR PEOPLE & BIODIVERSITY 247

4.8% reduction in the likelihood of obesity. Although the variables interacted

the authors suggest that land-use mix is an important determinant of walking
activity, and thus lower BMI, in their sample populations. In a follow-up study
accelerometers were deployed over a 2-day period to capture objective levels of
physical activity in 357 adults in the same city (Frank et al. 2005). Activity levels
were then compared to a walkability index derived from each household’s
physical location, the adjacent land-use mix, residential density and street
connectivity. The walkability index of these urban form factors was related to
levels of physical activity. Nielsen and Hansen’s (2007) survey of access and use
of green areas and their impact on obesity in 1200 Danish adults (18–80 years
old) provided similar results, indicating that access to a garden or short dis-
tances to green areas are associated with lower likelihood of obesity. The
authors suggest that the significance of distance to green areas is mainly
derived from its correlation with the character of the neighbourhood and its
conduciveness to outdoor activities and ‘healthy’ lifestyle modes.
The positive association between local green space provision and weight is
prevalent for all ages of society. Bell et al. (2008) note that African-American
economically disadvantaged children and youth exhibited an inverse associ-
ation between neighbourhood greenness and BMI scores. Their study examined
the lasting effect of green space provision by examining changes in the BMI
of children and youth (n ¼ 3842) in Indianapolis, USA. The results suggest that
children and youth in greener neighbourhoods were less likely to increase
their BMI scores over the two years of their study than their counterparts in
less green neighbourhoods. But associative links such as these are context-
dependent, notwithstanding the care and thought that has gone into study
designs. Garden and Jalaludin (2009) showed that urban sprawl in Sydney,
Australia, was associated with reduced activity levels and increased obesity
owing to an overreliance on cars for transport and poor neighbourhood
resources for shopping, community activities and the like. It appears that
where people are in the cityscape matters as well as their socioeconomic status,
gender and ethnicity, highlighting the need for spatially explicit models that
factor in geographic locations.

(iii) Improvements in psychological (mental) health and wellbeing

The mechanisms underpinning psychological responses to green spaces derive
from psycho-evolutionary theory (Ulrich et al. 1991) and relate to the capacity of
natural green spaces to act as ‘restorative places’ (Hartig et al. 2003). Attention
Restoration Theory (Kaplan & Kaplan 1989; Kaplan 1995) suggests that direct
and focused attention can be enhanced by direct contact with nature, which
replenishes an individual’s attention resource. In contrast, Ulrich’s (1984) work
on the restorative effects of nature, while in some ways similar to Kaplin’s,
emphasises an affective rather than cognitive response, and focuses on
248 J. SADLER, A. BATES, J. HALE AND P. JAMES

emotional and mental responses to taxing/threatening stimuli, not attention

deficits due to everyday humdrum activity (Hartig et al. 1991). Ulrich (1984)
highlighted the positive impact that a ‘green view’ can have on patients
convalescing in hospital after major surgery. In this ground-breaking study,
23 patients whose beds overlooked ‘natural scenery’ showed enhanced recovery
rates when compared to a control group (n ¼ 23) whose beds faced a brick wall.
Kuo (2001) studied attentional functioning and effectiveness in dealing with
major stresses in 149 urban public housing residents randomly assigned to
housing with or without nearby nature. Residents placed in housing with little
adjacent green space (nearby trees and grass) reported higher levels of procras-
tination and stress when facing major issues than their counterparts who were
inhabiting homes in greener areas, indicating that access to nature is a power-
ful means of coping with the daily demands of crushing poverty. Wells and
Evans (2003) considered whether exposure to nature in a sample of rural
children was related to their levels of stress. Their results suggest that children
living near nature were buffered against the potential impacts of life stress to a
greater extent than those with little nature nearby. Nielsen and Hansen (2007)
also show that Danish adults (18–80 years old) who have access to a garden or
short distances to green areas in the neighbourhood are associated with lower
self-reported stress levels. These studies suffer from the same associative links
so cannot be used to prove causality.
Studies of stress recovery are, however, amenable to experimental work that
provides stronger links between stress levels and human physiology. Ulrich
et al. (1991) subjected 120 people to a stressful movie and then exposed them to
videotapes of six different natural and urban settings. Data concerning their
recovery from the stressor were obtained from self-ratings of affective states
and a range of physiological measures including heart period, muscle tension,
skin conductance and pulse transit time. Findings from both measures con-
verged, indicating that recovery was faster and more complete in subjects that
were exposed to natural rather than urban environments. Hartig et al. (1991)
carried out quasi-experimental and true experimental work to assess the utility
of different theoretical models of restorative experience related to nature and
greenness. This illustrated that groups of people who vacationed in wilderness
situations showed much improved proofreading ability in comparison to con-
trol groups who vacationed elsewhere. In a second study, in the same paper,
they report that a group of people situated in a ‘natural’ rather than an ‘urban’
environment fared better in a range of attentionally fatiguing tasks.
Parsons et al. (1998) examined whether stress and recovery from stress varied
as a function of roadside environment in a sample of 160 college participants.
The participants were subjected to two stressors while being continuously
monitored for facial electromyographic (EMG) and autonomic (electrocardio-
gram, blood pressure and skin conductance) activity and then played a videotape
 IMPORTANCE OF GREEN SPACES FOR PEOPLE & BIODIVERSITY 249

that simulated a drive in one of four different environments. Participants who

viewed nature-dominated roadside environments exhibited less stress and
recovered from it more fully and faster than the control group who viewed
urban environments. Hartig et al. (2003) compared psycho-physiological stress
recovery and attention restoration in natural and urban field settings using
measurements of blood pressure, emotion and attention collected from 112
young adults. In the two-way experiments urban settings led to increased blood
pressure, heightened negative emotional responses and attention deficiencies.

Health inequalities
There is now a growing body of evidence that supports the notion that disad-
vantaged groups in society can gain more health benefits from green space
provision. Mitchell and Popham’s (2008) study factored in covariability of
access to green space by socioeconomic group and showed that people who
were more deprived and had good access to green space exhibited lower
incidence rates of all-cause and circulatory mortality than did those in areas
where green spaces were fewer. Importantly, this suggests that higher exposure
to green space could save 1328 lives per year in people suffering severe depriv-
ation. Other large epidemiological studies provide corroboratory results. De
Vries et al. (2003) showed that health effects of green space were more signifi-
cant in lower-income groups, and Maas et al. (2006) found that people
belonging to lower socioeconomic groups, the elderly, youth and higher-
educated (post-school) people seemed to gain more from access to green space.

Green space quality

One of the limitations of health/green space studies is that the analyses fail to
examine what it is about the green space that leads to improvements in health.
Is it the distance to the green space, its quality or its structure that matters, or
all three? Hillsdon et al. (2006) examined the relationship between access to
high-quality urban green space and the level of recreational activity in middle-
aged people (40–70 years). Their results show no clear linkages between the
distance to, quality of, or size of the green spaces and the amount of physical
activity recorded for the respondents. This is in contrast to other studies that
showed that proximity was important (Nielsen & Hansen 2007) and that access
to a large attractive public space is associated with increased probability of a
high level of walking (Giles-Corti et al. 2005). A recent study suggests that
people do appear to value and recognise nature in their local green spaces,
illustrating additional and important synergies between biodiversity in green
space and wellbeing. Fuller et al. (2007) analysed the linkage between biodiver-
sity in green spaces and psychological wellbeing. Plants, butterflies and birds
were sampled in 15 green spaces in the city of Sheffield during 2005. Linear
regression showed that the psychological measures were positively related to
250 J. SADLER, A. BATES, J. HALE AND P. JAMES

species richness and park area, and that local users can perceive species
richness reasonably well. Although the links found are illuminating, the
authors stress that they are not proof of causality.
In summary, the association of green space provision and usage to health
benefits is a positive one although it is not easily generalised. Notwithstanding
the attempts of researchers to use innovative sample design and careful mod-
elling of covariates, most studies are correlative and do not indicate underlying
mechanisms. Several researchers have argued that the distinctions between
people, place and health are contrived, and that there is a need to reconcep-
tualise (or reimagine) the relationship between the variables to emphasise their
complexity, history and relationality, and how these manifest themselves in
health outcomes and individual behaviours (Tunstall et al. 2004). This senti-
ment is shared by a growing number of health professionals (e.g. Frumkin
2003) who suggest that more emphasis needs to be placed on researching what
kinds of nature are experienced in green spaces. Put another way: what nature
is going to be viewed and how are users going to contact it? This should lead to
the formulation and definition of more meaningful health endpoints, rather
than the endless run of metrics and targets that are pervasive currently.
Frumkin (2003) also suggests that design features (e.g. amount and type of
vegetation, quiet areas for sitting, recreational amenities and so on) also need
to be carefully considered.

Synthesis: designing and managing green spaces

for people and wildlife
The use of habitat creation schemes as compensation for ecological features in
green spaces that are lost because of (re)development is commonplace, but
where the redevelopment schemes replace former derelict or brownfield sites,
these heavily managed landscaped open spaces are much less biologically
diverse than the pre-development landscapes they replace (Donovan et al.
2005; Figure 10.2). Much more careful thought, testing and design of these
schemes is needed, similar to the work carried out examining the biodiversity
value and possibilities for enhancement of garden ecosystems (Davies et al.
2009). Clearly, if agricultural land is developed for housing, as part of an
increasing urban sprawl, then it is possible that local diversity could increase
rather than decrease (Gaston et al. 2005). Where the removal of green space has
to occur, and remnants of the former landscape cannot be preserved, potential
still exists for ‘green engineering’ the built environment in a manner that
maximises its ecological function. A myriad of green technologies now exist
that can be used for this purpose, ranging from mitigation for particular
species, such as nest and roost boxes for birds, bees and bats, to broader-scale
initiatives such as permeable pavements, living walls and green (biodiversity)
roofs which may provide some biodiversity, ecosystem and social benefits
 IMPORTANCE OF GREEN SPACES FOR PEOPLE & BIODIVERSITY 251

Figure 10.2 Comparison of species richness of designed and derelict open spaces
in a redevelopment site in Eastside (Birmingham, UK). Site names on x axis are
as follows: Mill. Point ¼ Millennium Point; Pk. St. Gdns ¼ Park Street Gardens;
Ast. Cir. ¼ Aston Circus; Canal Ent. ¼ Canal Entrance; Dig. Br. Canal ¼ Digbeth
Branch Canal; Jenn. Rd ¼ Jennings Road; New Bond Street; Rea/Canal ¼ River Rea
and canal junction. Reproduced with permission from Donovan et al. (2005).

(Oberndorfer et al. 2007). There are, however, no systematic studies that test
their effectiveness in urban areas to mitigate the effects of lost green space.
The UK Royal Commission on Environmental Pollution (RCEP 2007) stated that
‘the natural environment should be at the heart of urban design and manage-
ment’ (p. 83). The wide range of policy documents at a national (Table 10.1) and
international level (EEA 2009) suggests that the political will now exists to move
towards a city future that includes more, better-designed and multi-functional
green spaces. For this to take place green spaces must be viewed as an important
component of the urban complex social–economic–ecological linked system.
Pickett et al. (2001) provide a conceptual framework which helps contextualise
the importance of urban green spaces, but its emphasis is on socio-ecological
linkages. Tzoulas et al. (2007) add to this greatly by articulating the relationships
between green spaces and public and ecosystem health, while others focus their
attention on creating models to predict the impact of urbanisation on biodiver-
sity (Williams et al. 2009).
While such frameworks are of considerable use in highlighting the complex
linkages between the diverse systems, and targeting areas of research need and
policy gaps, they do not necessarily provide concrete guidance for action on
the ground where there is a clear need for a stronger design framework
252 J. SADLER, A. BATES, J. HALE AND P. JAMES

(Hunter & Hunter 2008; Pickett & Cadenasso 2008). Niemelä and his colleagues
have consistently argued for more focus and emphasis on the planning process
(Niemelä 1999; James et al. 2009). Planners require spatially explicit data that
can be used to assess the roles that different green areas have in providing
people with ecosystem services (including biodiversity provision). There is a
growing body of scientific evidence to suggest that this is possible, at least in
terms of creating tools to simulate and model the impacts of fragmentation
(Andersson & Bodin 2009). What has been missing until recently is a means of
integrating spatially explicit biological and economic models that can be used
to find the most parsimonious land-use configurations (Polasky et al. 2008). We
suggest that, in some regions at least, the policy instruments needed to facili-
tate change are in place in the spatial planning system. What is required is the
local political will to implement them.

Future research directions

There is no doubt that urbanisation is a pervasive environmental issue that is
set to increase as the global population grows (Grimm et al. 2008). This will lead
to further fragmentation, isolation and degradation of green spaces in cities,
especially if urban sprawl is contained. This review has illustrated that green
spaces provide numerous benefits in terms of their value for biodiversity,
provision of ecosystem services, and benefits for public heath and wellbeing.
There is a pressing need therefore to place green spaces high on the political
agenda and work towards the use of policies that are already in place to protect
and enhance green space provision for people and wildlife. For this to occur,
several key challenges must be addressed (cf. James et al. 2009). First, the
physical science base needs strengthening. There is a need for: (i) more reliable
and quantified data on green space provision in cities; (ii) the creation of
baseline data from which loss and predicted loss can be estimated; (iii) esti-
mates of the changes in species richness and functional diversity as a result of
green space loss; (iv) the extension of landscape modelling to a wider portfolio
of ecosystem services; (v) more careful testing and monitoring of the efficacy of
designed green spaces that are created as part of ongoing urban changes and
development; and (vi) a greater emphasis on species-specific studies, perhaps
focusing on indicator species of conservation concern (Evans et al. 2009). There
are still large gaps in our autecological understanding of a range of species that
inhabit urban green spaces, even those that have evident and significant
relevance such as insect pollinators (Matteson et al. 2008).
Second, in terms of the social science, health and wellbeing and policy
arenas, emphasis needs placing on: (i) monitoring and assessing variability of
access to green spaces – few comparable studies exist here, but this lies at the
heart of current government and international policies concerning the quality
of life of urban citizens; (ii) strengthening the qualitative research that views
 IMPORTANCE OF GREEN SPACES FOR PEOPLE & BIODIVERSITY 253

green spaces as green places that are in part socially constructed – this should
help in the creation of better-designed spaces that maximise health and
wellbeing benefits; and (iii) the creation of models that are easily integrated
into planning systems.

Acknowledgements
This work and research has been funded by EPSRC grants EP/E021603 and EP/
007426/1 and an award from the Big Lottery Fund via the Open Air Laboratories
(OPAL) network. We are grateful for comments and discussions with colleagues
within our respective schools and the wider Urban Nature Community of Interest.

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CHAPTER ELEVEN

Integrating nature values in urban

planning and design
ROBBERT SNEP AND PAUL OPDAM

Introduction
In several chapters of this book it is emphasised that urban ecosystems are
under constant threat by ongoing urbanisation, by biological invasions and
by human use. Other chapters, however, highlight the specific character of
urban ecosystems (that by definition is indicated by the dominant role of urban
dynamics) and their importance to human wellbeing and quality of life. This
divergence illustrates two fundamentally different views of urban ecosystems:
as remnants of (previously extensive) natural ecosystems threatened by
ongoing human pressure, or as green patches with high use value to people
who work and live in the city landscape. From the first perspective urban
ecosystems are separated as much as possible from the urban system, and from
the second they are an essential part of that system.
These divergent views on urban ecosystems illustrate the type of debate that
can be expected from the variety of participants in an urban planning process.
They may hold opposing views and economic stakes, but have to develop a
common position on the desired future of the urban landscape, and what it
would take to achieve this. In this chapter we consider how both the
conservation (in terms of defending pieces of highly valued ecosystems against
undesired changes) and the enhancement (in terms of creating additional
ecological, social or economic value) of urban ecosystems can be integrated
into decision making on urban development.
We consider urban ecosystems at two spatial scales. At the site level we
distinguish a patch of a particular ecosystem type, interacting with its urban
context. At the level of the urban landscape we distinguish a pattern of ecosys-
tem sites, which may interact and thereby build an ecosystem network embed-
ded in the urban matrix. This network may also interact, at a regional scale,
with ecosystems situated outside the urban context. This distinction between
scales is essential, because decision making at the local level is different from

Urban Ecology, ed. Kevin J. Gaston. Published by Cambridge University Press.

# British Ecological Society 2010.
262 R. SNEP AND P. OPDAM

that at the city level, occurring on a different timescale, with different actors
using different criteria when identifying goals and measures. For example,
biodiversity in a local park is affected by decisions on local management taken
by the park manager. The size of the park, and the configuration of park sites
across the city, are the result of city planning by the local authority.

How is nature in the urban context conceptualised?

Urban ecosystems differ from the rest of the urban landscape because their
structure as well as their functioning is the result of an interplay between
human and natural processes. Green elements dominate the site of an urban
ecosystem, instead of stone, glass and concrete, and a significant part of the site
has established without the direct action of humans. So from this perspective,
nature is the opposite of human culture and technique. Planning for urban
ecosystems is then about deciding on the amount, character and functioning of
sites where natural processes dominate. The basis for this decision is value,
attributed by citizens, entrepreneurs or politicians.
In this chapter we distinguish two essentially different definitions of value:
the intrinsic value of natural assets, for example the value attributed by
humans to the occurrence of rare species, and the use value, the benefit
humans experience from urban ecosystems. Most current nature conservation
takes intrinsic value as a goal. From that perspective, conservation can be
defined as being a philosophy of managing the environment in a manner that
does not despoil, exhaust or extinguish ( Jordan 1995). Urban environments,
however, are so much altered that only few remnant patches may resemble the
natural situation. The major part of the urban landscape consists of built-up
areas or designed urban green spaces in which natural mechanisms (e.g.
ground and surface water flows, succession of vegetation) are predominantly
replaced by anthropogenic structures and management (e.g. sewer systems,
gardening). This affects the quality of the soil, water and air (e.g. Sukopp &
Starfinger 1999; Li et al. 2001; Penttinen et al. 2001) and the local climate in
terms of temperature and rainfall (Landsberg 1981; Oke 1982; Arnfield 2003).
The functioning of the urban ecosystem is therefore much more influenced by
human activities than in other ecosystems. Consequently, the prediction of, for
example, the distribution of plant and animal species cannot be achieved
predominantly from knowledge of natural driving-forces and disturbances.
Therefore, the planning and management of urban ecosystems cannot simply
draw on the experiences obtained in conservation planning in rural or natural
landscapes, and much effort should be put into defining what is conserved or
restored, with what purpose and against what reference.
Conservation of urban ecosystems may be legitimised by the occurrence of
endangered plant and animal species (e.g. Mörtberg & Wallentinus 2000), and
conservation measures may be aimed at sustaining the local populations.
 INTEGRATING NATURE VALUES IN URBAN PLANNING 263

Compared with rural and natural areas, the inherent value of urban environments
is often considered to be low, based on the assumption that only a few urban
species are listed as endangered. However, when urban areas have been thor-
oughly mapped for their biodiversity, it has become apparent that numerous
endangered species can be found and that urban species richness is much
higher than anticipated (e.g. Araújo 2003; Hope et al. 2003; Kühn et al. 2004).
Nevertheless, urban ecosystems are less appreciated for their inherent value by
policy- and decision-makers, and by the general public (e.g. Tucker et al. 2005).
A second justification for conserving urban ecosystems is that they bring
nature into the mindset of urban people. People may learn the importance
of nature and natural processes at first hand, and get to appreciate the value of
nature. According to Murphy (1988), ‘Our urban centers can be viewed as
bellwethers of our global environmental fate. Our success at meeting the
challenges of protecting biological diversity in urban areas is a good measure
of our commitment to protect functioning ecosystems worldwide. If we cannot
act as responsible stewards in our own backyards, the long-term prospects for
biological diversity in the rest of this planet are grim indeed.’ So, even if
restoring the natural situation is physically impossible, urban environments
can act as a demonstration area for nature conservation and restoration prac-
tices (see also McKinney 2002; Tucker et al. 2005).
The third justification for urban ecosystems is the services that they provide
to the quality of the physical environment for human life. As described in
earlier chapters (Chapters 9 and 10), urban ecosystems directly and indirectly
increase the quality level for people to live and work in cities in terms of, for
example, health, wellbeing, recreation opportunities, water retention and air
pollution filtering.
In the remainder of this chapter, we will summarise current conservation
practices, and subsequently present our views as to how nature conservation
can become part of the integrated planning of urban landscapes.

A review of conservation practices

Current support for conservation in urban societies
Because of the close contact between urban ecosystems and citizens, one would
expect these ecosystems to be a popular and broadly discussed issue in city
planning. However, based on a review of 217 papers published in the journal
Conservation Biology between 1995 and 1999, Miller and Hobbs (2002) state that
so far little attention has been paid to conservation in human settlements. They
argue that, for a variety of reasons, conservation has tended to focus on lands
with a relatively small human presence, often dominated by resource extrac-
tion and agriculture. They therefore call for extra attention to areas where
people live and work (Miller & Hobbs 2002). Their highly cited paper provides a
view on the conservation of wildlife and other natural values in urban
264 R. SNEP AND P. OPDAM

environments. Opposing views have been put forward in response to this paper.
Adams (2005), for example, illustrates that wildlife conservation in cities and
villages has a long history, and includes a range of research and conservation
activities taken in different countries throughout the world. Among other
initiatives, Adams refers to the UNESCO-MAB (Man and Biosphere) programme
which included the Urban-Forum in the United Kingdom, the Urban Wildlife
Trusts in several UK cities, the pioneer work on urban vegetation by Sukopp in
Berlin, Germany (Sukopp 1986), Durban’s Metropolitan Open Space System
(South Africa), the multiple-use green corridors in Singapore, the work of the
Australian Research Centre for Urban Ecology (ARCUE) in Melbourne, Australia,
and the Center for Urban Ecology of the US National Park Service. Adams shows
that over time numerous institutions, both public and private, have recognised
the value of urban nature, and therefore the need for conservation. This notion
was confirmed at the international mayors’ conference ‘Local action for bio-
diversity’ (Bonn, Germany, May 2008), where the importance of paying atten-
tion to biodiversity conservation in cities, the place where most people live,
was stressed. Adam’s review also suggests that urban nature conservation has
a distinct character. Education, human–wildlife interactions and the multi-
functional use of wildlife habitat seem to play a more prominent role in urban
nature conservation than in the conservation of other areas. Also, the ‘urban’
aspect means that many conservation initiatives are taken at the city level,
often in close cooperation with local governments and citizens. Urban conser-
vation practices are therefore more of a city than a regional or national event,
are on a highly applied level, and most times include a range of societal and
non-governmental stakeholders. This could explain why these initiatives are
rarely described in journals like Conservation Biology. Miller and Hobbs (2002) do,
however, bring up a good point by stressing the lack of interest by most
ecologists, and conservation biologists in particular, in the urban environment.

Current strategies
Traditionally, conservation and development of urban nature may start from
the perspective of water management, forest management, urban green man-
agement or wildlife management. For all, the perception of citizens towards
urban nature plays an important role. Based on literature and practical experi-
ence we here distinguish three types of nature conservation and development
practices in urban areas: traditional nature conservation, conservation and
development of multi-functional urban green and water, and conservation-
inclusive architecture (Table 11.1). This classification is based on the relation
between the zoning of the area, how citizens perceive that zoning, and – as a
result – the applied conservation or development practice. All three types of
practice can take place both at the city and site level, but in general only the
conservation and development of multi-functional urban green takes place at both.
Table 11.1 Three types of nature conservation and development practices in urban areas.

Conservation and development

Traditional nature conservation of multi-functional green and water Conservation-inclusive architecture

Zoning Conservation of biodiversity and natural Conservation of biodiversity and natural Conservation of biodiversity and natural
resources is a main goal of the area resources is only one of the goals of resources is not a goal of the area,
the area but only an extra value (on top of the
targeted (socio)economic or other
values)
Value The intrinsic value of nature is The use value of nature is emphasised; Both use and intrinsic value can be
emphasised; use value has less intrinsic value has less priority important, depending on the actual
priority practice
Type of Conservation measures are similarly Conservation practices are tuned with Conservation practices are tuned with
action applied as in other, ‘more natural’, the other green functions of the the architectural functions of the
areas outside the city zoning area zoning area
Appearance The zoning area is (looks like) a remnant The zoning area looks like The zoning area is a built-up area with
of a natural area ‘well-managed urban green or water’ extra attention for biodiversity
conservation
Location These areas are often located at the These areas are often located in These areas are located within the
margin of the urban landscape between the built-up areas within the built-up area of the urban landscape
urban landscape
266 R. SNEP AND P. OPDAM

The other two types of practice are implemented at the site level. In
Table 11.1 we describe the three classes in terms of zoning (how are the
areas labelled?), value (is the emphasis on intrinsic or user value?), type of
action (what is the character of the conservation or development actions?),
appearance (what does the area look like?) and location (in what part of the
city the action is taken?).
We now further examine these urban nature conservation and development
practices, using existing cases to illustrate what type of activities are included.

Traditional nature conservation practices applied

in urban landscapes
Although cities contain ‘typical urban habitats’ (e.g. Sukopp & Weiler 1988)
with associated species, such as the well-known feral pigeon Columba livia
(Murton et al. 1972), these species occur in cities because urban environments
may resemble their natural habitats (Wheater 1999). Plant and animal species
that have colonised urban areas have recognised the habitat opportunities that
cities offer and have adapted their lifestyle to the urban situation (e.g. Moller
2009). In terms of conservation, this means that urban environments –
although unique as habitat – are not the only places where a species is
occurring. If conservation of a particular species is targeted, this can also be
done in rural areas where implementing conservation measures is probably
easier and cheaper (e.g. more land is available for conservation). Conservation
in urban environments may therefore be much more based upon the demand
from citizens to protect local nature than considered from a nationwide
perspective on species conservation.
Nature conservation in cities and towns is often focused on (i) preserving
native vegetation, or (ii) conservation and restoration of wildlife habitat. We
here describe some examples of these practices.

(i) Preservation of native vegetation

Preserving native vegetation requires considerable efforts from green man-
agers, because urban environments often appear to favour exotic above native
species. Intensive management to protect native vegetation may only be pos-
sible (or fundable) if protecting this vegetation is of interest to citizens and
other parties. Here we provide some examples.
In a fragmented forest in Nishinomiya Shrine, southeastern Hyogo Prefec-
ture, Japan, active vegetation management was used to control the abundant
invasive plant species (e.g. Trachycarpus fortunei). In 2005, restoration measures
were undertaken to remove all individuals of T. fortunei from the forest,
resulting in increased canopy openness in the understorey. However, as the
understorey lacked seedlings and saplings of native species, re-vegetation may
additionally be necessary to accelerate re-establishment of native species in
 INTEGRATING NATURE VALUES IN URBAN PLANNING 267

this and other urban forest areas that are heavily invaded by exotic species
(Ishii & Iwasaki 2008).
In Wisconsin, USA, Wilcox et al. (2007) investigated whether the use of
herbicide with burning, clipping or seeding could reduce the cover of the
invasive perennial grass Phalaris arundinacea (reed canarygrass) and increase
the cover of native species in an urban wet prairie that receives stormwater
runoff. Although initially the use of herbicides decreased the abundance of
P. arundinacea and led to an increased cover of native species, two years after
treatment the exotic plant species recovered and its abundance was no differ-
ent from control plots. The researchers conclude that land managers should be
selective in allocating efforts to control monotypes of invasive species. Where
wetlands continually receive stormwater, the replacement of invasive mono-
types by native species will be difficult and the effort required might be
prohibitively expensive and ineffective in the long term. Highest priority
should go to sites that can best support native species restoration.

(ii) Conservation and restoration of wildlife habitat

Practices of wildlife conservation in urban areas resemble those in rural
environments. The conservation of specific species and their habitats is the
main goal of the measures taken, and ‘urban’ characteristics such as housing
development and intensive human presence are considered as threats rather
than as part of the dynamics of the urban landscape. These practices put the
emphasis on ‘conservation’ and ‘restoration’. For example, Brawn and Stotz
(2001) describe the case of the Chicago Wilderness initiative in which 60
public and private organisations undertake efforts to conserve and restore
habitats for breeding and migratory birds within the Chicago metropolitan
region, USA. They conclude that habitat protection in otherwise developed
metropolitan areas may offer significant conservation opportunities in
regions where natural habitat is scarce. Similarly, the London Biodiversity
Action Plan (LBAP) details plans to enhance the survival of rare and valued
species associated with some 15 London habitats such as woodland, heath-
land, railway line sides, private gardens, cemeteries and wastelands (Harrison
& Davies 2002). This action plan is part of the UK-wide Biodiversity Action
Plans (BAP), in which conservation of endangered plant and animal species
and their habitats are targeted. For the Greater London area, species like the
peregrine falcon Falco peregrinus, house sparrow Passer domesticus and black
redstart Phoenicurus ochruros are listed, and specific conservation measures
such as habitat restoration are defined (see the website http://www.lbp.org.
uk/londonap.html). These measures include contacting owners of suitable
buildings for construction of nesting sites, education of the public in the
species’ habitat requirements, and monitoring and studying the London
populations.
268 R. SNEP AND P. OPDAM

In South Africa, Gauteng’s last mountain wildlife ecosystem is being decimated

by ongoing urbanisation from cities like Johannesburg. Environmental NGOs
aim to protect the natural ecosystems on which the locally present birds of
prey (including the black eagle Aquila verreauxii) and other wildlife species
depend for their continued survival (http://www.blackeagles.co.za). In 2008,
several plans (including an ‘Urban Wildlife Reserve’) were developed to con-
serve raptors occurring at the edge of the cities. The proposed Urban Wildlife
Reserve aims to protect fauna and flora currently threatened by development,
by creating a unique sanctuary within the Johannesburg Metropolitan Area.

Conservation and development of multi-functional green and water

In urban green planning, design and management, there are two concepts that
address the conservation of natural values as part of multi-functional green or
water at the city level, those being the greenway concept and the concept of
urban stream rehabilitation. Both take the use value of the urban green or
water as a starting point.
Greenways are networks of land containing linear elements that are
planned, designed and managed for multiple purposes including ecological,
recreational, cultural, aesthetic or other purposes compatible with the concept
of sustainable land-use (Ahern 1995). ‘Multi-objective’ greenways address needs
of wildlife, flood damage reduction, water quality, education and other infra-
structure in addition to urban beautification and recreation (Searns 1995).
These greenways may also include streams and other water bodies (Asakawa
et al. 2004). In urban environments, greenways can support the occurrence of
wildlife. Bryant (2006) demonstrated the critical role of ecological greenways
and parks in addressing the need to conserve biodiversity. More specifically,
Mason et al. (2007) illustrated that, if appropriately designed, greenways may
provide habitat for neotropical migrants, insectivores and forest-interior spe-
cialist birds that decrease in diversity and abundance as a result of suburban
development. They recommend that ‘landscape and urban planners can facili-
tate the conservation of development-sensitive birds in greenways by minimis-
ing the width of the trail and associated mowed and landscaped surfaces
adjacent to the trail, locating trails near the edge of greenway forest corridors,
and giving priority to the protection of greenway corridors at least 100 m wide
with low levels of impervious surface (pavement, buildings) and bare earth in
the adjacent landscape’. In general, the extent to which urban open space
corridors meet recreational, habitat and wildlife needs is highly dependent
on the degree to which the interactive effects of these uses are identified and
incorporated into planning, design and management decisions (Briffet 2001).
For managers who aim to enhance compatibility between people and wildlife,
the challenge is to bridge the gap between established practice and a more
multi-functional approach. Ultimately the aim should be to put in place an
 INTEGRATING NATURE VALUES IN URBAN PLANNING 269

enlightened and sympathetic management regime that supports indigenous

biodiversity maintainable at an acceptable cost and is compatible with provid-
ing sufficient access to cater for a wide range of recreational activities.
Whereas greenways are networks of land planned, designed and managed
for multi-functional purposes, stream rehabilitation is more focused on
existing urban water networks with ecological roles that have over the years
been neglected. If the rehabilitation of ecological functioning is the goal, urban
streams may better support the intrinsic and use value of the urban ecosystem.
For example, Larned et al. (2006) illustrated the effects on invertebrates and
non-native macrophytes of transplanting patches of native macrophytes into a
230-m-long stream section. In this experimental urban water restoration pro-
ject, native macrophytes seemed to limit the spread of non-natives, which were
absent in the planted section by the second spring. Native macrophyte estab-
lishment did not enhance invertebrate communities as predicted; few inverte-
brate metrics differed significantly between planted and unplanted sections.
Pollution- and sediment-tolerant invertebrate taxa were abundant in both
sections, suggesting that invertebrate colonisation was limited by water quality
or sedimentation, not macrophyte composition. Survey respondents con-
sidered the stream to be visually and ecologically improved after rehabilita-
tion, and macrophyte establishment was generally considered positive or
neutral. According to Booth et al. (2004), successful stream rehabilitation
requires coordinated diagnosis of the causes of degradation and integrative
management to treat the range of ecological stressors within each urban area,
and it depends on appropriate remedies at scales from backyards to regional
stormwater systems. Recent studies of urban impacts on streams in Melbourne,
Australia, suggested that the primary degrading process in many urban areas is
effective imperviousness, the proportion of a catchment covered by impervious
surfaces directly connected to the stream by stormwater drainage pipes (Walsh
et al. 2005). This implies that even small rainfall events can produce sufficient
surface runoff to cause frequent disturbance through regular delivery of water
and pollutants; where impervious surfaces are not directly connected to
streams, small rainfall events are intercepted and infiltrated. Walsh et al.
(2005) showed, in a sample catchment, that it is possible to redesign the
drainage system to reduce effective imperviousness to a level at which the
models predict detectable improvement in most ecological indicators. Distrib-
uted, low-impact design measures are required that intercept rainfall from
small events and then facilitate its infiltration, evaporation, transpiration or
storage for later in-house use.

Conservation-inclusive architecture
In discussions of the opportunities for conservation and development of bio-
diversity and other ecosystem services in city environments, the focus has
270 R. SNEP AND P. OPDAM

mostly been on the non-built-up parts. It is only over the past 20 years that the
conservation value of buildings and other built-up structures present in urban
areas has begun to be recognised. This increasing attention has originated from
several directions, from architects aiming for ecological design (Todd & Todd
1994), from the emergence of ecological engineering as a discipline (Shijun
1985; Gattie et al. 2003) and from biodiversity conservation actions in urban
environments (e.g. the EU Countdown 2010 programme).
Over several decades, architects have developed an interest in integrating
ecology in the design of the buildings. This movement was called ‘green design’
(1980s), later ‘ecological design’ (1990s) and these days ‘sustainable design’,
representing an increasing broadening of scope in theory and practice (Madge
1997). Well-known examples of the ‘greening of buildings’ are ‘Le Mur Vegetal’
by Patrick Blanc (complex vegetation structures growing vertically, attached to
several buildings in Paris, France) and the green roof of the Chicago City Hall
(an impressive rooftop garden in the centre of Chicago; Velasquez 2005).
Whereas architects focus on buildings and combine visual appearance with
functional quality, engineers are more interested in innovative technical
systems that improve both human experiences and wellbeing, and environ-
mental qualities. According to Bergen et al. (2001), ecological engineering is
‘the design of sustainable systems, consistent with ecological principles, which
integrate human society with its natural environment for the benefit of both. It
recognises the relationship of organisms (including humans) with their envir-
onment and the constraints on design imposed by the complexity, variability
and uncertainty inherent to natural systems.’ In the built-up environment
engineers search for new ways to deal with water retention, air quality, energy
saving, noise reduction and other issues to improve their liveability. As the
ecological design of buildings may increase their sustainability, engineers have
developed new building techniques that are inspired by ecosystem functioning.
Just as for the architects, the concept of green roofs and living walls (Dunnett &
Kingsbury 2004) is an important part of that, but here functionality is much
more important than visual quality.
A third group that explores how buildings can support biodiversity and
other ecosystem services is wildlife conservationists. As in recent years the
conservation of biodiversity in the urban environment has received increasing
attention, so now buildings and other built-up structures are – apart from their
human functionality – also considered from a wildlife conservation point of
view. Because of their size, shape, material and location, conventional build-
ings may mimic habitat conditions for particular species that originate from
rocky and cliff environments. In addition, when ‘green design’ measures (green
roofs, living walls) are added to buildings, the range of habitat opportunities is
broadened, and more plant and animal species may find their habitat in or on
these man-made structures (Brenneisen 2003, 2006; Grant et al. 2003). In more
 INTEGRATING NATURE VALUES IN URBAN PLANNING 271

and more cities, wildlife conservationists actively encourage owners to

integrate specific conservation measures for wildlife (e.g. nest boxes for breed-
ing birds or bats) in their buildings (Baines 2000).

Incorporating nature into urban planning

Legitimacy of ecosystems as part of the city landscape
Spatial landscape planning is the organised process of attributing significance
to landscape functions, making decisions on values to be created or preserved,
and spatially organising and designing the physical structure of landscapes to
ensure the provision of landscape services in the short and long term. City
planning which includes ecological (‘green’) functions can be considered as
urban landscape planning. It is fundamentally different from conservation
planning, in that it considers a wide range of functions and values of multi-
functional landscapes. Because urban development is guided by human values
(Andersson 2006), urban landscapes are not planned primarily for rare species
and habitats (such as in conservation planning).
If remnant patches of former large natural landscapes in cities are con-
sidered as threatened sites to be protected from the devastating influence of
humankind, any conservation action is opposed to and conflicts with the
human socioeconomic system (as for example in Breuste 2004). In this chapter,
we take a different point of view, and consider ecosystems as part of an
ecological–social system. This implies that ecosystems (the ‘green’ elements
in the urban landscape) are considered as an intrinsic part of the urban system,
and that planning of ecosystems in the urban landscape is about the values
they could provide to the urban system as a whole. Therefore, although eco-
logical principles determined in rural landscapes are applicable to urban
landscapes (Savard et al. 2000), conservation planning rules as discussed in
the literature (Cabeza & Moilanen 2001; Margules & Sarkar 2007; Pressey et al.
2007) cannot simply be transferred to urban landscapes. We propose that a
transition from traditional conservation towards ecology as a cornerstone of
sustainable urban development (as described by Wu 2008) creates opportun-
ities to better position nature and biodiversity in urban planning.
The consequence of this view is that urban ecosystems are not planned and
designed for protected species or native vegetation, but for the ecosystem
services that the urban society cares for, at a level of intensity valued by that
society, and on which government and private enterprises choose to spend
money (Boland & Hunhammar 1999). Urban ecosystems are known to provide a
series of values to inhabitants, including human health, social cohesion, regu-
lation of air quality and temperature, stormwater regulation, nature percep-
tion and leisure activities ( Jackson 2003; Chiesura 2004; Tzoulas et al. 2007;
Miller 2008). Similar benefits were found in relation to employees and enter-
prises (Kaplan 2007; Snep et al. 2009). Biodiversity can be valued as a key factor
272 R. SNEP AND P. OPDAM

in well-functioning ecosystems which provide ecosystem services in the long

term (Hooper et al. 2005), as well as for the significance that humans attribute
to observing wild animals and plants in their living environment.

Planning at three levels of scale

Planning of ecosystems within the urban landscape occurs at multiple spatial
scales driven by the scale of influence of the decision-maker (Andersson 2006).
We distinguish three levels of spatial scale: national or regional scale
(depending on the scale at which governmental responsibilities for spatial
planning are established), the scale of the city and that of the site. At the first
level, strategic decisions are made about the density of buildings and infra-
structure in comparison to the area of green structures. As shown by Tratalos
et al. (2007), the contribution of ecosystems to the urban system is a matter of
the proportion of green area within the city landscape: the more ecosystem
area, the higher the level of ecosystem services. However, increasing the pro-
portion of the green component within the city limits means that the urban
zone expands further into the rural landscape (causing potentially detrimental
impacts on rural values, including nature reserves). It also implies that humans
have to cover greater distances between functional sites, which is potentially
detrimental to environmental quality, increases travel costs and the emission
of greenhouse gases, and may cause loss of social cohesion. Balancing these
trade-offs across different spatial scales requires a fundamental debate at the
highest level of spatial organisation, and might be organised by the govern-
ment and national planning authorities.
At the city level, the amount and spatial pattern of ecosystems (parks, green-
ways, amenities and watercourses) is determined. Ecosystems in urban land-
scapes are often small and narrow and embedded in a stony environment with
high-density road networks. Biodiversity and related ecosystem functioning is
limited by the impact of fragmentation effects. An effective strategy to improve
the values of ecosystems under pressure of fragmentation is to ensure that the
individual sites are linked in a network in which they support each other
(Opdam & Steingröver 2008) and increase the adaptive capacity of the ecosys-
tems (Opdam et al. 2006). Key characteristics of such networks include the total
network area, network density and the permeability of the urban space
between the sites (Opdam et al. 2003). These spatial characteristics are largely
determined at the level of city planning (Sukopp et al. 1995). This is also the case
for the overall pattern of urban expansion: for example, along transport corri-
dors or in satellite suburbs, or in concentric circles. The shape of the urban
zone is significant for the level of biodiversity in the city centre. Ecosystem sites
in the peri-urban zone may be functionally connected to the urban ecosystem
network (Snep et al. 2006), which supposedly increases the species diversity
within the city. An irregular shape with green wedges intruding into the urban
 INTEGRATING NATURE VALUES IN URBAN PLANNING 273

zone ensures a better connectivity between rural and urban ecosystem sites
than does a perfect circular shape. At this level of planning, the main actors are
the city planners and the governmental authorities, in dialogue with NGOs and
well-organised pressure groups.
At the site level, planning involves the identification of local targets and the
design of the site, including decisions on how to combine incompatible func-
tions spatially. The main players are governmental bodies, real estate enter-
prises, the local citizens and other interested parties. Here, the challenge is to
incorporate higher-level interests into the local decision making, for example
in the type of ecosystem which is developed. For example, does the site
functionally contribute to the larger network, or is it so different that it is
functionally isolated? A key issue is whether private partners are motivated to
invest in public services.
Scale levels are also relevant in funding conservation measures. In many
countries, nature conservation funding agencies do not consider the urban
environment as an area of interest. This means that urban nature projects may
not meet the requirements for (inter)national subsidies (e.g. because they lack
the minimum area size, the right label (‘conservation area’) or the presence of
sufficient target species). Urban nature measures are therefore often dependent
on local funding sources that are not really equipped to support biodiversity
conservation and development in the long term. This will limit opportunities
to insert wildlife conservation targets in local city planning, because creating
or conserving urban ecosystems will be considered as expensive extra costs that
inhibit the opportunities for economic expansion. However, if urban ecosystem
services are appreciated for their contribution to the economic and social
values of the urban system, local funding could become available. This oppor-
tunity is illustrated by the study of Snep et al. (2009), who showed that biodiver-
sity was not among the functions of the green office parks which were most
preferred by stakeholders, but that plant and animal species could profit from
the high preference for other urban green functions related to quality of
human life.

Goal setting
The chances for optimising the spatial pattern of ecosystem sites are much
greater if target setting is done early in the urban planning and design pro-
cesses. If conservation and development actions are undertaken once major
decisions concerning the layout of a specific area have already been completed,
it is more difficult to tune the conservation actions with the surroundings and
the (a)biotic conditions of the area, and consequently the urban ecosystem’s
functioning will be less effective. At the city level, a strategic vision of the
desirable role of ecosystems and the level of biodiversity and ecosystem services
is an important cornerstone in positioning ecosystems in negotiations about,
274 R. SNEP AND P. OPDAM

Table 11.2 Different starting points for integrating biodiversity values in urban planning
and design.

Ecological values are addressed Ecological values are

early in the planning and introduced afterwards,
development process in existing situations

Emphasis on Conditions for required Conservation measures for

species persistence of species are biodiversity depend fully on
targets inserted into the planning the opportunities offered by
process, and balanced with the existing green space and
other targeted urban values building conditions, e.g. in
parks
Emphasis on Minimal requirements for chosen Which ecosystem services can
ecosystem provision level of ecosystem be provided and at what level
services services are inserted into the fully depend on the
planning process; different opportunities offered by the
services can be combined in the existing conditions; no
same green structure and unite balancing of ecological,
different interest groups social and economic values
possible

for example, how to structure new suburbs, and how to restructure office
parks. Examples have been published in the scientific literature (e.g. Conine
et al. 2004; Li et al. 2005), and many more can be found on internet websites.
A vision may include a choice for target ecosystem types, target ecosystem
services and aspiration levels. For example, an important decision is whether
ecosystem types should be preferred that occur in the surrounding rural
countryside, to create an opportunity to link the urban ecosystem network to
the rural network and create a higher level of urban biodiversity. The vision
also emphasises where weak links in the network exist, so that any opportunity
that might occur in the dynamic urban landscape to place an ecosystem patch
in a reconstruction plan can be used. If ecosystem conservation and develop-
ment are not a part of the planning and design process, the emphasis is on
what is still possible within the constraints set by the design, and there is no
drive and no space to create more effective solutions (Table 11.2).
Legal aspects need to be considered at an early stage of planning. City
environments are multi-stakeholder centres, with highly dynamic property
ownership and land-use. Developments, even if they aim at creating opportun-
ities for ecosystems, may include destruction of habitat, and the presence of
protected plants and animals in the target area will provide environmental
non-governmental organisations with an argument to stop or delay develop-
ment projects. Again, a strategic vision on the quality of life that is well
 INTEGRATING NATURE VALUES IN URBAN PLANNING 275

established in the urban society may facilitate the debate about why and how
to adapt the metropolitan landscape to future needs.

Planning green and blue structures for ecosystem networks

Urban biodiversity and ecosystems are best conserved and developed in the city’s
green structure, because ecosystems are allowed more time to develop and
because the spatial configuration of ecosystem sites can better be optimised.
A key feature in urban ecosystems is spatial dynamics: urban development may
destroy sites, but at the same time may offer new opportunities. Ecosystem
networks (and greenways) offer a spatial structure that can better deal with such
dynamics. In efforts to optimise the functioning of this green structure for urban
biodiversity and ecosystem services, several key principles based on metapopula-
tion ecology need to be considered (Opdam & Steingröver 2008).

Size matters
Habitat size is an important factor determining urban biodiversity (Cornelis &
Hermy 2004; Donnelly & Marzluff 2004; Opdam & Steingröver 2008; Palmer
et al. 2008). Larger patches mean reduced edge effects (disturbances by adjacent
land-use), opportunities for species with larger area requirements, and larger
local populations of species. Species groups may differ in their response to
changing patch size (Godefroid & Koedam 2003). Mason et al. (2007) distin-
guished optimal width of greenways for the occurrence of different groups of
bird species, for example urban adaptors, forest interior and ground-nesting
species. Again, this emphasises the importance of goal setting. Also the func-
tional requirements of ecosystem services which are not based on biodiversity
may depend on the size of green elements, for example for climate regulation
(see Goméz et al. 2001 for a quantitative approach to relate human comfort to
the size of green zones in Valencia, Spain).

Large patches increase value for money

Verboom et al. (2001) showed that in an ecosystem network, a relatively large
patch allowed smaller total network areas to offer sustainable conditions for
metapopulations. Larger remnants of (semi-)native vegetation are the obvious
focal elements in any spatial green or blue structure. Measures to increase their
supportive role include enlargement of these sites and improved connectivity
to other sites (Opdam & Steingröver 2008; Palmer et al. 2008). While this may be
a difficult task in many older cities where space is limited (e.g. Fernández-
Juricic & Jokimäki 2001), it can be a valid approach within newer cities, such as
those in Australia, or within recently developed peri-urban areas.

Connectivity
Connectivity is a popular concept among planners and landscape architects,
but its application is still too often based on structural considerations, instead
276 R. SNEP AND P. OPDAM

of a functional analysis of ecosystem network structure. Improving connectivity

may be a solution to the total network area not being adequate for a persistent
population of a target species, or for raising the biodiversity level in a park
area in the city centre. Structural measures, such as a green corridor or
stepping stones that facilitate dispersal, may improve the spatial cohesion of
the network. Alternatively, measures that increase the carrying capacity of the
network patches (through improving habitat quality and enlarging sites) may
contribute to achieving the same end (Opdam & Steingröver 2008). In the
city landscape, road and water infrastructure may be used as a physical
template to improve connectivity for aquatic, marsh and terrestrial ecosys-
tems (Adams & Dove 1989).

City edge
The city edge is the most dynamic part of the urban landscape, with a constant
change in land-use. Owing to these dynamics temporary habitats often occur
that can have unique biodiversity levels, suggesting that a spatial development
vision for the city edge from the point of view of pioneer ecosystems may be
important. Also, in the process of urban sprawl, today’s city edge will be well
within the urban zone in a couple of decades. Therefore, any decision on
incorporating parts of today’s city edge into a future urban ecosystem network
has to be anticipated at an early stage of development.

Peri-urban areas
Rural habitat patches located adjacent to the city (peri-urban) may act as source
areas for inner city nature (e.g. ants in urban sites are more abundant where
natural areas are near; Pacheco & Vasconcelos 2007). Key features are size and
ecosystem type, as well as connectivity, for example supported by (rail)road
verges (Snep et al. 2006). Green wedges protruding into the city may increase
the functional connectivity between the inner city part of the ecosystem
network and the surrounding rural landscape. They also improve the liveabil-
ity of city environments for human beings (see for example Li et al. 2005).

Incorporating land-use dynamics in the ecosystem network

Because of its functional cohesion, an ecosystem network can stand a certain
level of turnover of patches without losing sustainability. However, there is a
relation between the turnover rate and the total network area required for
sustainability (A. J. A. Van Teeffelen et al., unpublished results). In ecosystem
types with a long development time, compensating loss of area by developing
an equal amount elsewhere is not sufficient as a compensation measure, since
it takes a long time before the newly developed patch is a functional part of the
network. If turnover is foreseen, the total area of the network should be
expanded to ensure that the network stays sustainable. This suggests that
 INTEGRATING NATURE VALUES IN URBAN PLANNING 277

within city landscapes it is efficient to keep part of the urban network stabilised,
and to connect this network with networks in the peri-urban zone. Pioneer
species often colonise new urban development lots. By including those oppor-
tunities in the urban planning process and connecting the pioneer habitats
with existing networks, added value for urban biodiversity conservation can be
obtained. For business parks, industrial estates and port areas, such an
approach has been elaborated into the ‘habitat backbone’ strategy, as applied
in the case of the natterjack toad Bufo calamita in the Port of Antwerp, Belgium
(Snep & Ottburg 2008).

Making use of the opportunities for temporary habitats

When urban green is addressed in city planning, only those green structures
are discussed that will last for many years. City environments, however, are
highly dynamic in land-use, creating numerous temporary opportunities for
the conservation and development of natural values. Locations within or at
the edge of the urban matrix that are not yet (re)developed may – for several
months or years – offer habitat for a wide range of pioneer plants and
animals, including threatened species (Breuste 2004). Typical areas for tem-
porary nature are located at older industrial sites, along (rail)road tracks and
at planned but not yet developed sites on the city edge (Straus & Biedermann
2006). By including those temporary opportunities in the city planning pro-
cess, an urban green structure with permanent and temporary elements can
be defined such that both types of green may strengthen each other. By doing
so, the opportunities for early-successional habitats are better utilised,
which will lead to an increase in the overall conservation value of the city
environment.

Planning and management of urban ecosystem sites

The effectiveness of measures at the site scale fully depends on how the spatial
pattern at a higher level supports the functional connectivity with other
ecosystem patches elsewhere. Local targets and measures have to be consistent
with the higher-level vision and strategies. We highlight here several issues
frequently discussed at the site level with opportunities to enhance nature
values in the planning and design process.

Business sites: contributing to common societal values

Business sites (e.g. business parks, industrial estates and distribution centres)
are currently not designed and managed to provide added values in terms of
sustainability to their (urban) surroundings. Snep (2009) explored options and
opportunities in planning, design and management of business sites to accom-
modate ecosystems as a source of ecosystem services to their surroundings. His
findings illustrate the significance that business sites can have for biodiversity
278 R. SNEP AND P. OPDAM

Figure 11.1 An office site design in Groningen, the Netherlands, in which specific
biodiversity conservation measures (such as a bat cave in the building and specific
vegetation for birds and butterflies in the green surrounding the building) are
incorporated in the site design. This site is actually (2008–11) being developed, including
the measures for plant and animal conservation. Artist’s impression: UNStudio and
Lodewijk Baljon Landscape architects, by order of DUO2.

conservation, based on their location, land-use and urban design. Snep shows
that by implementing different ‘green design’ measures, business sites may be
able to support biodiversity in adjacent areas, improve the liveability of neigh-
bouring residential areas and create a better quality for work at the business
sites (e.g. Figure 11.1). He demonstrates that, in terms of conservation value,
business sites are able to support the viability of threatened species, especially
those linked with early-successional vegetation (e.g. Snep & Ottburg 2008).
Implementing green roofs and opportunities for temporary habitats, and man-
aging the existing business site green in a more ecological way are among the
green design measures that could help to make use of the opportunities that
business sites potentially offer for the conservation and development of urban
ecosystems.

Buildings: green architecture

Built structures offer different opportunities for the conservation and develop-
ment of natural values. Green roofs are capable of supporting a whole range of
ecosystem services including water retention, energy savings, wildlife habitat,
 INTEGRATING NATURE VALUES IN URBAN PLANNING 279

Figure 11.2 Like the well-known Hanging Gardens of Babylon, this urban green design
in the inner city of Eindhoven, the Netherlands, illustrates how highly built-up areas
can still offer opportunities for conservation. The mix of plant species used at this
location are selected on both visual quality and habitat suitability for bees, butterflies
and birds. Photograph used with permission of Soontiëns Gardening Eindhoven.

sustainable use of building materials, and diminishing air pollution and urban
heat effects (Oberndorfer et al. 2007). Concerning their conservation value for
biodiversity, the substrate depth (not too shallow), soil origin (aim for natural
soils from the direct surroundings of the roof location) and vegetation struc-
ture (aim for variation in species mix and height of vegetation) of green roofs
are important factors (Kadas 2002; Brenneisen 2006; Dunnett et al. 2008;
Emilsson 2008). Under the right conditions, green roofs can have a significant
value for the conservation of invertebrates ( Jones 2002) and breeding birds
(Baumann 2006). Additionally, in some cases they can also increase wildlife
experience in urban environments (Figure 11.2).
With living walls (planted facades of buildings) there are also habitat
opportunities for wildlife, according to Köhler (2008). He specifically mentions
wood-adapted birds and spiders, beetles and others preferring thermophile or
synanthropic vegetation (species adapted to urban settings, including those
preferring warm temperatures; Klausnitzer 1993).
Finally, by including specific spaces in the actual building, birds, bats and
other wildlife may find nesting and hibernation opportunities. These days
there is a small industry that produces all types of ‘building blocks’ for housing
development (e.g. custom-made roof tiles with nesting places for birds) so that
these opportunities can be created on a large scale.
280 R. SNEP AND P. OPDAM

Figure 11.3 Small parcel of urban green in a highly built-up setting (business site).
Traditionally such an area will be developed and managed as lawn with some trees (left),
but with ecological management the same area could be designed in such a way that
both wildlife and citizens might much better appreciate this green site (potential
design, right). Reproduced with permission from Soontiëns Gardening Eindhoven.

Urban green management

Citizens appreciate wildlife in their direct living environment (with some
exceptions like gulls, starlings, feral pigeons), with diversity being more
important than density (Clergeau et al. 2001). To support a wide variety of
urban wildlife, great variation in the management of urban vegetation is
required. Offering a broad range of urban green types and using different
methods to manage the urban green will attract a high diversity of plant and
animal species. However, the management practice for a city’s public green is
usually rather straightforward, with cost-effectiveness and traditions in green
management being the main drivers. This results in a dominance by lawns and
street trees, and a shortage of shrub and brushwood vegetation. The latter two,
especially, include many opportunities for wildlife. If in the management of
the urban green particular attention is being paid to habitat opportunities for
wildlife, not only plant and animal species will benefit but also the people who
like to enjoy the city’s wildlife (e.g. Figure 11.3).

Gardens: private management for common values

Some recent visions of urban development focus on expansion within the city
limits, sacrificing existing urban green spaces (like gardens) for new housing or
business developments. Although a careful handling of the remnant open
space in peri-urban areas is in itself a good idea, one should also consider
the value of existing green for ecosystems and nature experience in cities
(Figure 11.4). Among urban green, gardens ‘constitute a considerable propor-
tion of “green space” in urban areas and are therefore of potential significance
for maintaining biodiversity and ecosystem service provision in such areas’
(Loram et al. 2007). For example, Wheater (1999) addresses the benefits of
 INTEGRATING NATURE VALUES IN URBAN PLANNING 281

Figure 11.4 Communal garden in ‘eco’ neighbourhood (De Kersentuin, Leidschenrijn,

the Netherlands): citizens share part of their gardens with their neighbours, thereby
creating semi-private green spaces that offer great places for both ecosystems and nature
experience.

garden plants (including many non-native plants) for producing nectar and
food (even in early spring and autumn) for bees, bumblebees, hoverflies and
birds. Baines (2000) mentions that gardens can optimally contribute to the
city’s biodiversity if managed in an ecological way, by so-called ‘wildlife
gardening’. Wildlife gardening can be broadly defined to encompass any
actions conducted in private or domestic gardens to increase their suitability
for wildlife, and thus includes the provision of a diversity of resources (e.g.
substrates, food, breeding and overwintering sites; Davies et al. 2009). Concern-
ing these actions, Gaston et al. (2005) observed that ‘whilst some methods for
increasing the biodiversity of garden environments may be very effective,
others have a low probability of success on the timescales and spatial scales
likely to be acceptable to many garden owners. If one of the functions of small
scale biodiversity enhancement is to develop and encourage awareness of
biodiversity and its conservation, then encouragement to conduct particular
activities must be balanced with a realistic appraisal of their likely success.’

Integrating nature values in urban planning and design

So far, city development and design has mainly been the territory of urban
planners, project developers and architects. Consequently, the attention and
budget have been focused on the built-up part of cities, with ‘green’ considered
as ‘accessories’ in architecture and as ‘necessary recreation area’ in between the
282 R. SNEP AND P. OPDAM

housing blocks. However, as we think of humans being part of the global

ecosystem, one could see the city as a local ecosystem where people, plants
and animals coexist, offering each other ‘ecosystem services’. From the view-
points of climate change, sustainability and urban ‘liveability’, such an idea
would offer humans a whole range of physical and social advantages (e.g. the
impact of urban green in city temperature buffering). Unfortunately, most of
us have probably not yet linked ecosystem functioning with the quality of
human city life. In particular, getting urban planners, project developers and
architects interested in the ecosystem perspective seems the largest challenge
(besides informing the general public). If in the future these urban profession-
als are willing to broaden their current point of view with the ecosystem
approach, new ways to combine ‘red’ and ‘green’ in the city will be asked for.
For now, this chapter provides a whole array of clues as to how to integrate
nature values in current urban planning and design.

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literature review. Landscape and Urban Areas Journal, 27, 354–65.
Planning, 81, 167–78. Wu, J. (2008). Making the case for landscape
Velasquez, L. S. (2005). Organic greenroof ecology an effective approach to urban
architecture: design considerations and sustainability. Landscape Journal, 27, 41–50.
CHAPTER TWELVE

Urban futures
MICHAEL L. MCKINNEY

In this chapter, I review some of the extensive literature on urbanisation to

examine the future of urban growth and its impact on the biosphere. I argue
that the relatively few ecological studies on urbanisation thus far have typically
focused on the environmental impact of cities at local spatial scales when we
also need to consider urban impacts at regional and global scales. By expanding
our view of urban impacts beyond describing local environmental impacts, it
becomes clear that cities can provide opportunities for biological conservation
at many spatial scales. For example, cities concentrate human populations and
produce economies of scale that can reduce per capita human impact on
the atmosphere and on regional watersheds, as well as on local air quality.
A ‘sustainable’ city is thus not just a city that sustains its own existence. Rather
it is a city that contributes to the long-term persistence of the biosphere and
indeed the global environment (McGranahan & Satterthwaite 2003). Ecologists
should therefore find and recommend ways that urban impacts can be reduced
on regional and global scales as well as the local scale. To that end, this chapter
devotes considerable discussion on ways to create more sustainable cities.

The future growth of cities

The proportion of humanity living in urban areas has consistently increased in
recent decades, from just 10% in 1900 to 50% in 2007. Put another way, the
twentieth century witnessed a ten-fold increase in urban populations. It has
been estimated that by 2025, five billion people will live in urban areas and
account for 65% of the global population (United Nations 2008).
More than 95% of this future net increase in the global urban population will
be in cities of the developing world. The greatest proportion of this growth is
expected to occur in Asia, where 71% of the global rural population currently
resides; the urban population of Africa, Asia, Latin America and the Caribbean
is now nearly three times the size of the urban population of the rest of the

Urban Ecology, ed. Kevin J. Gaston. Published by Cambridge University Press.

# British Ecological Society 2010.
288 M. L. MCKINNEY

world. Projections by the United Nations suggest that urban populations

are growing so much faster than rural populations that 85% of the growth in
the world’s population between 2010 and 2020 will be in urban areas, and
almost all of this growth will be in Africa, Asia and Latin America (United
Nations 2008).
This rapid urbanisation of developing nations has enormous consequences
for conservation planning because many of the developing nations are dispro-
portionately poor in monetary terms but are disproportionately wealthy in
biodiversity, especially of course those containing tropical rainforest habitats.
Roughly 72% of the urban population of Africa now lives in slums (Cohen
2006). The slum-dwelling proportion is 43% for Asia and the Pacific, 32% for
Latin America and 30% for the Middle East and Northern Africa (Cohen 2006).
Unfortunately, in many of the world’s poorest countries, the proportion of
urban poor is increasing faster than the overall rate of urban population
growth (United Nations 2008). This combination of increasing poverty and
urban population growth in nations that are rich in biodiversity is perhaps
the single greatest challenge to biological conservation today.
Another important urban trend is the growth of megacities, cities contain-
ing over 10 million people (Decker et al. 2000). At the beginning of the twenti-
eth century, just 16 cities in the world contained a million people or more.
Today, over 400 cities contain over a million people, and at least 25 cities
contain over 10 million people (Cohen 2006). Most megacities are found in
mid-latitude regions and are located near large river mouths within 100 km of
a coastline (Decker et al. 2000). The number of megacities will undoubtedly
grow, as will their impact on the environment at many scales. Grimm et al.
(2008) discuss the environmental importance of ‘megapolitan’ regions, consist-
ing of multi-city conglomerates covering hundreds of square kilometres.
Because of the enormous volumes of resources consumed by these areas, and
the enormous volumes of pollution that they discharge, these regions will have
an increasingly powerful influence on climate, watersheds, the oceans and
many other regional and global processes. However, urban conservation efforts
must also include smaller cities. Indeed, most urban population growth is
expected to occur in smaller cities with populations of less than one million.
In 2005, cities with populations under 500 000 accounted for 51% of urban
dwellers (United Nations 2008).
Finally, many urban areas, especially in the USA, have been trending toward
increasingly dispersed settlement patterns. Often referred to as ‘suburban
sprawl’ or ‘exurban’ growth, this has been driven by mass construction of large
residential developments of low-density housing beyond the urban periphery.
Low-density rural home development is the fastest-growing form of land-use in
the USA since 1950 (Hansen et al. 2005). Housing growth, in the form of both
suburban and rural sprawl, has been identified as one of the major threats to
 URBAN FUTURES 289

ecosystems in the USA (Lepczyk et al. 2007). This is also a harmful trend in other
regions. According to Liu et al. (2003), ‘In recent years, the number of house-
holds globally, and particularly in countries with biodiversity hotspots (areas
rich in endemic species and threatened by human activities), was more rapid
than aggregate population growth.’ This has increased per capita impacts on
biodiversity by increasing per capita consumption of wood for fuel, habitat
alteration for home building and associated activities, and greenhouse
gas emissions.

Cities as engines of biological loss: extinction,

eco-function and homogenisation
Urbanisation is, justifiably, often seen as a major cause of species extinction
and ecological damage. For example, urbanisation is the second-leading cause
of habitat loss and pollution in Canada (Venter et al. 2006), and it has been
documented as the second-leading cause of species endangerment in the USA
(Czech et al. 2000).
The increasingly harmful impacts of cities on the natural environment are
rooted in their origins. Cities are artificial habitats created primarily for one
species, and this requires greatly modifying, and often obliterating, whatever
natural habitats occur in the immediate vicinity. An obvious example is the
replacement of a local forest with asphalt and row housing. Such ‘construc-
tional’ urban impacts arise from the building activities of urbanisation.
Also, the maintenance of high human population densities in any city
requires the importation and consumption of massive amounts of resources
from many distant areas. As noted long ago by Odum (1963), cities are highly
heterotrophic entities which require huge inputs of matter and energy from
many distant sources. As these resources are exploited for urban consumption
(e.g. logging, mining, agriculture), natural habitats in those areas are inevitably
affected in negative ways. Conversely, vast quantities of waste from this con-
sumption are released into the air and water which transport these materials
as pollution into surrounding and distant regions. Such ‘maintenance’ urban
impacts occur from the consumption and pollution produced in maintaining
large urban human populations.
The accumulation of these constructional and maintenance impacts of cities
on the environment is of course increasingly large. Although urban area now
occupies less than 3% of the Earth’s terrestrial surface (Mills 2007), this area
produces roughly 78% of anthropogenic carbon emissions, and accounts for
60% of residential water use and 76% of wood used for industrial purposes
(Brown 2001). These numbers exemplify the extremely wide-ranging and
diverse impacts of cities on the global and regional atmosphere, the hydro-
sphere and the biosphere. In addition to these wide-ranging impacts at regional
to global scales, urbanisation has exceptionally severe local impacts. As
290 M. L. MCKINNEY

High
Urbanisation

Mining

Persistence
of change Agriculture

Logging

Low
High Low
Similarity to original natural habitat

Figure 12.1 The adverse effects of different human activities on natural habitat.
In terms of duration and intensity (deviation from the pristine natural condition),
urbanisation typically involves the greatest anthropogenic impact. Modified from
Marzluff and Ewing (2001).

discussed by Marzluff and Ewing (2001), urbanisation impacts on the local

environment are more intense and more enduring than any other common
human activity. Thus, the effects of logging, mining, farming and many other
anthropogenic human disturbances are typically less intensive and enduring as
a cause of habitat loss than urbanisation. Whereas logging, mining, farming
and most other land-uses have the goal of intermittent resource extraction,
urbanisation represents a more permanent (on ecological and human time-
scales) land-use with the goal of long-term habitation. Furthermore, our desire
to alter the land drastically, to fit our own aesthetic and functional needs,
produces extreme modifications of the pre-existing natural communities. For
instance, concrete and pavement, with zero primary productivity, would seem
to be a more extreme disturbance than occurs with logging, mining and
farming. These other activities at least generally allow soil and vegetation to
persist (see Figure 12.1).
In addition to habitat loss and pollution, urbanisation is also strongly asso-
ciated with many other anthropogenic causes of extinction such as non-native
species introductions and overharvesting. As transportation hubs, cities con-
tain a disproportionate number of non-native species which become major
sources for the export of non-native species into the surrounding region (von
der Lippe & Kowarik 2008). Urban consumers are also major markets for
products from illegally harvested species (Robinson 2001). This includes cities
in poor nations where local and regional species are often harvested for
‘bushmeat’ and other animal products which are increasingly sold in urban
areas (Edderai & Dame 2006).
 URBAN FUTURES 291

Although the impacts of urbanisation on ecosystem function (services)

are not nearly as well studied as its impacts on species losses, the existing
evidence generally confirms what might be intuitively inferred: that urbanisa-
tion degrades ecosystem function, especially with increasing urban density
(Chapter 3; Tratalos et al. 2007). For example, a study of an urbanisation
gradient in the Lake Tahoe basin of the USA showed that ecosystem services
became impaired after moderate amounts (30–40%) of development occurred
(Sanford et al. 2009). In this case, decline of ecosystem function was docu-
mented via its effect on ant communities, with the loss of decomposer and
aerator ants which, in turn, reduced soil productivity and rainwater
infiltration. Severe negative impacts of urbanisation are also seen in the eco-
system function of aquatic systems arising from drastic alteration of water flow
regimes, sedimentation and nutrient inputs (Roach et al. 2008). Fortunately,
there seems to be substantial variation in the negative relationship between
urbanisation and ecosystem function, suggesting that there is considerable
opportunity to maximise ecological performance (Tratalos et al. 2007).
A common consequence of the natural disruptions caused by urbanisation is
biological homogenisation (McKinney 2006). This can have several manifest-
ations. At the taxonomic level, the extirpation of local native species followed
by their replacement with widespread non-native species can produce hom-
ogenisation. Such a process is promoted by two basic dynamics of urbanisation
(McKinney 2006): (i) the creation of similar (artificial, human) habitats in many
places that are out of equilibrium with their natural matrix, and (ii) the
importation (deliberate and accidental) of non-native species that can survive
in these urban habitats. At the ecosystem level, urbanisation can also promote
functional homogenisation. A study of bird communities in France, for
example, showed that urbanisation induced community homogenisation and
that populations of specialist species became increasingly unstable with
increasing urbanisation of the landscape, increasing the functional similarity
of the bird communities (Devictor et al. 2007).

Cities as future centres of conservation

In view of the extensive environmental impacts caused by urbanisation, it may
seem counterintuitive to view cities in terms of conservation. There are, how-
ever, several basic reasons why urbanisation, where done in a more sustainably
planned way, can make substantial contributions toward nature conservation
(McKinney 2002). For one, the very fact that cities currently have such enor-
mous environmental impacts means that any reductions in those impacts are
positive steps toward sustainability. There are many ways (discussed below)
that the construction and maintenance of cities, ranging from housing to
transportation, can be redesigned to reduce their impact. Second, urban settle-
ments may have intrinsic properties that, of themselves, are the best way for
292 M. L. MCKINNEY

humans to minimise their impact on the natural environment. By concentrating

humans into small areas, cities reduce the human population density in other
locations. More importantly, this density concentration creates economies of
scale such that urban inhabitants have lower per capita impacts (e.g. resource
consumption, pollution) than rural human populations. This is because of a
physical infrastructure (e.g. mass transit, mass housing) that is more efficient on
a per capita basis. For example, total road surface, electrical cable length and fuel
stations scale more slowly than population size as a city grows (Bettencourt et al.
2007). Similarly, per capita consumption of motor fuel declines exponentially
with increasing human population density (Kennedy et al. 2007).
This potential of urbanisation to concentrate human impacts and ‘free up’
natural areas for preservation has a parallel in a debate about agricultural
impacts. Is wildlife preservation more effectively achieved by expanding
‘wildlife-friendly’ farming techniques or by using more intensive farming
methods that will more drastically alter farmland but will also reduce the area
of land that is farmed? While empirical data are sparse, available evidence
seems to suggest that (as with urbanisation) intensification may be the more
effective conservation strategy (Green et al. 2005).
Urbanisation can also promote nature conservation via the social and psycho-
logical impacts it has on city residents. Citizens of suburban and urban areas
tend to place a much higher value on species conservation than those living in
rural areas (Kellert 1996). Legislators from highly urbanised states and districts
tend to be more supportive of strengthening the Endangered Species Act in the
USA (Mehmood & Zhang 2001). This is especially true where local policies
directly promote conservation activities within the urban area such as landscap-
ing for wildlife, green roofs and the cultivation of native plants. There is a
psychological benefit for people in the cultivation of local biodiversity (Fuller
et al. 2007). This is a crucial point because any definition of ‘sustainability’ must
include the benefit to human welfare as well as biodiversity (McGranahan &
Satterthwaite 2003). Conversely, the disconnection from biodiversity in urban
environments can have a negative impact on humans as well as conservation, or
what Miller (2005) has called the ‘extinction of experience’.
These arguments imply that rethinking the way that cities are designed,
constructed and maintained could significantly reduce the total and per capita
anthropogenic impact on the natural environment, as well as maintain a high
quality of life for humans. What follows is a necessarily brief overview of some
fundamental ways to rethink cities in this way.

Reducing the urban footprint

Howard Odum (1971) was an early analyst of urban energetics, documen-
ting that modern cities consume 10–100 times more energy (per unit area)
than natural ecosystems. His use of input–output models documented the
 URBAN FUTURES 293

heterotrophic basis of urban habitats and their strong dependence on external

sources of matter and energy which are ‘metabolised’ by the city. In the past
few decades, two widely used approaches have evolved that have elaborated on
this insight: the ecological footprint and the mass balance (Kaye et al. 2006).
While neither of these approaches provides a complete accounting of the
environmental impacts of cities, they are useful heuristic models for describing
general impacts and, most importantly, ways to reduce urban impacts on the
environment. Because most of the matter and energy used by cities is imported
from distant locations, and because many wastes emitted by cities are then
transported to distant locations, the ecological footprint and mass-balance
models are useful ways to assess a city’s impact on the regional and global
environment.
The ecological footprint (EF) of a city is the area of land required to meet its
metabolic needs for resource consumption (inputs) and waste assimilation
(outputs). Its roots lie in earlier ideas, such as ‘ghost acres’ and similar concepts
developed by Borgstrom (1972) and Ehrlich (1968). Specifically, the EF is
the ‘total area of productive land and water ecosystems required to produce
the resources that the population consumes and assimilate the wastes that the
population produces, wherever on Earth that land and water may be located’
(Rees 2000, p. 371). Studies of many cities have documented that the EF is
orders of magnitude greater than the area actually occupied by a city (Olalla-
Tarraga 2006; Wackernagel et al. 2006). For example, Rees and Wackernagel
(1996) estimate Vancouver’s EF at more than 200 times its geographic area.
London’s EF is estimated to be 125 times its actual size ( Jopling & Girardet
1996). All of these studies confirm the comment by Rees (1997) that ‘cities as
presently conceived are incomplete ecosystems, typically occupying less than 1
per cent of the ecosystem area upon which they draw.’
One of the main attractions of the EF is its simplicity. The EF converts
several measures of consumption and waste assimilation into a single unit,
the amount of land required to support the consumption and waste. As a
result of this simplicity, the EF approach has become widely used and is
firmly entrenched in the scientific and popular literature. It is often used
by educators and even policy makers to illustrate the easily forgotten but
fundamental insight that urban impacts extend far beyond the boundaries of
any town or city.
On the other hand, the simplicity of the EF has led to several criticisms. Kaye
et al. (2006, p. 192), for example, argue that EF studies are ‘flawed because
they do not incorporate biophysical setting, population size and per capita
consumption rate into the footprint calculation. Until these factors are
included, footprints depict negative impacts of cities without accounting for
the probable efficiency of dense urban living.’ Several other criticisms are
summarised in Mcmanus and Haughton (2006), mostly echoing the problem
294 M. L. MCKINNEY

of oversimplification resulting in the neglect of several variables, resulting in a

number that is an incomplete metric of the overall impact of cities. Examples
of neglected parameters include water resources, geography (e.g. biophysical
settings noted above), environmental impacts within urban boundaries (such
as wetland degradation), and metrics of social and human welfare (Mcmanus &
Houghton 2006).
Perhaps the biggest problem from a purely analytical view is that the EF
simply does not include many variables that would provide a more thorough
assessment of urban impacts on the environment. For instance, the typical EF
calculation uses four consumption categories: energy use, the built environ-
ment (the land area covered by a settlement and its connection infrastructure),
food and forestry products. This is obviously a small subset of all resources,
determined by the practical requirements of data gathering. Similarly, EF
incorporates only a small subset of urban waste outputs, such as carbon diox-
ide, to be assimilated by natural ecosystems. These impact variables are then
converted into a single number (land area). While this is certainly an interest-
ing number, it obviously compresses many other dimensions of urban impacts
in a single value. The International Institute for Environment and Develop-
ment described the EF analysis, whereby all environmental impacts are aggre-
gated into a simple index, as ‘resource reductionism’ (Hammond 2000).
Despite the many criticisms of the EF, it has promoted much constructive
thinking about how to improve upon it, discussed in the next section. It has
also become a way to formulate specific proposals to reduce human impacts
(i.e. shrink the urban footprint). Wackernagel et al. (2006, p. 107) list these four
examples of ways to reduce the EF:

• use resource-efficient technology to reduce the demand on natural capital;

• reduce human consumption while preserving quality of life: for example,
reduce the need for fossil fuels by making cities pedestrian-friendly;
• lower the size of the human family in equitable and humane ways, so
that total consumption decreases even if per capita demand remains
unchanged; and
• invest in natural capital: for example, implement resource extraction
methods that increase rather than compromise the land’s biological prod-
uctivity, thereby increasing supply.

The first two of these examples relate directly to urban life. Cities need to be
constructed and maintained with more resource-efficient technologies (e.g.
buildings with improved energy and water conservation). Also, cities can be
redesigned in many ways to reduce consumption of many resources. In add-
ition to less reliance on automobiles to reduce energy consumption, an
example would be more area landscaped with native vegetation to reduce
energy and water consumption. Moos et al. (2006) used the EF as an urban
 URBAN FUTURES 295

planning tool to show that increasing the density of housing in residential

subdivisions can reduce the urban EF. In their words, ‘despite the challenges
associated with data collection and conversion, it is argued that EF has utility
for planners and urban designers because it enables assessment of built form
from an environmental consumption point of view’ (Moos et al. 2006, p. 195).
The EF has also become an excellent metaphor for public education, which
has led to specific policy recommendations to reduce the EF of some urban
areas. Wackernagel et al. (2006) cite several examples, such as California’s
Sonoma County Footprint project which has inspired every city in the county
to sign up for the Climate Saver Initiative of the International Council for Local
Environmental Initiatives (ICLEI) – also known as Local Governments for
Sustainability.
There have been several attempts to improve the EF by expanding the
parameters used in its measurement. For example, Satterthwaite (1997) sug-
gested including criteria to estimate human social welfare, such as health,
sanitation and recreational facilities. More commonly, many studies have
added biophysical parameters to increase the metrics of human impacts. The
amount of land needed to assimilate the nitrogen and phosphorus waste of
cities, for instance, is of the order of 400–1000 times the area of the cities (Folke
et al. 1997).

Reducing urban metabolism

Because of the problems noted above for the EF, many analysts favour the
‘metabolic’ model as a better method to assess the twin environmental impacts
of resource consumption and waste export of cities (Decker et al. 2000; Kaye
et al. 2006). This view examines mass balances which ‘quantify inputs and
outputs of elements or materials (or these values converted to energy units),
and yield information about whether a city is a source (input > output) or sink
(input < output) for the material and how the material is transformed by urban
activities’ (Kaye et al. 2006, p. 193).
Mass-balance studies show that cities act as a sink for many substances
imported from long distances. For example, building materials such as stone,
wood and metals are incorporated into the urban structure and then become
stored in landfills within or near the urban area (Decker et al. 2000). Indeed, it
has been estimated that less than 10% of urban material inputs are exported as
goods (Brunner et al. 1994). Thus, cities are net sinks for most materials. This is
especially true for rapidly growing younger cities, where the ratio of inputs to
outputs can be 10:1, whereas the ratio is closer to 1:1 in old cities experiencing
little growth (Brunner 2007). At the chemical scale, cities are sinks for many
elements such as nitrogen, phosphorus and, sometimes, metals (Kaye et al.
2006). A study of Gavle, Sweden, estimated that of the 330 tons of phosphorus
per year entering the urban system, only 35% (116 tons) leaves the system, half
296 M. L. MCKINNEY

of it in exported goods and half expelled into the Baltic Sea. The remaining 65%
is stored in the city, mainly as sewage sludge in landfills (Nilson 1995).
On the other hand, cities can also act as sources for some substances. These
are materials that are imported, often from far away, and then released into
the local and regional environment. Carbon, from fossil fuel combustion, is an
obvious example. A typical large city consumes of the order of 100 to 1000
petajoules (1 petajoule being 1015 joules) of energy per year to operate its
transportation, electrical and climate control infrastructures (Decker et al.
2000). As most of this energy is produced by fossil fuel combustion, cities are
a huge source of carbon dioxide (CO2). It was estimated, for example, that each
million residents of Sydney, Australia, produced about 48 148 tons of carbon
dioxide per day in 1990 (Newman & Kenworthy 1999).
‘Transformed’ inputs are converted to another form and then exported from
the urban system as waste (Decker et al. 2000). Consequently, cities are typically
local and regional sources for many of the transformed materials. Fuels, just
noted, are a good example in the transformation of hydrocarbons into CO2,
particulates and many other byproducts that become transported by air and
water to localities near and far. Food and water are two other major inputs that
become transformed and then largely exported into the environment. New
York City, USA, consumes 20 000 tons of food per day, with half being dis-
carded, mostly as organic waste into landfills (O’Meara 1999). Water is notable
as a transformed substance (mostly becoming converted into sewage) with the
greatest quantity of flux through the urban system. Cities consume an average
of roughly 1000 kt of water per day compared to 10 124 kt of fuel in a year
for Mexico City (Decker et al. 2000). Similarly, in 1990 each one million resi-
dents of Sydney, Australia, daily consumed over 490 000 tons of water but less
than 11 000 tons of fossil fuels (Newman & Kenworthy 1999). Most (over 75%) of
the water consumed in Sydney was exported locally as sewage.
As noted above, a key aspect of urban metabolism is that it has environ-
mental impacts at many spatial scales. It destroys habitat in many localities,
ranging from near to far away. Among the many examples would be the
habitat loss caused by the mining extraction of minerals and timber to supply
urban growth and maintenance, and the subsequent habitat loss produced by
the export, often by water and air, of polluting waste into natural habitats near
and far away from the city. Therefore, reducing the impact of cities on bio-
diversity requires reducing urban metabolism. Unfortunately for conservation
goals, the metabolism of cities has been increasing. The material turnover of a
modern city is about one order of magnitude larger than that in an ancient city
of the same size (Brunner et al. 1994). A review of urban metabolism studies
from eight metropolitan regions across five continents conducted since 1965
showed that most cities had increasing per capita (and total) urban metabolism
with respect to water, waste water, energy and materials (Kennedy et al. 2007).
 URBAN FUTURES 297

Reversing the trend toward increasing urban metabolism is clearly a basic

way for cities to help achieve global sustainability (Newman 1999). By reducing
resource inputs, cities not only can reduce the environmental burdens of
resource extraction and non-renewable resource depletion, they will also
reduce the environmental burdens created by the export of urban metabolic
waste and pollution. There are two (non-exclusive) ways for cities to reduce
their metabolic demands for materials: (i) increase the efficiency of use, and
(ii) increase the recycling of materials within urban areas. As energy cannot be
recycled, reducing the urban metabolic demand for energy must focus on
increasing efficiency. Another option will be increasing the use of renewable
energy, as long as the sources have relatively low environmental burdens.

Sustainable cities and global biodiversity conservation

Most of the ecological literature on urban biodiversity conservation has tended
to focus on ways to preserve biodiversity in localities within and adjacent to a
city (McKinney 2002). This literature often provides data that can help mitigate
the negative impacts of urbanisation on natural habitats, by preserving or
restoring those habitats. The basic goal is a ‘win–win’ scenario whereby human
needs can be met while minimising urban impacts on biodiversity conserva-
tion (Miller 2006). A common theme of this literature is the mitigation of local
environmental impacts. For example, one can promote the preservation or
reintroduction of native plants (and the animals that rely on them) in urban
ecosystems such as golf courses (Hammond & Hudson 2007) and residential
areas (e.g. Fetridge et al. 2008; Pennington et al. 2008).
Reducing local urban impacts in this way is clearly an extremely important
part of an overall strategy to preserve global biodiversity. This is especially
true if human settlements are often located in areas with a naturally high
richness of native species with high conservation value due to their endemism
and phylogenetic uniqueness. Furthermore, many urban areas still contain a
substantial number of threatened and other at-risk categories of species, des-
pite being occupied by humans for many years. Examples would include a
surprisingly large number of threatened bird species in cities in the UK (Fuller
et al. 2009) and plant species in urbanised areas of California, USA (Schwartz
et al. 2006).
This work of preserving local biodiversity in and around urban areas is
commonly discussed in the ecological literature because there is much that
ecologists can do to promote this aspect of global conservation. Ecologists have
a direct role in providing data that can help restore and preserve local biological
communities and ecosystems.
In contrast to this local emphasis, the approach I have taken here is to
emphasise the importance of the sustainable cities concept, and especially the
goal of reducing urban metabolic demands for its potential for contributing to
298 M. L. MCKINNEY

Table 12.1 Reduction in local and distant habitat alteration and pollution from urban
changes.

Clean (non-fossil-fuel) energy

Distant impact reduction: climate change, acid rain, smog, coal mining, oil extraction
and refining
Local impact reduction: smog, carbon monoxide
Reduced and recycled construction materials
Distant impact reduction: deforestation (lumber), oil extraction (plastic), mining
impacts (stone, metals)
Local impact reduction: landfill waste

the conservation of non-local biological resources. By reducing the demand for

vast quantities of natural resources that are extracted and imported from many
areas of the planet, there is a consequent reduction in the impacts to biota
affected by resource extraction in and pollution of those areas (Table 12.1).
For example, many cities have long exceeded or polluted local freshwater
supplies and must rely on water from very distant sources (UN-HABITAT 2003).
Dakar, the capital of Senegal, now needs to draw water from a lake 200 km
away (White 1992); Mexico City has to supplement local supplies with water
drawn from neighbouring basins that has to be pumped over 1000 metres in
height and drawn from up to 150 km away (Connolly 1999). Therefore, any
reductions in water demands in these (and most other) cities would reduce
these environmental impacts on the affected distant biota.
Another example would include the many distant impacts from the extrac-
tion and importation of vast amounts of construction materials (e.g. stone,
wood and metals). Mining of stone and especially metals involves moving
enormous volumes of rock. Most of this is open pit mining which typically
involves the complete removal of biological communities in the area. The large
overall impact of mining on the biosphere can be inferred from the estimate
that humans now move more rock and sediment on the planet than all the
natural processes (e.g. water, wind) combined (Meybeck 2004). In addition, the
mining and smelting of the ore produces very large amounts of air and water
pollution, with high concentrations of heavy metals and other very toxic
pollutants (Muezzinoglu 2003). Some of the impacts of mining on the local
and regional biota are well documented by ecologists. Examples would include
the effects on wildlife of gold mining in the USA and Australia (Donato et al.
2007), and copper mining in Poland (Kalisinska et al. 2004). Such cases typically
document mass mortality of vertebrates, especially waterfowl, caused by acute
toxicity of heavy metals and cyanide (used in ore extraction). In addition there
is widespread soil contamination leading to severe and long-term impacts on
native vegetation (Bech et al. 1997). Given that urban areas are the main
 URBAN FUTURES 299

markets for these materials, often for building construction (e.g. copper) and
other uses directly related to urban expansion, any reduction in the demand
would reduce the impacts of their extraction on those distant ecosystems.
The export of mahogany Swietenia macrophylla from the Amazon basin to
global urban markets outside of South America provides an illustration of
the benefits that could be derived from reducing urban wood consumption.
For each mahogany tree harvested, over 1 square kilometre of forest ground
vegetation is removed or damaged, resulting in the cumulative loss of thou-
sands of square kilometres of forest ecosystem (Verissimo et al. 1995).
Compounding this loss is the growing trend to convert the harvested forests
to cattle pasture. The growth of cities in China, combined with logging
restriction within the nation, has led to a rapid increase in the importation
of forest products from heretofore relatively undisturbed forest ecosystems
in more distant localities in Russia, Southeast Asia and Africa (Zhang & Gan
2007).
In addition to water and construction materials (stone, minerals, wood),
another major resource input into cities is energy, especially fossil fuels. The
extraction of these has severe impacts on the ecosystems where these resources
are found. The debate over petroleum drilling in tundra, ocean and other
fragile ecosystems has raised awareness of the impacts of oil extraction
(Kryuchkov 1993). The local ecological impacts from coal extraction and use
are probably even greater. Life cycle assessments that measure the effects of
mining, transport, combustion and waste disposal of coal show that all of these
have significant negative impacts on local and regional ecosystems in several
different localities (Cazplicka-Kolarz et al. 2004; Babbitt & Lindner 2008). Most
notable in terms of local biotic impacts is the mining of the coal body itself,
which typically involves extensive de-vegetation, and air and water pollution
impacts on local biota (Elberling et al. 2007). Acid rain and other regional air
pollution impacts must also be considered as urban impacts on ecosystems that
are distant from the point of use. Finally, the increasing global impacts of
climate change on the many ecosystems, especially those of high latitudes, also
exemplify impacts that are far removed from the urban areas that consume the
electricity produced by coal.

Redesigning cities for sustainability

Of course, merely pointing out that cities need to reduce metabolic demands is
a relatively easy task compared with applying these concepts. The way that
cities are constructed and maintained will need to be redesigned so that they
continue to meet human needs but also increase the efficiency of resource use,
recycle those resources and rely more on alternative, especially renewable
sources. Figure 12.2 summarises this general approach. What follows are some
specific examples of ways to implement the approach.
300 M. L. MCKINNEY

To reduce extraction To reduce waste

Increase efficiency of use
Waste capture
Recycle materials
Waste recycling
Alternative resources
City
resource extraction
Waste
timber
minerals
fossil fuels

Habitat loss Pollution

Overharvesting

Biodiversity loss (extinction)

Figure 12.2 Urban resource consumption (extraction) of natural resources and

consequent waste disposal are the underlying factors driving many species to extinction.
Often these species inhabit areas far removed from the city itself. Cities can reduce these
impacts (i.e. become more ‘sustainable’) by reducing their resource demands (via
increased efficiency, recycling, alternative resources) and reducing their waste output.

An obvious method to increase efficiency of resource use in cities is the

widespread adoption of more efficient technologies (e.g. compact fluorescent
and LCD lighting) by their inhabitants. This citizen-based approach has been
the focus of many educational campaigns to promote ‘green living’, such as the
Sonoma County Footprint project noted above (Wackernagel et al. 2006). The
goal in such cases is to encourage citizens to purchase hybrid cars, purchase
organic foods, improve home energy conservation and so on. However, a
second approach, at the scale of the city, provides another option for increasing
efficiency of resource use: redesigning the city itself (the urban form). It is
estimated, for example, that green design in the planning and building phase
of urban growth can have a large impact on the operational phase, reducing
energy demand by a factor of 5 to 10 (Brunner 2007). Similarly, cities that
concentrate people at higher population densities are more efficient in terms
of transportation and housing. There is an exponentially inverse relationship,
for instance, between per capita energy consumption for transportation and
population density in cities (Newman & Kenworthy 1991).
Recycling materials will also be an important way to reduce the urban
metabolic demand for consumption of external resources. Future cities will
need to rely less on ‘linear’ metabolic flows and more on ‘cyclic’ metabolic
flows (Doughty & Hammond 2004). This would entail increasing the recycling
 URBAN FUTURES 301

of organic and inorganic materials within urban areas, which would simultan-
eously reduce both consumption of resources and the export of waste and
pollution. Ore mining will be replaced by urban mining, resulting in less
demand for energy and resources and fewer emissions (Brunner 2007).
One problem with the ‘sustainable city’ concept is that it can foster the idea
that there is a universal solution to the problem of urban impacts on the
environment when, in fact, cities are often quite different in terms of their
natural and social contexts. For example, some cities are currently ‘less sustain-
able’ than others. Wealthier cities such as Canberra, Chicago and Los Angeles
have between 6 and 9 times the carbon dioxide emissions per person of the
world’s average and 25 or more times that of cities such as Dhaka (Nishioka
et al. 1990). Motor fuel use per capita in cities such as Houston, Detroit and Los
Angeles, with among the world’s highest consumption levels, is 100 times or
more that of most cities in low-income nations (Newman & Kenworthy 1999).
Average waste generation per inhabitant in cities can vary more than 20-fold
when comparing urban citizens in high-income nations (who may generate
1000 kg or more of waste per person per year) with those in some of the lowest-
income nations (who may generate less than 50 kg per person per year) (Hardoy
et al. 2001). In Los Angeles, only 10% of solid waste is recycled compared with
over 60% for Tokyo (Decker et al. 2000).
In addition, the geographic location of a city can affect its pathway to
sustainability. An emphasis on water conservation is important for cities in
arid regions but may be much less so for cities located in areas with abundant
water supplies, such as many tropical regions. Similarly, the age, or develop-
mental stage, of a city can affect its pathway to sustainability. In young, rapidly
growing cities, the ratio of materials input to materials output can exceed 10:1
whereas this ratio is closer to 1:1 in mature cities (Brunner 2007). An implica-
tion is that it may be harder to promote materials recycling in younger cities
because of the large amount of imported primary resources.

Urban sustainability as win–win ecology

Discussions about sustainable development have often invoked sustainability
as a ‘win–win–win’ scenario for its potential simultaneously to optimise eco-
nomic, environmental and social goals (Gibbs & Deutz 2005). In the context
here, we can focus on sustaining global biodiversity and especially the preser-
vation of endangered native species in the face of urbanisation. Rosenzweig
(2003) has promoted the use of ‘win–win’ scenarios to maximise species preser-
vation. This involves establishing and maintaining habitats to conserve species
diversity in places where people live, work or play.
Win–win scenarios are critical for the success of redesigning cities in ways
that reduce their environmental impacts because they also include ways to
meet human economic and social needs. As noted above, most of the ecological
302 M. L. MCKINNEY

literature on urban nature conservation focuses on such scenarios at local

scales. Green roofs, naturalised golf courses and residential landscaping for
wildlife are examples (when done properly) of this type of win–win scenario
because they meet human aesthetic, recreational and other needs while also
promoting biodiversity conservation goals.
A main goal of this chapter has been to extend the spatial scale of these
ecological win–win scenarios beyond the urban area itself. A benefit of this
approach is to address one of the major criticisms of the win–win scenario,
which is that it is often applied in incomplete ways and focuses on small-scale
conservation problems (Sorice 2005). For example, urban win–win scenarios
are often discussed in terms of designed habitats that maximise local biodiver-
sity, such as optimising the configuration of green spaces (Colding 2007).
However, the maximised biodiversity is usually defined simply as species rich-
ness, regardless of whether it includes non-native species or if the native
species have any conservation value. Conversely, many studies have focused
on ways to promote ecological services in urban habitats with the cultivation of
designed ecosystems in allotment gardens, cemeteries, city parks (Andersson
et al. 2007) and green roofs (Oberndorfer et al. 2007). However, most such
proposals do not encourage, or simply omit mentioning, the importance of
conserving native species, especially those native species at risk. Indeed, restor-
ation ecology has often included non-native species in its efforts to restore or
enhance ecosystem functions ( Jones 2003).
By focusing only on the local city ecosystem, efforts to increase urban species
richness and urban ecosystem services therefore neglect the critical criterion of
global sustainability noted at the beginning of this chapter. The long-term
health of the biosphere hinges on preventing the global extinction of endemic
native species which are found in the vicinity of urban areas. Because human
settlements tend to be found in areas with many native species of high conser-
vation value, proposals to increase urban species richness or ecological services
would be incomplete without including rare, endemic and at-risk native
species. In a sense, this enhances the ‘win–win’ scenario of simultaneously
meeting ‘human needs and ecological function’ or ‘human needs and local
biodiversity’ into a ‘win–win–win’ scenario whereby ‘global biodiversity conser-
vation’ is added to either of the preceding combinations.
The conservation importance of globally threatened species is not, of course,
a novel insight, but I do suggest that it is a neglected dimension in much
of the literature on urban ecology. Among the relatively few discussions of
urban biodiversity conservation that do address global conservation in urban
ecosystems would be the study by Lawson et al. (2008) showing that urbanis-
ing habitats, despite years of conservation neglect, still contain many plant
populations of high conservation value, and that conservation planning for
global biodiversity preservation should allocate relatively more resources to
 URBAN FUTURES 303

protecting urban-associated plant taxa. Similarly, Turner et al. (2005) note that,
while residential ecosystems of Halifax, Nova Scotia, are species-rich, the dom-
inance of exotic species reduces the ecological integrity of those communities.
Pennington et al. (2008) discuss the importance of urban riparian zones for
globally threatened migrating Neotropical migrant birds. Finally, Milder et al.
(2008) propose that residential suburban developments explicitly consider
threatened natural resources including rare species and ecological commu-
nities in their design, something that traditional ‘conservation developments’
have tended to ignore.
There are also a few studies that indicate a role for rare or declining native
species in restoring ecosystem services. Fiedler et al. (2008) discuss evidence that
native plants can be as useful in restoration as much as widely recommended
non-natives. Winfree et al. (2007) show evidence that native bees can provide
pollinator functions to replace those of the declining exotic honeybee in
the USA.
There is yet one more dimension to this ‘win–win–win’ scenario. It is possible
to conceive of urban conservation initiatives that not only meet human needs,
promote local species richness and preserve global biodiversity but in addition
reduce other distant urban impacts. An example would be an urban wetland
that benefits human needs for recreation, aesthetics and water purification,
promotes local species richness, provides habitat for globally threatened avi-
fauna, and also reduces impacts on the overall watershed by reducing urban
water demand and the total pollution load on the watershed. Furthermore, as
wetlands are net carbon sinks, urban wetlands may have potential for amelior-
ating anthropogenic impacts on climate change (Erwin 2009).
Again, it is not a novel insight that well-designed urban ecosystems can
simultaneously meet several human needs as well as several conservation
goals, both local and global. Indeed, it seems likely that discussions of ‘sustain-
able cities’ will increasingly converge on this approach for reasons of practical-
ity, efficiency and economics (e.g. Choi et al. 2008; Martinez & Lopez-Barrera
2008). My point herein is that the current conservation and urban sustainabil-
ity literature is highly fragmented in its approach, focusing on just one or two
objectives of these potential ‘win–win–win–win’ scenarios that can potentially
achieve several objectives if a larger spatial scale of planning is attempted.

Summary and conclusions

The urbanisation of our planet will clearly continue for many years to come.
Efforts to preserve what remains of our biosphere must, consequently, increas-
ingly focus on ways to reduce the impact of cities. Indeed, one could argue
that, because cities have the potential to concentrate human populations
into relatively small areas as well as reduce per capita resource consumption
and per capita pollution via economies of scale, then well-designed (and
304 M. L. MCKINNEY

maintained) cities may be the best opportunity our species has for a sustainable
coexistence with the natural biosphere.
However, if this enormous conservation potential for cities is to be attained,
conservation planning must extend far beyond the very local scales at which
most urban ecology is carried out. We must begin to focus much more atten-
tion on the many impacts of cities on distant areas at all spatial scales, ranging
from resource consumption of water, minerals, energy and biological products
to waste and pollution impacts on the hydrosphere and atmosphere at regional
and global scales. ‘Sustainable’ cities of the future must sustain the biosphere
as well as local and regional ecosystems.
This chapter has centred on the ecological footprint and especially the mass-
balance (metabolic) approach because they provide well-established metrics for
the resource inputs and waste outputs of cities. From the mass-balance view, it
seems clear that there are many opportunities to reduce per capita human
impacts by finding ways to reduce urban metabolic demands. By reducing per
capita resource consumption and waste production, most per capita environ-
mental impacts are simultaneously decreased.
Such a policy of ‘source reduction’ of the waste stream is commonly discussed
in the resource economics and management literature (Fullerton & Wu 1998)
but it is not a common theme in urban ecology. One reason is that much of the
work needed to reduce urban metabolism is simply outside the realm of what
ecologists have traditionally studied: green technologies that increase efficiency
of resource use, recycling those resources after use or utilising more renewable
resources are of more interest to engineers than ecologists (Montalvo 2008).
There are, however, two crucial reasons that reducing urban metabolism
should be of interest to urban ecologists. One is that it is, arguably, the best way
of preserving what remains of the most pristine ecosystems on the planet.
Urban consumption is the main driver of resource extraction, even in areas
that are far removed from the city itself. Conversely, urban waste and pollution
is the main driver of climate change and other processes that also affect
ecosystems that are far removed from the urban sources that produce them.
Another reason that urban metabolism should be of interest to urban ecolo-
gists is that ecologists can play a significant role in doing something about the
problem. Much ecological literature on cities has focused on the conservation
of local native biota, especially via the restoration or preservation of native
species habitat. However, ecologists can also help design urban ecosystems that
reduce urban impacts at scales beyond the peri-urban fringe, such as artificial
wetlands that reduce demand on the regional watershed or providing habitat
for globally threatened species. Much of this work has the potential for simul-
taneously achieving multiple goals (‘win–win–win’ scenarios) whereby human
welfare is improved, economic costs are reduced, and both local and global
urban impacts are decreased as well.
 URBAN FUTURES 305

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biodiversity potential and ecosystem Ecology Letters, 11, 1105–13.
services. Landscape and Urban Planning, Zhang, J. and Gan, J. B. (2007). Who will
83, 308–17. meet China’s import demand for
Turner, K., Lefler, L. and Freedman, B. (2005). forest products? World Development,
Plant communities of selected urbanized 55, 2150–60.
Index

Locators in bold refer to major content

Locators in italic refer to figures/tables
Locators for headings which also have subheadings refer to general aspects of that topic only.

abundance, species 55–7, 93, 99, 157; ants 62, 240

see also human population density; anxiety 205; see also health/wellbeing;
population mood states
accessibility, green spaces 208–9, 222, aquatic systems see water
237–8, 244–5, 249 Aquila verreauxii (black eagle) 267
Accessible Natural Greenspace Standards architecture 269–71, 278–9; see also planning
(ANGSt) 222, 237–8, 244–5 and design
adaptation to urbanisation 55, 75, 95, 98, ARCUE (Australian Research Centre for
124–5; see also genetics; plasticity; Urban Ecology) 264
wildlife species arousal states, urban/rural comparisons 60
ADHD (attention deficit hyperactivity Asclepias syriaca (common milkweed) 126
disorder) 206 Asia, future of urbanisation 287
administrative boundaries 10–11 atmospheric chemistry 44–5, 46; see also air
Aedes spp. (mosquito) 62 quality standards
aesthetic value 209–10 at-risk species 297, 303
Africa 147 Atta sexdens (leafcutter ants) 62
aggression, urban 209 attention, concentrated 202, 247
agriculture 36, 290, 292 attention deficit hyperactivity disorder
Ailanthus altissima (tree of heaven) 123, 128 (ADHD) 206
air quality standards 46; see also atmospheric Australia
chemistry biodiversity 91, 92, 93, 94–5, 99,
air-conditioning 38 100–1, 103
alien species 65, 106, 120–2, 129–30; drivers of biodiversity 107
see also wildlife species; urban/rural exercise, humans 207, 247
comparisons exotic/native species 101, 108
biodiversity, spatiotemporal patterns 100–1 feeding urban wildlife 101–2, 155–6
biotic homogenisation 128–9 landscaping 191–2
cities as sources of 126–8 Australian Research Centre for Urban Ecology
ecological characteristics 122–3 (ARCUE) 264
effect of cities on 290 avoiders, urbanisation 98
feeding urban wildlife 157
garden management 149 Bangkok 143, 152
invasion pathways 125–6 BAP (Biodiversity Action Plans), UK 267
typical urban species 123–5 bats 93, 104, 239, 242
urban/rural comparisons 67 bees 240, 242
amphibians 71–2, 93, 239, 242 beetles 122, 239, 242
ANGSt (Accessible Natural Greenspace Beijing 13, 24, 45, 178
Standards) 222, 237–8, 244–5 biodiversity 3, 35, 36, 46, 136, 268–9;
animals, biodiversity 183, 187, 239–8 see also spatiotemporal patterns,
anthropogenic noise 59–62, 67, 141 biodiversity
anthropogenic species 125 crisis 134, 161
antioxidants 63 developing nations 288
310 INDEX

biodiversity (cont.) central city, definitions/

future of urbanisation 289–91, 297–9 conceptualisations 176
gardens, private 147 chemistry, atmospheric 44–5, 46; see also air
genetic structure of populations 75 quality standards
geographic location 21 Chicago 175, 209
global contexts 302–3 Chicago City Hall 270
green spaces 141, 144–5, 238–43, 239 Chicago Wilderness initiative 267
and health/wellbeing 110, 139–41, 205, Chicorum intybus (common chicory) 123
249–50 China 138, 148
land-use policies 182, 183 cities see also individual cities by name;
planning and design 274 sustainable cities; urbanisation
socioeconomics 158–9, 160, 186–7 compact 25, 292, 300, 303; see also human
urban ecosystems 263 population density; urban/suburban sprawl
values, nature 271–2 conservation planning 272–3
Biodiversity Action Plans (BAP), UK 267 ecology 4
BIOLFLOR database 125 edge 276
biophilia 138 future growth of 287–9
biotic homogenisation see homogenisation as sources of alien species 126–8
birds see also feeding urban wildlife; wildlife citizen science 136
watching classification, green space 233–6
biodiversity 183, 239 classrooms, outdoor 210, 223
genetic structure of populations 72–3 climate change 243, 303, 304
predation pressure 67–8, 193 Climate Saver Initiative 295
reproductive success 67–8 coalescence, urban 12
spatiotemporal patterns of biodiversity cognitive functioning, and green space 206
90, 93–5 coleoptera (beetles) 122, 239, 242
urban adaptations 55 colonisation, and genetics 69
birdsong, and urban noise 59–62 Columba livia (feral pigeon) 57, 266
Birmingham 144, 236, 244 Commission for Architecture and the Built
blood pressure 213–14, 218 Environment (CABE) 231
blue space 204–5, 268–9, 275–7 communication mechanisms, animal 59–62
Boa Vista, Brazil 148 compact cities 25, 292, 300, 303;
body mass index 246–7 see also human population density;
body size in animals, urban adaptations intensification; urban/suburban sprawl
57–9 comparative studies, biodiversity 89, 93–7;
bottlenecks, genetic 69 see also wildlife species, urban/rural
Brazil 148, 152 comparisons
Britain see United Kingdom; see also individual competition, intraspecific 59, 60
cities by name composition of urban areas 25–8, 27–8, 29
broad-scale patterns of species richness concentric model of urban development
88, 89–93, 106 12, 176
Buenos Aires 24 connectivity/linkage 242–3, 275–6;
building/construction materials 290, 298–9 see also corridors
business sites, planning and design 277–8 conservation 3, 135; see also planning
butterflies 122, 240, 242 and design
biodiversity 107, 110–11, 161
CABE (Commission for Architecture and the current strategies 264–6
Built Environment) 231 current support 263–4
Cairo 16, 24 developing nations 288
Calcutta 24 feeding urban wildlife 157
California 295 funding 273
Canada 11, 234 future of urbanisation 287, 291–2
car transport 25 global contexts 302–3
carbon cycles 41–4 green/blue space 141–2, 268–9
carbon dioxide emissions 38, 44–5, habitats 267–8
289, 296 people–nature interactions 160
Cardinalis cardinalis (northern cardinal) 193 sustainable cities 297–9
carotenoids 63 traditional practices 266–8
Carpodacus mexicanus (house finch) 60, 193 in urban ecosystems 261, 262, 263–4
Caruelis tristis (American goldfinch) 193 construction materials 290, 298–9
cellular automata 12 contaminants 125; see also alien species
Center for Urban Ecology, US 264 control of nature 147
 INDEX 311

core–periphery spatial model 12, 176 ecology of inequality see inequality

corridors, green 125, 126–7, 147, 242–3, ecology of prestige 189, 191
268–9; see also connectivity/linkage; economics 174, 178; see also socioeconomic
invasion pathways status
corticosterones 62–3 ecosystem services/functions 35–6, 35–6,
cost-benefits, environmental 35; 46–7; see also ecological footprint
see also ecosystem services atmospheric chemistry 44–5, 46
cost-effectiveness, planning and design 280 at-risk species 303
crime 209 biodiversity 110
crisis carbon cycles 41–4
biodiversity 134, 161 future of urbanisation 291
of overaccumulation 177 green space 243–4
cultural differences 138, 148, 160–1, 174, 191 green spaces 142
planning and design 174
DDT 192 species interactions 46
decision-making see planning and design temperature 36–8
definitions/conceptualisations urban 2, 20
ecological footprint 293 vegetation, urban 185
ecosystem services 243 water flow 39, 40
green infrastructure 234 water quality 38–9
green space 202 ecosystem types, urban 22, 21–3
health/wellbeing 203, 204 ecosystems, urban 271–2
open space 233, 234 EEA (European Environment Agency)
people–nature interactions 137–9 236–7, 244
social organisation of cities 174, 176 EF (ecological footprint) 20–1, 292–5, 294–5
species richness 89 Empidonax virescens (Acadian flycatcher) 59
urban 89 energy metabolism, cities 292, 295–7, 299,
urban ecology 1 304; see also fossil fuel consumption
urbanisation 10–11 engineering, ecological 250, 270
values 262 England see United Kingdom; see also individual
deforestation 179 cities by name
Delhi 24 environmental activism 110
demographic traits, urban/rural wildlife 66–8; environmental altruism 138
see also human population density; infant environmental disamenities 179
mortality; population epidemiological studies 204–5, 245–6,
depression 205; see also health/wellbeing; 246–7
mood states Erithacus rubecula (European robin) 61
deprivation see inequality, socioeconomic escapes, garden 125, 151; see also alien species
status ethnic minorities 191
design see land-use policies, planning and design Europe 95, 142, 236–7; see also individual
developed nations, form/structure of cities 25 countries by name
developing nations 25, 287–8 European Environment Agency (EEA) 236–7,
development practices see planning and design 244
Dhaka 24 exercise 207–9, 223, 246–7
diet, urban animal populations 63, 193 exotic species see alien species
dilution effects 66 experiments, field 4
disease risks to wildlife, urban/rural exploiters, urban 98
comparisons 64–6 extinction of experience 135, 160, 292
dispersal, plant/animal 74, 99 extinction of species 120, 289–91, 297
distance-decay function 25
disturbance 98, 145–6, 158 Falco peregrinus (peregrine falcon) 267
diversity, cities 301 FD (functional diversity) 242;
diversity, genetic see biodiversity, genetics see also biodiversity
drivers of biodiversity 105, 106 fear of humans, urban animal
populations 62–3
ecological feeding urban wildlife 62, 66, 135
design 270; see also planning and design biodiversity, spatiotemporal patterns
engineering 250, 270 101–2, 108
footprint (EF) 20–1, 292–5, 294–5; negative effects 156, 193
see also ecosystem services/functions people-nature interactions 155–7
outcomes, social organisation 181, 182, social organisation of cities 192–3
184–8 survival 68
312 INDEX

Ficus carica (common fig) 128 green engineering 250, 270

field experiments 4 green exercise 207–9
flooding 39–40; see also runoff green infrastructure 234
food availability, urban wildlife 58; green living 300
see also feeding urban wildlife green management/planning/design 270,
fossil fuel consumption 38; see also energy 280–1
metabolism; mining green roofs 250, 270, 278–9
founder effects 69 benefits 37–8
fractal models 12 win–win scenarios 302
France, biodiversity 99 green space 25–8, 230–1; see also health/
frogs 99 wellbeing; vegetation
functional diversity (FD) 242; see also accessibility 208–9, 222, 237–8, 244–5
biodiversity aesthetic value 209–10
funding, conservation 273, 275 biodiversity 100, 141, 144–5, 238–43,
future of urbanisation 287, 303–4 239–8
cities, future growth of 287–9 classification/typology 233–6
conservation policies 291–2 conservation 268–9
ecological footprint reduction 292–5 ecosystem services 243–4
energy metabolism 292, 295–7 future research needs 252–3
extinction/biodiversity loss 289–91 naturalistic design 145, 146
sustainable cities 287, 297–9, 299–301 outdoor classrooms 210
win–win scenarios 297, 301–3, 304 people–nature interactions 139–43,
143–4, 144–6
galls 240 planning and design 250–2, 275–7
gardens, private policy 231–3
benefits 154, 155 provision/quality 136–7
escapes 125, 151; see also alien species quality 136–7, 249–50
exotic/native species 149 quantification 236–7
green management 280–1 restoration projects 215–16
management 149–52, 152–3 safety perceptions 145
people–nature interactions 146–9 social networking 206–7
genetics standards 237–8
adaptation 54, 75, 95, 98, 124–5 water flow 41; see also runoff
bottlenecks 69 Green Spaces, Better Places (Green Spaces
intraspecific variation 53; see also wildlife Taskforce) 231
species, urban/rural comparisons green technology 304
structure of populations, urban/rural green walls 270, 278–9
comparisons 68–75, 72–3 green wedges 276
gentrification 179, 185 greenways 268–9; see also connectivity/
geographic location 14, 21–3, 301 linkage; corridors
Germany grid mapping techniques 123
alien species 120–1, 122, 126, 127–8 GST (Green Spaces Taskforce) 231
biodiversity 95, 96 Guangzhou, China 139, 141, 143
biotic homogenisation 129 guidelines see also standards
typical urban species, native/alien 124 feeding urban wildlife 157
ghost acres 293 Open Space Strategies: Best Practice
Gibrat’s law 23 Guidelines 231
global contexts 287
distribution of urbanisation 21–3 habitat conservation see conservation
future of urbanisation 302–3 habitat creation 250–2
social organisation of cities 179–80 habitat fragmentation 36, 179
goal setting, planning and design 273–5 biodiversity 242
golf courses 302 frogs 99
gradient analysis 3–4 genetic structure of populations 69, 74
biodiversity 89, 97–9 planning and design 252
limitations 184–5 habitat heterogeneity 107, 108
typical urban species, native/alien 123 habitats, alien species 124
grain size Halifax, Nova Scotia 303
alien species 121 health/wellbeing 204, 221
biodiversity 90 active participation in nature 217–21
green architecture 269–71, 278–9; aesthetic value 209–10
see also sustainable cities benefits of green space 202–3, 203–6, 245–9
 INDEX 313

and biodiversity 110, 139–41, 205, 249–50 immune system 63, 65

cognitive functioning 206 Index of Multiple Deprivation 158–9
and conservation 292 individuality, expression of 189
crime/aggression/violence 209 inequality 180, 182, 186, 187, 188;
effect of cities on 2 see also socioeconomic status
exercise 207–9, 223 accessibility of green spaces 245
functional engagement with nature 215–17 ecological outcomes 181, 184–8, 191
gardening benefits 154, 155 health 249
inequality 249 infant mortality, history 13
infant mortality 13 information processing, and health/
levels of engagement with nature 210 wellbeing 202
outdoor classrooms 210, 223 insects 46, 62, 71, 240; see also ants; butterflies
people–nature interactions 158–9, 160 intensification 236, 241, 242; see also compact
pollution 45 cities; human population density; urban/
psychiatric morbidity 203–4, 247–9 suburban sprawl
social networking 206–7 interdisciplinary research 3, 134, 161
urbanisation 12, 35 International Council for Local
vegetation, urban 185 Environmental Initiatives (ICLEI) 295
wellbeing 204 intraspecific variation see genetics, plasticity,
wildlife watching 211–14, 215 wildlife species, urban/rural
heart disease 45 comparisons
heat island effects 36, 37, 41, 122; intrinsic value 262
see also temperature introductions see alien species
animal communication mechanisms 60 invasion pathways, exotic species 125–6;
body size in urban areas 58 see also alien species; corridors
physiology of urban populations 62 invertebrates, biodiversity 183, 239–8, 242
timing of reproduction 66–7 island biogeography 241
heavy metal tolerance in plants 74
Helianthus spp. (sunflower) 157 Japan 11, 246
herbicides 192 Junco hyemalis (dark-eyed juncos) 61
heterogeneity, landscape/habitat 107, 108;
see also homogenisation Karachi 24
hierarchical status, definitions/ Koelreuteria paniculata (goldenrain tree) 128
conceptualisations 174 Kuala Lumpur 143
history; see also temporal patterns
green spaces 142–3 Lake Tahoe basin 291
health/wellbeing 245 land cover, urban areas 17–20
social organisation of cities 173–4 land-use
urbanisation 12–14 dynamics 276–7
homogenisation, biotic 96–7, 128–9, 291; policies 21–3, 107, 181–4, 231–3, 236;
see also heterogeneity see also planning and design
hospitals, views of nature 212, 248; landscapes
see also wildlife watching fragmentation see habitat fragmentation
householder behaviour see also feeding heterogeneity 107, 108;
urban wildlife see also homogenisation
and biodiversity 101, 109 landscaping 191–2, 302; see also planning
ecological outcomes 191–4 and design
wildlife watching 211 Las Vegas 16
housing policies 230–1 lattice models 12
human population density (HPD); LBAP (London Biodiversity Action Plan) 267
see also compact cities; intensification; Le Mur Vegetal 270
population; urban/suburban sprawl leap-frog development 12
and biodiversity 89–90, 91, 92, 105, 106 Leicester 236, 244–5
and conservation 292 linkage see connectivity, corridors
future of urbanisation 303 literature searches
sustainable cities, designing 300 conservation, current support 263
hydrology see water species richness 239–8
hypercities 24 Liverpool 230–1
hypersegregation 180 Living Places – Cleaner, Safer, Greener (ODPM) 231
local action for biodiversity 264
ICLEI (International Council for Local local ecology 4
Environmental Initiatives) 295 Local Governments for Sustainability 295
314 INDEX

local green spaces 208–9, 222, 237–8, levels of engagement with 210
244–5 wildlife watching 135, 211–14, 215
local resource use 298 nature conservation see conservation
logging 289, 290, 299 nest boxes 250
lognormal distributions 23 nest predation 67–8, 193
Lombard effect 59 Netherlands 141, 204–5
London 20, 144, 178, 236, 244 New Delhi 143
London Biodiversity Action Plan (LBAP) 267 New York 24
longitudinal studies, health/wellbeing 204 New Zealand 11, 96
Lophophanes cristatus (crested tit) 192 Nicotiana glauca (tree tobacco) 123
Los Angeles 143 nitrous oxide 44–5
Luscinia megarhynchos (nightingale) 59 noise, urban 59–62, 67, 141
Lythrum salicaria (purple loosestrife) 126 nutrient cycling 183–4

macroecology 4 obesity 208, 231, 246–7

mammals 73, 93, 242 observational studies 4
management, private gardens 149–52; Ohio 152–3
see also planning and design Open Space Strategies: Best Practice
Manchester 236, 244 Guidelines 231
Manila 24 Örebro, Sweden 144
Marchetti’s constant 178 Osaka–Kobe 24
mass-balance approach 292, 295–7, 299, 304; outdoor classrooms 210, 223
see also fossil fuel consumption oxidative stress 63
megacities 24, 288 ozone 44–5
Melospiza melodia (song sparrow) 60
Mesopotamia, ancient 13, 173 parasites, wildlife 64–6
metabolic model 292, 295–7, 299, 304; park users 205; see also green space
see also fossil fuel consumption Parus major (great tit) 59–60
metacities 24 Parus montanus (willow tit) 192
Metropolitan Open Space System, Passer domesticus (house sparrow) 58, 128, 267
South Africa 264 patch size see also spatial scales
Mexico City 24 alien species 121
microclimates, urban 122 biodiversity 108
migration, bird 193, 303 people–nature interactions 144
mining 290, 298–9; see also fossil fuel planning and design 275
consumption peat extraction 151
mismatch hypothesis 221–2 people–nature interactions 134–7, 161;
mitigation strategies 297 see also green space; nature;
mood states 205, 214, 218–19 wildlife watching
mortality, infant 13 definitions/conceptualisations 137–9
Moscow 24 diversity of 160–1
multi-disciplinary research 3, 134, 161 feeding urban wildlife 155–7
Mumbai 24 garden management 149–52, 152–3
gardening benefits 154, 155
Naja spp. (cobras) 58 gardens, private 146–9
Nakaru, Kenya 152 green spaces, access 143–4
Nanjing 16
natal habitat preference induction 74 health/wellbeing 158–9, 160
nation-level conservation planning 272 interdependence 159–60
native species; see also wildlife species, outcomes 144–6
urban/rural comparisons shaping of urban environments 158
and biodiversity 108 peri-urban areas 276
conservation 266–7 Perth, Australia 152
ecosystem services 303 pesticides 192
typical urban species 123–5 Phalaris arundinacea (reed canary grass) 267
Natural England 237 phenology 38, 66–7
naturalistic design, green space 145, 146 phenotypic plasticity 53, 75; see also wildlife
nature see also green space; people–nature species, urban/rural comparisons
interactions; wildlife watching Phoenicurus ochuros (black redstart) 267
active participation in 217–21 Phoenix 46
conceptualising in urban context 262–3 photosynthesis 63–4
functional engagement with 215–17 physical activity 207–9, 223, 246–7
 INDEX 315

physiology, wildlife species, urban/rural Prunella modularis (dunnock) 55

comparisons 62–4 psychiatric morbidity 203–4, 247–9
pictures of nature 212–14; see also wildlife
watching quality
planning and design 261–2; green space 136–7, 249–50
see also conservation of life 223, 263
and biodiversity 110
business sites 277–8 racial groups 191
conceptualising nature in urban context RCEP (Royal Commission on Environmental
262–3 Pollution) 251
connectivity/linkage 275–6 reaction-diffusion models 12
ecological design 270 recycling 300–1
ecosystem services 174 releases, alien species 125
ecosystems as part of city landscape remnants
271–2 patches 271
gardens, private 280–1 urban ecosystems 261
goal setting 273–5 reproduction, timing of 66–7
green architecture 269–71, 278–9 reproductive success, urban/rural wildlife
green engineering 250, 270 comparisons 67–8
green management 280–1 reptiles 58, 72, 239, 242
green/blue space 231–3, 250–2, 275–7 resource efficiency 294–5, 298
green technology 304 respiratory disease 45
land-use dynamics 276–7 restoration, green spaces 215–16
land-use policies 21–3, 107, 181–4, 231–3, Rio de Janeiro 24
236 road networks 178
landscaping 191–2, 302 alien species corridors 126–7
shaping of urban environments 158 biodiversity 93, 99, 242
size factors 275 Royal Commission on Environmental
spatial scales 261–2, 272–3 Pollution (RCEP) 251
sustainable cities 299–301 runoff 39–40, 41, 122
temporary habitats 277 rural habitats, effect of cities on 2, 20, 29–30,
urban ecosystem sites 277–9 289, 304; see also ecological footprint;
values, nature 262, 271–2, 281–2 ecosystem services; wildlife
plants see green space, tree cover, vegetation
plasticity, phenotypic 53, 75; see also wildlife safe-nesting zone hypothesis 67
species, urban/rural comparisons safety considerations, feeding urban
policy, green space 232, 233; see also planning wildlife 156
and design safety perceptions, green space 145, 159, 209
pollination/seed production 46, 67 sampling bias, biodiversity 107–8
pollution 122 São Paulo 24
atmospheric chemistry 44–5 scale of urbanisation 14–21
disease risk 64, 65 Scotland 154; see also United Kingdom
heavy metal tolerance in plants 74 seasonal patterns, biodiversity 99;
pesticides/herbicides 192 see also phenology
stream rehabilitation 269 seed production 46, 67
urban animal populations 63 segregation of cities 174–7
urban plant populations 45, 63–4 selection pressures, urban/rural
polycentric development 12, 176 comparisons 74
ponds, garden 136 self-esteem 214, 215, 217, 219
population 14–16, 17, 18; see also abundance; Senecio inaequidens (South African ragwort) 126
human population density sensescapes 137
factors in urbanisation 21–3 Shanghai 16, 24, 45
reduction 294 shantytowns 180
power, political 176–7 Sheffield
precipitation patterns 40–1, 122 accessibility of green spaces 143, 244–5
predation pressures biodiversity 101, 158–9, 249–50
birds 67–8, 193 deprivation 158–9
body size in urban areas 58–9 feeding urban wildlife 101–2, 156,
primary productivity 106 157, 192
prisons, wildlife watching 212 garden management 149–50, 151, 152–3
proximity to green space 208–9, 222, 237–8, gardens, private 146
244–5, 249 land-use policies 236
316 INDEX

Sheffield (cont.) Spermophilus beecheyi (Californian ground

quantification of green space 236 squirrel) 60
safety perceptions, green spaces 145 spiders, biodiversity 240
visiting green spaces 139–41 sprawl see urban/suburban sprawl
Singapore 104, 264 squatter settlements 180
sinks, material/energy, cities as 295–6 stage of city development 301
site-level conservation planning 273 standards, green space 222, 237–8, 237–8,
size factors see also patch size; spatial scales 244–5; see also guidelines
alien species 121 stomatal density, urban plant
biodiversity 90, 241 populations 64
planning and design 275 stowaways, alien species 125
urbanisation 23–4 stream rehabilitation 269
slum dwellers 14 stress physiology
Social Determinants of Health, World Health human, and green space 248–9
Organization 244 urban animal populations 62–3
social networking, and green space structure of urban areas 21–8
206–7 study types 3–4
social norms 151–2, 189 suburbanisation 25, 26, 176, 179;
social organisation of cities 172–3, 194 see also urban/suburban sprawl
ecological outcomes 181, 182, 184–8 Sukopp, Berlin 264
feeding urban wildlife 192–3 sulphur dioxide 44–5
future research needs 194 survival, wildlife species, urban/rural
global contexts 179–80 comparisons 68
history 173–4 Sus scrofa (wild boar) 55
householder behaviour 191–4 sustainable cities 30, 139, 202
land-use policies 181–4 and conservation 292
other social dynamics 189–91 design 270, 299–301
spatial organisation/boundaries 174–7 future of 287, 297–9
temporal patterns 177–9 Sweden 205–6
socioeconomic status see also inequality Swietenia macrophylla (mahogany) 299
biodiversity 101, 109, 158–9, 186–7 symbolic representation of self 189
definitions/conceptualisations 174
ecological outcomes 184–8 target-setting, planning and design
gardens, private 148–9 273–5
people–nature interactions 160–1 Tasmania 151–2
spatial organisation of cities 176–7 temperature 36–8; see also heat island
urban ecology 3 effects
soils 21–3, 42–4, 122, 183–4 animal communication mechanisms 60
Solidago canadensis (Canadian golden-rod) 123 body size in urban areas 58
Sonoma County Footprint project physiology of urban populations 62
295, 300 urban 122
source reduction, waste 304 temporal patterns; see also history;
South Africa 107, 267 spatiotemporal patterns
Spain 93 biodiversity 102–5
spatial organisation of cities 174–7 social organisation of cities 177–9
spatial scales, urban ecosystems 261–2, 272–3, study types 4
302–3; see also patch size temporary habitats 277, 278
spatiotemporal fixes 178 testosterone levels, urban/rural
spatiotemporal patterns, biodiversity comparisons 60
88–9 theory of island biogeography 241
broad-scale patterns 88, 89–93, 106 thermodynamics, and body size in urban
comparative studies 89, 93–7 areas 58; see also heat island effects;
drivers 105, 106 temperature
future research needs 110–11 thermophilous species 123
gradient analysis 89, 97–9 timber usage 289, 290, 299
temporal patterns 102–5 timing of biological events (phenology)
urban-centric patterns 89, 99–102 38, 66–7
spatiotemporal patterns, social organisation timing of vocalisations 60–1
of cities 187–8 Tokyo 24, 204
species interactions 46 topography 21–3
species persistence, broad-scale patterns 91 Toronto 20
species richness see biodiversity Town and Country Planning Act (1990) 233
 INDEX 317

Trachycarpus fortunei (windmill palm) 266–7 Urban Task Force Report 231
trade-off mechanisms, body size 58 Urban Wildlife Trust, UK 264
transformed inputs 296 urbanisation 10
transport policies 21–3, 178 composition 25–8, 27–8, 29
travelling, wildlife watching 212 definition/conceptualisation 10–11
tree cover 38; see also green space; vegetation ecological footprint 20–1
carbon cycles 41–2 factors 21–3
and social organisation of cities 172–3 form/structure 25, 26
Turdus merula (blackbird) 60, 64 geographic location 14, 21–3
typology, green space 233–6 history 12–14
land cover 17–20
UNESCO-MAB programme 264 population/household numbers 14–16,
United Kingdom see also Sheffield and other 17, 18
individual cities by name routes to/processes of 11–12
biodiversity 95–6, 136 scale 14–21
cities as sources of invading species 128 size factors 23–4
garden management 149–50, 151 structure of urban areas 21–8
gardening benefits 154 sustainable cities 30
gardens, private 147 urbanoneutral species 123
green space access 143–4 urbanophilic species 123
green space classification/typology urbanophobic species 123
233–6 urban/suburban sprawl 25, 179–80;
green space policy 231–3 see also suburbanisation
green space quantification 236–7 definitions/conceptualisations 176
landscaping 191–2 future of urbanisation 288–9
Royal Commission on Environmental health/wellbeing 222, 247
Pollution 251 impact on green space 230–1
United States USA see United States
biodiversity 90, 93, 100–1, 103, 104
biotic homogenisation 129 values, human 262, 271–2, 281–2
feeding urban wildlife 101–2, 155–6, 192–3 vandalism 223
garden management motivations 152 vegetation, urban 37–8; see also alien
gardens, private 147 species; green space; native species;
green spaces history 142–3 tree cover; wildlife
health/wellbeing and exercise biodiversity 95–7, 108–9, 182, 186–7
behaviour 246–7 carbon cycles 41–2
people–nature interactions 138 conservation of native 266–7
social networking 206–7 genetic structure of populations 70–1
social organisation of cities 187 pollution 45, 63–4
Sonoma County Footprint project 295 social organisation of cities 185–6
urban adapted species 55, 75, 95, 98, 124–5; urban adaptations 55
see also genetics; plasticity; urban species; views of nature 248; see also wildlife watching
wildlife species violence, urban 209
urban-centric patterns in biodiversity vision, planning and design 273–5
89, 99–102 Vitamin G project 205
urban ecology
definition/conceptualisation 1 walkability index 247
emergence of 1–3 water 296
future research needs 6 availability and urbanisation 21–3
perspectives in 3–4 effect of cities on usage 289, 291
wicked problems in 5–6 flow, ecosystem functions 39, 40
urban ecosystems hydrology 122–3
conservation 263–4, 264–6, 266–8, 268–9 quality, ecosystem functions 38–9
planning and design 277–9 sustainable cities 298
remnant/designed 261 wellbeing see health/wellbeing
spatial scales 261–2, 272–3 West Nile virus 66
Urban Forum, UK 264 wicked problems in urban ecology 5–6
Urban Parks Forum 244 wildlife feeding see feeding urban wildlife
urban sensitive species 95 wildlife species, urban/rural comparisons
urban species 123–5; see also alien species; 53–4, 55, 75–6; see also alien; native; urban
native species; urban adapted species; adapted species
wildlife species abundance 55–7
318 INDEX

wildlife species, urban/rural comparisons timing of reproduction 66–7

(cont.) typical urban species 123–5
body size 57–9 wildlife watching 135, 211–14, 215, 248;
communication mechanisms 59–62 see also green space; nature;
demographic traits 66–8 people–nature interactions
disease risk 64–6 win–win scenarios 297, 301–3, 304
future research needs 75–6 wood usage 289, 290, 299
genetic structure of populations 68–75, 73 workplace wildlife watching 211
physiology 62–4 World Health Organization 244
reproductive success 67–8
survival 68 Zipf’s law 23